VOLUME 86, NUMBER 319 DECEMBER 12, 1984 Echinodermata of the Middle Ordovician Lebanon Limestone, Central Tennessee by Thomas Edgar Guensburg MU ——— i — — ~ ыр — Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York, 14850 U.S.A. PALEONTOLOGICAL RESEARCH INSTITUTION Officers PRERISENE a a ccce eee en ee WILLIAM A. OLIVER, JR. VICE-PRESIDENT + с ак Ie ee з=; pres WILLIAM P. S. VENTRESS MOC жула e C e qu LU Mec NE. HENRY W. THEISEN "EREASUBENR A MOL тыные Кон ы ы Co SEM ie Плана ROBERT E. TERWILLEGAR ЕЕЕ ЕОР Уул ЫЫ m IUUD ri САТСЫЗ Тонм L. Сме а а со ана леви aa I а Oe M Vo Nd pM E сее PETER К. HOOVER LEGAL COUNSEL och на ози eg es a ee ЕР HENRY W. THEISEN Trustees BRUCE М. BELL (to 6/30/87) WILLIAM A. OLIVER, JR. (to 6/30/86) RICHARD E. BYRD (to 6/30/86) Тонм РОЈЕТА, JR. (to 6/30/85) Тонм L. CISNE (to 6/30/85) JAMES E. SORAUF (to 6/30/85) J. THOMAS DUTRO, JR. (to 6/30/87) ROBERT E. TERWILLEGAR (to 6/30/87) LEE В. GIBSON (to 6/30/86) Немву МУ. 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Hoover Director Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York 14850 U.S.A. 607-273-6623 | | | | ! | | | | YOLUM -OLUME 86, NUMBER 319 DECEMBER 12, 1984 Echinodermata of the Middle Ordovician Lebanon Limestone, Central Tennessee by Thomas Edgar Guensburg Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York, 14850 U.S.A. Library of Congress Card Number: 84-62635 Printed in the United States of America Allen Press, Inc. Lawrence, KS 66044 U.S.A. | | | | | | | | | | | | | CONTENTS page NUS TEAC VERE eur О кк ле өз. к сак cr 5 ИЛЛА АН ee ee er RM NM 9 Соате НЕ 5 The Lebanon Limestone Geologic and Paleogeographic Setting ................... 6 Sttatipraphic Бе Орн e ee do Ее E ШИЮ re c m mE 8 Fauna and Flora Fauna other than Echinoderms ....................... 11 ЛОТА Жн пъ T TRE ee ROUES 12 Echinoderm Ranna daroa a а а cae 12 DeposınonalEenvironmertt s 77. 5. 7 aas 13 Taphonomy of an Echinoderm Assemblage .............. 14 AS andk@orTolationnenann ya tee NES ete музон кие FERRUM 16 Systematic Paleontology Introductiong on dieto eol. op PL. TRE Emo a. osea 19 Philosophical Considerations ........................... 19 Abbreviations of Repository Institutions ................. 20 Subphylum Crinozoa Class Crinozoa Subclass Camerata Order Diplobathrida Suborder Zygodiplobathra Family Cleiocrinidae Gons Гето Us ata аканин 20 Suborder Eudiplobathrina Superfamily Rhodocrinitacea Family Reteocrinidae GENUS Кетен изискани фа сиам 25 Family Anthracocrinidae Genus Gustabilicrinus, n. gen. ........ 33 Family Rhodocrinitidae СТЕП ААО ИИ ao ros 36 Genus Diabolo rinus ze ar 3 kamılvaunknown o eI ME LI 38 Order Monobathrida Family Glyptocrinidae Genus Abludoglyptocrinus ............ 39 Subclass Inadunata OE DSD AN ыл уук узы а з кише RA 40 Superfamily Homocrinacea Family Homocrinidae Genus Apodasmocrinus .............. 40 Superfamily Calceocrinacea Family Calceocrinidae CONUS СОЛО ОНИН ASA 41 Superfamily Cincinnaticrinacea Family Cincinnaticrinidae Subfamily Cincinnaticrininae (Сели опон FEA + 42 Genus COMBINA mann ne 43 Subfamily Tryssocrininae, n. subfam. ... 46 Genus Tryssocrinus, n. gen. .......... 46 Superfamily Anomalocrinacea Family Anomalocrinidae Genus Anomalochinus «eec ie te 48 Superfamily Myelodactylacea Family Tornatilicrinidae, n. fam. ......... 49 Genus Tornatilicrinus, n. gen. ........ 50 Order Hybocrinida Family Hybocrinidae GENUS DOCUS Ea adoos 51 Order Cladida Suborder Cyathocrinina Superfamily Gasterocomacea Family Carabocrinidae Genus Согаростиикга ER 33 Family Porocrinidae Сели” БОЛОМ nn а ia 53 Superfamily Dendrocrinacea Family Dendrocrinidae Genus Quinquecaudex ............... 55 Family Cupulocrinidae j Genus Gupulockinusi. d ioi a а 55 Class Paracrinoidea Order Comarocystitida Family Amygdalocystitidae Genus Oklahomacystis .............. 56 Class Rhombifera Order Dichoporata Superfamily Glyptocystida Family Glyptocystidae Genus Ланашсууияи а. 59 Family Pleurocystidae Genus Praepleurocystis .............. 60 Сепо есу arica o. seii. 62 Class Asteroidea Order Uncertain Suborder Pustulosina Family Hudsonasteridae CONOS H NUSONI а пе а 63 Order Uncertain Family Schuchertiidae Ч GENUS 5 СССРа ол а а e. 64 Order Uncertain Suborder Uractinina Family Urasterellidae GenussSalterasier ye nen Sasse 65 Class Edrioasteroidea Order болова ај u ee ска 66 Suborder Cyathocystina Family Cyathocystidae (еп са нову ис en M 66 Order Edrioasterida Family Edrioasteridae Genus AITO USIP. ре саз ee и 66 Class Echinoidea Order Bothriocidarida Family Bothriocidaridae срив ЛООНУ a ac 67 Genus Unibothriocidaris ............. 68 Genus Neobothriocidaris ............. 69 Class Unknown Holdfasts and other skeletal elements................ 69 Appendix Local Еее Die aa del 71 СЕТЕВОЕ CUM o eo м... 71 а СЪД она ME Ue LL IRA IUS 71 Праја coun: вина увери E QU уг re en 94 LIST OF ILLUSTRATIONS Text-figure Page 1. Map of study area showing localities where Lebanon echmoderms were: collected... ure scene ro se le 6 2. Three lithostratigraphic sections of the Lebanon Lime- оти bep tenute cec у ey A clie ers ac 9 3. Cleiocrinus laevus Springer, drawing of lower part of ИИ cue etu pn aN АНИ aR EOS qu ps 25 4. ?tegmen of ?Cleiocrinus sp., tracing of portion of speci- TTT ol te qe e. ME tl оты зе Ndi oi NEM E 24 5. Gustabilicrinus plektanikaulos n. gen. and sp., tracing Of cap пи DOPE a ee nee 34 6. Doliocrinus monilicaulis n. sp., plate diagram of cup, showing arm branching pattern `...................... 43 7. Columbicrinus crassus plate diagram of cup ........... 45 8. Tryssocrinus endotomitus n. gen. and sp., plate diagram ОГЕШ хт ее ei mer on gt es 47 9. ?Anomalocrinus antiquus п. sp., camera lucida drawing Ol cup айй parts Ol als: ~ оси с анале pe nye ton есе nae 48 10. Tornatilicrinus longicaudis n. gen. and sp., plate dia- gram Gi pardal pown os. oes ср аа geese es 50 11. Oblique view restorations of typical thecal plates in Okla- homacystis trigonis n. sp., and Oklahomacystis bibrachi- QUAS ата о eg mea у= ооо a rere ee 56 12. Tanaocystis sprinklei n. sp., composite partial plate dia- "ip ru ere cy Ru aca aa низу A ee 59 13. Praepleurocystis ranaformis п. sp., plate diagram of ОББ Ae. озуну еей Жу ашакы зай ра ЧИДЕ 61 14. Praepleurocystis ranaformis п. sp., plate diagram of distal pari of veutaldaue e o ЫЙ ыры ro ee Ee de 61 15. Amecystis nanus n. sp., plate tracing of adoral part of Fal pico чин пале utes eee, VES SI E eas aha ee 63 16. Unibothriocidaris kieri n. sp., tracing of partial peri- stome region Ul рага туре еее 68 LIST OF TABLES Table Page 1. Correlation of selected Champlainian (Black River and ЕТО НЕ А Me EE ТУ КГ? 17 2. Occurrence of Lebanon Limestone echinoderm taxa and related species in Eastern North America ......... 18 3. Measurements of specimens of Reteocrinus from the Leb- O a ОНЫ Ur recu 2 4. Summary comparison of species of Reteocrinus 28,29 5. Measurements of specimens of two species of Gustabili- СИТИ Ue BEM te EL nen 34 6. Measurements of four specimens of Archaeocrinus sny- ANS RA A Ne 7. Measurements of three specimens of ?Diabolocrinus sp. А 38 8. Measurements of three specimens of Abludoglyptocrinus IA AR зле ен 9. Measurements of ten specimens of Columbicrinus cras- ОНО а Би 44 10. Measurements of six specimens of Tryssocrinus endoto- ИШ ДӨЙ, АШКЫ у ete nee 4 11. Measurements of three specimens of Cupulocrinus ..... 56 12. Explanation of Acronyms used in the Taxonomy Sec- ДО (орат oe DR лу с кулу ИН foldout inside back cover ECHINODERMATA OF THE MIDDLE ORDOVICIAN LEBANON LIMESTONE, CENTRAL TENNESSEE By THOMAS EDGAR GUENSBURG Getty Oil Company Three Park Central, Suite 700 1515 Arapahoe Street Denver, Colorado 80202 U.S.A. ABSTRACT Representatives of six classes of echinoderms, the Crinoidea, Paracrinoidea, Rhombifera, Asteroidea, Edrioasteroidea, and Echinoidea are present in the Middle Ordovician Lebanon Limestone of central Tennessee. Crinoids are the most abundant and | diverse class present and include Ше following new taxa: Tornatilicrinidae, new family; Tryssocrininae, new subfamily; Torna- | tilicrinus, Tryssocrinus, and Gustabilicrinus, all new genera; Cleiocrinus springeri, Reteocrinus polki, R. variabilicaulis, R. fenes- tratus, Gustabilicrinus plektanikaulos, G. latomium, Archaeocrinus snyderi, Abludoglyptocrinus gregatus, Tornatilicrinus longi- caudis, ?Anomalocrinus antiquus, Tryssocrinus endotomitus, Doliocrinus monilicaulis, Porocrinus lebanonensis, and Hybocrinus | bilateralis, all new species. Reteocrinus Billings, 1859, is revised and Traskocrinus Kolata, 1975, is suppressed and reassigned to Reteocrinus. Columbicrinus crassus Ulrich, 1925, is redescribed based on more complete new material and Praecursoricrinus Frest, Strimple, and McGinnis, 1979, is suppressed and reassigned to Columbicrinus Ulrich, 1925. Of other echinoderms, Oklahomacystis trigonis (Paracrinoidea); Praepleurocystis ranaformis, Amecystis nanus, and Tanaocystis sprinklei (Rhombifera); Schuchertia darwini (Asteroidea); Unibothriocidaris kieri and Bothriocidaris vulcani (Echinoidea) are new. Previously unknown features include crinoid and paracrinoid columns and holdfasts and a partial peristome of Unibothriocidaris Kier, 1982. Echinoderms indicate a middle to late Blackriveran age for the Lebanon, an interpretation in approximate agreement with ages based on conodonts and brachiopods. Other major faunal groups in the Lebanon are brachiopods, bryozoans, and arthropods. Dominant lithologies include pellet biocalarenites, brachiopod, bryozoan, echinoderm biocalcirudites, calcisiltites, and calcilutites. Paleontologic and sedimentologic evidence indicates the Lebanon was deposited in a moderate-energy normal marine shelf environment ranging from mean low water to 20 to 30 m in depth. INTRODUCTION In recent years, several monographic and smaller works have greatly enhanced our knowledge of Middle Ordovician echinoderms. Most notable among these are Brower and Veinus (1974) on echinoderms from Chazyan and Blackriveran rocks of east Tennessee; Kolata (1975) on both Blackriveran and Trentonian echinoderms from northern Illinois and southern Wis- consin; Brower and Veinus (1978) on the Blackriveran and Trentonian faunas from the Minneapolis-St. Paul area, Minnesota; and Sprinkle (1982e) on Blackriveran faunas from the Arbuckle Mountains area, south-cen- tral Oklahoma. These studies add to prior knowledge of Middle Ordovician echinoderms that was in large Part a result of the efforts of Billings (1854, 1857, 1858, 1859), who studied Trentonian echinoderms from the Ottawa area, Ontario, and Springer (1911), who stud- ied similar Trentonian faunas from the Kirkfield area, Ontario. Over 300 well-preserved and commonly almost complete specimens were collected for this study. These, augmented by a few previously collected specimens, Constitute a rich and diverse Blackriveran echinoderm fauna from the Lebanon Limestone of central Ten- nessee. This fauna adds significantly to available in- formation on Middle Ordovician echinoderms, per- mits a detailed analysis of the functional morphology of many forms, and aids in interpreting the phyloge- netic history of many groups. The new fauna includes at least 32 genera in six classes: the Crinoidea, Para- crinoidea, Rhombifera, Asteroidea, Edrioasteroidea, and Echinoidea. The purposes of this study are threefold: 1) to de- scribe all known Lebanon Formation echinoderms and document their stratigraphic positions; 2) to determine the paleoenvironmental setting of the Lebanon Lime- stone; 3) to realize the regional biostratigraphic poten- tial of echinoderms. ACKNOWLEDGMENTS I am grateful to a number of people for help during this study, which is based on my Ph.D. dissertation at the University of Illinois. I would especially like to thank my advisor, Daniel B. Blake, for his many help- ful suggestions and ideas, and his critical reading of the manuscript. Albert V. Carrozzi, Ralph L. Langen- 6 BULLETIN 319 heim, Philip Sandberg, and Daryl Sweeney, all of the University of Illinois, and Dennis R. Kolata, of the Illinois State Geological Survey, read the manuscript, offered suggestions, and served on my dissertation committee. James Sprinkle, University of Texas, and Ronald Parsley, Tulane University, and Dennis Kolata reviewed a revised manuscript, offered many sugges- tions, and assisted in other ways. All of the above increased the quality of the resulting monograph. Thomas E. Bolton of the Geological Survey of Can- ada (GSC), Ottawa, Ontario, Canada; Porter M. Kier of the United States National Museum (USNM, USNM S), Washington, D.C.; David L. Meyer of the Univer- sity of Cincinnati (UC), Cincinnati, Ohio; Eugene S. Richardson of the Field Museum of Natural History (FM(UC)), Chicago, Illinois; and Harrell Strimple of the University of Iowa (SUI), Iowa City, Iowa, ar- ranged for the loan of specimens by their respective institutions. I would like to thank my parents Ed and Kay Guens- burg, and my wife Susan for both financial and moral support through many trying times in the last few years. THE LEBANON LIMESTONE GEOLOGIC AND PALEOGEOGRAPHIC SETTING The study area is the Central Basin of Tennessee, a topographic feature occupying much of central Ten- nessee. Nashville, the only major population center in the basin, is situated on its northwest edge (Text-fig. 1). The Central Basin is approximately oval with di- mensions of approximately 120 by 80 km; the major axis trends north-northeast. The basin is surrounded by the topographically higher Highland Rim. Struc- turally, the basin is underlain by the deeply dissected Nashville Dome, which is the southern continuation of the Cincinnati Arch and Jessamine Dome of Ohio and Kentucky. The Cumberland Saddle separates the Jessamine and Nashville domes. Strata generally dip at low angles away from the central part of the Nash- ville Dome in all directions, and small scale folding and faulting upon the dome is widespread. Silurian through Carboniferous strata are exposed around the edges of the dome, along the Highland Rim, and on resistant inliers. Ordovician strata form most of the bedrock topog- raphy within the Central Basin itself, and in an up- faulted outlier along the southern edge of the Nashville Dome, where younger strata are eroded. The Lebanon Limestone is less resistant than the over- and under- lying formations and therefore tends to form topo- graphic lows. Soil cover, however, is thin over much of the Lebanon Limestone outcrop belt so that natural exposures are abundant and form what are locally known as glades. Most natural exposures are deeply eroded, however, and many exposed fossils are exten- sively damaged by dissolution. Fresh exposures of the Lebanon Limestone, mainly quarries and roadcuts, are also numerous; 12 of these have yielded the echino- derms studied here (see Appendix). During the Middle Ordovician time, the Central Ba- sin region of Tennessee was a southern part of the large cratonic carbonate shelf that occupied most of present- day eastern North America. The continental margin was approximately 200 to 300 km to the south and east and the shallow shelf extended to the north and west from the Central Basin. To the east of the Central Basin in what is now the Sequatchie Valley of Ten- nessee, and to the north on what is now the Jessamine Dome of Kentucky, the Lebanon Limestone and equivalent-age rocks (Camp Nelson Formation) indi- cate supratidal, intertidal, and subtidal environments (Cressman and Noger, 1976; Bennedict, 1974). The Central Basin was approximately 20 to 25 degrees lat- itude south of the equator during the Middle Ordo- vician (Bambach, Scotese, and Ziegler, 1980, p. 30). 7 Nc z " > A © ®/DAVIDS у р › ( 4 1 > › AE ! ENS (2 1 x | WILLIAMSON AL CANNON pee | I^ А Y E / ne | ри MAURY чара © Columbia (С e ге | | ( bu ? į BEDFORD E 7-656 ) | (а) ) m | M | А Shelbyvilleo «2661 Lewisb eld чой Feo Жн \ 1 І ха MARSHALL ~ „~ fae | \ ERAN rp Saft ena | GILES | Wn ди rn. A а | [= => Е 0 10 20 30 kilometers ? © Pulaski | TENNESSEE ~ == ше 2652 (65) | | А | CENTRAL BASIN STUDY AREA —— y] | | \ 1 INDEX МАР Text-figure 1.- Мар of study area showing localities where Leb- anon echinoderms were collected. Location of study area is show? on inset map of Tennessee. Heavy line = maximum extent of Leb- anon outcrop belt; dashed line on inset = approximate boundary of the Central Basin/Nashville Dome area. LEBANON LIMESTONE ECHINODERMS: GUENSBURG 7 STRATIGRAPHIC SETTING The Lebanon Limestone is part of the lower to mid- dle Champlainian Stones River Group. In the Central Basin of Tennessee, this sequence is divided into five formations including, in ascending order, the Mur- phreesboro Limestone, Pierce Limestone, Ridley Limestone, Lebanon Limestone, and Carters Lime- stone. The Lebanon Limestone is a relatively argilla- ceous, thin- to medium-bedded interval between the more massive Carters Limestone and Ridley Lime- stone (Wilson, 1949). The Lebanon averages approx- imately 28 m thick in the Central Basin (Wilson, 1949). The Ridley - Lebanon contact as exposed is con- formable (Wilson, 1949, p. 37; pers. observ.). Over most of the study area (including locs. Z-652, Z-658, Z-665, and Z-667) the top of the Ridley Limestone is marked by a 1.5 to 2.0 m thick pelletoidal biocalca- renite. Wilson (1949, pp. 43, 54) postulated an unconfor- mity between the Lebanon Limestone and the Carters Limestone, and cited several mostly indirect lines of evidence to support this conclusion. The upper mem- ber of the Lebanon Limestone and lower part of the Overlying lower member of the Carters Limestone both generally thicken away from the axis of the Nashville Dome. This was interpreted by Wilson to have resulted from slight uplift and erosion of the dome immediately following Lebanon deposition and subsequent Carters deposition first about the dome edges, then over the entire area (resulting in a thicker section of Carters Limestone around the margins of the dome). Some beds occurring at or near the top of the Lebanon were Observed to have a “pinkish peripheral zone” indicat- ing oxidation; this was interpreted as evidence for sub- aerial exposure of the Lebanon before Carters depo- Sition. The contact between the Lebanon and Carters is abrupt. A local angular unconformity between the Lebanon and Carters formations was believed to have been observed by Wilson (1949, p. 56) on the far south- ern flank of the Nashville Dome along Richland Creek, 1.6 km south of Milky Way Station, Giles County, Tennessee. I have found Wilson’s arguments unconvincing for the following reasons. As pointed out by Hofstetter (1965b, p. 57), the thinning of the Lebanon could as readily have been caused by differential rates of de- Position. Beds with a pinkish cast are typically calca- renites or calcirudites. They occur at many levels in the Lebanon Limestone and are most common near the middle of this formation, not concentrated at or Near the top, as was indicated by Wilson. There is also no evidence (e.g., vadose crusts, desiccation cracks, or local relief on top of the Lebanon) that the oxidation of these horizons occurred during times of subaerial exposure. An abrupt contact need not indicate an un- conformity. The abruptness of the Lebanon - Carters contact is due to the rapid change from many argil- laceous and (or) dolomitic interbeds in the Lebanon to fewer of these in the Carters; the change is accen- tuated by weathering. I visited the site of the above stated local angular unconformity and found no an- gular relationship between the Lebanon and Carters formations. This exposure is poor, and the Carters Limestone is deeply weathered. G. L. Bennedict and L. Alberstadt visited the locality several times and were also unable to discern an angular relationship between the Lebanon and Carters (Bennedict, oral commun., 1981). I found no indication of an-unconformity at better nearby exposures (e.g., loc. Z-652). Bassler (1932) believed a considerable diastem (last- ing from early Chazyan to Blackriveran) separates the Lebanon and the Carters (called **Carters Member of the Lowville Formation” by Bassler, 1932) limestones. Others, including Wilson (1949) and Cooper (1956), have ignored Bassler’s published interpretation. Bass- ler apparently based his conclusion on faunal differ- ences alone (particularly corals) because he states (p. 58) that the Lebanon and Carters limestones “‘appear to be in conformable relationship.” Neither Cooper (1956, chart 1), using brachiopods, nor Sweet and Berg- ström (1976, р. 133), using conodonts, were able to recognize a significant diastem between the Lebanon and Carters formations. During this study, two guide fossils characteristic of the lower Carters Limestone, Foerstephyllum halli Nicholson, 1879, and Streptelas- ma profundum (Conrad, 1843) were collected in the upper Lebanon Formation. Based on brachiopods, conodonts, and corals, therefore, the Lebanon Lime- stone appears to be only slightly older than the Carters Limestone. No physical evidence of an erosional unconformity between the Lebanon and Carters formations, such as widespread scoured or corroded surfaces, basal lag de- posits, or weathering features such as dissolution or vadose crusts, were observed during this study. In- stead, the contact investigated at several localities in this study (locs. Z-651, Z-653, Z-654, Z-657, and Z-662, among others) proved to be a change in bedding. Con- tact recognition in fresh exposures (such as at loc. Z- 651) can be difficult and somewhat arbitrary (Pl. 16, fig. 4). The change in bedding results from the fewer argillaceous and (or) dolomitic interbeds in the Carters. Because of basic lithic and paleontologic continuity, I believe that no major unconformity or diastem exists between the Lebanon and Carters formations. The pri- mary lithologic difference between the two units is de- 8 BULLETIN 319 creased clastic input during Carters deposition; the causes for this change have not yet been identified. Wilson (1949) subdivided the Lebanon Limestone into three rock-stratigraphic subunits informally termed “lower,” “massive,” and “upper” members. The lower member is thin-bedded and averages approximately 18 m thick. The massive member is near the middle of the Lebanon; it is massive and averages approxi- mately 1.6 m thick. The upper member is again thin- bedded, and averages approximately 10.6 m thick. Where the massive member thins (it is approxi- mately 0.5 m thick at loc. Z-651 [Р1. 16, fig. 1]) and where other massive strata occur locally in the Leba- non section, it can be difficult to recognize (Hofstetter, 1965b; Bennedict, 1974; pers. observ.). Because of these subtleties, Hofstetter (1965b, p. 15) referred to the massive member as the “massive weathering member” rather than the “massive member,” and Bennedict (1974) did not use the three member concept of Wil- son. Bennedict, however, studied only three exposures and one well core in the Central Basin in detail, and his emphasis was on lithofacies analysis and not stra- tigraphy. I found the massive member at all sections where the middle interval of the Lebanon Limestone is exposed; although the member is difficult to discern in places. The middle interval was located by visual observation of upper or lower formation contacts, from elevations and contacts on published geologic quad- rangle maps, and by identification of the Sowerbyella - Diplograptus horizon (see discussion below). The mas- sive member is typically a single bed in the north and central parts of the Central Basin. It is approxi- mately 1.6 m thick at locality Z-651 near Nashville but only 0.5 m thick at locality Z-654 near Columbia, Maury County. In the southernmost part of the study area, south of Pulaski, Giles County, the member is approximately 3 m thick and consists of several thick to thin beds. Lithologically, the middle massive bed- ded member is characteristically a bioturbated pellet biocalcarenite (lithofacies 2 under Lithology); locally it contains oncolites in a calcilutite matrix. The mem- beris whitish brown to whitish pink in fresh exposures, which contrast with the grey adjacent strata. Because the massive member is continuous over the entire Cen- tral Basin and is readily recognizable over most of this area, it is here utilized as а marker unit. The massive member is also identified by the pres- ence immediately above it of a zone containing the graptolite Diplograptus multidens Elles and Wood, 1907, and abundant brachiopods of the species Sow- erbyella lebanonensis Bassler, 1935. This interval is here referred to as the Sowerbyella - Diplograptus Zone of the upper member. It is located from 0.8 to 3.0 m above the massive member. Graptolites have not been found at any other horizon in the Lebanon and Sow- erbyella Jones, 1928, does not occur above this zone. This zone locally contains prominent megaripples near its base and it is more argillaceous than under- and overlying strata. Stratigraphic columns of three thick, well-exposed sections are presented in Text-figure 2. The columns are approximately aligned in a north- south direction across the Central Basin; the farthest- separated sections are approximately 120 km apart. Although the columns differ in detail, the overall se- quences including the positions of the middle massive bedded member and Sowerbyella - Diplograptus Zone are similar. LITHOLOGY General features. — The Lebanon consists principally of thin- to medium-bedded (rarely thick-bedded) light to dark grey limestone with thin interbeds of calcareous shale or dolomite (bedding classification after Ingram, 1954). Beds are typically between 5 and 20 cm thick but attain a maximum thickness of 170 cm; deeply- weathered exposures appear more thinly-bedded than fresh ones. Interbeds range from approximately 1 mm to 2.5 cm thick. Dolomitic interbeds weather to light buffor yellow. Beds are subplanar, irregular, wavy (rip- pled), and nodular; their lateral extent 15 from a meter or less, to (rarely) kilometers. Stylolites are common. The Lebanon becomes more argillaceous toward the south and west sides of the Nashville Dome, and, in addition, clay content decreases near the formational LEGEND Argillaceous limestone Pim — — T —j with prominent soft 2] Indistinct bedding LS [— 1 clay - dolomite interbeds ee x Calcarenite/ N cross bedded calcarenite SS M Prominent megaripples I | Limestone, dolomitic ji limestone; textures © varied; bedding regular 7a I to slightly irregular Oncolites Ripple (wavy) * ae Well preserved bedding echinoderms Nodular bedding; irregular, shaly Text-figure 2.—Three lithostratigraphic sections of the Lebanon Limestone. The localities are near Nashville, in the north (Z-651); near Pulaski, in the south (Z-652); and east of Columbia, approxi- mately midway between the first two (Z-654). See Text-figure 1 and the Appendix for more detailed locality information. LEBANON LIMESTONE ECHINODERMS: GUENSBURG а Ee Ss l = \ 5 5 \ Е 3 5 = aay $3E Е i E Ф © Р = \ NI : \ \ |, ET Р. [[ | \ 210 e (| 28 n IH PTT I mE те à 38 ШИ! ale IRAE : = | IHR. ТШШ ШШ : || | ШП eatin tos | 3 ЈЕВ: El а о > ES > | $8 | % = | ёга 5 2 c ET d % = | дей > б = EIN = E e ЕЗ | 56 | 8 | [<> [| Е © 500 © 5 7 УШ ER 2 || || = © | үү © V E: 7 | | A Ridley Limestone covered 10 BULLETIN 319 contacts and in the massive member. Dolomite ap- pears to increase toward formational contacts. Lithofacies. — Bennedict (1974) divided the Lebanon into five end member lithofacies based primarily on petrographic characteristics; four of these occur in the Central Basin and one occurs only in the Sequatchie Valley in eastern Tennessee. The same lithofacies can be recognized on the basis of field observation, hand samples, and limited thin-section and acetate-peel analysis. Bennedict’s Central Basin lithofacies, there- fore, are here summarized with a few modifications. The carbonate classification used is that of Carozzi (1972) and differs from that of Bennedict, who used a combination of the classifications of Dunham (1966) and Folk (1959). Lithofacies 1 (Pl. 15, fig. 4) is a poorly- to well-sorted fine to coarse pelletal and pelletoidal calcarenite (pel- letal or pelletoidal grainstone of Bennedict, 1974). The matrix is typically sparite but can be mud (calcilutite). The color is medium to dark brownish-grey, occasion- ally brownish-white. External bedding typically is thin, occasionally medium or thick. Internally, beds are thinly- to thickly-laminated or cross-laminated, but occasionally structureless and bioturbated. Pellets and pelletoids range in diameter from approximately 0.1 to 1.0 mm. Many larger pelletoids show faint shapes within them and appear to be micritized skeletal frag- ments. Some pellets could be fecal in origin, as indi- cated by their relatively constant diameter. Fossils are broken, abraded and rounded, and many have distinct micrite coatings. They are concentrated in stringers or scattered more or less homogeneously through the sed- iment. Common fossil groups include ostracods, gas- tropods, trilobites, brachiopods, and bryozoans. An- gular to subrounded intraclasts of pellets and calcilutite are locally common. Oncolites are locally common and are constructed of tubes of the blue-green algae Gir- vanella Nicholson and Etheridge, 1880. They nucleate around bioclasts or intraclasts. Horizontal and vertical burrows are locally common. Lithofacies 2 is an oncolitic biocalcirudite with a sparite or calcarenite matrix (oncolithite of Bennedict, 1974). The calcarenite matrix is composed either of pelletoids or skeletal fragments (mainly of brachiopods and echinoderms). Most oncolites are 5-10 mm in diameter but are rarely greater than 30 mm in diam- eter. They are composed of a skeletal grain or pellet nucleus surrounded by concentric laminae of Girva- nella tubes. This lithofacies is rare and occurs in lenses or pods in beds comprised primarily of lithofacies 1 and 4 (PI. 15, fig. 5). Bennedict reported it from a few horizons in exposures in the southwestern part of the Central Basin in Maury and Giles counties. I have also collected a sample from locality Z-651 near Una, Da- vidson County, in the northwest part of the Central Basin. Lithofacies 3 (Pl. 15, fig. 6) is a bioturbated biocal- cilutite (massive micrite of Bennedict, 1974). Bedding is thin to medium and irregular or nodular. Beds are internally unstratified to finely-laminated and of me- dium-grey to whitish-grey color. Much of the calcilutite appears to be comprised of micritized microscopic fos- sil fragments. Megafossils typically are uncommon but locally can be rare to abundant. Fossils commonly are whole and unabraded and can be exquisitely preserved. Common forms include ostracods, particularly Eole- perditia Swartz, 1949, gastropods, and trilobites. Less commonly, a more diverse fauna is present, often in lenses of bioaccumulated limestone surrounded by less fossiliferous micrite. These lenses contain brachiopods (particularly strophomenids), bryozoans, bivalves, sponges and echinoderms. Two types of bioturbation are recognizable. The first is represented by discrete dolomite-filled burrows. The dolomite weathers more rapidly than the adjacent limestone, yielding irregular, often intersecting tunnels. The second consists of swirl structures caused by orientation of fine fossil debris and is usually accompanied by mottling of the calci- lutite (micrite) matrix. Lithofacies 3 is common throughout the Central Basin of Tennessee. One to 2.5 m thick intervals consisting primarily of this lithofacies occur in the lower Lebanon Limestone at localities Z- 652 and Z-655 and beds of this lithology also occur in the Sowerbyella - Diplograptus Zone in Maury County (е.г., at loc. Z-654). Lithofacies 4 (Pl. 15, figs. 2, 3) ranges from a well- sorted, fine to coarse biocalcarenite with a sparite ma- trix, to a well- to poorly-sorted biocalcirudite with sparite or micrite matrix [bioclastic lithologies of Ben- nedict (1974) including grainstones and packstones]. The color is medium to dark grey, and may have a pink or reddish cast. The former rock type is thin- to medium-bedded and beds are finely- to coarsely-lam- inated, occasionally cross-laminated. The latter li- thology is thin- to medium-, rarely thick-bedded ex- ternally and coarsely laminated to thin-bedded internally. Cross-bedding and graded bedding are com- mon. Beds of both rock types commonly occur as lens-shaped, small, temporary channels or more widely- extended subtabular beds, many characterized by rip- ple marks. Larger ripple marks to small scale dunes (Pl. 16, fig. 2) are associated with the calcirudites and have maximum wavelengths ranging from 0.8 to 1.0 m, and a ripple index (wavelength + amplitude) of 10 to 20. Smaller ripples are typically associated with the calcarenites and usually have wavelengths of 15 to 25 LEBANON LIMESTONE ECHINODERMS: GUENSBURG 11 cm. Fossils are most commonly extensively abraded and rounded, and when identifiable typically consist of echinoderms and brachiopods. Calcirudites typi- cally contain whole unabraded to slightly abraded fos- sils, mainly brachiopods, that may form coquinites. Oncolites, pellets and pelletoids are commonly abun- dant. Intraclasts of lithofacies 1, 3, or 4 are locally com- mon in the calcirudites and uncommon in calcarenites. The intraclasts typically are highly angular, and range in size from coarse sand to (rare) pebbles over 8 cm in diameter. Dolomite-filled trace fossils are also lo- cally common, especially in the calcarenites. Lithofa- cies 4 is common throughout the Central Basin but is most common on the west side of the major axis of the Nashville Dome. Mixing and rapid changes of lithofacies over short lateral and vertical distances are common (Pl. 15, fig. 5) and many samples are varied, indicating a complex interweaving of the various subenvironments implied by the lithofacies. As also noted under “Stratigraphic Setting”, the massive member consists primarily of lithofacies 1 (pellet and pelletoidal calcarenites). Both upper and lower members are comprised predomi- nantly of lithofacies 1, 3, and 4. FAUNA AND FLORA Fauna Other than Echinoderms The Lebanon contains an abundant and diverse invertebrate shelly fauna dominated by bryozoans, brachiopods, and echinoderms. Taxonomic studies of Lebanon Limestone fossils are few, however. The most comprehensive works on these fossils to date are by Cooper (1956), who included Lebanon brachiopods as Part of his larger work on Chazyan brachiopods and Coryell (1921) who studied Lebanon bryozoans as a Portion of his Stones River Group bryozoan work. Hofstetter (1965a) reported the graptolite Diplograptus multidens Elles and Wood, 1907; Bassler (1935) briefly described the brachiopod Sowerbyella lebanonensis Bassler, 1935, and two ichnofossil species; and Troost (in Wood, 1909), Springer (1911), Schuchert (1912), and Ulrich (1925), each described or reported an echi- noderm species (for details see “Echinoderm Fauna” and “Systematic Paleontology"). Safford (1869), Cor- yell (1921), Bassler (1932), and Wilson (1949) all pro- Vided faunal lists for the Lebanon Limestone, but taxo- попис revisions of many taxa, especially bryozoans, Corals, sponges and arthropods, are needed for such lists to be meaningful. A large macrofossil collection Made during this study contains over 120 species, many of which are previously unrecorded and undescribed. At least 10 genera of trilobites were collected, of which only three were included in previous faunal lists. Be- cause of these problems, no compendium of species from the Lebanon Limestone is provided here. Ma- crofossil groups are discussed below, however, and rel- ative abundances (1.е., abundant, common, uncom- mon, rare) of taxa within groups are based upon field observations. Abundances given are rough approxi- mations only. Abundant indicates that many speci- mens of the taxon in question can be collected from most exposures in the Central Basin of Tennessee. Common indicates that several specimens can be col- lected from each exposure in the study area. Uncom- mon indicates fossils that are widespread over the study area, but occur in small numbers and require intensive search to find. Rare fossils are those of which only a few specimens were found; these usually occur only at one or two localities. Field impressions of abundances gathered here are biased in favor of specimens that tend to be most visible on bedding planes or that easily weather free, such as brachiopods, bryozoans, trilo- bites, and echinoderms. Whole fossils or well-pre- served fossils were usually collected in favor of fossils that were broken or poorly preserved. A deliberate effort was made to collect rare or unusual specimens and, therefore, actual numbers of specimens in collec- tions are also biased. Sponges.— Poriferans are common in the Lebanon Limestone; there are a minimum of four genera, in- cluding 1 stromatoporoid. Calcisponges are uncom- mon in the lower member of the Lebanon Limestone at localities Z-651, Z-652, Z-654, and other non-listed localities. One calcisponge is tentatively identified as Zittelella Miller, 1889 (Bassler, 1932). Bennedict (1974) found abundant pyritized hexacton sponge spicules in the Sowerbyella - Diplograptus Zone of the upper member, Lebanon Limestone. Corals. — Corals are usually rare to locally common (atthe top ofthe Lebanon Limestone at locality Z-651). At least two genera of tabulates including Foerste- phyllum Bassler, 1941, and Tetradium Dana, 1846, and two kinds of solitary rugosans including Strepte- lasma Hall, 1847, are present. Fragments of conularids are rare in shaly interbeds. Bryozoans.— Bryozoans are abundant and diverse; at least 20 species occur in the Lebanon Limestone. Ptylodictyoids (bifoliate cryptostomes) are probably numerically most abundant. The thin ribbon-like (ba- sally articulated and non-articulated) genera such as Stictopora Hall, 1847, and Escharopora Hall, 1847, are most abundant. Large species with ribbon-like, fo- liose, and reticulate growth modes also occur. Other common to abundant cryptostomes include arthro- 12 BULLETIN 319 stylids, of which articulated genera such as Arthrocle- ma Billings, 1865, and Ulrichostylus Bassler, 1952, are most common. Fenestrate bryozoans are represented by common phylloporinids, of which at least two species are represented. Trepostome bryozoans are common and diverse. Most forms have a massive or low en- crusting growth habit; a few species with erect dendroid growth habit are also present. Common trepostome genera (Bassler, 1932) include Homotrypa Ulrich, 1882, Nicholsonella Ulrich, 1889, Ceramoporella Hall, 1851, and Monticulipora d'Orbigny, 1849a. Both cystoporate and cyclostome bryozoans were rarely encountered. Brachiopods.—Inarticulate brachiopods are rare to uncommon. At least three genera are represented, in- cluding Acanthocrania Williams, 1943 (two species), Conotreta Walcott, 1889, and Lingulops Hall, 1872. Articulate brachiopods are abundant; Cooper (1956) lists 22 species and 18 genera. Strophemenids are the most abundant order and include Strophemena de Blainville, 1825, Oepikina Salmon, 1942, and Sow- erbyella Jones, 1928. Orthids are also abundant, par- ticularly Doleroides Cooper, 1930; other common or- thids include Pionodema Foerste, 1912, Hesperorthis Schuchert and Cooper, 1931, Skenidioides Schuchert and Cooper, 1931, and Multicostella Schuchert and Cooper, 1931. Abundant rhynchonellids and spirifer- ids include Rostricellula Ulrich and Cooper, 1942, and Zygospira Hall, 1862, respectively. Mollusks.—Mollusks are common to locally abun- dant. Most occur as poorly preserved molds or spar- filled casts in fine-grained matrices, making identifi- cation difficult. Gastropods are locally abundant and diverse. Common genera (from Bassler, 1932) include Maclurites Le Sueur, 1818, Subulites Emmons, 1842, Phragmolites Conrad, 1838, Lophospira Whitfield, 1886, and others. Bivalves are rare; one unidentified species, at least, is represented. Cephalopods are un- common; forms include a large endocerid and small nautiloids. Trilobites.—Trilobites are common and diverse al- though complete or near complete specimens are very rare. At least 10 genera were identified during this study. The most common genera appear to be ?Се- raurus Green, 1832, Isotelus DeKay, 1824, Encrinurus Emmrich, 1844, Шаепиз Dalman, 1827, Thaelops Conrad, 1843, and Dolichoharpes Whittington, 1949. Other faunal elements.—The graptolite, Diplograp- tus multidens Elles and Wood, 1907 (see Hofstetter, 1965a), is abundant in and confined to the Sowerby- ella - Diplograptus Zone, upper member, Lebanon Limestone. One specimen of a possible dendroid grap- tolite was found in the upper member at locality Z-653. Cornulitids are rare. Scolecodonts (annelid jaw parts) are common and locally abundant. Trace fossils. — Burrows аге common to abundant in the Lebanon Limestone. Most within limestone beds tend to retain their shape whereas those in shaly in- terbeds are typically flattened by compaction. Most appear to be horizontal or subhorizontal irregular feed- ing structures. Feeding burrows (Phycodes Richter, 1850) are locally common as are vertical dwelling bur- rows. Other trace fossils include Cruziana d’Orbigny, 1842, and several unidentified trails or burrows. Ben- nedict (1974) has noted possible fungal borings in bra- chiopod fragments from biocalcirudite lithofacies (lithofacies 4). Flora Calcareous algae are common in the Lebanon Lime- stone. Several types were observed in thin sections or reported from thin sections by Bennedict (1974), in- cluding the blue-green alga Girvanella Nicholson and Etheridge, 1880, which is locally abundant, and the red alga Solenopora Dybowski, 1877, and two or more green algae including Vermiporella Stolley, 1893, which are uncommon but widespread. Fragments of Nidulites Salter, 1851, are uncommon in hand samples, and a single specimen each of a calathid alga and an unde- termined large (?)alga were found. Echinoderm Fauna Disarticulated echinodermal debris is one of the dominant constituents in the Lebanon Limestone; articulated specimens are uncommon overall and only locally common, however. Approximately 300 partial to complete echinoderms were collected during this study. Most articulated specimens were collected from the lower member between 4 and 12 m above the contact between the Lebanon and Ridley limestones. Other specimens are from the upper member, partic- ularly the Sowerbyella - Diplograptus Zone. No spec- imens were found in the massive member. Specimens occur both as isolated individuals and in clusters. Ap- proximately 80 percent of the echinoderms studied are from localities Z-651, Z-652, Z-654, Z-654a, and Z- 655; approximately one-third of all specimens were found ina shaly interbed at locality Z-654a (see below). Disarticulation of fossils precluded determination of stratigraphic ranges within the Lebanon Limestone for most species. Archaeocrinus snyderi n. sp., Cleiocrinus tessellatus (Troost, in Wood, 1909), Quinquecaudex sp. A, and Porocrinus lebanonensis n. sp. range throughout most of the Lebanon Limestone. Prae- pleurocystis ranaformis n. sp. was not found above the Sowerbyella - Diplograptus Zone, upper member, and Praepleurocystis sp. cf. P. watkinsi (Strimple, 1948) was not observed at or below this horizon. LEBANON LIMESTONE ECHINODERMS: GUENSBURG 13 Four species of echinoderm previously described from the Lebanon Limestone of central Tennessee in- clude three crinoids and a single sea star. Previously described crinoids include Cleiocrinus tessellatus (Troost, in Wood, 1909), Cleiocrinus laevus Springer, 1911, and Columbicrinus crassus Ulrich, 1925. Schu- chert (1915) referred a Lebanon Limestone asteroid to Hudsonaster narrawayi (Hudson, 1912). At least 40 Species and 32 genera of Lebanon Limestone echino- derms were identified during this study. The complete list of known Lebanon Limestone echinoderm species 18 as follows: Class Crinoidea Cleiocrinus tessellatus (Troost, їп Wood, 1909) Cleiocrinus laevus Springer, 1911 Cleiocrinus springeri п. sp. Reteocrinus variabilicaulis n. sp. Reteocrinus polki п. sp. Reteocrinus fenestratus п. sp. Reteocrinus sp. cf. R. variabilicaulis Reteocrinus sp. Gustabilicrinus plektanikaulos n. gen. et sp. Gustabilicrinus latomium n. sp. Archaeocrinus snyderi n. sp. Abludoglyptocrinus gregatus n. sp. ?Diabolocrinus sp. A Rhodocrinitid sp. A Rhodocrinitacid sp. A Apodasmocrinus sp. cf. A. daubei Warn and Strimple, 1977 Cremacrinus sp. cf. C. punctatus Ulrich, 1886 Columbicrinus crassus Ulrich, 1925 Doliocrinus monilicaulis n. sp. Tryssocrinus endotomitus n. gen. et sp. Tornatilicrinus longicaudis n. gen. et sp. Anomalocrinus antiquus n. sp. Hybocrinus bilateralis n. sp. Carabocrinus sp. Porocrinus lebanonensis n. sp. Cupulocrinus sp. cf. C. gracilis (Hall, 1847) Cupulocrinus sp. Quinquecaudex sp. A Class Paracrinoidea Oklahomacystis trigonis n. sp. Oklahomacystis sp. aff. О. trigonis n. sp. Class Rhombifera Tanaocystis sprinklei n. sp. Praepleurocystis ranaformis n. sp. Praepleurocystis sp. cf. P. watkinsi (Strimple, 1948) Amecystis nanus n. sp. Class Asteroidea Hudsonaster sp. cf. H. narrawayi (Hudson, 1912) Salteraster sp. cf. S. grandis (Meek, 1872) juvenile Salteraster sp. Schuchertia darwini n. sp. Class Edrioasteroidea ?Edrioaster sp. A ?Cyathocystis sp. indeterminate isorophid Class Echinoidea Unibothriocidaris kieri n. sp. ?Unibothriocidaris sp. Bothriocidaris vulcani n. sp. ?Neobothriocidaris sp. The Lebanon Limestone echinoderm fauna is dom- inated by camerate and disparid inadunate crinoids. Dominant camerate genera include Archaeocrinus Wachsmuth and Springer, 1881, Reteocrinus Billings, 1859, Gustabilicrinus n. gen. (Anthracocrinidae), and Abludoglyptocrinus Kolata, 1982; thick-plated rho- docrinitids are conspicuously scarce (only ?Diabolo- crinus sp. А 15 known). The most often encountered and widespread disparid inadunate is Columbicrinus Ulrich, 1925. Pleurocystid rhombiferans and amyg- dalocystid paracrinoids are also locally found. DEPOSITIONAL ENVIRONMENT The lithology, primary sedimentary structures, trace fossils, flora, and fauna of the Lebanon Formation in- dicate a well-lit, normal marine, subtidal environment of moderate energy. Depth ranged from mean low water to 20 or 30 m. Sediment was primarily locally-derived, biogenically-precipitated calcareous debris with minor amounts of clay derived from outside the depositional area. The origin of the lime mud is largely unknown but some is micritized fossil skeletal fragments. Both physical and biological processes were important in- fluences on fabric and texture in the Lebanon Lime- stone. Abundant broken and well-rounded current laminated or bedded bioclastic debris and local on- colites indicate considerable reworking of sediment be- fore final burial. Transport mechanisms included waves (perhaps storm-generated), as indicated by abundant symmetrical ripples and megaripples, and poorly chan- nelized bottom currents, as indicated by lenses of coarse to fine, poorly- to well-sorted, often cross-laminated or cross-bedded bioclastic debris (Bennedict, 1974; pers. observ.), and current ripples. Intervals dominated by calcilutites and few associated bioaccumulated lime- stones were likely areas of quieter water deposition, perhaps in subtle topographic lows or where significant baffling from plants or animals occurred, because low current energies would have allowed lime mud to settle. Bioturbation was important, particularly in pel- let and pelletoid calcarenite and calcilutite microfacies, and in the shaly interbeds. Bioturbation produced some of the common mixed lithologies and pellets in the Lebanon Formation, and also contributed to the break- age of fossils. Firm, perhaps cemented substrates were common in the Lebanon Limestone; these substrates are primarily developed on calcarenites and calcirudites and the as- sociated substrate topography varies from irregular to 14 BULLETIN 319 subplanar. Evidence indicating firm or cemented sub- strates includes holdfasts or entire colonies of ptylo- dictyoid or trepostome bryozoans, encrusting hold- fasts, attachment discs of echinoderms (crinoids and edrioasteroids), and vertical borings (Trypanites Mag- defrau, 1932). In addition, intraclasts of various lith- ologies including calcarenites are common, indicating partial cementation before reworking. Insofar as can be compared, the fauna of the firm substrates is similar to the faunas of “hardgrounds” reported by Palmer and Palmer (1977) from the Trentonian Galena Group of Iowa, and by Brett and Liddell (1978) from the Trentonian Bobcaygeon Limestone near Kirkfield, On- tario. Depositional conditions apparently varied only slightly with time. The massive member is lithologi- cally similar to the upper Ridley Limestone and pos- sibly the lower Carters Limestone (immediately below and above the Lebanon, respectively, and represents a brief time of decreased clastic input and dominant pellet and pelletoid deposition. The Sowerbyella - Dip- lograptus Zone was deposited during times of slightly increased clastic input. The estimated paleobathymetric range of mean low water to roughly 20 to 30 m is derived from the com- bined overlap of known physical and biological indi- cators present in the Lebanon Limestone (see Benne- dict and Walker, 1978, p. 583 for a summary table of Paleozoic bathymetric indicators). The most diagnos- tic such indicators in the Lebanon Limestone are mega- ripples, oncolites, and an abundant flora containing blue-green, red, and green algae. TAPHONOMY OF AN ECHINODERM ASSEMBLAGE Observations.— A remarkably large and diverse fau- nule of relatively complete invertebrates, including many echinoderms, was recovered from a dolomitic shale interbed in the lower member of the Lebanon Limestone along Interstate Route 65 near Columbia, Tennessee (loc. Z-654a). The taphonomy of this ho- rizon is here evaluated in an attempt to better under- stand this unique occurrence, and the living and death conditions of the echinoderms. The well-preserved echinoderms were collected along a 7 m horizontal interval exposed in a highway cut. The distribution of the highly fossiliferous part of the interbed probably was restricted. Although the in- terbed continues along the west side of the highway for at least 150 m, no complete specimens were ob- served beyond the 7 m interval. Further, only four less complete specimens from the same horizon were col- lected on the east (opposite) side of the highway. А1- though some specimens had weathered free of the ma- trix, most material was found on small quarried slabs. Because accessible bedding surfaces were discontin- uous and relatively few, only approximately 1.5 m? of slab surface could be collected and it was impossible to reconstruct a large surface area. The fossil-bearing interbed averages 1 to 3 cm thick. The well-preserved fossils occur in the thickest part of the interbed, in which the lithology is non-laminated, burrow mottled, dark grey, dolomitic shale. Dolomite content increases toward both upper and lower con- tacts of the interbed and is the dominant constituent immediately adjacent to both contacts. The dolomite is authigenic, consisting of discrete well-formed rhombs displacing and surrounded by clay. The interbed clay mineralogy was not determined. The interbed also con- tains abundant small fragments of both organic debris and calcareous skeletal elements (Pl. 15, fig. 1). Small amounts of finely-disseminated pyrite and occasional secondary gypsum occur along weathered surfaces. Directly below the interbed is a poorly-sorted intra- clastic oncolite biocalcirudite with a sparite and mi- crite matrix (Pl. 15, fig. 1). The interface between the shaly interbed and the calcirudite below is sharp but poorly exposed. Poor exposure is a result of the large amount of weathering-resistant dolomite in the lower part of the shaly interbed that adheres to the interface below. The interface is slightly irregular and covered with bioclastic debris. No sign of intensive scouring 15 present. A few of the fossils within the shaly interbed are impressed into the calcirudite, indicating this layer was not completely lithified at the time of deposition. The overlying bed is a poorly-sorted biocalcirudite with a calcilutite matrix and has a sharp interface with the interbed below. The distribution and orientation of fossils within the interbed is important to understanding their taphon- omy. Whole fossils and fossil debris in the interbed are concentrated just above the lower interface; fossils rarely contact the interface itself. Thus material in the interbed is crudely graded. No fossils in the interbed were found attached to the underlying calcirudite. Large and small fossils of various taxa are mixed indiscrim- inately, with two exceptions (discussed below). The distribution of fossils is uneven, with some slabs hav- ing several specimens, whereas roughly equal areas of other slabs have few. Fossils apparently are not cur- rent-oriented. Long crinoid columns, with crowns at- tached, often cross one another at nearly right angles. A few fossils are preserved in orientations that clearly do not represent living positions. For example, trepo- stome bryozoan fragments lie on crinoid crowns, and one large sea star, Salteraster sp. cf. S. grandis (Meek, 1872) is inverted over a crinoid crown. Only two fossils LEBANON LIMESTONE ECHINODERMS: GUENSBURG 15 in the interbed were obviously in life position: small specimens of Doleroides Cooper, 1930, commonly are preserved in small clusters of approximately 10 to 15 specimens and distal columns of Gustabilicrinus n. gen. are often found wrapped around the columns of other crinoids. The condition of the fossils, particularly that of the echinoderms, varies greatly. A few crinoids are essen- tially complete skeletons with articulated crown, col- umn, and holdfast. Most specimens, however, are par- tially dismembered and (or) slightly disarticulated. Dismemberment occurs in different ways and varies in extent. For example, many crinoids and paracrin- oids are complete except for arms or arm tips whereas others are reduced to parts of cups with attached arms, column sections, or isolated arms with attached partial to complete pinnules, or armlets. Dismembered parts, particularly thin-plated cup fragments, are common. These can be broken across, rather than along, plate boundaries. Specimen completeness generally is relat- ed to organism type. The relatively sturdy disparid inadunates, including Tryssocrinus n. gen. and Colum- bicrinus Ulrich, 1925, and the stout-plated camerate Reteocrinus Billings, 1859, are typically little dismem- bered or disarticulated. Most specimens of Hybocrinus Billings, 1857, and Oklahomacystis Parsley and Mintz, 1975, possess complete calices (thecae), but their col- umns and arms are partial or missing. Thin-plated crinoids, however, such as the camerate Gustabilicri- nus n. gen. and the inadunate Carabocrinus Billings, 1857 typically were reduced to partial cups, crowns, or isolated column segments. Fragments of dismem- bered and disarticulated specimens are isolated, with- out missing parts lying nearby, suggesting disruption occurred before final burial. In addition to being dis- membered and slightly disarticulated, many echino- derms are in distorted positions. No fossils from the interbed show signs of pre-burial mechanical abrasion. Sand- or silt-sized calcareous fossil debris is com- mon in the interbed. Identifiable fragments primarily are smaller pieces of the more complete fossils: Dol- eroides Cooper, 1930, and echinoderm plates are par- ticularly common. Subhorizontal feeding burrows are common in the interbed. They are light grey rather than the dark grey of the surrounding rock and vary from approximately 5 to 10 mm in maximum diameter. Burrows rarely disrupt fossils even though they often occur among them. A few burrows are filled with bioclastic debris. Many fossils and all burrows were flattened by com- paction. Burrows average five times wider than high; assuming that these were originally circular, this shape Suggests considerable compaction occurred after the time of burrowing (presumably shortly after deposi- tion). Crinoid columns and thick-plated calices such as those in the disparid inadunates are typically un- crushed, and occasionally have small-scale slickensides caused by compaction of adjacent clay. Some specimens, though well preserved overall, show corrosion on surfaces that contact the subjacent lime- stone bed. Corrosion is most prevalent in thin-plated ог pore-bearing genera, including Cleiocrinus Billings, 1857, Carabocrinus, and Oklahomacystis. Associated finely-disseminated pyrite suggests the corrosion took place in reducing microenvironments. Interpretations.—The interbed fossils of sessile or- ganisms were torn from their attachment sites. They are often dismembered but only slightly disarticulated and none of the fossils are attached to the substrate below; holdfasts are always broken or torn loose. Thus, the fossils must have been transported before burial. The Lebanon Limestone depositional environment apparently was well oxygenated and presumably con- tained many scavenging organisms. The exceptional completeness of many echinoderms and other interbed fossils reflects rapid deposition, shortly after detach- ment from life sites. Taphonomic studies of modern echinoderms particularly well illustrate this. For ex- ample, Liddell (1975) and Meyer (1971), among oth- ers, have studied postmortem decay and disarticula- tion of modern crinoids and found this to be rapid, requiring from one to six days for complete disarti- culation under well-oxygenated conditions. Delicate skeletal structures such as cirri or pinnules can be lost only a few hours after death whereas more rigidly ar- ticulated elements such as cup plates require much longer. The apparent lack of mechanical size sorting and current orientation of fossils, as well as concurrent deposition of nonlaminated clay and other fine parti- cles indicate quiet water deposition. Coarser particles, including the fossils, are concentrated near the bottom of the interbed, indicating specific gravities and settling velocities at the time of deposition averaging slightly greater than clays in the interbed. After burial, the fossils remained largely undisturbed by burrowing. The most reasonable causal agent for the interbed is a storm. A storm could provide sufficient energy to detach, dismember the creatures and transport them with accompanying fine sediment. A rapid energy de- crease such as occurs after a storm passes would ac- count for the absence of both current orientation and size sorting, as well as the occurrence of the fossils in a single thick clay blanket and the crude graded bedding of the interbed. The rapid deposition under such cir- cumstances would help explain the exceptional pres- ervation. The faunule is thus considered to be a trans- 16 BULLETIN 319 ported death assemblage (sensu Craig and Hallam, 1963). The postulated tropical Lebanon Limestone pa- leoenvironment was shallow enough to be affected by major storms (see “Depositional Environment”), and the paleolatitude of 20 to 2598 compares well with latitudes of worldwide modern tropical storm tracks (Jordan, 1966). Few observations concerning the living relationships of the interbed assemblage can be made because of its disrupted nature. Biases acting upon the assemblage may have included selective removal, transport, and mixing of various life assemblages. Evidence from fos- sils in the interbed, as listed below, suggests a single life assemblage is represented. 1. There is no apparent size sorting of fossils. 2. The most abundant and diverse megafossils in the Lebanon Formation are the brachiopods and bryo- zoans. In contrast, brachiopods and bryozoans in the interbed have low diversities but high abundances. (See bryozoans and brachiopods under “Fauna other than Echinoderms,” herein, for partial listing of many species from the Lebanon Limestone.) Approximately seven species of bryozoans, including two arthrostylids, two bifoliates and three trepostomes, and a single brachio- pod species (Doleroides sp.) were found in the interbed. Mixing of the Lebanon Limestone assemblages would presumably have produced a much greater diversity of these groups. 3. The life modes of the various interbed fossils is compatible insofar as can be determined. Nearly all interbed echinoderms appear to have lived fixed to a firm substrate. An exception, the sea star Salteraster sp. cf. S. grandis (Meek, 1872), was probably vagrant. At least four crinoid genera possessed small discoid en- crusting holdfasts. Doleroides was pedically attached and bryozoans were either attached by smaller ce- mented holdfasts (e.g., arthrostylids, bifoliates) or sim- ply were cemented to or rested on the substrate (e.g., trepostomes). Other bottom-dwelling groups present in the interbed include at least one unidentified trilo- bite, annelids (represented by scolecodonts), and con- ularids (fragments), all of which probably could have occupied a firm substrate. Organisms thought to have inhabited soft bottoms, such as crinoids with rootlike holdfasts and strophomenid brachiopods such as Oep- ikina Salmon, 1942 and Strophomena de Blainville, 1825, are absent from the interbed. Strophomenid brachiopods are abundant in the Lebanon Limestone as a whole. The occurrences of distal columns of Gustabilicrinus n. gen. coiled about columns of other crinoids, and small specimens of Doleroides Cooper, 1930, in clus- ters, are preserved remnants of probable living occur- rences of these organisms. The Doleroides cluster at- tachment sites (?algal) are no longer preserved and apparently decayed after burial. AGE AND CORRELATION Information from both brachiopods and conodonts suggests that the Lebanon Limestone belongs in the upper part of the Middle Ordovician Blackriveran Stage. Based on their conodont faunas, Sweet and Berg- stróm (1976, р. 133) suggested the following as ap- proximate lateral equivalents to the Lebanon Lime- stone (see Table 1): the upper Pooleville Member of the Bromide Formation, Oklahoma, the Camp Nelson Formation, central Kentucky, the Glenwood Forma- tion of Тома and Minnesota, and the Watertown For- mation of the type Blackriveran section in New York. Based on brachiopods, Cooper (1956, p. 102 and chart 1) assigned the Lebanon Limestone to his Wilderness Stage [early Trenton or latest Blackriveran] and listed the Witten Formation (uppermost Ottosee Group) of east Tennessee, possibly the Rockland (Selby, in part) Formation of New York, and the Grand Detour For- mation of Illinois as approximate lateral equivalents (see Table 1) A single graptolite, Diplograptus multidens Elles and Wood, 1907, has been identified in the Lebanon Lime- stone. This species ranges from the Blackriveran through the middle Trentonian- (Kirkfieldian) age rocks (Sweet and Bergstróm, 1976, p. 134) in North America and also occurs in the upper Dalbyan and lower Mol- dan stages in Sweden. These two Swedish stages have been correlated with the Idaverian and Jóhvian stages in Estonia (Jaanussen, 1976, p. 320). The Lebanon Limestone echinoderm fauna is sim- ilar to several previously described Blackriveran and classical Trentonian faunas. Although most similari- ties are at the generic level, many species are clearly closely related as well. The Blackriveran Bromide Formation of Oklahoma (see Sprinkle, 1982a) contains a large number of genera and probably one species (Apodasmocrinus daubi Warn and Strimple, 1977) in common with the Lebanon Limestone (see Table 2). In addition, two Lebanon Limestone species, Archaeocrinus snyderi n. sp. and Columbicrinus crassus Ulrich, 1925, appear to be closely related to Archaeocrinus subovalis Strimple, 1953, and Columbicrinus sulphurensis (Frest, Strim- * The Trenton Stage, retained here, has in recent years been sub- divided by some workers into three stages; Rockland, Kirkfield, and Sherman, from oldest to youngest. In addition, the latest Trenton, or Cobourg Substage, is considered by these workers as the basal Edenian Stage. These new terms are here included in parentheses for further clarity when possible. | | | LEBANON LIMESTONE ECHINODERMS: GUENSBURG 17 Table 1.—Correlation of selected Champlainian (Black River and Trenton) units (modified from Sweet and Bergstróm, 1976, and Cooper, 1956). The Hull Member of the Ottawa Formation is equivalent in age to the Kirkfield and Bobcaygeon limestones (not shown) of the Kirkfield, Ontario area. REGIONS-UNITS Ф| ш ШЕЕ «|< Ш РІ SOUTH CENTRAL EAST TENNESSEE- | SOUTHEAST ONTARIO- on | б) | CENTRAL TENNESSEE АЦЕ — CENTRAL KENTUCKY VALLEY OKLAHOMA SOUTHWEST VIRGINIA | NORTHERN NEW YORK E COBOURG u z 45 МЕМВЕВ 5 2 EE a Е | SHERMAN FALL |5 2 ales 3 ES MEMBER 20 5 >> © кє = ш о | 2 oo X =| CURDSVILLE Е CURDSVILLE © 5. | HULL MEMBER | 2 MEMBER ш МЕМВЕВ < с = Е ш < DECORAH E SUBGROUP d SELBY FORMATION = ш = CARTERS d 3 LIMESTONE > 5 GRAND DETOUR WITTEN WATERTOWN Е : Е g FORMATION FORMATION FORMATION = 5 < MIFFLIN WARDELL Gier 2 LEBANON а. FORMATION POOLEVILLE FORMATION CAMP NELSON E е LIMESTONE 5 MEMBER = ЕОВМАТ!ОМ = ДЕ RIDLEY E = BENBOLT S = LIMESTONE GLENWOOD FORMATION z MOUNTAIN LAKE | © FORMATION ә < ш о 3| 2 PIERCE = е ш е LIMESTONE © о ш pos a > MURPHREESBORO > S LIMESTONE 2 с ple, and McGinnis, 1979), respectively, from Ше Bro- mide Formation. Camerate crinoids, rhombiferans, and echinoids, most closely resemble forms occurring in the upper or Pooleville Member of the Bromide For- mation whereas the long-armed disparid inadunates better correlate with forms from the lower or Mountain Lake Member. Overall, the closest resemblance seems to be with Pooleville Member forms. Lebanon Limestone echinoderms are also similar to those of the Blackriveran Platteville Group and over- lying early Trentonian (Rocklandian) Decorah Subgroup of the upper Mississippi Valley (Kolata, 1975; Brower and Veinus, 1978; see Table 2). Similarities are greatest with the Platteville Group because Cu- Dulocrinus gracilis (Hall, 1847), Abludoglyptocrinus Kolata, 1982, Quinquecaudex Brower and Veinus, 1982, Praepleurocystis Paul, 1967a, ?Tanaocystis Sprinkle, 1982b, and Unibothriocidaris Kier, 1982, are known only in this interval. The mid to late Trenton faunas from the Hull, Sher- man Falls, and Cobourg members of the Ottawa For- Mation and equivalent units, southeast Ontario, are Similar to the Lebanon Limestone faunas (Table 2). Most species from the two areas, however, do not ap- Pear to be closely related. Both rhombiferans and para- crinoids from the Trenton faunas are generically dis- tinct. The Lebanon Limestone fauna shows few similari- ties with nearby Blackriveran to (?)Chazyan-age for- mations of the Ottosee Group (Benbolt, Hogskin, Lin- colnshire, and Wardell formations) and related rocks in east Tennessee and southwest Virginia (Table 2). The east Tennessee faunas are characterized by thick- plated rhodocrinitids such as certain species of Ar- chaeocrinus Wachsmuth and Springer, 1881, and Dia- bolocrinus Wachsmuth and Springer, 1897, hybocrin- ids (of which Hybocrinus Billings, 1857, is most common), and platycystitid paracrinoids, which are represented by Platycystites Miller, 1889, and allied forms. Of these, only Hybocrinus is present in the Le- banon Limestone. The stratigraphic distribution of echinoderms within other formations of central Tennessee and adjacent areas aids in determination of the age of the Lebanon. The Pierce Limestone is the second unit below the Lebanon in the Stones River Group of the study area. Three echinoderms identified thus far from this unit are Paleocrinus sp. (Bassler, 1932), Hybocrinus punc- tatus (Miller and Gurley, 1895), and Diabolocrinus sp. (Brower and Veinus, 1974); all three taxa are known BULLETIN 319 Table 2.—Occurrence of Lebanon Limestone echinoderm taxa and related species in Eastern North America. egion; approximate age; stratigraphic Classes uuu Genera Species Upper Mississippi Valley (from Kolata,1975) Blackriveran | Trentonian | Galena Gr. Platteville Gr. | including | Decorah Subgr. Arbuckle Mountains South Central Oklahoma (from Sprinkle,1982e) Blackriveran Bromide Fm. Mountain | pootevitie Mbr. Lake Mbr. East Tennessee- Southwest Virginia (mainly from Brower and Veinus,1974) Chazyan-Blackriveran Ottosee Group; Benbolt Fm. and other fms. Southeast Ontario (from Wilson,1946; Springer 1911) Trentonian Ottawa Fm. and other fms. Central Tennessee (Original, herein) Blackriveran Lebanon Limestone Crinoidea Abludoglyptocrinus A. spp. A. gregatus* A. charltoni* Archaeocrinus A. spp. A. snyderi* A. subovalis* Apodasmocrinus A. spp. A. daubi Carabocrinus spp. Cleiocrinus С. $рр. С. laevus* С. bromidensis* Columbicrinus С. crassus * С. sulphurensis* Cremacrinus С. spp. С. punctatus* C. sp.*? Cupulocrinus C. spp. C. gracilis Doliocrinus spp. Hybocrinus spp. Porocrinus Р. spp. Р, lebanonensis * P. elegans * Quinquecaudex spp.|* · - Reteocrinus Spp. Paracrinoidea Oklahomacystis spp. Rhombifera Tanaocystis spp. Praepleurocystis зрр. - · + Amecystis spp. Asteroidea Hudsonaster spp. Salteraster spp. Schuchertia spp. Edrioasteroidea Edrioaster spp. Cyathocystis spp. Echinoidea Unibothriocidaris Y sp3 U. bromidensis* U. kieri* Bothriocidaris B. spp. B. solemi* B. vulcani* . Neobothriocidaris spp. | | | ! 1 Undescribed specimen of Kolata (oral commun., 1981). 2 Cremacrinus sp. of Kolata (1975, p. 24). 3 = genus and species unknown of Kolata (1975, p. 68). * Denotes closely related species within the genus. LEBANON LIMESTONE ECHINODERMS: GUENSBURG 19 from the Ottosee Group of east Tennessee and similar forms occur in the Mountain Lake Member of the Bromide Formation, Oklahoma. The Curdsville Lime- stone Member of the Lexington Limestone of Кеп- tucky extends to the edges of the Central Basin, where it is only 0.5 to 3 m thick and is referred to as the Curdsville Member at the base of the Hermitage For- mation. The Hermitage is the second formation above the Lebanon and is the basal unit of the Nashville Group, which is separated from the Stones River Group by an unconformity (Wilson, 1949). The Curdsville, therefore, is significantly younger than the Lebanon. The Curdsville Member in Kentucky has produced a diverse echinoderm fauna that, among other species, includes Amygdalocystites florealis, Billings, 1854, Cu- Pulocrinus jewetti Billings, 1859, Reteocrinus alveola- tus Miller and Gurley, 1894, Carabocrinus radiatus Billings, 1859, and Paleocrinus angulatus Billings, 1857, all of which are also known from the mid to late Trenton of Ontario. Based on echinoderms from the Lebanon Forma- tion, and from overlying and underlying formations, the Lebanon seems to be middle to late Blackriveran in age. The Lebanon Limestone faunas are clearly older than those of the Trentonian of Kentucky and Ontario and younger than most of the Ottosee Group of east Tennessee and the Mountain Lake Member of the Bro- Mide Formation, Oklahoma. This age is in approxi- Mate agreement with correlations based on conodonts (Sweet and Bergstróm, 1976), and is approximately equivalent to slightly older than correlations based on brachiopods (Cooper, 1956). SYSTEMATIC PALEONTOLOGY INTRODUCTION The classification, arrangement, and terminology of echinoderms used herein are as listed below. Class di- Vision and arrangement only is from Sprinkle (1980, Р. 26). Crinoid classification follows part T ofthe Trea- tise on Invertebrate Paleontology except that the Ar- Chaeocrinidae is rejected (see Kolata, 1982) and the Classification of the Disparida is accepted, but with Teservations (see Remarks, Order Disparida). In ad- dition, the term “‘interradial,” rejected in the Treatise, 15 retained herein for use with certain camerate cri- Noids. Paracrinoid classification and terminology fol- low Parsley and Mintz (1975). Glyptocystid rhombi- ferans are classified and described after Sprinkle (19825), and pleurocystid rhombiferans after Parsley (1 970), except that quotes are deleted from certain oral and radial plate acronyms. Sea stars (Asteroidea) are Not classified to order (see Remarks, Class Asteroidea); Spencer and Wright (1966) were followed for familial and generic classifications and the terminology is after Blake (1973; oral commun., 1980-1981). Edrioaster- oids were classified and described following Bell (1976, 1980). Echinoid classification and terms are from Dur- ham (1966) and Kier (1982). Because crinoids are numerically the most common and taxonomically the most diverse echinoderm class in the Lebanon Limestone, this group was the most intensively studied. Many species fit well into pre-ex- isting classifications; a few species and higher level taxa, however, required emendation or redescription. Suprageneric revision of the Disparida, although need- ed, was not undertaken during this study. Nearly all non-crinoid echinoderms were readily incorporated into existing classifications. One sea star specimen, however, was described. Sea star orders were not re- vised, although this also is badly needed. Exact synonymies are given for the various previ- ously described taxa. Only the most important syn- onyms are typically listed; in particular, bibliographic (such as Bassler and Moodey, 1943) and other short references are commonly omitted. Explanations of acronyms used in the taxonomic section are given in Table 12. PHILOSOPHICAL CONSIDERATIONS The modern biological concept of a species is based on an organism’s having the potential to interbreed and produce fertile offspring in natural conditions. Modern metazoan species are thus nearly always readi- ly defined (clines and ring species are problems). This concept is, of course, untestable using fossils and fossil species are defined by comparative methods utilizing preserved parts. Modern relatives aid in development of criteria for recognition of fossil species. The species concept in paleontology is therefore somewhat arbi- trary but nevertheless does offer a reasonable approx- imation of biological species boundaries. Further, this system is amenable to criticism and revision. (For a detailed discussion of fossil species and their relation- ship to modern species, see Raup and Stanley, 1978.) Species concepts adopted in this monograph are those established by early workers and generally followed by modern students of fossil echinoderms. The echino- derm skeleton, due to its unique articulated construc- tion and complexity, serves as an excellent and primary taxobasis within this diverse group. In few (perhaps no) other phyla is the nature of the overall morphology, soft part anatomy, and functional morphology so well revealed by hard (preservable) parts alone. Indeed the nature of the skeleton is a primary taxobasis for mod- ern echinoderms as well as for fossils, and studies of 20 BULLETIN 319 recent species confirm the overall usefulness of the skeleton for differentiating species as well as higher taxa (see, e.g., Clark, 1915; Fisher, 1911). Characters most commonly of taxonomic value are: thecal (or body) shape; position and nature of openings and append- ages, and plate morphology, number, arrangement, size and ornament. New species (and higher level taxa) are herein based on clear consistent differences from pre- existing taxa in one or more of these characters, though plate ornament alone is generally not a good taxobasis. The overall goal of the following taxonomic section is to approximate, as well as possible, biological species for the Lebanon echinoderms. I consider my taxonom- ic assignments to be near the midrange in the spectrum of modern workers’ species philosophies, between those known either as “lumpers” or “splitters.” Fossil echinoderms are rarely complete, and speci- mens are few. This is a major hindrance to their study. The typical excellent preservation, and relatively large assemblages of Lebanon Limestone echinoderms offer a rare opportunity for such detailed observations. ABBREVIATIONS OF REPOSITORY INSTITUTIONS All specimens collected during this study are repos- ited in the collections of the Department of Geology, University of Illinois. Acronyms and addresses of oth- er cited specimen repositories are listed below: BMNH: Burpee Museum of Natural History, Rockford, IL Walker Museum Collection, Field Mu- seum of Natural History, Chicago, IL GSC: Geological Survey of Canada, Ottawa, Ontario, Canada FM (UC): MCZ: Museum of Comparative Zoology, Har- vard University, Cambridge, MA SUE University of Iowa, Iowa City, IA we University of Cincinnati Geology Mu- seum (Kopf Collection), Cincinnati, OH UI X: University of Illinois, Urbana, IL USNM: United States National Museum, Wash- ington, DC USNM S: United States National Museum (Springer Collection), Washington, DC Subphylum CRINOZOA Matsumoto, 1929 Class CRINOZOA Miller, 1821 Subclass CAMERATA Wachsmuth and Springer, 1885 Order DIPLOBATHRIDA Moore and Laudon, 1943 Suborder ZYGODIPLOBATHRA Ubaghs, 1953 Family CLEIOCRINIDAE Miller, 1889 Genus CLEIOCRINUS Billings, 1857 Type species. —Cleiocrinus regius Billings, 1857, p. 276. Cleiocrinus tessellatus (Troost) Plate 1, figures 3, 5—9, 11—17, 19—21 Campanulites tessellatus Troost, 1849, p. 419 (nom. nud.). Cleiocrinus tessellatus (Troost). Wood, 1909, p. 100, pl. 7, fig. 11. Diagnosis.—A species of Cleiocrinus characterized by large, elongate, steep-sided conical to funnel-shaped cup; cup sides diverging at approximately 10 to 30° from vertical at level of IBrr, then flaring slightly, usu- ally at level of IIBr, ,, to approximately 25 to 35°. Cup plates flat to slightly convex. Exospires of parallel rows of tiny pores forming subrhombic to subpolygonal pat- terns at plate surfaces; small central area of plates usu- ally without pores. Description.—Cup moderately large, conical, steep cup sides diverge at approximately 10 to 30° from ver- tical at level of IBrr,, then usually flare slightly to 25 to 35° approximately at level of IIBrr, з, flare can reach 45°, at least, adorally. Lower cup shape varies from round, without median ray ridges to subpentagonal, with low median ray ridges; ridges usually disappear by ШВгг. Lower cup plates thickened, tightly articu- lated, possibly ankylosed, sutures indistinct. Lower anal series can be slightly protuberant (UI X-5902; Pl. 1, fig. 20). Cup plates usually flat, can be slightly convex; sutures usually flush, can be slightly impressed. Exospires externally of subequally-spaced parallel rows of tiny pores; pore rows perpendicular to, bisected by, and abutting across plate sutures; forming sub- rhombic or other subpolygonal patterns on plate sur- faces. Pore rows lead internally to slit-like channels (Pl. 1. figs. 5, 7, 12, 16); channels oblique to plate surfaces (usually about 45°). Alternating thick and thin vane-like walls separate channels; thin walls extend only a short distance internally so that channels sep- arated by thin walls merge below (Pl. 1, fig. 16), thick walls extend below to main body of plate, except at plate margins. Channels open along plate margins into interarticular canals. Canals formed equally by орро5- ing articulating plates, expanding toward plate corners; canal can be continuous across facet face forming low inverted V-shape in lateral view, or divided at center of facet face into two separate halves. Canals connected to coelomic cavity via subcircular to suboval pores at inner plate corners. Inside of cup with wide, radially-positioned grooves; these extending axially down each ray, and ray bifur- LEBANON LIMESTONE ECHINODERMS: GUENSBURG 24 cation. Pores, connecting to exospires, open on ridges between grooves (Pl. 1, fig. 13). External plate surfaces with indistinct stellate pattern formed by non-pore-bearing plate centers, narrow rays radiating to plate corners; in some specimens addi- tional rays extend to midpoints between plate corners; rays abut across adjacent plates. Plate centers small, usually approximate plate outline in shape, slightly convex to flat; can have fine, faint, subconcentric ridges and grooves, small shallow central depression divided by low vertical ridge. Uncommonly, specimens have non-pore-bearing centers, no distinguishable rays. Ar- ticulation surfaces between cup plates finely crenulate did. fig). IBB five, wider than tall in one specimen where visible, confined to basal concavity; axial canal large, Subpentagonal. Columnal articular surface forms faint to prominent notch in basal concavity; either smooth or with faint radially disposed crenulae. BB five, small, apparently subpentagonal to suboval, wider than tall; alternating with RR that form a circlet of ten plates, at least; circlet projects over, below IBB. One specimen (UI X-5902) appears to have supernumerary plates intercalated into basal-radial circlets. RR small, sub- pentagonal, wider than tall, widest distally, much smaller than IBrr. IBr, , axillary, wider than tall. Suc- ceeding axillaries known on IIBrr, , and ПВг,. IIBrr Slightly wider than tall to nearly as tall as wide. Prox- imal anal plates known in one specimen (UI Х-5902; PL. 1, fig. 21); subhexagonal, arranged in vertical row. Anal plates 1, 2 (counting from base of cup) wider than tall, anal plate 3 about as wide as tall. Distal cup, free arms, tegmen known. Column long, tapering proximally; wide to very wide in preserved columnals, ratio of height to width varies from approximately 1:5 to 1:8; columnals 7 to 9 mm in diameter, cup column articular facets 4 to 5 mm in diameter indicating proximal column of this width. Columnals straight-sided, smooth to pitted (pitted sur- faces could be weathered). Columnals of pentameres; Pentameres articulate laterally along crenulate margin; Zygum divided into outer narrow crenularium, wider attenuated inner areola. Areolae with vermicular ridges and grooves, confluent with crenulae ridges and grooves. Shallow, triangular, smooth-bottomed notch- €s interrupt crenulae, areola ridges and grooves; notch- €s bisected by pentamere articulations. Lumen large, Subpentagonal, pentagon corners at mid-points of pen- tameres. Holdfast unknown. Discussion. —Cleiocrinus regius Billings, 1857, and С. magnificus Billings, 1859, both from the latest Tren- tonian Cobourg Member, Ottawa Formation, Ontario, differ most conspicuously from C. tessellatus in that the former has well-developed median ray ridges and external exospires of short channels (slits), whereas the latter has a larger, wider, more rapidly expanding cup. C. sculptus Springer, 1911, from the middle Trenton- ian (Kirkfieldian) Curdsville Limestone, Kentucky, and С. springeri n. sp. from the Lebanon Limestone, both differ in having highly ornamented cup plates with distinctly raised plate centers. The poorly known C. ornatus Kolata, 1982, from the Blackriveran Pooleville Member, Bromide Formation, Oklahoma, differs in having exospires of rows of channels rather than pore rows. Deeply weathered C. tessellatus plates show a pattern of channels similar to that in C. ornatus (com- pare pl. 16, figs. 11, 12 of Kolata, 1982, with Pl. 1, fig. 16). C. laevus Springer, 1911, also from the Lebanon Limestone, C. perforatus Hudson, 1911, from the Cha- zyan Valcour Limestone, New York, and C. bromi- densis Kolata, 1982, from the Bromide Formation all have exospires of pores along sutures only. Three crown fragments of C. tessellatus from differ- ent localities were found associated with similar col- umn segments of large diameter; no other identifiable echinoderms were associated with these specimens. Further, the morphology is similar to that of specimens of С. regius (Springer, 1905, pl. 1, fig. 3). These column fragments are therefore considered to belong to C. tes- sellatus; they indicate a heteromorphic, proximally ta- pering column. Final confirmation, however, must await discovery of an attached crown and column. Cleiocrinus libanus Safford, 1869 (p. 285; nom. nud.), was placed in synonymy with C. tessellatus by Bassler and Moodey (1943, p. 365). Safford’s material is now missing (Alberstadt, oral commun., 1980), making ver- ification of Bassler’s identification impossible. Types and occurrence. — The holotype, USNM 39910, is from an unspecified horizon in the Lebanon Limestone along the Duck River near Columbia, Mau- ry County, Tennessee. Twenty-one hypotypes include UI X-5894 through 5911, 5917; USNM 166816; and USNM 42271. UI X-5894 is from the upper member at locality Z-653. UI X-5895, 5899, 5902, and 5903 are all from the lower member at locality Z-651 and UI X-5904 through 5907 are from the upper member atlocality Z-651. UI X-5896 is from the lower member at locality Z-654a and UI X-5908 is from the lower member at locality Z-654. UI X-5898, 5899, and 5909 are all from the lower member at locality Z-652. UI X-5900 and 5901 are both from the lower member at locality Z-656 and UI X-5910, 5911 are from the lower member at locality Z-658. USNM 42271 is from an unknown horizon in the Lebanon at Shelbyville, Ten- nessee, and USNM 166816 is from an unspecified ho- rizon in the Lebanon northwest of Woodbury, Cannon 22 BULLETIN 319 County, Tennessee. Figured specimens are Ше holo- type USNM 39910 and hypotypes UI Х-5895, 5897, 5898, 5900, 5902, 5903, 5908, 5917; USNM 42271 апа 166816. Cleiocrinus springeri new species Plate 2, figures 4, 8, 9 Etymology of name.—The specific nomen honors Frank Springer, well-known student of fossil echino- derms. Diagnosis.—A species of Cleiocrinus characterized by a steep-sided, conical cup. Cup plates highly or- namented, having high relief; exospires externally of rows of parallel, discontinuous slit-like channels form- ing subrhombic to subpolygonal patterns on plate sur- faces; cup plate margins upraised slightly, centers pro- tuberant, without slits; ridges radiate off plate center protuberances toward plate corners, most abut across plates. Description.—Cup moderately large, tall, funnel- shaped (Pl. 2, figs. 8, 9); expansion angle approximately 10° from vertical at level of IBr,, expanding to approx- imately 20 to 30° at level of ШВгг. Basal concavity wide. Cup plates with prominent ornament (PI. 2, fig. 4), high relief, plates of lower part of cup thick, tightly sutured (perhaps fused), plate boundaries obscure. Exospires of discontinuous channel-like slits separated by irregular vane-like walls. Slits perpendicular to, bi- sected by, or abutting across sutures; forming sub- rhombic or other subpolygonal patterns on plate sur- faces. Slits increase in size, decrease in frequency proximally on cup; approximately 60 slits per 5 mm on IIIBr,, approximately 30 slits per 5 mm on distal IBr, margin. Plate margins forming elevated ridges along plate sutures. Plate centers protuberant, without slits, roughly approximating plate outline in shape; ridges radiate from protuberances toward and often abut across plate corners; smaller ridges commonly radiate from protuberances at right angles to plate faces. IBB apparently five, boundaries obscure. BB fve, alternate in circlet of ten plates with RR; apparently small, shape uncertain. RR five, small, much smaller than Brr; each with wide medial vertical ornamenting ridge, single depression on either side. IBrr, apparently axial, pentagonal, much wider than tall. IIBr, axillary; IIBrr wider than tall. Proximal ШВгг wider than tall; cup unknown beyond ШІВг,. Anal plates known through anal plate 10, form vertical series. Anal plates 1 to 5 wider than tall; anal plates 5 to 10 taller than wide. Tegmen, arms, column unknown. Discussion.—The new species closely resembles C. tessellatus (Troost, in Wood, 1909), also from the Leb- anon Limestone, in cup shape and plate arrangement. The new species differs most conspicuously, however, in possessing prominent cup plate relief including cen- tral protrusions and upraised margins. С. sculptus Springer, 1911, from the Trentonian Curdsville Lime- stone of Kentucky differs from C. springeri n. sp. in having a much wider basal concavity and non-raised plate margins. The new species differs from all other species in possessing discontinuous pore slits and high plate relief. The E ray side of the holotype was buried facing downward, touching the bedding plane below. This area has undergone differential dissolution, obliterat- ing plate relief (Pl. 2, fig. 8). The degree of dissolution decreases gradually away from the E ray and is in- versely proportional to distance from the bedding plane. Type and occurrence.— The holotype and only known specimen, UI X-5918, a partial cup, 15 from the lower member, Lebanon Limestone, at locality Z-654a. Cleiocrinus laevus Springer Plate 2, figures 1-3, 6, 7; Text-figure 3 Cleiocrinus laevus Springer, 1911, p. 44, pl. 5, figs. Па-с. Cleiocrinus tessellatus (Troost). Bassler and Moodey, 1943, p. 365. Cleiocrinus tessellatus (Troost). Ubaghs, 1978b, fig. 174, nos. 6-8. Diagnosis.—A species of Cleiocrinus characterized by a rapidly expanding funnel-shaped cup (PL 2, figs. 1, 2, 6, 7); expansion angle of about 10 to 20° from vertical at level of BB-RR circlet, increasing to about 60 to 70° at Шат. Exospires of rows of pores along plate sutures, about 18 pores per 5 mm at level BB- RR circlet, increasing to about 37 pores per 5 mm at level of Шат. Plates nearly flat with very fine, irreg- ular, anastomosing, concentric ridges and grooves. BB- RR circlet forms thin attenuated collar extending downward over proximal column. Emended description. — Aboral cup rapidly expand- ing funnel (Pl. 2, figs. 1, 2, 6, 7), expansion angle about 10 to 20° from vertical at level of BB-RR circlet, in- creasing to about 60 to 70° at level of IIIBrr. Exospires of rows of sutural pores (Pl. 2, fig. 3), about 18 pores per 5 mm at level of BB-RR circlet, increasing to about 37 pores per 5 mm at level of IIBrr; pores extend internally to interarticular canals. Two canals per plate facet, each extends internally and laterally from near midpoint of facet to nearest corner. Canals connect at plate corners to large pores, which open in inside of cup. Inside of cup with wide grooves extending lon- gitudinally out each ray, and ray bifurcation. Large pores enter inside of cup in high areas between grooves. Lower cup obscurely subpentagonal, with faint median ray ridges. Plates nearly flat except for ray ridges, with fine irregular anastomosing subconcentric ridges and LEBANON LIMESTONE ECHINODERMS: GUENSBURG ра grooves (Pl. 2, fig. 3). Brr can have shallow central depression divided by vertical ridge, faint grooves ex- tending in stellate pattern away from plate centers. Basal concavity large, subround in outline. IBB five (damaged in available material), deeply in- set in basal concavity, wider than either BB or RR; column articular facet with fine crenulae; circlet un- dulant, with highs adjacent to RR. Narrow shallow channel between articular facet, BB-RR circlet. BB, RR each five, alternate in circlet of 10 plates, forming attenuated collar projecting downward over anterior- most column; lower edge of collar with RR projecting slightly farther downward than BB. BB subpentagonal, usually wider than high, can be about as wide as high; size variable, average slightly larger than BB. RR wider than high, subpentagonal. Small wedge-shaped ?su- pernumerary plate below E ray R in holotype. IBr, ; axillary, IBr, axillary in В and D rays of holotype. IBrr much wider than high, lateral margins usually diverge rapidly distally to accommodate rapid cup expansion. IBrr, inverted V-shape. IIBrr; „4 axillary; IIBrr much wider than tall. Secundaxillaries subpentagonal, usu- ally with slight indentation along lower margin. Prox- imal ШВгг wider than tall. Anal plates form single vertical row; anal plates 1 to 5 much wider than tall, anal plate 6 wider than tall. Distal cup, tegmen, arms, column, holdfast unknown. Discussion.—C. laevus is distinguished from most other species of Cleiocrinus by its exospire openings of sutural pores. Both C. bromidensis Kolata, 1982 from the Blackriveran Bromide Formation, Oklahoma and C. perforatus Hudson, 1911 from the Chazyan Valcour Limestone, New York have similar exospire morphologies. The former differs from C. /aevus in having a relatively smaller basal concavity, a less at- tenuated BB-RR circlet, and a more regular arrange- Text-figure 3.— Cleiocrinus laevus Springer, drawing of lower part ОЁ cup. Shows CD interray (right), D ray (center), and E ray (left) of holotype (USNM 50045); 25 = possible supernumerary сир plate, approximately x4. ment of fine concentric ridges and grooves on cup plates. The latter species is known only from a few patches of apparently deeply weathered cup plates; it appears to differ from C. /aevus in possessing fewer, larger sutural pores. Types and occurrence.—Only the holotype, USNM 50045, was available for study and is refigured here. Springer (1911) lists the specimen as being from an unspecified horizon of the Lebanon at Shelbyville, Bedford County, Tennessee. columns of Cleiocrinus species Blate figures: 2, 10,25 Remarks.—Two column fragments are here assigned to Cleiocrinus sp. Both were found as isolated speci- mens. Generic assignment is based on great similarity of morphology to fragments assigned to C. tessellatus (Troost in Wood, 1909) from the Lebanon and С. regius Billings, 1859, and C. magnificus Billings, 1859, both from the Trenton of Ontario (Billings, 1859, pl. V, figs. 1, 3; Springer, 1905, pl. 1. figs. 1-3, 12). The zygum morphology of column fragments assigned to C. tessellatus is distinctive and unlike that of any other known Lebanon crinoid (see Cleiocrinus tessellatus de- scription herein). The fragments assigned here have similar zygal morphology but differ in other details. UI X-5916 (Pl. 1, figs. 10, 25) differs from column segments assigned to С. tessellatus in having a sub- pentagonal column outline, a smaller column diameter (6.8 mm), a slightly smaller columnal height to width ratio of about 1:10, and columnals with slightly con- cave latera at angles of subpentagonal outline. UI X-5915 (Pl. 1, figs. 1, 2) differs in having a wider col- umn diameter (about 12 mm), a subpentagonal column outline, a much smaller columnal height to width ratio of about 1:20, and small lens-shaped ossicles interca- lated at irregular intervals between columnals, ap- proximately at mid-points between lateral articula- tions of pentameres. The column in C. regius is heteromorphic and it is probably also heteromorphic in C. tessellatus (see Remarks under C. tessellatus); one or both fragments could therefore belong to the latter species, and the differences in morphology could reflect only position along the column. UI X-5915 bears a strong similarity to distal column in C. regius (com- pare Pl. 1, figs. 1, 2 with pl. 1, fig. 3e of Springer, 1905). It is also possible that the fragments belong to another Lebanon species of Cleiocrinus, either C. springeri n. sp. ог С. laevus Springer, 1911. The column is un- known in both of these rare species. These column fragments are therefore not specifically assignable. The two column fragments also differ in that UI X-5915 has lateral pentamere articulations at the an- 24 BULLETIN 319 gles of the subpentagonal column, whereas the artic- ulation points of UI X-5916 are midway between the angles. Types and occurrence.—Figured specimens UI X-5916 and UI X-5915 are from the lower member, Lebanon Limestone, at localities Z-656 and Z-652, respectively. holdfasts of ?Cleiocrinus species Plate 1, figures 4, 18 Remarks.— Four encrusting holdfasts are here ten- tatively assigned to Cleiocrinus. All are large, varying from 30 to 43 mm in maximum diameter. The spec- imens have a low to high cone-shaped upper surface; UI X-5914 is nearly round in outline, UI X-6012 is suboval, and UI X-5912 and UI X-5913 (Pl. 1, fig. 18) are subround with short lobate radicles. Upper surfaces are composed of very thick, flat or near flat, tightly sutured, flush, subpolygonal plates. UI X-5914 (Pl. 1, fig. 4) and UI X-6012 have a thin lower disc with numerous radiating septa extending upward into the holdfast lumen. All specimens have a large holdfast lumen; none of the specimens has a well-preserved column facet. The column lumen diameter was ap- parently large, approximately 10 to 20 mm. Generic assignment is based on large size and similar morphology to holdfasts of Cleiocrinus regius Billings, 1859, from Trenton rocks of Canada (Springer, 1905, pl. 1, figs. За-Зе). Upper surfaces in all species as well as the basal disc morphology in UI X-5914 and UI X-6012 are also very similar to those of Cleiocrinus sp. from the middle Trentonian Bobcaygeon Lime- stone, Ontario (Brett and Liddell, 1978, p. 339, fig. 6). The distalmost column lumen in the Lebanon speci- mens appears to have been very wide (about 10 to 20 mm), indicating a wide distal column. Cleiocrinus is the only Lebanon echinoderm known to have pos- sessed a distal column and lumen of such large di- ameter. The Lebanon specimens are also similar to those of Brett and Liddell in having variable mor- phology of the holdfast, depending on the substrate topography. UI X-5914 (Pl. 1, fig. 4) is attached to a relatively flat substrate and is circular in outline with- out short lobate extensions. In contrast, UI X-5913 (PL 1, fig. 18) has short lobate extensions that conform to the irregular topography upon which it attached. Types and occurrence.— UI X-5913 is from the (?)upper member of the Lebanon Limestone at locality Z-661; UI X-5912 and UI X-5914 are from the lower member, Lebanon Limestone at localities Z-656 and Z-658, respectively; and UI X-6012 is from the upper member, Lebanon Limestone, approximately one m above the massive member at locality Z-651. UI X-5913 and UI X-5914 are figured. ?tegmen of ?Cleiocrinus species Plate 1, figures 22-24; Text-figure 4 Description. — Fossil shape low, conical, with round- ed subcentral apex; outline subcircular. Fossil con- structed of tightly sutured, possibly locally fused array of at least two basic types of small plates that enclose internal network of tunnels (Pl. 1, fig. 22). Plates rel- atively massive toward cone apex, thinning toward periphery. Tunnels ovate to subcircular in cross-sec- tion, radiating from apex and bifurcating at least three or four.times distally, approximately (?)five tunnels near apex and approximately 100 about periphery. Tunnels decrease in diameter from approximately 1.0 mm at center apex to 0.5 mm near periphery; spaced at intervals equal to or slightly wider than tunnel widths. Cone apex damaged, features obscure. Small subcir- cular area developed adjacent to apex constructed of small closely packed, thick plates (Pl. 1, fig. 24). Exposed surface other than small subcircular area of above constructed of two plate types (Text-fig. 4). Туре 1 arranged in alternating to nearly opposing biseries that covers tunnels. Type 2 arranged in opposing bi- series of attenuated wall-forming plates above and be- tween tunnels. Type 1 plates transversely elongate, dumbbell-shaped, abutting medially above tunnels along a straight or more commonly zigzag surface. Type 1 plate medial constrictions producing longitudinal rows of sutural pores along each row of the biseries; pores apparently opening into tunnels below. Type 2 plates forming peaked walls; each T-shaped as seen from above; each abuts laterally along short arm with opposing type 2 plate, forming H-shaped pair; oppos- ing plate pairs connected longitudinally in series 10 form ladder-like pattern with deep subrectangular to Text-figure 4. —?tegmen of ?Cleiocrinus sp., tracing of portion of specimen. Type 1 plates (P1) cover tunnels, articulate laterally with type 2 plates (P2); approximately x4.5. | ) | ) | LEBANON LIMESTONE ECHINODERMS: GUENSBURG 25 suboval depressions in gaps between plates. Type 1 plates tightly abut type 2 plates approximately above lateral margins of tunnels; articulation location alter- nates from mid-point of each type 2 plate longitudinal wall to point of contact between adjacent longitudinal walls of type 2 plates. Remarks.—The radiating pattern of enclosed tun- nels in this fossil is unlike other echinoderms, but re- sembles the internal ambulacral tracts in many cam- erate crinoids in which the ambulacral grooves and ambulacral plates are found below the outer plate sur- face along the inner side of the tegmen. Some camerates have subcircular tubes comprised of numerous plates that extend from the arm bases along the inside of the tegmen and probably contained the food grooves (Ubaghs, 19785, p. T179). Tunnels in the ?tegmen de- scribed here are similarly thought to have been food grooves. Sutural pores between type 1 plates are in- ferred to have been communication avenues (for res- Piration, nutrients, or nervous tissue?) to supply tissues lining the tunnels. The small subcircular area of closely Packed plates adjacent to the central area might, by analogy with camerate crinoids, have been the anal Structure. The large number of equally spaced tunnels arrayed about the margin of the ?tegmen likely reflects the arrangement of arms about the calyx. Among the known Lebanon crinoids only Cleiocrinus possesses such an arm arrangement. The large size of the ?tegmen is also compatible for Cleiocrinus. Final evaluation Must await discovery of an associated dorsal cup and tegmen. The exposed side of this fossil is presumed to be internal because of the exposed complicated frame- work containing numerous pores, deep depressions, and attenuated walls. Channels were revealed by pol- ishing a vertical surface broken through the side of the Specimen. Plates are generally tightly sutured, possibly fused. Gaps in the specimen most often occur as frac- tures through plates rather than along their margins. The morphology and preservation of the specimen, therefore, suggest that it formed a rigid structure that Offered protection and support for the network of ra- diating tunnels at the top of the calyx. The specimen Shows no sign of significant distortion due to compac- tion. The occurrence of this ?tegmen as an isolated entity is unusual. A partial explanation may be found in the fact that the edges of the specimen are thin, indicating a zone of relative weakness. The adoral part of the cup and arm bases in Lebanon species of Cleiocrinus have never been found despite the relatively large number of lower cup portions, suggesting fragile construction near the arm bases. The ?tegmen morphology is superficially suggestive of the inside of the aboral cup in Cleiocrinus. It differs most conspicuously in its slightly concave shape (which would literally require the specimen to be turned inside out if it were a cup), the absence of a central axial canal at the apex, which would be the column position if the fossil were an aboral cup, and the presence of internal tunnels rather than the open grooves typical of Cleio- crinus cups (Pl. 1, fig. 22). Types and occurrence.— The figured single speci- men, UI X-5919, is from the lower member, Lebanon Limestone, approximately 8 m above the base of the Lebanon at locality Z-652. Suborder EUDIPLOBATHRINA Ubaghs, 1953 Superfamily RHODOCRINITACEA Roemer, 1855 Family RETEOCRINIDAE Wachsmuth and Springer, 1885 Diagnosis.—Calyx obconical, bilaterally symmetri- cal across the A/CD plane. Calyx with strong median ray ridges. Arms uniserial, pinnulate or non-pinnulate. Anal area with prominent sagittal anal series. Inter- brachial areas depressed, filled with numerous small plates. Discussion.—Ubaghs (1978a, p. T414) lists three genera in the Reteocrinidae; Reteocrinus Billings, 1859, and Traskocrinus Kolata, 1975, from the Middle Or- dovician and Gaurocrinus Miller, 1883, from the Up- per Ordovician. As discussed under Remarks on Reteocrinus, Traskocrinus is here synonymized with Reteocrinus. Gaurocrinus differs from Reteocrinus in being pinnulate and in not having either interbasal or interinfrabasal gaps. In addition, Gaurocrinus has pro- portionately smaller IBB and fewer arm branchings. Genus RETEOCRINUS Billings, 1859 Type species. —Reteocrinus stellaris Billings, 1859. Diagnosis. — А reteocrinid with steep-sided to widely flaring convex-sided cup; sides of cup with greatest curvature approximately at level of radials. Major cup plates large, deeply cleft (i.e., multilimbed), forming prominent reticulated framework with interbasal gaps (spaces between major cup plates), small gaps or depressions on primary plate surfaces in interinfra- basal areas. Interbasal gaps, possibly interinfrabasal gaps (when present), interbrachial areas, occupied by small, irregular, loosely articulating plates. Secondary plate fields depressed below level of primaries: appar- ently embedded in flexible integument in life (Kolata, 1975, p. 47; Brower, 1974, p. 23). Posterior crown of large pentaxial basal followed by prominent sagittal anal series extending nearly to top of, and confluent 26 BULLETIN 319 with, tegmen where known. Tegmen vaulted, sac-like, flexible; not known to extend beyond arm tips. Arms non-pinnulate, branching isotomously at least three times. Column round or pentagonal; where known, lumen usually wide, quinquelobate. Articulae crypto- symplectial. Description. — A reteocrinid with small to large adult crown, cup nearly straight-sided, steeply conical to widely flaring, convex-sided with greatest curvature approximately at level of RR. Major cup plates deeply cleft (i.e., multilimbed), forming prominent reticulate framework, with gaps in interbasal areas, with depres- sions or small gaps in interinfrabasal areas; plates evenly rounded, somewhat flattened, or slightly distended. Keels may radiate from plate centers onto plate limbs, and may abut keels from adjacent plates. Other surface ornamentation can be smooth, reticulated, striated, or warty. RR and Brr can bear lateral projections that extend into interbrachial areas. IBB five, relatively large, open U-shaped in species with interinfrabasal depressions, low x-shaped in species with interinfra- basal gaps. One small plate can occupy interinfrabasal gaps. BB five, large, x-shaped; commonly higher than wide, wider distally than proximally. CD B pentiaxial with distal vertical extension supporting primanal. Each B articulates with two iBB, two RR, numerous small interbasal, iBr plates. BB form sides of interbasal gaps, bottom of interbrachial areas, upper margin of inter- infrabasal depression or gap. Interbasal gaps generally large, filled with small, depressed plates similar to those in interbrachial areas. RR five, with low, inverted Y-shape, each articulates with two BB, IBr,, small in- terbasal, interbrachial plates. RR form upper margin of interbasal gap, lower margins of interbrachial area. Posterior of prominent tall sagittal anal series extend- ing nearly to top of tegmen where known. Proximal anals usually resemble primibrachs in size, shape, but can be smaller. Anal series generally symmetrically disposed in CD interray, confluent with tegmen, bor- dered on either side by depressed interbrachial fields. Arms non-pinnulate, branching isotomously at least three times. IBrr, с, or rarely RR, axillary. Interbra- chial areas depressed, filled with small, loosely artic- ulating plates, confluent with tegmen; apparently imbedded in a flexible integument in life. Tegmen vaulted, sac-like, not known to extend beyond arm tips; web-like extensions connect to sides and ventral surfaces of arms (Kolata, 1975, p. 47). Anal opening on tegmen near top of anal series (Kolata, 1975, p. 47; Pl. 5, fig. 4). Column round or pentagonal, ?homeo- morphic or xenomorphic, usually broad. Columnals heteromorphic, often thin, highly varied in some species, can be constructed distally of pentameres. Ar- ticulations cryptosymplectic or rarely ankylosed; lu- men quinquelobate where observed, terminations ra- dially disposed. Column can terminate as multiplated subconical encrusting holdfast. Discussion. — Reteocrinus is now known from 10 species, three of which are new. Characters most useful in species recognition include shape of aboral cup, plate ornament, nature of interinfrabasal areas, shape of IBB, number of IBrr, and column features. Char- acters less consistently useful are the shape and relative size of the RR and BB, size of the interbasal gaps, size and shape of the anals, and nature of the iBrr plates. Known species of Reteocrinus are compared in Table 4. Terminology for the lateral projections found on the RR and Brr of many Reteocrinus species varies among authors. These have been called /ateral buttresses (Springer, 1911, p. 11), sharp girders (Raymond, 1931, p. 203), and /ateral spines (Kolata, 1975, p. 45). Be- cause these are apparently homologous structures, they are here all referred to as /ateral projections and are differentiated by adding modifiers (e.g., wedge-shaped, hemispherical, etc.). This new terminology is used be- cause none of the previously used terms describe all of the shapes seen. Traskocrinus Kolata, 1975 was first described as a monotypic genus whose principal diagnostic character was considered to be the presence of large interbasal gaps filled with small plates similar to those present in the interbrachial areas. In reexamining type and topo- type specimens of all species of Reteocrinus, I found proportionately large interbasal gaps similar to those in Traskocrinus in all species except R. depressus Ko- lata, 1982, and R. rocktonensis Kolata, 1975, and rel- atively smaller gaps in these species. This finding dif- fers from that of Wachsmuth and Springer (1883, p. 262-263, fig. 1; 1897, p. 178), who thought that Ке- teocrinus had interbasal depressions rather than gaps. Subsequently, Ubaghs (19782, p. T414) described Re- teocrinus as generally having spaces (i.e., gaps) in the infrabasal areas. I have identified plates in the inter- basal gaps similar to those in the interbrachial areas in Reteocrinus stellaris Billings, 1859 (the type species of Reteocrinus), R. alveolatus Miller and Gurley, 1894, R. polki n. sp., and R. variabilicaulis n. sp. I further believe that most or all of the remaining species of Reteocrinus had such plates because the interbasal gaps and adjacent cup plates are similar among all species of Reteocrinus. Incomplete preservation or highly in- durated matrices in the available material precludes conclusive determination of this point in R. depressus Kolata, 1982, R. elongatus Raymond, 1931, R. fenes- tratus n. sp., R. rocktonensis Kolata, 1975, and R. spi- LEBANON LIMESTONE ECHINODERMS: GUENSBURG 27 Table 3.—Measurements (їп mm) of specimens of Reteocrinus from the Lebanon Limestone. | Cup height Cup width Column di- \ Crown at tops at tops IB IB B B R R ameter at | height of RR of RR Height Width Height Width Height Width base of cup / | R. fenestratus 1 UI X-5690 - 9.5 8.9* 2.9 3.6 3.9 3.4 3.9 3.4 ен ) UI X-5695 = = ан 2.9 3.4 3.2 2.9 3.2 2.9 2 | R. variabilicaulis UI X-5805 39 6.3 9.5* 1:6 2.9 3.0 255 2:6 2948 4.3 | UI X-5411 42 TES) 1d 27 4.0 4.0 3.4 3.4 3.8 6.8 \ UI X-5412 28 11 4.9* 1.2 2.1 2% 2.0 2.1 2.0 3.7 UI X-5409 20 4.6 64 12 2: 2.6 22 2.4 2.8 4.0 | R. polki | UI X-5790 41 10 20* — - ek 4.8 4.8 5.0 10.4* | UI X-5417 22 52 9* 12 2 2.6 2.2 2.4 2.8 4.0 | UI X-5413 _ = = 2.8 4.9 4.4 4.2 = a ~ К. sp. | UI X-5416 23 5 - 1:5 2.8 9 2.4 2.6 2:9 327 ) К. sp. cf. | К. variabilicaulis UI X-5414 30 12 9* hd 2:7 - - 2.8 2.8 3.8 AAA AAA Сы: —.— ee ee * = estimated. * = аБпогта!. позиз Kolata, 1975. Each of these species is known from one to three specimens. Other characters used in the definition of Trasko- crinus include the number of IBrr per arm and crown Size. Primibrach number in Traskocrinus lies within the range found within two species assigned to Reteo- crinus, R. spinosus and R. rocktonensis (see Table 3). Primibrach number is a variable feature and has been Used as a species characterization in Reteocrinus (Ko- lata, 1975, p. 41; Raymond, 1931, p. 203; herein). The Crown of the one known specimen assigned to Trasko- Crinus is about 50% larger than that of any specimen traditionally assigned to Reteocrinus. Crown size among largest specimens of known species of Reteocrinus Tanges from about 25 to well over 50 mm, a factor Over 100% (Table 4). Size, therefore, seems to provide 4 weak generic criterion. Because of these similarities, Tjudge Traskocrinus and Reteocrinus to be congeneric and Traskocrinus, therefore, is herein synonymized With Reteocrinus. The ontogeny of Reteocrinus is poorly known. The Maximum range in crown size of a Reteocrinus species, MR. polki n. sp., is from 22 to 60 mm. Significant size ranges are also known from R. alveolatus Miller and Gurley, 1894 (see Springer, 1911, pl. 1, figs. 1-5) and - variabilicaulis п. sp. (Table 2). Growth in Reteo- crinus was anisometric. Smaller specimens typically differ from the larger ones in that multilimb extensions of Primary cup plates, cup gaps and (or) depressions, and plate ornamentation, particularly the lateral pro- jections on the RR and Brr, are disproportionately small. In species having a pentagonal column, the col- umn increases in angularity with size. Branching in- creases at an essentially constant rate later in ontogeny and therefore branches are fewer in number in smaller specimens. Identification of smaller specimens can be difficult because features that develop late in ontogeny are often those most useful in species recognition. Intraspecific variation is important in Reteocrinus. As summarized under the descriptions, all characters used for species definition were found to vary within populations (e.g., R. polkin. sp. and R. variabilicaulis n. Sp.). In addition to the three new species described here, the type species, Reteocrinus stellaris Billings, 1859, is herein redescribed. Remarks concerning the descrip- tions of several other species, including R. alveolatus Miller and Gurley, 1894, R. elongatus Raymond, 1931, R. mahlburgi (Kolata, 1975), and R. spinosus Kolata, 1975, are also provided. Specimens of R. alveolatus and R. elongatus are refigured. Range and occurrence.— Middle Ordovician, ?Whiterockian, Blackriveran through Trentonian. North America; U.S.A. (Illinois, Kentucky, ?Nevada, Oklahoma, Tennessee, Wisconsin), Canada (Ontario, Quebec). Mode of life. — Опе complete specimen of В. varia- bilicaulis n. sp. including crown, column, and holdfast, plus nearly complete specimens of R. polki n. 5р. and R. spinosus were available for study. R. variabilicaulis 28 BULLETIN 319 Table 4.—Summary comparison of species of Reteocrinus. Crown height? width of cup Maximum Width vs. height of (in mm) at Nature of known crown cup (in mm) at middle middle of Position of first interinfrabasal Species size in mm of RR' RR arm branching areas stellaris >45 w>h 3 IBr, d alveolatus >45 w>h 4 IBr,_, g depressus 22 w>h 2 RR-IBr, very small d elongatus Зи м > ћ 4 IBr, 5 small а fenestratus - w>h - - Б тай Бигот 90 ()м> 8 4 IBr,_, d polki 60 w>h 3 IBr3_»4 d rocktonensis 23 w>h 3 IBr, ; g spinosus 21 -— 4 ТВг. 6 а variabilicaulis 45 w>h 4 Шг, 4 а mutli-limbed fold plates - = = = 5 of Sprinkle, 1971 12 = estimated due to flattening; d = depressions, g = gaps. and R. spinosus attached to the substrate by means of a dorsally polyplated encrusting holdfast. The holdfast was floored in R. variabilicaulis by an attachment disc. Total column length varies according to species; it is short in R. variabilicaulis, moderate in К. polki and R. spinosus; and long in R. alveolatus. A short, narrow column segment adjacent to the holdfast in R. varia- bilicaulis was probably flexible, because it is divided into loosely articulated pentameres. Flexibility is also suggested by the relatively strong curvature of pre- served pentamere segments. Short distal columns in R. polkiand R. spinosus are comprised of many small, irregular, subpolygonal, loosely articulated plates: these were also probably flexible. The proximal to medial columns in nearly every known species of Reteocrinus in contrast appear to have been only slightly flexible, as is indicated by the pervasive straight to slightly curved column attitudes and more importantly, the tight cryptosymplectial (or rarely intermittently an- kylosed) articulations of the columnals. The columns in living R. variabilicaulis, R. spinosus, and R. polki were rigid except for movement at the short flexible distal sections. This overall column morphology is functionally very similar to the Lebanon Limestone disparid inadunates Columbicrinus crassus Ulrich, 1925, and Tryssocrinus endotomitus n. gen. et sp. The crown, particularly the interbrachial areas, ap- pear to have been pliant in Reteocrinus (Brower, 1974, p. 23; Kolata, 1975, p. 47; Ubaghs et al., 1978, p. T285). Interbasal, interbrachial, and tegmen plates are subpolygonal with thin attenuated lateral margins, un- like the planar (zygosynostosial) lateral margins that are found in most other camerate taxa. These plates were presumably imbedded in and held together by a flexible integument. Study of specimens of R. alveolatus from Ontario shows that the interbasal-interbrachial plates in these specimens are pierced by small holes arrayed around the plate centers (Pl. 4, fig. 7). Pierced plates were not found in other species of Reteocrinus, but those of R. alveolatus have scalloped margins so that gaps аге рге5- ent between plates. Extensive distribution of these pores about the interbasal, interbrachial, and tegmen regions suggests that the holes mark the sites of respiratory structures. Reteocrinus stellaris Billings Plate 4, figures 1-3 Reteocrinus stellaris Billings, 1859, p. 64, pl. 9, figs. 4a—c; Wachs- muth and Springer, 1882, pp. 262-263, fig. 1; Wachsmuth and Springer, 1897, р. 178, pl. 9, figs. За-с; Springer, 1911, p. 10, pl. 1, figs. 6, 7; Ubaghs, 1978a, p. T414, fig. 223, nos. la, b. Emended diagnosis.—A species of Reteocrinus with aboral cup high, convex-sided, slightly wider than high at level of RR, relatively short arms, wide interray areas; keels radiate from axes along limbs of IBB, BB, RR; distal RR through Brr ornamented with sublon- gitudinal fine ridges, no lateral projections; interinfra- basal depressions medium-sized (relative to other species of Reteocrinus), much higher than wide, open below; IBr, axial. Proximal column round, columnals thin with narrow rounded epifacets. Emended description.—Crown moderate to large averaging approximately 45 mm high among know? | | ——— — LEBANON LIMESTONE ECHINODERMS: GUENSBURG 29 Ў | Table 4.—Continued. Plate ornamentation | Width/height Keels on cup Nature of surficial Locations of lateral ) of IBB plates ornament projections on arms Column shape | w>h present longitudinal striations none Round w=h present reticulate and (or) thick RR-IIBrr (not al- Pentagonal discontinuous striations ways present) у> absent smooth none Round w>h absent smooth КК-ПВгг Round w=h absent smooth to nearly smooth RR-IIBrr (?) Round w>h absent reticulate, warty none = | w>h absent none or faintly reticulate distal ПВгг- Pentagonal \ arm tips | w>h present fine longitudinal IBrr-(?) ПВгг Subpentagonal striations (faint) à | w>h present reticulate IBrr-IIBrr Subpentagonal | wh absent reticulate RR-IIBrr or Round to sub- IBrr-IIBrr pentagonal wh present longitudinal striations at least RR (anals) == = = с a Specimens; cup high, convex-sided, slightly wider than high at level of radials; arms relatively short, interray areas relatively wide. Keels radiate from plate axes along limbs of RR, BB, IBB; abut at plate articulations. Sutures flush. RR through Brr ornamented with sub- longitudinal fine ridges, usually four or five per plate, ridges abut across plate articulations. Surface of prox- imal cup can have additional small surface irregular- Шев. Interinfrabasal depressions moderately large, deep, Much higher than wide, open below. Interbasal gaps large, subround. IBB large, up to two times wider than high. BB large, Usually about as high as wide but variable. RR large, about 1.5 times higher than wide. Interbrachial-inter- basal areas with depressed fields of small subround to Subpolygonal plates having coarse stellate ornament (РІ. 4, fig. 2). IBr; axillary; IBrr much taller than wide, becoming less so distally. IIBrr, , axillary, IIBrr higher than wide, becoming less so distally. Brr beyond IIBrr typically about as high as wide, variable. Arms branch at least four times in larger specimens. Proximal anals as much as five times higher than wide, becoming less 80 distally; distal anals unknown. Tegmen apparently Sac-like, not extending beyond arm tips; poorly known. Only proximal 3 cm of column known; round, not tapering. Columnals very thin; epifacets narrow, mar- 8115 rounded to attenuated; nodals separated by thin internode of several internodals, middle internodal is Priminternodal. Articulae cryptosymplectial, lumen Quinquelobate. Remarks.—See Table 4 for comparison with other Species of Reteocrinus. Types and occurrence.— All original type material Was available for study. GSC 1525 is here figured and designated the lectotype. GSC 1525b-e are designated paralectotypes and GSC 1525b is figured. GSC 1525f was not part of the hypodigm and is therefore desig- nated a topotype. The specimens are from the late Trenton Cobourg Beds, Ottawa Formation, at Ottawa, Ontario, Canada. Reteocrinus alveolatus Miller and Gurley Plate 4, figures 5—7, 9, 14, 15 Reteocrinus alveolatus Miller and Gurley, 1894, pp. 46—47, pl. 11, fig. 22; Springer, 1911, pp. 10, 11, pl. 1, figs. 1-5; Ubaghs, 19782, p. T414, fig. 223, nos. 1c, d. Remarks.— The original description was based on a single individual from Kentucky. This specimen is coarsely silicified and lacks most details of plate or- nament. Well-preserved material assignable to this species was later reported from Ontario (Springer, 1911). Additional Ontario specimens from approxi- mately the same horizon and at a locality near Spring- er's are in the University of Cincinnati collections. I examined the holotype and referred specimens from Ontario and found no significant differences among them. The following additions and corrections should be added to the largely accurate descriptions provided by Miller and Gurley (1894) and Springer (1911). The cup is high, with sides slightly convex in type specimen, slightly convex or possibly straight-sided in Canadian specimens. Interinfrabasal gaps are small, suboval; in- terbasal gaps are large and subcircular. IBB, BB, RR possess an interconnecting network of prominent keels radiating from plate centers; the keels are offset from limb axes. RR to IIIBrr typically have small to large wedge-shaped lateral projections. 30 BULLETIN 319 The following additions were observed in Canadian specimens only; their absence in the holotype probably results from coarse silicification. Crown plate orna- ment is variable; can be on RR and Brr only, it consists of few subparallel sublongitudinal fine ridges to many irregular, sinuous, sublongitudinal, anastomosing fine ridges; the ridges typically abut across plates (Pl. 4, fig. 5). Interinfrabasal, infrabasal, as well as interbrachial fields are filled with small, irregular, subpolygonal plates; the plates have coarse stellate ornament ridges and numerous pores arrayed about plate centers be- tween ridges (Pl. 4, figs. 7, 15). The column widens slightly near the crown; it is subpentagonal to round in small specimens and consists of nodals separated by an internode of six to eight columnals. Internodes typically have a medially disposed priminternodal flanked by secundinternodals. The column can also possess discontinuous longitudinal ridges along the pentagonal angles. Epifacets are narrow, margins at- tenuated, rounded, to nearly straight sided. Articulae are cryptosymplectial. Types and occurrence.—The holotype FM(UC)-6045 is here refigured; it is from the middle Trentonian (Kirkfieldian) Curdsville Limestone, Mercer County, Kentucky. Referred and figured specimens in the Kopf Collection at the University of Cincinnati are from the middle Trentonian (Kirkfieldian) Hull Limestone, Kirkfield Quarry, Victoria County, Ontario, Canada; these are UC-35991, UC-36514, and UC-36006. Oth- er referred specimens from the Hull Limestone near Kirkfield, Ontario, [USNM(S)] were not examined. (The Hull Limestone in the Kirkfield area is now re- defined as Bobcaygeon and Kirkfield limestones.) Reteocrinus elongatus Raymond Plate 4, figure 4 Reteocrinus elongatus Raymond, 1931, pp. 203-204, pl. IV, fig. 5. Remarks.—The following emendations are here made to Raymond’s brief but accurate description. In- terbasal gaps are large and interinfrabasal depressions are small and shallow. RR, IBrr, and IIBrr have small wedge-shaped lateral projections. [Raymond (1931, p. 203) referred to these as “sharp girders”.] The column is round, tapering gradually away from the crown. It is composed of thin alternating nodals and internodals; nodals variably sized; largest columnals are intermit- tently distributed along column (usually every fourth or fifth columnal). Nodal epifacets are narrow with rounded margins. The downward-projecting frills on proximal colum- nals inferred by Raymond were caused by differential weathering (Pl. 4, fig. 4) and are artifacts of preser- vation only. Types and occurrence. — The holotype and only spec- imen, here refigured, is MCZ 3370 [formerly 3994] and itis from 2.5 mi east of Cumberland Gap, Tennessee. Raymond lists the specimen as occurring in the Ce banon” Limestone, but this term is no longer applied to rocks east of Sequatchie Valley in Tennessee. The specimen could be from the Upper Ottosee Group, possibly the Witten or Wardell formations. Reteocrinus fenestratus new species Plate 2, figures 10-13, 18 Etymology of name. — fenestratus (L.) = window, and refers to the open nature of the cup in this species. Diagnosis.—A species of Reteocrinus characterized by medium-sized crown for genus with steep, nearly straight-sided, cone-shaped cup, smooth or faint retic- ulate ornamentation on cup plates; relatively large high IBB, small interinfrabasal gaps, small wedge-shaped lateral projections on RR, IBrr, ПВгг, and with low rounded hemispherical to small wedge-shaped lateral projections on proximal anals. Column differentiated into at least proximal and medial parts; proximal со- lumnals with narrow sharp epifacets, farther distal co- lumnals straight-sided or with narrow straight-sided epifacets. Description. —Moderate-sized crown for genus with steep-sided cone-shaped cup, slightly higher than wide at level of radials. Major cup plates slightly flattened, with smooth or faintly reticulate ornament. RR, IBrr, IIBrr with short lateral wedge-shaped projections, proximal anals with low hemispherical to small lateral wedge-shaped projections. Small typically subtrian- gular interinfrabasal gaps; interbasal gaps large. IBB slightly wider than high, forming relatively large part of sides as well as base of cup. BB largest plate in cup, slightly higher than wide. RR large, higher than wide. Anal plate 1 as much as four times higher than wide, succeeding lower anals becoming slightly higher than wide. IBrr, 4 axillary (Pl. 2, fig. 11); IBr, higher than wide, succeeding IBrr becoming only slightly higher than wide. IIBr, axillary, IIBrr slightly higher than wide. Arms beyond IIBrr unknown. iBrr very small, with irregular stellate ornamentation. Column ob- scurely (?)pentagonal or round, columnals thin, ер!- facets narrow or absent, known over proximal (to ap” proximately 20 mm below cup) and medial (at least 75 mm beyond proximal) portions. Proximal colum- nals with narrow attenuated epifacets; thicker, wider nodals alternate with two to three thinner narrow intet- nodals; internode can have priminternodal, secundin- ternodals. Medial column an irregular series, generally of two to three similar straight-sided internodals sep” arated by thicker nodals with gently convex or straight- LEBANON LIMESTONE ECHINODERMS: GUENSBURG 31 Sided epifacets (Pl. 2, fig. 18); nodals apparently consist of two to three ?ankylosed thin columnals, the median ?fused columnal often slightly wider, protruding. Ar- ticulae cryptosymplectial. Remarks. —See Table 4 for comparisons. Types and occurrence.—The holotype, UI X-5690, is from the lower Lebanon at locality Z-651. Paratypes UI X-5695 and UI X-5696 are from the lower Lebanon at localities Z-658 and Z-659, respectively. All speci- mens are figured, and UI X-5690 and UI X-5695 were measured. Reteocrinus mahlburgi (Kolata) Traskocrinus mahlburgi Kolata, 1975, pp. 46-47, text-fig. 14, pl. 8, figs. 1, 4-5. Remarks. —This species is well described except for the inference of the presence of interinfrabasal gaps (Kolata, 1975, text-fig. 14; remarks under Traskocri- nus, р. 46). The IBB are missing in the type and only known specimen. The lower margins of BB are de- Pressed, meeting medially at vertices; this morphology 15 typical of species of Reteocrinus that lack interin- frabasal gaps (R. polkin. sp., R. stellaris Billings, 1859, К. elongatus Raymond, 1931). In contrast, species pos- Sessing interinfrabasal gaps have rounded inverted notches along their lower medial margins (R. spinosus Kolata, 1975, R. rocktonensis Kolata, 1975, R. alveo- latus Miller and Gurley, 1894, R. fenestratus n. sp.). Types and occurrence. — The holotype, BMNH Pk- 48, is from the Blackriveran Grand Detour Formation, Northern Illinois. Reteocrinus polki new species Plate 2, figures 5, 17; Plate 3, figures 1-4, 11 Etymology of name.—The specific name honors James Knox Polk, eleventh President of the United States, whose home is near the collecting locality. _Diagnosis.—A species of Reteocrinus with convex- Sided aboral cup, twice as wide as high at level of RR, Primary cup plates smooth or very faintly reticulate, Slightly distended or evenly rounded, with narrow Shelves on latero-ventral margins adjoining interbasal 8aps, interbrachial areas; interinfrabasal depression; Mterbasal gaps large, typically wider than high, IBrr3,, axillary; Brr beyond ПВгг with prominent sharp wedge- Shaped lateral projections. Column pentagonal, colum- nals extremely thin, with cryptosymplectial articula- tions. Description. — Adult crown large (see Table 3); aboral Сир convex-sided, twice as wide as high at level of RR. ajor cup plates with large multilimbed extensions; *venly rounded or slightly distended, smooth or with faint reticulate ornament (PI. 3, figs. 1, 3). Brr can have faint longitudinal ridges on proximal and distal mar- gins. Sutures flush. Major cup plates with narrow shelves on ventro-lateral margins adjoining interbasal gaps, interbrachial areas. Interinfrabasal depressions small (Pl. 2, fig. 5), depressions shallow in small specimens (UI Х-5417), larger, deeper, subtriangular in large specimens (UI X-5790). Interbasal gaps large for genus, typically wider than high. Brr beyond proximal ШВгг with prominent dorso-ventrally flattened wedge-shaped lateral projections; projections diminish in size away from IIIBrr (Pl. 3, fig. 1). IBB five, at least two times wider than high. BB, RR largest plates in cup. RR five, about as high as wide. iBr, interbasal plates thin, small, irregularly subpo- lygonal, with thickened centers; arranged in loose mo- saic; ornamented with fine ridges emanating in irreg- ular stellate pattern from plate centers (Pl. 3, fig. 1). iBrr become thinner, more weakly calcified distally. IBrr,;, axillary; IBr, much higher than wide, IBr, ,, , only slightly higher than wide. IIBrr; , axillary. Brr beyond proximal ПВгг usually about as high as wide, variable; having prominent dorsoventrally-flattened wedge-shaped lateral projections (Pl. 3, fig. 1). Projec- tions diminish in size away from IIBrr. Arms branch isotomously at least five times in adults. Ambulacral grooves apparently wide. Coverplates small, appar- ently of median biseries of small wedge-shaped plates (ambulacrals) flanked on either side by a single row of plates (adambulacrals). Proximal anals resemble IBr, but can be much higher than wide. Tegmen sac-like, extending approximately to level of arm tips on one specimen (Pl. 3, fig. 1), covered by small, thin sub- polygonal to suboval plates confluent with iBrr; plates become thinner, more weakly calcified distally. Web- like extensions of tegmen extend to ШВгг. Column pentagonal, much longer than crown height, at least 14.8 cm in holotype UI X-5790; heteromor- phic, tapering slightly away from crown. Proximal and medial column of extremely thin columnals bearing very narrow rounded epifacets; alternating columnals with slightly more prominent epifacets; largest epifa- cets on columnals at variable intervals, generally every fourth or fifth columnal. Distal column of numerous tiny tightly fitting subequant plates forming mosaic (Pl. 3, fig. 11). Remarks. — Reteocrinus polki n. sp. is compared with other species of Reteocrinus in Table 4. Types and occurrence. — The holotype is UI X-5790; paratypes are UI X-5410, 5413, 5414, 5417; UI X-5413, 5417 and 5790 are figured and measured; UI X-5410 is figured. All specimens are from locality Z-654a. 32 BULLETIN 319 Reteocrinus spinosus Kolata Reteocrinus spinosus Kolata, 1975, pp. 45—46, text-fig. 13, pl. 8, figs. 38. Remarks.—Kolata’s description is accurate except for the inference of interinfrabasal gaps. Additional preparation of the holotype and only known specimen shows the specimen had interinfrabasal depressions. Types and occurrence.—The holotype, UI X-4881, is from the Blackriveran Grand Detour Formation, northern Illinois. Reteocrinus variabilicaulis new species Plate 2, figures 14-16, 20; Plate 3, figures 5-10, 13; Plate 4, figures 12, 13 Etymology of name.—variabilis (L.) = variable; caulis (L.) = tail, referring to the highly variable col- umn morphology of this species. Diagnosis.—A species of Reteocrinus characterized by a medium-sized (for genus) elongate crown with steep nearly straight to slightly convex-sided cup, cup slightly wider than high at level of RR, multilimbed extensions of cup plates relatively short. Small inter- infrabasal depressions, large rounded lateral projec- tions on RR to ПВгг, two to four IBrr per ray, and moderately large interbasal gaps. Column heteromor- phic, relatively short; proximal, medial column with large round to subpentagonal nodals separated by vari- able internode series; nodals typically of two or three ankylosed columnals. Distal column of pentameres ta- pers rapidly toward subconical, polyplated, encrusting holdfast. Description. —Crown moderate-sized (Table 3), slen- der. Aboral cup steeply conical, sides nearly straight to slightly convex, slightly wider than high at level of RR. Major cup plates relatively compact with short lateral multilimbed extensions, evenly rounded to slightly flattened, with ornament faint reticulate to smooth. ОЛ X-5409 with faint keels on limbs of CD В. UI X-5412 and UI X-5694 possess supernumerary major cup plates (Pl. 2, fig. 20; Pl. 4, fig. 13). RR-IIBrr possess large subhemispherical lateral projections (Pl. 3, figs. 8, 10); projection size decreases away from IBr, „2. Interinfrabasal depressions small, can be open below. Interbasal gaps medium-sized for genus, typi- cally taller than wide to about as tall as wide, variable. IBB medium-sized, approximately twice as wide as high; often with small low node projecting from me- dial-adoral surface toward interbasal gap. BB five, higher than wide, widest distally. CD B pentaxial, higher than wide. RR five, large, slightly higher than wide. Two to four IBrr per ray. IBr, , generally higher than wide, succeeding IBrr about as high as wide, variable. IIBr, typically axillary. Interbasal, iBr plates small, irregularly polygonal, with prominent ridges that gen- erally radiate in stellate pattern from near plate centers. Proximal anals higher than wide, slightly narrower than IBrr. Tegmen poorly known, apparently sac-like, not extending to arm tips. Proximal tegmen covered with slightly smaller plates than interbrachials, with less pronounced or no stellate ornamentation. Arms uni- serial, long, branching isotomously at least five times in bigger specimens. Column short, about two-and-one-half times crown height, weakly tripartite, heteromorphic, highly varied along length among specimens. Column slightly thick- er to subequal in thickness from crown to about middle column, then tapering rapidly to holdfast. Articulae pentagonal, cryptosymplectial; lumen quinquelobate. Proximal, medial column sections composed of in- tervals of several internodals separated by single no- dals; proximal columnals of both series more or less compressed. Nodals thick to thin; compound, typically of two to three ankylosed columnals; outline circular or obscurely pentagonal, with wide epifacets, straight to rounded margins (Pl. 3, figs. 5, 6). Straight-sided epifacets (Pl. 3, fig. 6) sculpted with irregular ridges, grooves encircling columnal; rounded epifacets smooth or pitted in distal medial column region. Internodes variable with five to eight columnals; columnals either low, subpentagonal throughout or approximate middle internodal is slightly (Pl. 3, figs. 5, 9) to greatly enlarged [i.e., nodiform (РІ. 3, fig. 9)]; can obscure adjacent low internodals. Low subpentagonal internodals typically having low, thin sharply attenuated epifacets. Distal column narrow, rapidly tapering, of projecting round- ed pentameres, circlets of which alternate slightly in size. Larger circlets replace nodals in sequence, smaller circlets replace internodals; circlets compressed dis- tally, thin. Holdfast encrusting, subconical, low, surface smooth; dorsally of tightly sutured (?fused) polygonal plates; ventral disc present. Remarks.—See Table 4 for comparison of R. var- iabilicaulis with other species of Reteocrinus. The an- kylosed columnals forming the nudinodals in R. var- iabilicaulis are an unusual feature among crinoids. Also of note in this species is the great individual variation in column morphology. UI X-5415 (PI. 2, fig. 16) is tentatively assigned to this species. It is a distorted crown and proximal col- umn with badly corroded arm tips differing from other specimens assigned to this species in having thinner more attenuated lateral projections on the IBrr and IIBrr. I cannot be certain of the column shape, but the LEBANON LIMESTONE ECHINODERMS: GUENSBURG 33 columnal morphology is similar to that of UI X-5408 (Pl. 2, fig. 15) and UI Х-5694 (PI. 4, fig. 13). Types and occurrence. —Six specimens are designat- ed types and are figured; the holotype, UI X-5411, and paratypes UI X-5408, 5409, 5412, 5805, and 5694, are all from the lower member, Lebanon Limestone at locality Z-654a. UI X-5409, 5411, 5412, and 5805 are measured. Referred, figured, and measured specimen UI X-5415 is from the lower member, Lebanon Lime- Stone, at locality Z-651. Reteocrinus species cf. R. variabilicaulis new species Plate 3, figure 12 Remarks.—A single small crown and proximal col- umn, UI X-5414, is tentatively assigned to R. varia- bilicaulis. It differs from other specimens ascribed to this species principally in possessing a relatively nar- Tower, more gradually tapering column; also columnals are thinner with narrower lower epifacets. I suspect the differences are due to ontogenetic and intraspecific Variation. Types and occurrence. — The single figured and mea- sured crown, UI X-5414, is from the lower member, Lebanon Limestone, at locality Z-654a, where it co- Occurs with both R. variabilicaulis and R. polki. Reteocrinus species Plate 2, figure 19 Remarks.—The single small partial crown with Proximalmost column herein described (see Table 3 for measurements) is most similar to R. fenestratus, but differs in lacking interinfrabasal gaps, possessing four or five IBrr instead of (three or four, and having more angular lateral projections on distal cup plates (Pl. 2, fig. 19). Interinfrabasal gaps could develop later in ontogeny similar to interinfrabasal depression de- velopment in R. polki n. sp. and R. variabilicaulis n. Sp. Small size supports this possibility. The other dif- ferences could be a result of intraspecific variation. The specimen is also similar to specimens assigned to R. variabilicaulis, but the former differs in possess- ing thin homeomorphic proximal columnals, four to five IBrr instead of two to four, and more angular, in Some cases subrectangular (Pl. 2, fig. 19) lateral pro- jections on distal cup plates. It is possible, again, that the differences are due to intraspecific and ontogenetic variation. Because of lack of information concerning ontoge- netic and intraspecific variation, therefore, the speci- men cannot be assigned although I suspect the speci- men belongs to either R. fenestratus n. sp. or R. variabilicaulis n. sp. Types and occurrence.—' This single specimen, UI X-5416, is figured and measured, and is from the lower member approximately 8 m below the massive mem- ber at locality Z-651. Family ANTHRACOCRINIDAE Strimple and Watkins, 1955 Genus GUSTABILICRINUS new genus Etymology of name.—gustabilis (L.) = appetizing; crinum (L.) = lily. The first element suggests one pos- sible cause for the rarity of intact calyxes in this genus, a condition that probably is the result of relatively fragile construction. Type species. — Gustabilicrinus plektanikaulos n. sp. Diagnosis. — Anthracocrinidae with subconical cup, slightly higher than wide, median ray ridges faint to absent; sutures moderately depressed to flush; plates flat to slightly convex. IBB, aboral portion of BB form deep cylinder-like basal invagination, tightly appressed over proximal column. BB highly reflexed, form at- tenuated lower cup margin. All interrays apparently of nearly equal width; lateral interray consists of large interradial followed by two moderate-sized iBrr,; CD interray poorly known, of small primanal followed by two rows with two plates per row. Fixed Brr bifurcate isotomously twice; four arms per ray. Free arms uni- serial, cuneate. Column heteromorphic, large, round, with slight gradual distal taper. Remarks. — Features which place Gustabilicrinus n. gen. in the Anthracocrinidae are the deep basal con- cavity with concealed IBB, iBrr separated from tegmen by fixed pinnules, fixed Brr and pinnules forming web- like extensions of the cup at the arm bases, and CD interray apparently equisized with lateral interrays. The new genus resembles Anthracocrinus Strimple and Watkins, 1955, but the latter differs in possessing 15 arms and a wide CD interray whose plates connect with the tegmen. Hercocrinus Hudson, 1907, and Deo- crinus Hudson, 1907, differ in possessing a globular cup and apparently 10 arms (Hudson, 1907, pp. 121, 125, pls. 8, 9); Hercocrinus also has biserial arms. Rheocrinus Haugh, 1979, differs most conspicuously in possessing biserial arms and BB and RR alternating in a “bicirclet”” along the lower edge of the cup. Gustabilicrinus plektanikaulos new species Plate 3, figures 1, 3; Plate 5, figures 2—5, 12, 13, 18, 20-22; Text-figure 5 Etymology of name.—plektanos (Gr.) = curly; kau- los (Gr.) = stem. This epithet refers to the commonly coiled column of the species. Diagnosis. — A species of Gustabilicrinus character- 34 BULLETIN 319 Table 5.—Measurements of specimens of two species of Gustabilicrinus n. gen. (in mm). G. plektanikaulos G. latomium UI X-5952 UI X-5943 UI X-5791 UI X-5962 UI X-5961 Cup height 24 187 — 6.5 11.4* Cup width 20° 11 — @ 27 - В height 3.8 3.8 3.1 Sel - 1.9 2.0 3.4 32) B width 4.3 845 Sei 3.6 — 1.4 175 3:2 DIO R height 3.8 4.0 4.1 305 2,6 2.4* 0.9 Ii DAI 9 R width 4.3 4.1 4.8 33 32 3.0 1.6 1.6 Sul 2.8 iR height 4.8 4.4 5.2, 2.8 2.9 U 1.6 1.8 2.8 2.8 iR width 4.2 4.6 4.2 3.0 3.1 9i 13 1.3 2.4* 2:2 Column diameter at base of cup 4.1* 559 — 1.9 43 * = estimated; multiple measurements are of several plates. ized by a moderate-sized to large, convex-sided cup about as high as wide; lateral interrays of relatively small iR followed by rows of two, then two or one ¡IBrr; ¡RR not contacting Brr above IBr,; IBrr slightly wider than high to wider than high; proximal column nodals with smooth, rounded latera. Description. — Dorsal cup medium to large (Table 5), subconical, slightly higher than wide, cup sides slightly convex; deep cylinder-like basal concavity; concavity extends upward into coelomic cavity approximately to level of IBr, or IBr,, appressed tightly against and concealing proximalmost column (Pl. 5, fig. 4). Cup plates flat to slightly convex, can have depressed cen- ters; smooth or with fine pits; sutures flush to mod- erately depressed, depressed particularly at triple junc- tions, between adjacent BB, RR. Median ray ridges faint, low, wide, occasionally absent; where present gradually disappearing distally at IBrr or IIBrr. Dorsal surfaces of fixed IIIBrr to proximal five or six IVBrr can have faint longitudinal abutting keels. IBB five, forming ventral half of basal concavity, large, taller than wide, pentagonal, sides diverge slight- ly distally away from column articular facet. BB five, articulate with two IBB, two BB, two RR, one iR; sharply reflexed medially, V- or Y-shaped in transverse section, form dorsal half of basal concavity, attenuated lower cup margin. RR five, pentagonal, large; each in contact with two BB, two iRR, IBr,. IBrr, smaller than RR, hexagonal, wider than high; rarely pentagonal, ax- illary. IIBrr, axillary, pentagonal, wider than tall, much smaller than RR. IIBr, wider than high, irregularly pentagonal to hexagonal, smaller than IBrr. IIBr, pentagonal, axillary; typically slightly smaller than IIBr,, can be nearly the same size, bears two arms; IIIBr, hexagonal; ПІВг, pentagonal or quadrangular; IIIBrr, axillary, bear a single fixed pinnule, one IVBr. IVBrr, ; non-pinnulate, IVBr, can also be non-pin- nulate. Succeeding Brr with single pinnules on alter- nating sides of arms. Two or three illBrr, aligned in vertical row, decreasing greatly in size distally. One to four iIIIBrr; aligned in vertical row. Interrays narrow, typically of eight to 10 plates; not reaching tegmen (Text-fig. 5). ЖЕ heptagonal, large, as large as or slightly larger than RR; followed by rows of two, then two or less commonly one subequal-sized iIBrr. One interray of UI X-5952 with unusually tall iR followed by row with two unequal ilBrr (Pl. 5, fig. 2), right iIBr of this row much taller than left iIBr, abuts iR along smaller facet than left ШвВг. One interray of UI X-5943 has first row above iR with right iIBr taller, wider than left iIBr (Pl. 5, fig. 12); right bordering ray with IBr, axillary. CD interray not defined; ap- Text-figure 5.— Gustabilicrinus plektanikaulos n. gen. and 5р. tracing of cup, anterior view, in holotype UI X-5952. Interray areas (IRR, iIBrr) are stippled (x3). | | | | | — - — — rn Á — LEBANON LIMESTONE ECHINODERMS: GUENSBURG 35 parently of approximately equal width to other inter- rays. Tegmen poorly known, of many small plates with single spinose projections. Arms 20, long, length greater than 1.5 times height of cup, tapering slightly proximally, more rapidly dis- tally; cuneate, uniserial, densely pinnulate, free above IVBr, ¿. Pinnules closely spaced, large, flattened, with gradual distal taper; approximately 10-12 pinnulars per pinnule. Column round, heteromorphic, long, greater than three times as long as crown height; distally tapering, commonly coiling (Pl. 5, fig. 21); of closely spaced thin nodals alternating with few thin internodals near crown; gradually changing to widely spaced nodals alternating with approximately four to six (most commonly five) internodals distally. Nodals with wide epifacets, rounded margins; radially arranged low ridges on up- рег, lower epifacet surfaces, do not reach margins (Pl. 5, fig. 22). Internodals with slightly convex, irregular latera. Articulations symplectial, crenularium narrow, lumen large, subpentagonal. Remarks.—See remarks under Gustabilicrinus la- tomium n. sp. for comparison with С. plektanikaulos п. sp. Mode of life. —Several distal columns of Gustabili- crinus plektanikaulos n. sp. were found coiled about columns of Tryssocrinus endotomitus n. gen. et sp., Reteocrinus polki п. sp., and Hybocrinus bilateralis n. Sp. Tryssocrinus endotomitus, Н. bilateralis, and prob- ably R. polki were attached to the substrate by а hold- fast. Gustabilicrinus plektanikaulos then, maintained its position by prehensile attachment to the columns Of crinoids that were fixed to Ше substrate. Types and occurrence.—The holotype is UI X-5952 and paratypes are UI X-5758, 5769 through 5773, 5791, 5792, 5802, 5811, 5940, 5943 through 5951, 5953 through 5960, 6048, 6049, 6051, 6054. UI X-5952, 5943, and 5791 are figured and measured; UI X-5758, 5773, 5792, 5951, and 5959 are figured. All Specimens are from the lower member of the Lebanon Limestone at locality Z-654a. Gustabilicrinus latomium new species Plate 4, figures 8, 10, 11, 16, 17; Plate 5, figures 1, 6 Etymology of name.—latomium (L.) = stone quar- Ty. This epithet refers to the collecting locality. Diagnosis.—A species of Gustabilicrinus with steep, Nearly straight-sided cup; IBrr much wider than tall, Wider to nearly as wide as interray areas; iRR relatively large, abut IIBrr,. Presumed CD interray of small primanal followed by rows in series of two, two plates. Column round; nodals serrated along epifacet margins. Description.—Cup small (Table 5), subconical, slightly higher than wide, cup sides straight, nearly vertical at level of BB, slightly convex, flaring above BB. Basal concavity deep, cylinder-like, extends into cup cavity to level of IBr,, appressed tightly against, concealing proximalmost column. Cup plates flat to slightly convex, ornamented with fine pits, ridges and grooves; ridges and grooves can abut across sutures. Sutures flush; indentation in BB at junction with lower edge of each R. Median ray ridges faint, low; more prominent distally on cup. IBB not observed. BB five, taller than wide, artic- ulate with two IBB(?), two BB, two RR, one iR; sharply reflexed medially, Y-shaped in transverse section, form lower half of basal concavity, attenuated lower cup margin. RR five, pentagonal, wider than high, large, in contact with two BB, two iRR, one IBr,. IBr, smaller than RR, much wider than high, subquadrangular to hexagonal; IBrr, axillary, pentagonal, much wider than high. ПВгг wider than high; ПВг, hexagonal, can be pentagonal; IIBr, axillary, pentagonal, bears two IIIBr,. IIIBr, pentagonal; IIIBr, axillary; each bears one fixed pinnule, one IVBr. IVBrr,, without fixed pinnules; succeeding Brr poorly known. Two to four iIIBr rows of single plate per row; ilIBrr decrease in size distally. One or two illIBrr between fixed ШВгг. Lateral interrays narrow, not contiguous with teg- men, of approximately three to five plates; iR large, in contact with one B, two RR, four IBrr, two IIBrr, two ог less commonly one iIBrr. Presumed CD interray known in UI X-5962; narrow, apparently not reaching tegmen (Pl. 5, fig. 1) small (?)primanal not extending above mid-level of IBr, followed by at least two rows of two plates. Tegmen poorly known, of small plates having single spine-like projections (Pl. 4, fig. 16). Arms poorly known, apparently 20, long; Brr wider than tall, cu- neate uniserial, one pinnule per Br; pinnulars taller than wide, laterally compressed. Column round, heteromorphic; proximal column of closely spaced nodals, distal column with nodals sep- arated more widely. Nodals with wide epifacets; epi- facets serrated, with numerous distally projecting short spinose processes (PI. 4, fig. 10). Articulae symplectial, lumen moderately large, round. Remarks. —Gustabilicrinus latomium n. sp. closely resembles Gustabilicrinus plektanikaulos n. sp., also from the Lebanon Limestone, but the latter can be distinguished by its relatively smaller iRR that do not abut IBr, or IIBr,, relatively taller IBrr (compare Pl. 4, fig. 17 with Pl. 5, fig. 20), and nodals with smooth 36 BULLETIN 319 rounded latera. G. latomium is also similar to Anthra- cocrinus primitivus Strimple and Watkins, 1955, from the Blackriveran Pooleville Member, Bromide For- mation, of Oklahoma but the latter has 15 arms instead of 20 and relatively wider interrays. Types and occurrence.—The holotype, UI X-5961, and a single paratype, UI X-5962, are the only known specimens. Both are figured and measured; they are from the lower member, Lebanon Limestone, at lo- cality Z-651. Family RHODOCRINITIDAE Bassler, 1938 Genus ARCHAEOCRINUS Wachsmuth and Springer, 1881 Type species. — Glyptocrinus lacunosus Billings, 1859, p. 261. Diagnosis (modified from Kolata, 1982, p. 181).— A rhodocrinitid crinoid with a high subconical cup, IBB small, hidden in basal concavity; BB large, form- ing lower half of basal concavity; lateral interrays of iR followed by two iBrr; CD interray widest, of prim- anal, followed by row of three ¡IBrr; arms 10, biserial branching isotomously several times; column large, round. Archaeocrinus snyderi new species Plate 5, figures 7-11, 14-17, 19; Plate 6, figures 3, 5, 6 Etymology of name.—The specific epithet honors Edward M. Snyder who assisted the writer with field work, offered suggestions, and who discovered some of the type material. Diagnosis.—A species of Archaeocrinus character- ized by a large, relatively high cup with generally smooth flush plates; faint median ray ridges, large basal con- cavity; column wide, long, nodals with smooth round- ed epifacets along proximal, medial column. Description. — Aboral cup large (Table 6), height ap- proximately equal to width, widest about level of IIBr,; cup sides moderately to slightly convex; circular in basal view; constricted slightly below level of free arm bases. Basal concavity large, funnel-shaped, of mod- erate depth, subpentagonal to subpentastellate in out- line, with faint to prominent surrounding rim, involv- ing IBB, BB. Median ray ridges low, narrow, faint, often discontinuous; extending from BB to conver- gence at RR; then extending distally along IBrr, di- verging at axillary IBrr, extending along IIBrr to bases of free arms. IBB five, small, pentagonal, generally wider than tall, restricted to basal concavity; proximal rim projects upward into body cavity for short distance, forms col- Table 6. — Measurements of four specimens of Archaeocrinus sny- deri n. sp. (in mm). UI X- UI X- UI X- UI X- 5732 5740 5743 5750 Cup height — 37.8 313" DSO Cup width - 37.6 SI 27:55 В height 8.3 ти 6.9 6.2 B width 10.8 9.2 8.3 7.8 R height 10.0 8.8 MR Да R width 10.7 9.1 yo» 8.2 iR height 10.8 9.7 US 7.6 iR width 11.4 10.0 8.0 7.5 Column diameter at base of cup 6.7 5:7 4.7 * — approximate (due to crushing). lar continuous with column lumen. Column articular facet forms narrow, crenulate shelf. BB five, heptagonal (B ray B can be octagonal) slightly wider than high; inflected medially forming lower cup margin; articu- lating with two IBB, two BB, two RR, one iR. RR five, A, B, Eray RR inverted pentagons, D ray R hexagonal, Cray R pentagonal or hexagonal; about as high as wide; typically articulating with two BB, two iRR, and one IBr. IBr, typically hexagonal, about as high as wide, nearly as large as RR. IBr, generally heptagonal, about as high as wide, smaller than IBr,, axillary. ПВгг be- come progressively smaller distally, IIBr, usually slightly taller than wide, hexagonal; followed at least by four fixed IIBrr. Second IIBrr of same ray separated by hexagonal Вг which is followed by rows of two, then three iIIBrr. iRR large, as large as RR, heptagonal, about as high as wide to slightly wider than high; fol- lowed by approximately five or six ШВг rows; row ar- ticulating iR usually with two plates, then followed by two rows with three plates each. iIBrr diminish in size distally. CD interray slightly wider than other inter- rays, with more numerous plates. Primanal large, usu- ally hexagonal. UI X-5740 with two roughly equisized pentagonal plates occupying primanal position, both abutting B below (Pl. 5, fig. 15). Primanal followed by row with three iIBrr, then row with three or four (rarely five) iIBrr (Pl. 6, fig. 5) and five or six additional rows with five to eight plates per row. Anal series not dis- tinct. Tegmen unknown. Arms poorly known; ten, long, slender, gradually tapering, uniserial, free above ПВг;, branching isoto- mously at least three times, with evenly rounded dorsal surfaces. Free IIBrr wedge-shaped, articulating along sides of arms with modified first pinnulars, which are incorporated, fixed, into primary arm structure (Pl. 5, fig. 9); fixed pinnulars give rise to free pinnulars along zig-zag suture pattern. Free IIIBrr and beyond unise- rial, slightly cuneate becoming non-cuneate, without | | | | | | | == < T. mo = | | LEBANON LIMESTONE ECHINODERMS: GUENSBURG 37 fixed first pinnulars. Free Br articular surfaces crenu- late. Pinnules begin with initial free ПВг, given off alternatively from arms one per Br. Pinnules long, del- icate, laterally compressed with attenuated aboral mar- gin; ambulacral groove narrow, covered by alternating biseries of tiny coverplates; coverplates meet medially along zigzag suture. Pinnulars taller than wide, many per pinnule. Column long, greater than 13.5 cm in SUI 46910, wide, heteromorphic; regions intergrade gradually. Proximal, medial column greater than 13.5 cm in length in SUI 46910, diameter subequal. Proximal column of thin closely-spaced nodals separated by four or five internodals; thinner, thicker nodals alternate along col- umn length (Pl. 5, figs. 8, 17). Medial column with equisized nodals widely separated by as many as 12 internodals (Pl. 5, figs. 7, 19). Distal column rapidly tapering, much narrower than proximal, medial re- gions, of relatively thick nodals separated by four to six internodals. Nodals thin, with wide epifacets, me- dial column nodals with rounded margins; thin prox- imal nodals with sharp, attenuated margins; distal col- umn nodals with low, evenly spaced distolaterally nodelike projections (Pl. 5, fig. 10). Internodals straight- Sided, without epifacets. Columnal articulations cren- ulate symplectial; lumen large, circular. Remarks.— Archaeocrinus snyderi n. sp. is similar to Archaeocrinus subovalis Strimple, 1953, from the Blackriveran Pooleville Member, Bromide Formation of Oklahoma (see Kolata, 1982, p. 183 for redescrip- tion of this species). The latter species differs slightly in having a relatively smaller basal concavity, cup sides with a greater expansion angle, relatively smaller BB, RR and iRR, fine pustules or radiating ridges on some lower cup plates, and proximal column nodals with Short, distally curving spines. Archaeocrinus buckhorn- ensis Kolata, 1982, from the Upper Mountain Lake Member, Bromide Formation of Oklahoma, has a sim- ilar high cup with low median ray ridges, and thin smooth flush cup plates, but differs most conspicuously in possessing a relatively much smaller basal concavity and smaller RR and BB. Types and occurrence.—The holotype UI X-5740, and paratypes UI X-5717 through 5739, and 5741 through 5749, are from the Sowerbyella - Diplograptus Zone of the upper member, Lebanon Limestone, at locality Z-654. Paratype UI X-5750 is from the lower member of the Lebanon Limestone at locality Z-651. Paratype SUI 56910 is from the upper member of the Lebanon Limestone approximately 3 m below the Car- ters Limestone at locality Z-662. UI X-5740, 5743, and 5750 are figured and measured; UI X-5737, 5739, 5741, 5742, 5744, and 5745 are figured; UI X-5732 is Measured. Rhodocrinitid species A Plate 6, figures 7, 10, 11 Remarks.—One poorly preserved partial crown, UI X-5752, represents this species. The aboral cup is ap- parently bowl-shaped with a basal concavity and low faint median ray ridges. Cup plates are smooth and thin; sutures are flush. IBB are unknown. BB are large and reflexed, indicating a basal concavity. RR are poor- ly preserved and apparently large, IBr, is axillary. In- terbrachial areas are moderately wide and reach the tegmen. Arms are free above ПВг;; ilIBrr are present. Free arms are biserial and branch isotomously a short distance above the cup (PI. 6, fig. 10). Brr are cuneate, wider than tall with medial transverse indentations and slightly impressed sutures (Pl. 6, fig. 7). The column is large with thin closely-spaced nodals having wide epi- facets and rounded margins. This specimen most closely resembles Archaeocrinus Wachsmuth and Springer, 1881, as far as can be ob- served. The Brr with medial transverse indentations and impressed sutures distinguish this form from the only known Lebanon archaeocrinid, Archaeocrinus snyderi n. sp. The specimen is too poorly preserved to allow generic and specific assignment. Types and occurrence.—The figured single speci- men, UI X-5752 is from the lower member, Lebanon Limestone, at locality Z-651. Genus DIABOLOCRINUS Wachsmuth and Springer, 1897 ?Diabolocrinus species A Plate 6, figures 1, 2, 4 Remarks.—This species is represented by four small crowns with proximal columns (Table 7). The calyx is globular. The cup is bowl-shaped, about as wide as high to slightly wider than high with small basal con- cavity. Cup plates are smooth in most specimens, but may have faint radiating ridges in UI X-5794, sutures are flush. Median ray ridges are faint; interray areas are slightly depressed. IBB, if present, are very small. Five large, hexagonal BB are about as high as wide, and articulate with two BB, two RR, one iR and possibly one IB; BB are in- folded, forming the basal concavity. The basal con- cavity rim is slightly thickened. The five RR are large, pentagonal, typically wider than tall in larger speci- mens and about as wide as tall in smaller specimens and some larger specimens. RR articulate with two BB, two КК, and one IBr,. RR through IBrr decrease in size distally. IBr, is hexagonal, wider than tall, and smaller than RR. IBr, is pentagonal, wider than tall, 38 BULLETIN 319 axillary, and smaller than IBr,. Lateral interrays reach the tegmen; they consist of a large hexagonal to hep- tagonal interradial followed by rows of two, and then two or three ilBrr. The CD interray is known in UI X-5853; it is continuous with tegmen, wider than lat- eral interrays (PI. 6, fig. 4). The primanal is taller than wide, hexagonal, followed by small plates arranged in rows of three, then five plates. The tegmen is unknown. Ten arms, with length twice cup height, are cuneate, uniserial, and pinnulate. Arms are free above IBr, or IIBr,. Brr are wider than tall in larger specimens and about as wide as tall in smaller specimens. Pinnules are long, one per Br and laterally compressed; pinnules begin on the interradial side of ПВг,. Pinnules are denser, longer in larger specimens. The column is round, heteromorphic and distally tapering; proximally, the column consists of thick no- dals with wide epifacets and convex margins closely spaced between a variable internode of few thin narrow internodals. Columnals in the distal column of UI X- 5767 are taller than wide, straight-sided and cylinder- like. IBB were not seen in any of the specimens; however, they are inferred to have been present because: 1) the RR are separated all around by iRR; 2) the BB have straight lower facets suggesting articulation with another circlet of plates; and 3) column articular facets on the BB are apparently absent. These specimens are provisionally assigned to Dia- bolocrinus. They resemble other known species of Dia- bolocrinus in having a similar basal concavity, basic cup plate arrangement, and arm number. They differ in possessing a relatively higher cup, few if any sup- plementary plates in the interrays, non-protuberant arm bases, relatively thinner arms, and a smaller known crown size. Among Diabolocrinus species, the forms described here are most like D. oklahomensis Kolata, 1982, from the Pooleville Member, Bromide Forma- tion, Oklahoma. Both forms have simple interrays of an iR followed by two iBrr, small crown size, cuneate uniserial arms, and no or faint median ray ridges. The Oklahoma species differs in having cup plate ornament of sinuous ridges, a relatively wider cup, and thicker arms with protuberant arm bases. The specimens assigned here are immature, or a pae- domorphic species that did not significantly exceed in size the largest preserved specimen (Table 7). Sug- gesting early ontogenetic stages are: small size; IBr, and ПВг, are the last fixed Brr; relatively large RR, and few iBrr (see Brower, Lane, and Rasmussen, 1978, pp. T245, T253). If the specimens are immature, they have not developed many of the potentially diagnostic features found in adults. However, the relatively sim- Table 7.—Measurements of three specimens of ?Diabolocrinus sp. A (in mm). UI X-5853 01 X-5767 UI X-5794 Dorsal cup height 3:85 27 3.3 Dorsal cup width 4.1* AH - В height 1.6 0.9 0.9 В width 7 0.9 1.3 К height 1.3 0.7 1.1 R width 1.9 0.8 19 Агт 1епрїһ 10.0% 4.9 OR Column diameter at base of cup 09 97 (E * — estimated. ple cup plate arrangement and arm number of Dia- bolocrinus does not differ significantly from that seen in these specimens and this genus could have given rise to a paedomorphic form such as this species. Be- cause of the above uncertainties, these specimens are not assigned to species and only provisionally assigned to genus. Types and occurrence. — Four specimens were avail- able: UI X-5767, 5794, 5795, and 5853. UI X-5767 and 5853 are measured and figured; UI X-5794 is mea- sured. All specimens are from the lower member, Leb- anon Limestone, at locality Z-654a. Family unknown Rhodocrinitacid species A Plate 6, figures 8, 9 Remarks.—'This species is represented by one ex- tensively corroded calyx, UI X-5753, and a cup plate, UI X-5754. The cup is large, bowl-shaped, probably slightly wider than high (?), widest at the level of IIBr;. Median ray ridges are prominent and narrow. Low broad ridges radiate from centers of distal iRR and ilIBrr; ridges abut across plates. Plates are flat to slight- ly convex and ornamented with very fine parallel ridges and small pustules (Pl. 6, fig. 9); ridges are perpendic- ular to and abut across plate boundaries; pustules can also be arranged in rows that are perpendicular to and abut across plate boundaries. Plate sutures are flush. IBB and BB are unknown. RR are large, apparently pentagonal; the median ray ridges on the RR form an inverted Y shape. RR are separated by iRR. IBrr are taller than wide; IBr, is hexagonal; IBr, is axillary. There are approximately four fixed IIBrr. IIBr, has а small ridge bifurcating off the larger ray ridge; the small ridge then passes upward toward the tegmen. IIBrr are apparently separated by iIIBrr. Interrays are wide and reach the tegmen. They consist of a large octagonal iR followed by ? three iIBrr. The tegmen is highly arched and acuminate with a distal (?)anal opening. Convex LEBANON LIMESTONE ECHINODERMS: GUENSBURG 39 Table 8.—Measurements of three specimens of Abludoglyptocrinus gregatus n. sp. (in mm). UI X-5964 UIX-5969 UI X-5998 Cup height E75 16.5 — Cup width 16.5* 14* - В height ASA 1.6 1.5 B width 3.0 2.6 DES R height 3.4 2.8 В R width 4.9 4.4 4.7 Column diameter at base of cup S: 2.4 2:07 * — estimated. lobes extend from the sides of the tegmen toward the free arms. This species is assumed to be dicyclic because RR are separated by iRR. This inference, the strong ray ridges, and interrays contacting the tegmen all indicate this species is a rhodocrinitacid. It resembles Parar- chaeocrinus Strimple and Watkins, 1955, of the Rho- docrinitidae, in having a row of three iIBrr following the iR, prominent ray ridges, and a small secondary ridge branching from the primary ray ridge on IIBr). The aboral cup differs, however, in being higher than in any known species of Pararchaeocrinus and having different cup plate ornament. Insufficient material pre- cludes generic and specific assignment. Types and occurrence. — UI X-5753 is figured and from the upper member, Lebanon Limestone, at lo- cality Z-653; UI X-5754 is from the Sowerbyella - Diplograptus Zone of the upper member, Lebanon Limestone, at locality Z-654. Order MONOBATHRIDA Moore and Laudon, 1943 Family GLYPTOCRINIDAE Zittel, 1879 Genus ABLUDOGLYPTOCRINUS Kolata, 1982 Type species. — Glyptocrinus charltoni Kolata, 1975, p. 49. Emended diagnosis. — А glyptocrinid with 10 uni- Serial arms; arms free above IIBrr, 4, median ray, anal ridges, present; cup ornamented with small pustules and ridges or unornamented, smooth. Remarks.— The original generic diagnosis is here al- tered slightly to encompass the new species. At the time of the original diagnosis, only species with free arms above IIBrr, ; were known; the free arms in A. ®atus n. sp. arise above ШВг, 6. Abludoglyptocrinus gregatus new species Plate 6, figures 12-18 Etymology of name.—gregatus (L.) ^ to collect, or assemble. This refers to the occurrence of the hypo- digm in a single “garden,” ог pod. Diagnosis. — A species of Abludoglyptocrinus char- acterized by a bowl-shaped cup, slightly higher than wide; median ray, anal ridges low, rounded; BB small, much wider than tall; lateral interrays narrow, iR fol- lowed by rows of two, and one or two ШВгг. Description. — Cup moderate-sized (Table 8), bowl- shaped, slightly higher than wide to nearly as wide as high. Cup plates flat, can be slightly convex, smooth flush sutures. Median ray ridges low, evenly rounded, become more prominent distally. BB five, small, much wider than high. Sharp rim projects laterally from lower margin of each B; rims form flat subcircular surface lateral to column articular facet (Pl. 6, figs. 13, 18). RR five, largest plates in cup, wider than tall, heptagonal, contacting two BB, two RR, two iRR, one IBr. IBr, hexagonal or pentagonal, wider than high. IBr, axillary, wider than high to about as high as wide, pentagonal or hexagonal. IIBr, through ПВГ; в fixed, wider than high; HBr, with fixed pinnule branching off interray side; small ridge runs longitu- dinally, medially along each fixed pinnule, ridge abuts median ray ridge at IIBr,. ilIBrr small, arrangement variable: can be in rows of: one, two, and two; one, one, and two; one, one, and one; two, one, and two plates. Lateral interrays flat to slightly depressed, rel- atively narrow (Pl. 6, figs. 13, 18), of 10 to 12 plates at least; iR large; followed by rows of two or rarely one, then two or less commonly опе iIBrr. CD interray wide, median anal ridge low, wide, of six or seven plates (including primanal) in longitudinal series, grad- ually diminishing in size distally, extending to tegmen. Ridge bifurcates aborally, extends faintly onto C,D ray RR. Primanal large, heptagonal to hexagonal; plates above primanal arranged in rows of three or four, four, and then six plates. Tegmen poorly known, of many small subpolygonal convex plates. Arms ten, longer than cup height, uniserial, densely pinnulate (Pl. 6, fig. 16). Brr evenly rounded, cuneate, much wider than high; articulation surfaces between Brr crenulate. Pinnules slender; pinnulars taller than wide, compressed. Column round, long, heteromorphic. Proximal col- umn of thin nodals closely spaced between thin inter- node; nodals with wide epifacets, attenuated margins; internode of few thin straight-sided secundinternodals subequally disposed on either side of single primin- ternodal (Pl. 6, fig. 15); priminternodals resemble no- dals but thinner with narrower epifacets; internode can be of few straight-sided internodals only. More distal column of thin nodals widely separated by eight to 17 thin internodals. Nodals with wide epifacets; margins rounded with numerous low distally (?) curving low projections (Pl. 6, fig. 17). Internodals with straight 40 BULLETIN 319 sides, no epifacets. Articulations symplectial; distal lu- men pentastellate; angles rounded. Remarks.— Abludoglyptocrinus gregatus n. sp. is similar to A. charltoni (Kolata, 1975), from the Black- riveran Platteville Group of northern Illinois. The lat- ter species differs in possessing a slightly wider, lower cup, wider lateral interrays; and in having lateral in- terrays arranged in one, two, three plates per row pat- tern instead of a one, two, two or one, one, two pattern. The two species are remarkably similar in all other respects. A. ornatus (Billings, 1857) from the Trenton Ottawa Formation at Ottawa and Kirkfield, Ontario, differs in possessing fine ridges on cup plates; the ridges ra- diate from plate centers in stellate patterns. Abludo- glyptocrinus insperatus (Rowley, 1904) from the lower Silurian Edgewood Limestone, Pike County, Missouri, differs in having larger, taller BB with elevated Y-shaped ray ridges and the iRR have a single centrally-located pustule. А. pustulosus (Kolata, 1975) from the Platte- ville Group, northern Illinois, has a narrow conical cup and prominent pustulose plate surfaces in areas between prominent median ray ridges. A. laticostatus Kolata, 1982, from the Pooleville Member, Bromide Formation, Oklahoma, is also similar to the new species but differs most conspicuously in possessing highly el- evated median ray ridges and pustulose ornamentation in depressed areas between ridges. Types and occurrence. — The holotype, UI X-5964, and paratypes UI X-5963, 5965 through 5973, and 5994 through 6003 are from the upper member, Leba- non Limestone, at locality Z-653. UI X-5964, 5969, and 5998 are figured and measured; UI X-5995 through 5997 are figured. Columnals probably referable to this species occur at many localities and horizons in the Lebanon Limestone. Subclass INADUNATA Wachsmuth and Springer, 1885 Order DISPARIDA Moore and Laudon, 1943 Remarks. — The classification of the Disparida used in the Treatise on Invertebrate Paleontology (Moore et al., 1978) and by most modern workers is that of Moore (1962b), who divided this diverse group into super- families based on planes of symmetry within the cup. Symmetry planes are determined using the number and position of biradials. Recently, Frest, Strimple, and McGinnis (1979, p. 412) criticized this approach. They cite examples among the Disparida of 1) a genus CAbyssocrinus Strimple, 1963) having variable num- bers of biradials, 2) lineages, based on other characters, such as cup shape, that apparently lose infraradials (Apodasmocrinus Warn and Strimple, 1977 - Difficil- icrinus Frest, Strimple, and McGinnis, 1979), and 3) genera that have simple RR, which are shown to be comprised of fused biradials because the plates retain signs of sutures. Frest, Strimple, and McGinnis also incorporate Difficilicrinus into the Homocrinacea even though the genus has compound RR in the B and E rays only, rather than in B, C, and E rays, as typical of homocrinaceans (sensu Moore, 1962b). Tryssocri- nus n. gen. from the Lebanon supports the views of Frest, Strimple, and McGinnis, in that this genus closely resembles Daedalocrinus Ulrich, 1925 (Trentonian, Ontario) in cup shape, and column and arm mor- phology but differs in the number and position of bi- radials. (Tryssocrinus has biradials in the C and E rays; Daedalocrinus has them in the B, C, and E rays.) In accordance with the concepts of the Treatise, the two genera would be placed in different superfamilies even though they are similar in other characters. Position and number of compound radials, therefore, appears to be a more variable feature than previously thought and as a result the superfamilial classification based essentially on this feature alone apparently does not reflect natural groupings of the disparids. Revision of the Disparida is needed and suprageneric classification should incorporate such other characters as cup shape, arm and column morphology as well as number and position of biradials. Such a revision, however, de- mands extensive research that goes far beyond the scope of this study and therefore the Treatise classification has been used here for the Lebanon disparids. Superfamily HOMOCRINACEA Kirk, 1914 Family HOMOCRINIDAE Kirk, 1914 Genus APODASMOCRINUS Warn and Strimple, 1977 Type species.—Apodasmocrinus daubei Warn and Strimple, 1977. Apodasmocrinus species cf. A. daubei Warn and Strimple, 1977 Plate 7, figures 6, 14, 15 Apodasmocrinus daubei Warn and Strimple, 1977, pp. 93-96, text- figs. 21, 22а-с; Warn, 1982, pp. 78—81, pl. 4, figs. 13-30. Remarks. — A single well-preserved crown, with col- umn attached and largely buried in a hard matrix is referred to this species. The Lebanon specimen is sim- ilar to the type and referred specimens from the middle and upper Mountain Lake Member, Bromide For- mation, Oklahoma (see Warn, 1982). It differs from most Oklahoma specimens in possessing shorter RR. Oklahoma specimens are described as having simple RR approximately 1.25 times taller than wide (Warn, | | | | | ¿(REE - —— "au" WM > LEBANON LIMESTONE ECHINODERMS: GUENSBURG 41 1982, p. 80) whereas the D ray simple R in the Lebanon Limestone specimen is approximately 1.25 times wid- er than tall. This feature is variable among Oklahoma specimens, however, and Warn (1982, pl. 4, figs. 15— 16) figures a specimen with approximately the same simple R width to height ratio (1.25) as the Lebanon specimen. The Lebanon Limestone specimen appar- ently also differs from Oklahoma specimens in pos- sessing a subround rather than subpentagonal proxi- malmost column. As far as can be compared, the Lebanon specimen otherwise closely resembles those from Oklahoma. These differences are considered in- sufficient for erection of a new species. The Lebanon Limestone specimen possesses a col- umn known over a greater length than other specimens of A. daubei; 7.5 cm is preserved, and it appears to be nearly complete (Pl. 7, fig. 14). The column is hetero- morphic and gradually changing. It is of moderate length, greater than 2.5 times the approximate crown height; wide, about two-thirds as wide as the cup proxi- mally; tapering gradually away from the second nodal below the cup. The columnals are apparently solid Proximally within 1 cm of the crown, divided into pentameres from | to 6.5 cm beyond the crown, and apparently solid in the remaining distal 1 cm of pre- served column. The medial column from approxi- mately 3 to 5.5 cm below the crown has longitudinal Striations adjacent to and symmetrically disposed across lateral pentamere articulation surfaces: there are two Striations per articulation surface proximally and in- creasing to four distally. Within the first cm from the crown, nodals are subround to subpentagonal with wide rounded latera. They gradually become rounded, pen- talobate from approximately 1.5 to 6 cm below the crown with lateral pentamere sutures in the depressed regions between the lobes. Lobes become lower, less protruding distally. Internodes vary in thickness from much thinner than adjacent nodals near crown, to slightly thicker than adjacent nodals in the medial col- umn region, to slightly thinner than adjacent nodals in the distal column region. There are typically three internodals per internode in the medial column and these consist of two pentagonal, nearly straight-sided secundinternodals separated by a thicker wider sub- Pentalobate periminternodal. Distal internodes are composed of a single convex internodal. The distal- most 1 cm of column as preserved is composed of Subequal-sized and -shaped pentagonal columnals that are slightly convex-sided and have slightly impressed Sutures between adjacent columnals. Articulae appear to be cryptosymplectial. No holdfast is visible. Measurements (in mm).—Cup height = 4.6; cup width = 4.5; DE B height = 1.5; DE B width = 2.1; DR height = 2.2; DR width = 2.8; EiR height = 1.2; E iR width = 2.4; E superradial height = 2.0; Е super- radial width = 2.3; column diameter at base of cup = 2% Types and occurrence.—The single figured апа теа- sured specimen, UI X-5714, is from the Sowerbyella - Diplograptus Zone, upper member, Lebanon Lime- stone, locality Z-652. Superfamily CALCEOCRINACEA Meek and Worthen, 1868 Family CALCEOCRINIDAE Meek and Worthen, 1868 Genus CREMACRINUS Ulrich, 1886 Type species. — Cremacrinus punctatus Ulrich, 1886, pp. 106-107, fig. 1. Cremacrinus species cf. C. punctatus Ulrich Plate 7, figures 1, 2,7 Cremacrinus punctatus Ulrich, 1886, pp. 106-107, fig. 1; Sardeson, 1928, p. 35, pl. 1, figs. 6-8; Moore, 1962a, pl. 21, figs, 3a, b; Brower and Veinus, 1978, pp. 467-470, pl. 18, figs. 1—10. Remarks.— Two specimens are assigned to this species, UI X-5891 and UI X-5892. UI X-5892 (Pl. 7, figs. 1, 2) is a crown with distal arms missing; UI X-5891 (Pl. 7, fig. 7) is a crown and proximal column largely buried in hard matrix. UI X-5891 differs from UI X-5892 in having more tumid cup plates, and a thicker E ray; the two specimens are otherwise similar insofar as can be compared. Kolata (1975, pp. 21, 23) has noted considerable intraspecific variation among specimens of Cremacrinus arctus Sardeson, 1928, from the Platteville Group, northern Illinois. Both Kolata (1975, pp. 21, 23) and Brower and Veinus (1978, p. 459) studied ontogeny in Cremacrinus arctus from northern Illinois and the Minneapolis-St. Paul area of Minnesota, respectively. They found that cup and arm plates in this species become relatively thicker, more tumid in older, larger specimens. Because of these stud- ies, it is here considered that the differences between the Lebanon specimens are probably due to intraspe- cific and (or) ontogenetic variation. It is possible, how- ever, that the specimens belong to two closely related species. UI X-5892 differs in no significant way from spec- imens assigned to Cremacrinus punctatus Ulrich, 1886, from the early Trentonian Decorah Shale, Minneap- olis, Minnesota, and Escanaba, Michigan whereas UI X-5891 is also similar except for the unusually thick E ray (Pl. 7, fig. 7). Cremacrinus sp. described by Ко- lata (1975, p. 24) from the Blackriveran Grand Detour Formation, northern Illinois, closely resembles UI 42 BULLETIN 319 X-5891 in possessing a thick E ray. The Illinois spec- imen differs slightly from both Lebanon specimens in possessing thicker arms other than in the E ray as well as slightly coarser punctate ornament. Kolata noted the resemblance of his specimen to C. punctatus. Types and occurrence. — UI X-5891 is from the low- er member, Lebanon Limestone, at locality Z-652 and UI X-5892 is from the (upper member of the Leb- anon Limestone at locality Z-651. Both specimens are figured. Superfamily CINCINNATICRINACEA Warn and Strimple, 1977 Remarks.— The description and diagnosis of the su- perfamily by Warn and Strimple (1977, p. 28) is ac- cepted as given except for additions made necessary by new forms described herein. Radials are typically of regular pentagonal shape with one vertex pointed downward, but can be other shapes such as asymmet- пса! hexagons, heptagons (as in, e.g., Tryssocrinus n. gen.). Branching of the arms beyond the initial isoto- mous bifurcation can be isotomous, alternating het- erotomous, or endotomous heterotomous (Tryssocri- nus n. gen.). Modes of life.—Two Lebanon Limestone cincinna- ticrinaceans, Tryssocrinus endotomitus n. gen. et sp. and Columbicrinus crassus Ulrich, 1925, are known from complete composite reconstructions. They had similar life modes that differ significantly from that postulated for Cincinnaticrinus Warn and Strimple, 1977 (Warn and Strimple, 1977, p. 53). The latter genus apparently broke free (autotomized) from an en- crusting discoidal type holdfast and as an adult was free living, trailing a long column as a stabilizer. Trys- socrinus and Columbicrinus as adults had long columns (approximately 15 to 20 cm in Tryssocrinus endoto- mitus and approximately 40 to 50 cm in Columbicrinus crassus) that were attached by a dorsally multiplated holdfast (“Podolithus” type of Sardeson, 1908, and type 1C of Lewis, 1982). The proximal eight- to nine- tenths of the column in these forms is composed of solid columnals or pentamere circlets that are tightly articulated and are typically preserved intact with little or no curvature. The articulae are cryptosymplectial and the lumen is small. The distalmost column in these species is composed of five relatively loosely-articu- lated vertical series of plates; the plates alternate with laterally adjacent plates forming zigzag sutures; the lu- men is large, the articulae are cryptosymplectial. The distal columns were often disarticulated or broken be- fore burial and typically show more curvature than the proximal columns. The evidence suggests, therefore, that orientation of the crown in Tryssocrinus and Co- lumbicrinus was accomplished mainly through flexure of the distal column and that the long semi-rigid prox- imal column section enabled the crown to reach higher feeding levels. Family CINCINNATICRINIDAE Warn and Strimple, 1977 Subfamily CINCINNATICRININAE Warn and Strimple, 1977 Genus DOLIOCRINUS Warn, 1982 Type species. — Doliocrinus pustulatus Warn, 1982. Emended diagnosis. — Cincinnaticrininae with a rel- atively broad low conical to subconical nearly straight- to slightly convex-sided cup; cup plates tumid to slight- ly convex, smooth or with coarse pustulose ornament, sutures impressed to flush; AB ray B symmetrical pen- tagonal, other BB typically slightly asymmetrical pen- tagonal; compound RR relatively short, about as tall as wide to slightly taller than wide; small iBrr between IBrr,, proximal AB iR, abuts both A and B ray RR along short sutures; arms non-pinnulate, isotomously branching. IBrr, axillary. Remarks. — The generic diagnosis provided by Warn (1982) is emended to accommodate the new species described here and to more fully differentiate the genus. Doliocrinus closely resembles Isotomocrinus Ulrich, 1925, in the nature of the primary cup plate arrange- ment and arm branching pattern. The former differs, however, in possessing a relatively wider cup and shorter compound RR, in having small iBrr (particu- larly prominent in the AB interray), and in having the first arm bifurcation on IBrr, (in D. monilicaulis, at least) instead of IBrr, ,, в. Doliocrinus monilicaulis new species Plate 7, figures 8, 9; Text-figure 6 Etymology of name.— monilis (L.) — necklace, string of pearls, caulis (L.) = stem; the specific epithet refers to the nature of the column in this species. Diagnosis. — A species of Doliocrinus characterized by a moderate-sized crown with smooth, flush cup plates; round proximal column with smooth colum- nals. Description. — Crown moderate-sized (see Measure- ments); cup steeply conical but with rapidly diverging nearly straight sides, about as high as wide, widest distally. Cup plates smooth, slightly convex, sutures flush. BB five, subequal-sized, pentagonal, about as high as wide to slightly higher than wide; AB interray B symmetrical, other BB slightly asymmetrical because of slightly larger facets where abutting biradials. RR | | | | — —MÀ C —— ——" СР Se LEBANON LIMESTONE ECHINODERMS: GUENSBURG 43 five, two biradials (C, E rays), three simple (A, B, D rays). Radial facets deeply arcuate, not as wide as RR. Biradials much taller than simple RR, extend nearly to top of IBr, in rays with simple RR, sides notched for insertion of vertices from simple RR. Simple RR large, about as tall as wide, asymmetrical heptagons; A, B ray RR mutually abut across straight suture. E ray iR inverted symmetrical pentagon, slightly taller than E ray sR; E ray sR obscure hexagon, slightly wider than E ray iR. C ray biradial only partly exposed, slight- ly shorter than E ray biradial; iR shorter than sR, also Shorter than E ray iR. Small polygonal iBrr fill V-shaped gaps between IBrr,, sRR; extend farthest proximally in AB interray where hexagonal iBr fills notch between distal A, B ray RR, then followed by two irregular polygonal plates (Pl. 7, fig. 9). Tegmen Obscured, apparently highly vaulted. Anal series un- known; anal tube apparently of irregular, small polyg- onal plates. Arms long, approximately four times as long as cup height, slender, non-pinnulate, branching isotomously four to five times. Brr with evenly rounded dorsal sur- faces; Brr beyond IBrr can be ornamented with faint, irregular, sublongitudinal ridges; IIBrr, and ПВгг, pairs appressed; some IBrr, IIBrr with narrow ventrolateral Shelves, shelves can abut across Brr. IBrr widest at Proximal IBrr,, tapering to IBrr,, then expand again at axillary IBrr;; IBr, fixed, A, E ray IBrr, wider than high, B ray IBr, about as high as wide; IBrr, typi- A Text-figure 6.— Doliocrinus monilicaulis n. sp., plate diagram of Сир, showing arm branching pattern. Dashed lines indicate not ex- Posed; from holotype UI X-5716, x2.5. cally slightly wider than high, variable. IIBr, ,, 4 axil- lary; ПВгг approximately as tall as wide, variable. IIIBrr, „5 axillary; IIIBrr taller than wide, variable. IVBrr, VBrr typically taller than wide; IVBrr numer- ous along each arm branch. Proximal 4.5 cm of column known; round; distally expanding. Columnals smooth; single nodals alternate with single much thinner, narrower internodals; ex- ternal expression of point of abutment between nodals and internodals smooth curve (without distinct topo- graphic break) (Pl. 7, fig. 8). Nodals thick, distally en- larged; proximally with low, rounded, evenly curved latera; distally becoming bulbous, bead-like, with wide evenly curved latera. Distalmost two nodals with faint longitudinal indentations indicating ?fused penta- meres. Internodals thin, straight to slightly concave- sided, nearly as wide as nodals proximally, becoming much narrower than nodals distally, forming narrow necks between bulbous nodals. Lumen small, possibly pentagonal. Distal column, holdfast unknown. Remarks. — Doliocrinus monilicaulis n. sp. has a similar cup plate arrangement to D. pustulatus Warn, 1982, from the Lower Echinoderm Zone, Mountain Lake Member, Bromide Formation, Oklahoma. The Oklahoma species is easily distinguished, however, by its coarse pustulose plate ornament, impressed cup su- tures, and non-bulbous pustular columnals, and rela- tively shorter simple RR. Types and occurrence. — The figured and measured holotype and only specimen, UI X-5716 is from the lower member of the Lebanon Limestone at locality Z-654a. Measurements (in mm). — Height is first, width sec- ond for each pair of plate measurements. Crown length = 23.8*; cup height = 4.5; cup width = 5.2*. BB: АВ = 1.7, 1.7; ВС = 1.7, ; EA = 1.7, 1.5. RR: А = 2.1, 2.1; B = 1.9, 2.1; C sR = 1.4, -; E iR = 1.6, 1.4; E sR = 1.5, 1.8. Column diameter at base of cup = 2.3 [* = estimated]. Genus COLUMBICRINUS Ulrich, 1925 Type species. — Columbicrinus crassus Ulrich, 1925, p. 91, fig. 8. Emended diagnosis. — Cincinnaticrininae with ten long pinnulate arms, Brr beyond bifurcation uniserial, cuneate. BB nearly as tall as or taller than undivided RR, obscurely hexagonal in side view. IBrr much shorter than RR. Remarks. — Ulrich (1925, p. 91, fig. 8) described Со/- umbicrinus from a single cup and proximal arms through IBrr and parts of IIBrr in A and B rays (Pl. 7, figs. 3, 4). The only locality and horizon data avail- able for Ulrich's specimen is “lower half of the Leb- BULLETIN 319 Table 9.—Measurements of ten specimens of Columbicrinus crassus Ulrich (in mm). USNM 89826 UI X- UI X- UI X- UI X- UI X- UI X- UI X- UI X- UI X- (holotype) 5704 5705 5706 5707 5708 5709 5711 5712 Sys) Cup height 9.5 12.0 ТӨ 8.0 10* 9.0 — 6.7 69° 6.6 Cup width 9.3 TRZ 10.8 8.5 10* 8.3 8.3 8.0 7.0 78 АВ В height 52S 6.8 6.3 4.5 — — — 4.0* - - AB B width 4.7 6.0 De 4.2 - - 4.2 4.0 - 3.4 AR height 5:3 6.3 6.3 47 — 5.0 4.1* 4.0 — 32 AR width 5.0 5.8 5.8 4.9 - Sio) 4.7* 4.7 - 4.0 BR height 3.9 6.7 6.3 4.7 — — 4.0* 4.4 - 3.9 ВЕ width 5.0 6.3 6:2 4.8 — — 4.7* 4.3 — 4.0 C iR height 3.4 5.0 4.7 38 4.0 E 555 35 25 2.8 CiR width 5.9 6.0 6.1 4.5 5.4 — 4.3 4.5 Bos 35 С sR height 35 o 35 2.5 Jal — 2 DIS - 23 C sR width 4.8 6.5 5-5 4.7 929 - 4.6 4.4 8:597 4.0 D R height 5:5 6.8 6.3 4.3 993 — 4.8 4.3 3.4* 4.0 D R width Sl 750 7.3 597 6.5 — 6.0 Sl 4.1 4.3 E iR height 347 5.0 4.5 383 - 4.1 3:7 3:2 Dae 3.0 E iR width 4.8 6.3 SY 4.7 — 5:5 4.4 4.3 Bo 305 E sR height да 3,0 3.0 23 - Dall 2.0 1.9 2.21 1.9 E sR width 50 5.8 => 4.5 - 5.0 4.5 3.8 3:08 Sul Width of column at base of cup 5195 6.0 6.3 4.5 — 4.3 — 52 3.8 — Crown height — — 73 60* 85 3/5; 40* 597 — 48 * = estimated. ** = mean value. anon Limestone, Columbia, Tennessee.” Warn and Strimple (1977, p. 31) considered Columbicrinus un- recognizable because the type lacks distal arms and column. Frest, Strimple, and McGinnis (1979, p. 408) followed this view and named a new genus, Praecur- soricinus, based on more complete material from the upper Mountain Lake Member, Bromide Formation, of Oklahoma. The authors noted that Praecursoricri- nus has the same basic cup and proximal arm plating arrangement as Columbicrinus. (See remarks under C. crassus for a comparison of the two forms.) Their cri- teria for taxon recognition are not clear, however; in the same paper (p. 413), these authors base a new genus, Difficilicrinus, on the cup alone, material less complete than the holotype of Columbicrinus. I have collected 15 additional specimens that I believe to be conspecific with Ulrich’s type. Twelve of these speci- mens were found in the lower member and one in the upper member of the Lebanon, within 10 km of Co- lumbia (loc. Z-654); two specimens are from the lower member and one is from the upper member near Nash- ville, Tennessee (loc. Z-651). Several of the new spec- imens are complete crowns with proximal column. All specimens with cup and proximal arms have plate ar- rangements essentially identical to those of the holo- type of Columbicrinus. Further, there are no significant arm structure variations among the more complete new specimens. The holotype of Columbicrinus crassus is in the mid-size range of the new specimens. No other known crinoid in the Lebanon could be easily confused with Columbicrinus. 1 therefore consider Columbicri- nus to be readily recognizable and Praecursoricrinus Frest, Strimple, and McGinnis, 1979, is a junior syn- onym; it is therefore suppressed. Praecursoricrinus sul- phurensis Frest, Strimple, and McGinnis, 1979, 15 herein reassigned to Columbicrinus sulphurensis. Columbicrinus crassus Ulrich, 1925 Plate 7, figures 3-5, 10-13, 16; Plate 8, figures 1, 2, 4, 6-10, 17; Text-figure 7 Columbicrinus crassus Ulrich, 1925, p. 91, fig. 8. Columbicrinus crassus Ulrich. Moore, 1962b, p. 13, fig. 5, no. 5. Unrecognizable crinoid. Warn and Strimple, 1977, р. 31, pl. 2. figs. 6—7. Columbicrinus crassus Ulrich. Moore et al., 1978, р. Т549, fig. 342, nos. 4a, b. Unrecognizable crinoid. Frest, Strimple, and McGinnis, 1979, p. 408. Diagnosis.—Columbicrinus with moderate to large crown, up to 85 mm in height; BB about as tall as simple RR; simple RR nearly as wide as high to wider than high; sRR much wider than high, shorter than iRR; IBrr, 3 axillary; proximal column round to ob- scurely pentagonal. Emended description.—Crown moderate to large (Table 9). Cup higher than wide, maximum cup width at RR tops approximately 2.2 times as broad as ргох- | | | A O а LEBANON LIMESTONE ECHINODERMS: GUENSBURG 45 imalmost column; steeply obconical with sides dis- tinctly to slightly convex; slightly oval in transverse Section, variable, with major axis from C ray or BC interray through EA interray, minor axis from AB in- terray through D ray. A, B ray side of cup typically indented at level of RR, IBrr,. Sutures deeply to slightly impressed, plates smooth, thick. BB five, subequal, higher than wide to approxi- mately as high as wide, symmetrical to slightly asym- metrical hexagons with one vertex each pointing up and down, lower vertex obscure; sides diverge slightly distally; upper shoulders typically slightly unequal, with Shoulders bordering iRR largest. RR five, biradials in C, E rays, simple in A, B, D rays; biradials extend slightly beyond simple RR in both aboral and adoral directions; R facets fully as wide as RR. A ray R sym- metrically pentagonal to obscurely hexagonal (obscure vertex along side to accommodate E ray biradial), typ- ically slightly higher than wide, can be approximately as high as wide. B ray R asymmetrically hexagonal, approximately as high as wide, R facet tilted slightly away from C ray (to accommodate anal series). C ray iR symmetrical inverted pentagon, wider than high; higher than, typically slightly narrower than C ray sR. C ray sR upright asymmetrical pentagon, much wider than high; R facet wider than facet for X,, slightly Narrower than other R facets. D ray R very large, widest Plate in cup, wider than high, asymmetrical, obscurely Octagonal to heptagonal; vertices on plate sides form Wide angles; abuts X,. E ray iR inverted symmetrical pentagon, wider than high; higher, slightly wider than E ray sR. E ray sR quadrangular, much wider than high, nearly straight-sided, or sides expanding slightly distally. Anal series short, incurved; slightly narrower than adjacent arms proximally, tapering rapidly distally; of Six or (?)seven plates in some specimens, apparently as few as four in others. X, inverted subpentagon, low- E D с В А Text-figure 7.—Columbicrinus crassus Ulrich, plate diagram of Cup. Holotype USNM 89826, х 2.5. er, lateral margins curved, inserted in notch formed by upper left shoulder of C ray sR, upper right shoulder of D ray R. X, „ s quadrangular, wider than high to about as wide as high. Arms ten, uniserial cuneate, pinnulate; long, ap- proximately six to ten times longer than cup height; slender, tapering slightly, gradually over most of length, rapidly near distal tips. IBr, ,, ; axillary; IBrr distally tapering (primaxil can expand slightly), stout, thick. IBrr, wider than high, IBrr, much higher than wide when axillary, approximately as high as wide when non-axillary; primaxil slightly constricted laterally, distally. IIBrr subquadrangular proximally, becoming wedge-shaped, cuneate beyond IIBr, 10; up to 90 per arm. Pinnules given off alternately from IIBr, „4 and beyond, inserted into notches on lateral sides of Brr. Pinnules large, long, laterally compressed, each with as many as 12 subrectangular pinnulars. Column heteromorphic, long, greater than 30 cm long in UI X-5710 (distal portion illustrated on Pl. 8, fig. 17); narrow, from one-half to five-eighths as wide as cup near cup; round to obscurely pentagonal over proximal, medial column; distally becoming subpen- talobate and finally pentagonal by distal column. Col- umn typically expands for 4 to 8 cm, then tapers slight- ly toward holdfast, finally expands slightly for approximately last 0.5 cm before holdfast. Proximal, medial column of solid columnals, medial distal col- umn (3 to 7 cm of column length) of pentameres, dis- talmost column of longitudinal plate rows. Column length, columnal size, shape variable among specimens (compare Pl. 7, figs. 5, 11, 12 and Pl. 8, fig. 9). Nodals large, thick; only slightly wider than adjacent inter- nodals with slight convex latera near crown, becoming wider than adjacent internodals with prominent con- vex rounded latera distally. Distal nodals subpenta- lobate in outline, pentamere articulations in vertical notches between lobes; can have one or two faint stria- tions on either side of distal lateral pentamere artic- ulations. Internodes low, straight or slightly convex- sided, typically not as thick to rarely slightly thicker than adjacent nodals; typically one columnal per in- ternode, occasionally up to three internodals with mid- dle priminternodal intermediate between typical no- dals and internodals in size. Nodals and internodals become compressed distally with internodes eventu- ally being concealed between abutting nodals; grade transitionally into distalmost column. Distalmost col- umn of distally thinning, greatly compressed penta- meres, with low convex latera, abut adjacent rows along irregular to zigzag sutures. Proximal, medial column lumen small, quinquestellate, vertices correspond to pentamere articulations distally. 46 BULLETIN 319 Holdfast discoidal, dorsally multiplated; plates small subpolygonal, tightly abutting along flush sutures; col- umn facet 3 to 4 mm in diameter among three speci- mens. Remarks. —Columbicrinus crassus resembles C. sul- phurensis (Frest, Strimple, and McGinnis, 1979) from the upper (Oklahomacystis Zone) Mountain Lake Member, Bromide Formation, Oklahoma, but the lat- ter can be distinguished most readily by its smaller crown, typically narrower cup, relatively taller BB, which are taller than simple RR, relatively narrow sim- ple RR, which are taller than wide, and by its more angular pentagonal proximalmost column. The relatively large number of specimens of Co- lumbicrinus crassus display intraspecific variation, particularly in the column and less so in the cup. Two specimens from the upper Lebanon, UI X-5711 and UI X-5712, differ slightly from those in the lower Leb- anon; UI X-5711 (Pl. 8, fig. 1) possesses a relatively wider proximal column and UI X-5712 (Pl. 7, fig. 16) possesses BB in which lower margins project laterally slightly from the proximal column. All specimens are considered to be within the possible range of intraspe- cific variation. The column in this species was reconstructed using composite information from several incomplete col- umns. Approximate equivalent positions along the col- umn were determined using overlap among the various specimens. UI X-5710 is a nearly complete column and probably is the same specimen as partial crown UI X-5715. The holdfast in this species is similar to those described as ““Podolithus” by Sardeson (1908), and type 1C of Lewis (1982). Types and occurrence.—The holotype (USNM 89826; here refigured and measured) is from the lower Lebanon Limestone, near Columbia, Tennessee. Fif- teen other hypotypes are referred to this species. Three hypotypes are from locality Z-651: UI X-5704, and 5714 are from the lower member, Lebanon Limestone, and UI X-5711 is from the upper member approxi- mately 3 m below the top of the Lebanon Limestone. UI X-5712 is from the Sowerbyella - Diplograptus Zone of the upper member, Lebanon Limestone, at locality 7-654. The remaining hypotypes are from the lower member, Lebanon Limestone at locality Z-654a, and are UI X-5705 through 5710, 5713, 5693, 5715, 5768, and 5815. USNM 89826, UI X-5704, 5705, 5707, 5708, and 5711 through 5713 are figured and mea- sured; UI X-5693, 5714, and 5768 are figured; UI X- 5706 and 5709 are measured. Subfamily TRYSSOCRININAE new subfamily Diagnosis. —Cincinnaticrinidae with steeply conical dorsal cup. Unequal-sized compound RR taller than fused RR. X, large, deeply set in dorsal cup, resting on C ray iR; anal series long, armlike. Remarks.—The new subfamily agrees with other cincinnaticrinaceans in having compound RR in C, E rays, a conical dorsal cup, and initial isotomous arm bifurcations. Differences include the deeply inserted X, resting directly on the C ray iR, endotomous arm branching beyond the initial bifurcation, and wide, often asymmetrical, pentagonal, hexagonal, or heptag- onal RR. Other subfamilies within the superfamily Cincinnaticrinacea are the Cincinnaticrininae and Othneiocrininae (Atopocrininae). In cincinnaticri- nines X, abuts C ray sR, arm branching is isotomous or alternating heterotomous, and RR are generally in- verted pentagons. Othneiocrinines differ principally in having X, branch from IBr,, a less steeply conical cup, and alternating heterotomous branching beyond the initial bifurcation. The Tryssocrininae is similar in several ways to the Daedalocrininae of the superfamily Homocrinacea. Arm branching, cup shape, proximal anal series and stem morphologies are all similar. Under the scheme of classifications of Moore et al. (1978), and Warn and Strimple (1977), these similarities are subordinate to cup plate arrangement, particularly the number and position of compound RR (= bilateral symmetry planes of Ubaghs, 1953). Cup plate differences include the facts that X, rests on C ray sR, BB are proportionately much lower, and, most importantly, the B ray R is compound in Daedalocrinus Ulrich, 1925. Genus TRYSSOCRINUS new genus Etymology of name.—tryssus (Gr.) = delicate, dain- ty (refers to the crown) + crinum (L.) = lily (the stan- dard crinoid suffix). Diagnosis.—Tryssocrininae with wide, pentagonal BB, sutures flush, plates smooth; endotomously branched armlets after the initial isotomous bifurca- tion. Tryssocrinus endotomitus new species Plate 8, figures 3, 5, 11-16, 18-20; Plate 9, figures 1, 2, 6, 7, 14, 15; Text-figure 8 Etymology of name.—endo (Gr.) = within, inside + tomos (L.) = part (refers to the armlet branching in this species). Diagnosis.—As for genus. Description.—Dorsal cup smail (see Table 10), ap- proximately twice as wide as high, widest at level of IBrr,, steeply conical, nearly straight to slightly con- vex-sided, cup height from base of BB to top of E ray sR varies from 4.1 to 5.5 mm. Plates smooth or with slightly irregular surfaces, sutures flush. ЕЦ, 582 LEBANON LIMESTONE ECHINODERMS: GUENSBURG 47 Table 10.—Measurements of six specimens of Tryssocrinus endotomitus n. gen. and sp. (in mm). Ux ОХЕ UI X- UX 5824 Ul xX UI X- 5810 5816 5820 (holotype) 5828 5831 Cup height! = 4.1 4.6 4.5 4.8 4.7* Cup width? Sal 4.3 4.8 4.5 Sal 4.9* АВ В height = 1.6 2.1 1.8 = 1.9 AB B width = 1.9 1.9 1.9 20 1.8 АВ height - Xi 2.3 201 2.5 2.3 A R width == 252 2.8 23 2.6 2.3 BR height a 28] 2.3 D 2.4 2.3 ВК width = 259 2.3 2 2:7 272 C iR height 2.6 21 2.3 2.4 2.5 2:9 C iR width 2.8 2.3 25 23 2.9 2.5 C sR height 167 is 1.3 1.4 1.6 1.5 C sR width 227, 1.9 1.9 1.9 229] 2.0 DR height ES E 212) 2.5 2.4 28 DR width ES 25 255 2:5 3.1 3.0 E iR height == 1.8 1.8 1.9 1.9 1.8 E iR width E 1.9 1.9 2.0 2.0 1.9 E sR height = 1.1 12 1.3 1.6 1.4 E sR width E 1.9 2. 292, 2:3 24 Width of column at base of cup 3.0 2.6 2/3 2 3.3 2.6* ! = measured from base to top at E ray sR. * = measured at level of top at E ray sR. * — estimated. Basals five, symmetrically pentagonal, vertical mar- gins usually diverging slightly upward, can be nearly Straight-sided; central lower margin can have small downward projecting vertex, CD База! averaging Slightly wider than other subequal-sized basals. RR five, three simple (A, B, D rays), two compound (C, E rays); compound RR slightly taller than simple RR. Radial facets slightly concave. A ray R about as tall as Wide, asymmetrically hexagonal, side adjacent to E ray With vertice to accommodate compound R. B ray R about as high as wide, asymmetrically hexagonal, pos- terior (towards C ray) side with vertice to accommo- date compound R. C ray iR symmetrically hexagonal, Wider than tall; C ray sR much smaller than C ray iR, displaced away Кот CD interray to accommodate deeply set large X,, pentagonal to obscurely hexagonal; R facet displaced away from CD interray. D ray R asymmetrically heptagonal, wider than high with one Vertice on either side to accommodate compound RR. E ray iR about as high as wide, symmetrical inverted Pentagon; E ray sR symmetrically hexagonal, much Wider than tall. Anal tube tall, narrow, non-tapering as far as known, approximately two-thirds arm length in one specimen; Consists of dorsal anal series, lateral and ventral thin small polygonal plates apparently higher than wide lat- erally, wider than high ventrally. X, large, approxi- mately equal to C ray sR, set deep in cup; resting below on C ray iR, abuts D ray R on lower left, C ray iR on right side; upper margin extends slightly to far beyond upper margin of C ray sR. Anals beyond X, narrower, much smaller than X,, narrow slightly distally; subquadrangular to obscurely hexagonal, first four to six wider than high, gradually becoming higher than wide more distally. Arms ten, long, thin, as much as nine or ten times longer than cup height (PI. 8, fig. 3). Initial bifurcation Text-figure 8.— Tryssocrinus endotomitus n. gen. and sp., plate diagram of cup. Drawn primarily from paratype UI X-5824; dashed lines indicate approximate position of anal tube, approximately x6. 48 BULLETIN 319 isotomous followed by numerous endotomous armlet bifurcations. IBr, ,, ; axillary; IBrr, fixed in rays with fused RR, enlarged, laterally abut compound RR. IBrr taper distally. Armlet bifurcations every fourth or fifth Br proximally, increased to every other Br distally; axillary Brr wider than intervening non-axillary Brr. Armlets long, slender, pinnule-like, becoming shorter distally; unbranched to rarely bifurcating once isoto- mously (only one example observed); as many as 34 armlets per arm. Column heteromorphic, gradually changing, of al- ternating nodals, internodes for most of column length (PL 9, figs. 1, 6, 7, 14, 15); long, 3.5 to 5 times as long as crown height, greater than 14.5 cm long in UI X-5808; narrow, approximately one-third cup width at base of cup, tapers slightly for approximately 1 to 2 cm below crown, then expands slightly for most of column length, then tapers slightly again for one-third to one-fifth column length, finally expands slightly again for approximately the last 1 cm before holdfast; round proximally, gradually changing to pentalobate, then to pentagonal by distalmost column. Columnals solid for approximately first half of column, of pentameres for last half of column. Nodals thick near crown, about as wide as internodals, straight or slightly convex-sided; gradually becoming much wider than internodals with wide rounded pentalobate latera along distomedial col- umn regions. Notches between lobes mark lateral pen- tamere articulation facets. Internodes straight or nearly straight-sided; of single thin columnal near crown, much thinner than adjacent nodals; increasing to typically three (maximum five) internodals at approximately two-thirds distance to holdfast; internode up to twice as thick as adjacent nodals. Internodes of medial/distal column can have fine faint longitudinal ridges (Pl. 9, figs. 6, 14). Nodals, internodals become compressed, nodals become relatively smaller merging with inter- nodals, pentameres become wedge-shaped toward dis- talmost column region. Distalmost column of five lon- gitudinal plate rows; plates alternate laterally with plates of adjacent longitudinal rows, form zigzag suture pat- terns (Pl. 9, fig. 15) between plate rows. Plates of dis- talmost column thin, wedge-shaped, straight-sided, subequal in size. Lumen quinquestellate, small proxi- mally, moderate in size distally; vertices correspond with sutures of pentameres. Articulae cryptosymplec- tial. Holdfast small, 6 mm in diameter in paratype UI X-5826; column articular facet large, approximately 4.5 mm in diameter in same specimen; dorsally poly- plated, discoidal, encrusting. Dorsal plates small, sub- polygonal, tightly interlocking along flush sutures; ven- tral surface with thick (?)single plate. Types and occurrence. — Holotype: UI X-5808; Paratypes: UI X-5759 through 5766, 5797, 5798, 5804, 5806, 5809, 5810, 5812, 5813, 5816 through 5831, 5941, 6047, 6050, 6052. All are from the lower mem- ber of the Lebanon Limestone at locality Z-654a. UI X-5816, 5824, and 5831 and measured and figured; UI X-5810 and 5820 are measured; UI X-5764, 5804, 5808, 5809, 5819, 5823, and 5826 are figured. Superfamily ANOMALOCRINACEA Wachsmuth and Springer, 1886 Family ANOMALOCRINIDAE Wachsmuth and Springer, 1886 Genus ANOMALOCRINUS Meek and Worthen, 1865 Type species.— Anomalocrinus incurvus Meek and Worthen, 1865. ?Anomalocrinus antiquus new species Plate 9, figure 17; Text-figure 9 Etymology of name.—antiquus (L.) — ancient, old (refers to the early occurrence of this species). Diagnosis.— А species of Anomalocrinus character- ized by a small, low bowl-shaped cup with small slight- ly unequal-sized, slightly upflared BB; relatively long, thin arms with tall thin Brr; and IBrr,, IIBrr, axillary. Description. — Crown small, dorsal cup low, bowl- shaped, widest at tops of RR, pentagonal in outline, approximately 2.5 times as broad as column articular facet. Cup plates smooth, sutures flush. BB five, small, unequal in size, pentagonal, wider to much wider than tall, sides rapidly diverging, upflared only slightly; BC, CD, DE BB subequal-sized, appar- ently larger than AB, EA BB. RR five, two compound (C, E rays), three simple (A. Text-figure 9.—?4nomalocrinus antiquus n. sp., camera lucida drawing of cup and parts of arms (as seen from below, based on holotype, UI X-5852). Dashed line marks position of partial column as preserved, approximately x4. LEBANON LIMESTONE ECHINODERMS: GUENSBURG 49 B, Drays). Compound RR taller, narrower than simple RR. R facets at ends of short distolaterally projecting necks; necks much narrower than RR, round in trans- verse section (Pl. 9, fig. 17). Simple RR largest plates in cup, wider than tall; D ray R obscurely octagonal (counting distal margin as one side); A, B ray RR sub- hexagonal. iRR narrower, shorter than sRR, symmet- rical inverted pentagons, wider than tall, wide-angled lower vertice. sRR wider than, and abutting adjacent iR, simple RR along straight sutures; C ray sR abuts Жү; Х, probably wider than tall, inserted into broad, shallow, notch in CD interray; notch formed by upper left shoulder of C ray sR, upper right shoulder of D ray R; distal margin apparently gently curved. Arms poorly known, distal arm described from D ray only. Arms thin, at least three times longer than сир height, initial branching isotomous, succeeding branchings heterotomous, ramules in one known ray branch beyond IIBrr at irregular intervals. Brr round in cross section; Br pair immediately above axillaries well separated, rapidly diverging (Pl. 9, fig. 17). IBrr, axillary, higher than wide, widest distally. IIBrr, ax- illary, higher than wide. IIIBrr as much as five times taller than wide. Two apparent ramules visible on dif- ferent arm branches, branch from ШВг,, IIIBr,5; con- sist of few Brr (two or three), Brr much higher than wide. Column unknown; small fragment attached to crown Suggests proximal columnals were thin. Remarks. —?Anomalocrinus antiquus n. sp. is broad- ly similar to the type species, А. incurvatus Meek and Worthen, 1865, from the Maysvillian Fairview For- mation at Cincinnati, Ohio. The latter species can be distinguished primarily by its thicker, shorter arms, and relatively shorter primibrachs. Some specimens of 4. incurvatus possess an extra quadrangular plate in the basal circlet and a vertically divided D ray radial; neither of these features are present in the type and only known specimen of A. antiquus. The new species could be generically distinct, but is retained in Апот- alocrinus until better preserved material is found. Type and occurrence.— The figured holotype, UI X-5852, is from the upper member of the Lebanon Limestone at locality Z-653. Superfamily MYELODACTYLACEA Miller, 1883 Family TORNATILICRINIDAE new family Type genus.— Tornatilicrinus n. gen. Diagnosis. — Myelodactylacea with slender, elongate Crown; cup of moderate height, biradials probably in all five rays (B ray R unknown), anal series resting on, extending directly above left shoulder of C ray sR; A, D, E ray iRR of different heights, all projecting above upper margin of C ray iR, below C ray sR; R facets fully as wide as RR; rays five, arms 10, equal-sized; column of pentameres. Remarks.— The new family is monogeneric. It is as- signed to the Myelodactylacea because it possesses bi- radials in all five rays or possibly a single biradial in the C ray only, depending on terminology used (see discussion below). Difficulties concerning this classi- fication arise from the nature of present disparid clas- sification which, as previously discussed under re- marks for the order Disparida, is based primarily on position and number of biradials and the difficulty of recognition of biradials in Tornatilicrinus. Recognition of biradials in all but the C ray is uncertain because 1) cup and arms merge without distinctive break, and 2) plates herein designated as sRR in the А, D, E rays appear almost the same as succeeding IBrr and extend for about half their heights to slightly different heights beyond the top of the C ray sR. This last feature is due primarily to iR heights in these rays that are distinctly unequal, with A > D > E. The only known specimen of Tornatilicrinus is slightly disarticulated with BB and RR circlets sepa- rated and rays slightly ajar. The first two plates in each ray (here designated iRR, sRR) remain tightly ap- pressed, however, thus suggesting a strong connection. Lateral facets of the first two plates (iR, sR) of the A ray are visible with the zygosynostosial articular facet of the first plate (iR) continuing across to the second (sR), then narrowing and disappearing distally. The second plate (sR) thus appears to have been a rigid part of the cup structure. The upper margins of the second ray plates (sRR) in the A, D, E rays of Tor- natilicrinus are at only slightly different levels. Mor- phologic evidence, therefore, supports the designations ofsRR and iRR (and classification) adopted here. How- ever, the designation of these second ray plates as SRR rather than fixed Brr remains uncertain and arbitrary due to their intermediate position between free arms and cup. (See Ubaghs, 1978b, p. T119 for a discussion concerning this difficulty.) Further, no consistency ex- ists for resolving similar problems in other disparids. Plates with similar positions to the second ray plates (КК) in the Tornatilicrinidae are termed sRR in Еи- stenocrinus Ulrich, 1925 (Eustenocrinidae) whereas they are fixed Brr in Herpetocrinus Salter, 1873 (Mye- lodactylidae). Three other families are presently included in the Myelodactylacea; the Myelodactylidae, Iocrinidea, and Eustenocrinidae. The Myelodactylidae differs from the new family in having specialized bilateral columnals arranged in a doubly recurved coil, with elongate cirri 50 BULLETIN 319 that protect Ше crown. The crown has five unequal- sized rays or only four rays, compound R in the C ray only. Crown similarities include X, resting on, extend- ing upward from left shoulder of C ray sR, and initial arm bifurcation well above the cup. The Iocrinidae differs in having a relatively wider cup, five undivided equal-sized RR with prominent distal articular facets, and an anibrachial (not a biradial) plate above the C ray R from which the anal series and the C ray branch (Ubaghs, 1978b, p. T118). Eustenocrinids are similar in having compound RR in all five rays, but differ significantly in having X, supported directly by C ray sR (no arm in C ray) or by the left side of IBr,. Beyond the Myelodactylacea, Tornatilicrinus n. gen. is most similar to /bexocrinus Lane, 1970, of the Ho- mocrinacea. Dorsal cups are similar in shape and broadly similar in plate arrangement; both have X, resting on the left shoulder of C ray sR only. Proximal columns are similarly of thin pentamere circlets, and both have 10 arms. Ibexocrinus, however, does not have compound RR in A, D rays (this difference from Tornatilicrinus is not great, however, as it is the result of only minor proportional differences in R, IBr plate heights). Arm branching occurs much closer to the cup in Ibexocrinus than in Tornatilicrinus;, armlets are al- ternating heterotomous in /bexocrinus, endotomous in Tornatilicrinus. That Tornatilicrinus is a primitive disparid inadu- nate is supported by its simple, generalized cup plate arrangement and column of pentameres. In order to fit present disparid classification it is placed in the Myelodactylacea although it is similar in many ways to Ibexocrinus of the Homocrinacea. Tornatilicrinus might indicate a close relationship between at least some of the disparid superfamilies [Anomalocrinacea, Cincinnaticrinacea, Homocrinacea, Myelodactylacea in part] than has previously been believed, but its phy- logenetic relationships are obscure. Genus TORNATILICRINUS new genus Etymology of name.—tornatilis (L.) = beautifully rounded, turned (refers to the elegant crown) + cri- num (L.) = Шу. Type species.— Tornatilicrinus longicaudis n. sp. Diagnosis.— Tornatilicrinidae with a small crown having a slightly wider than tall cup, no distinct break between arms and cup, initial isotomous arm bifur- cation well above dorsal cup at IBr; „„ в, in single known example, few (approximately three) armlets branched endotomously on IIBrr; proximal column pentagonal. Tornatilicrinus longicaudis new species Plate 9, figures 3—5, 9, 12, 13, 20; Text-figure 10 Etymology of name.—longis (L.) = long + caudis (L.) = stem (refers to the long column in this species). Diagnosis. — As for the genus. Description. — Crown small, narrow; cup steeply con- ical, sides slightly convex, slightly wider than tall, wid- est at top of cup, approximately 1.6 times as wide as proximalmost column, subpentagonal in transverse outline at level of BB, round at level of RR. Cup plates smooth, slightly convex to flat, sutures flush. BB five, symmetrical to nearly symmetrical penta- gons, slightly wider than high, sides subvertical to slightly diverging; facets bordering C ray iR slightly longer than other distal facets. RR five, probably all compound, B ray R unknown; straight or nearly straight-sided, C ray R apparently widest. R facets fully as wide as RR in all but C ray. IRR symmetrical in- verted pentagons, of differing heights; heights range T 1 П mm m E E Ї | F Sm. NN Text-figure 10.—Tornatilicrinus longicaudis n. gen. and sp., plate diagram of partial crown (based on holotype UI X-5755). Distal parts of some arms not shown, dashed lines indicate not exposed or missing. Magnification approximately x 5.5. LEBANON LIMESTONE ECHINODERMS: GUENSBURG 51 from A, D, E, С in order from tallest to shortest; A, D, E ray КК higher than wide, C ray iR approximately as tall as wide, much lower than other iRR. A, D, E ray sRR quadrangular, wider than high; C ray sR sub- pentagonal, about as tall as other sRR, not extending as far distally as other sRR, with divergent left side and deep asymmetrical concavity in upper left shoul- der for insertion of X,. Anal tube tall, narrow, widens distally to near tops of arms, termination unknown; of dorsal armlike anal series, lateral plates small, thin, ventral plates much wider than high. Anals rectangular, varying greatly in height along series, becoming wider distally, narrower than adjacent Brr proximally; becoming wider than adjacent Brr distally, distal anals approximately one- quarter to one-third wider than proximal anals. X, higher than wide, left side slightly convex, lower mar- gin asymmetrical, highly convex. X, about as high as wide, X, е taller than wide. Beginning with Х,, short much wider than tall plates intercalated into and typ- ically alternating with taller plates (Pl. 9, fig. 5). Arms long, approximately four times longer than Сир height, tapering gradually away from cup; arm bundle approximately as wide as cup when folded to- gether. Brr dorsally flattened; Brr other than Axx rect- angular, sides subparallel. IBrr wider than tall; IBrr,, at least, axillary, much wider than high, about as wide as IBrr,; IBrr, approximately two-thirds width of IBrr,. Initial arm bifurcation isotomous. IIBrr about as high as wide to slightly wider than high, variable. Two or three short armlets branch endotomously from IIBrr; branching begins on IIBrr,, at least, occurs every third ог fourth IIBrr thereafter. Column incompletely known, long, narrow, > 20 cm in holotype (Pl. 9, figs. 3, 4, 20); tapers slightly for approximately 4 mm away from crown, then gradually €xpands distally; pentagonal over most of known length, apparently rounded distally. Columnals of straight-sid- €d thin to moderately thick pentameres, thicknesses variable, generally thicken distally; thick and thin co- lumnals can alternate along length; some columnals 6 to 8 cm below crown incomplete in holotype, penta- meres lens-shaped, not abutting laterally. Exteriors of Pentameres typically with rounded upper and lower margins, form thin discontinuous notches between co- lumnals. Apparent far distal column segment (Pl. 9, fig. 9) of five vertical series of subhexagonal plates; Plates much wider than high, correspond in position to pentameres of proximal column. Each plate series alternates with adjacent series forming zigzag suture between series. Proximal, medial column lumen pen- tagonal, vertices correspond with vertical pentamere Sutures and are midway between angles of column. Articulae symplectial; crenularium petaloidal (Pl. 9, fig. 12), each “petal” corresponds with pentamere. Remarks. —*' Dendrocrinus modestus" Safford, 1869 (nom. nud.), 18 tentatively referred to this species. Saf- ford's only known specimen is apparentlv lost; how- ever, two casts of the specimen (Pl. 9, fig. 12) are pre- served as USNM 52273. The casts do not show great detail, but do show similar cup shape, arm branching, and size to the holotype of Tornatilicrinus. The casts also indicate a B ray R similar to other RR. Types and occurrence. — The figured and measured holotype, UI X-5755, 1s from the lower member of the Lebanon Limestone at locality Z-655. Two column fragments, UI X-5756 and UI X-5757, were associated with the holotype. UI X-5757 is figured. The referred casts, USNM 52273, are from an unspecified horizon in the Lebanon Limestone at Lebanon, Tennessee. Measurements of UI X-5755 (in mm).— Crown height = 21%; cup height = 3.1; cup width = 3.5% CD B height — 1.4; CD B width — 1.7; A iR height — 1.8; A sR height — 1.1; C iR height — 1.5; C sR height — 1.4; D iR height = 1.8; D iR width = 1.6; D sR height = 1.0; D sR width = 1.6; E iR height = 1.4; E iR width — 1.6; E sR height — 1.0; E sR width — 1.6; column diameter at base of cup — 2.2 [* — estimated]. Order HYBOCRINIDA Jaekel, 1918 Family HYBOCRINIDAE Zittel, 1879 Genus HYBOCRINUS Billings, 1857 Type species. — Hybocrinus conicus Billings, 1857, p. 275; 1859, р. 29, pl. 2, figs. 2a, b. Hybocrinus bilateralis new species Plate 9, figures 8, 10, 11, 16, 18, 19; Plate 10, figures 1—11, 13—16, 18, 21, 22 Etymology of name.—bi (L.) = two, lateralis (L.) = at the side (refers to the crown, which is nearly bilat- erally symmetrical through the A/CD plane). Diagnosis.—A species of Hybocrinus characterized by a large subglobular calyx with tumid plates; sub- pentagonal as viewed from oral surface; plate surfaces pitted; A ray markedly thicker than others; anal X projects slightly above level of adjacent RR, upper margin with faint to distinct grooves. Description. — Calyx relatively large in largest spec- imens, maximum height measured from posterior over 20 mm; subglobular, distinctly asymmetrical with pos- terior protuberant in large specimens, calyx more con- ical in small specimens (PI. 9, fig. 16); posterior margin slightly higher than anterior margin; subpentagonal as viewed from oral surface. Major calyx, arm plate sur- faces finely to coarsely pitted (Pl. 10, figs. 9, 10); pits 52 BULLETIN 319 irregular, from three to six per mm; pits sometimes weathered, obscure. Some specimens with fine alter- nating grooves or striations along and perpendicular to calyx plate margins; grooves and striations abut across sutures (P1. 10, fig. 10). Grooves, striations 1 mm long, rapidly disappear toward plate centers. Su- tures slightly impressed, dorsal cup plates slightly tum- id. Upper margins of anal X with weak to strong grooves; commonly pass confluently to upper lateral margins of RR, lateral surfaces of BB (Pl. 10, figs. 3, 18). BB five, asymmetrical pentagons subequal in size; shape variable, nearly always taller than wide, or about as tall as wide. BB of CD, DE interray largest, other BB slightly smaller, shorter. Radial circlet plates largest in cup, plate size increases from A ray R toward RA, D ray R. A, B, D, E ray RR subpentagonal (assuming upper margins as one side), RA hexagonal; height-to- width ratio of RR variable. Small, subpentagonal C ray R rests on right shoulder of RA. R facets large, about one-third R width in B, C, D, E rays, one-half R width in A ray, nearly round; mounted on low protuberant necks in B, C, D, Erays, on high prominent protuberant neck оп A ray R (Pl. 10, fig. 5). Radial facets directed slightly laterally in B through E rays; markedly laterally in A ray. Ambulacral furrows deep, narrow. Anal X rests on upper left shoulder of RA, taller than wide to about as tall as wide, widest distally; upper margin convex, elevated slightly above level of adjacent RR, with faint to distinct grooves. Tegmen nearly flat, ambulacral furrows radially dis- posed about central oral opening; CD interray area slightly larger than others. Four anterior OO similar- sized, subquadrate, cover AB, BC, DE, AE interrays; distolateral margins notched, form elongate slits con- fluent with ambulacral furrows, extend to within | mm of oral opening where OO abut. Slits in C, D rays bordered posteriorly by small plates. CD interray oral moderate-sized, suboval to hemicircular, forms bridge between oral opening, CD interray area; with small circular hydropore located atop conical mound. Am- bulacral furrows extend from arms along slits, then above proximally abutting OO, converge above cir- cular oral opening. Ambulacral cover plates of DE and BC ray pairs merge lateral to oral opening, form keeled triradiate medial suture pattern above oral opening, proximal OO. Proximal ambulacrals keeled, of larger transversely elongate plates, smaller wedge-shaped plates intercalated medially. CD interray area appar- ently with valvular anal pyramid surrounded by small loosely connected ossicles. Anal pyramid plates ap- parently elongate, triangular (Pl. 10, fig. 11). Arms short, narrow (except А ray arm), uniserial, non-branching, with gradual distal taper, averaging 1.3 times longer than calyx height in two specimens. Á ray arm stout, much wider and deeper than C and D arms which are slightly wider and deeper than B and E arms. Brr variable, usually taller than wide, can be about as tall as wide. Ambulacral cover plates on arms slightly wider than tall, subtrapezoidal to subrectangular, meet medially along keeled nearly straight to slightly zigzag suture; approximately seven ambulacral plate pairs per Br. Column heteromorphic, short, length about equal to crown height, narrow, usually tapering slightly away from crown, then expanding near holdfast (Pl. 9, fig. 8; Pl. 10, fig. 22). Columnal morphologies change grad- ually along column length. Approximately proximal third of column of thin columnals having low, rounded nodose latera; nodes can abut with those of proximal- distal columnals forming faint longitudinal ridges (Pl. 9, fig. 10). Medial column columnals irregularly mul- tipartite, thinner than proximal columnals, latera sim- ilar to proximal column. Distal column of small irreg- ular subequidimensional subpolygonal plates tightly abutting to form irregular mosaic; plates with central nodes (Pl. 9, fig. 10). Holdfast small, low, encrusting, discoidal, apparently of a single solid ossicle. Remarks.—The new species resembles Hybocrinus tumidus Billings, 1859 from the “Trenton Limestone”, Ottawa, Ontario, Canada. Both have a similar sub- globular calyx, tumid calyx plates, and X, with grooved upper margin elevated above adjacent radials. H. bi- lateralis п. sp. can be differentiated, however, by 15 larger size, in having the A ray much thicker than the other rays, and in having a relatively smaller, narrower anal X. Hybocrinus nitidus Sinclair, 1945, from the Mountain Lake Member, Bromide Formation, Okla- homa resembles the new species in having an enlarged A ray. The Oklahoma species can be distinguished by its subcircular calyx outline as seen from above, less protuberant arm articular facets on RR, and pustular plate ornament (Sprinkle, 1982d, p. 121). Hybocrinus bilateralis also resembles H. pristinus Billings, 1858, from the “Chazy Limestone,” Montreal, Canada, 15 smaller, has coarse granulate ornament, and apparently has equisized arms. Coarseness of ornamental pitting is highly variable among specimens of Н. bilateralis (Pl. 10, figs. 9, 10). This variation is believed to be intraspecific because no other correlative variable morphological features were seen, and pitting coarseness is gradational among specimens. Types and occurrence. — Holotype: UI X-5807; para- types UI X-5799, 5858 through 5869 (all from the lower member of the Lebanon Limestone at locality Z-654a). UI X-5799, 5807, 5861-5864, and 5867-5869 are figured. LEBANON LIMESTONE ECHINODERMS: GUENSBURG 53 Order CLADIDA Moore and Laudon, 1943 Suborder CYATHOCRININA Bather, 1899 Superfamily GASTEROCOMACEA Roemer, 1854 Family CARABOCRINIDAE Bather, 1899 Genus CARABOCRINUS Billings, 1857 Type species. — Carabocrinus radiatus Billings, 1857, p. 275; 1859, p. 31, pl. 2, figs. За-е. Carabocrinus species Plate 10, figures 12, 17, 20, 24, 25 Description. — Crown large, cup shape unknown; plates thin. Fine, sharp, radiating ridges extend from, perpendicular to, cup plate margins toward plate cen- ters where many meet. Ridge intersections form chev- rons (Pl. 10, fig. 17), ridges abut across adjacent plate faces; ridge frequency averages approximately 0.63 mm. Respiration "rhombs" (see Sprinkle, 1982c, p. 145) large, involving RR, OO (РІ. 10, figs. 20, 25), of many thin folds, elevated above general plate surfaces. RR with at least 20 folds per “rhomb”. IBB poorly known, sides rapidly diverging. BB taller than wide, widest distally. C ray B heptagonal. RR large, subhexagonal, approximately 20 mm wide, 18 mm tall in one specimen; sides subparallel. R facets horseshoe-shaped, much taller than wide; ambulacral groove deeply cleft with narrow sides. Arms of moderate length, branching isotomously initially, heterotomously thereafter, up to seven branchings per arm (Pl. 10, fig. 20). Brr uniserial; first one or two Br pairs beyond axillaries tightly abutting; typically subcuneiform. Proximal Brr elongate dor- Soventrally (transversely), becoming subround by ap- Proximately mid-arm-length. Each Br lateral margin With pair of sharp ridges; each ridge borders an im- Dressed suture between Brr; similar structures of ad- Jacent Brr form series of wedge-shaped grooves (be- tween lateral sides of Brr) bordered by sharp ridges (Pl. 10, fig. 20); grooves appear to be covered by small plates. These covered grooves appear to be contiguous With respiratory “rhombs”. IBrr, axillary; IIBrr, „ ; axillary. Column poorly known; long, narrowing slightly dis- tally, round. Proximally of thin, straight-sided radially (externally) undulating columnals (Pl. 10, fig. 12). Dis- tally, much wider than tall, of thicker, approximately €quisized columnals; divided into pentameres (Pl. 10, fig. 24). Laterally adjacent pentameres slightly offset, Sutures canted, form discontinuous in-echelon series extending along column length. Lumen pentalobate Proximally, pentastellate distally, Articulae symplec- tial; fine crenulae disposed radially about outer margin of articular surfaces. Remarks.—Lebanon carabocrinids are represented by several mostly fragmentary and corroded speci- mens. UI X-5857 (Pl. 10, fig. 25) differs from other specimens in having faint thecal plate ridges. The fre- quency, distribution and size of the ridges, where ob- servable, however, is the same as in other specimens; respiratory “rhomb” morphology is also similar. This specimen has possibly undergone localized dissolution along surfaces of the thecal plates (associated speci- mens of Hybocrinus Billings, 1857, and Cleiocrinus Billings, 1857, also from locality Z-654a, show such partial obliteration of fine surface detail). Column frag- ments assigned to this species were found in a single stratum associated with thecal plates belonging to Ca- rabocrinus sp. No other crinoids were discovered in this stratum. The size of the fragments seem reasonable for the associated thecal plates. Final confirmation, however, must await discovery of attached cup and column. All Lebanon carabocrinid specimens are ten- tatively assigned to this species. Carabocrinus sp. apparently most closely resembles Carabocrinus sp. from the Blackriveran part of the Chickamauga Limestone, Alabama (Butts, 1926, pl. 31, figs. 2, 3) and Carabocrinus sp. from the Blackriv- eran Grand Detour Formation, northern Illinois (Ko- lata, 1975, pp. 30, 31, figs. 1, 2). АП three agree in having taller than wide BB and large goniospires that are elevated above general plate surfaces. The Lebanon species differs in having fewer, more widely-spaced ridges. This species also resembles Carabocrinus tread- welli Sinclair, 1945 (see Sprinkle, 1982c, for redescrip- tion of this species) from the Mountain Lake Member, Bromide Formation, Oklahoma, but this species differs in having fewer thecal plate ridges. Known Lebanon specimens of Carabocrinus are too poorly preserved to allow specific assignment. Overall poor preservation of this species probably results from large size and relatively fragile construction. Types and occurrence. — UI X-5856, 5857 are from thelower member ofthe Lebanon Limestone at locality Z-654a; UI X-5855 is from the lower member, Leb- anon Limestone, at locality Z-651; UI X-5978, 5980, 5991 through 5993 are from the lower member, Le- banon Limestone, at locality Z-652. UI X-5855 through 5857, 5981, and 5991 are figured. Family POROCRINIDAE Miller and Gurley, 1894 Genus POROCRINUS Billings, 1857 Type species. — Porocrinus conicus Billings, 1857, p. 2779; 1859, p. 34, pl. 2, figs. 5a-d. 54 BULLETIN 319 Porocrinus lebanonensis new species Plate 10, figures 19, 23; Plate 11, figures 3-5 Etymology of name.—Lebanon (Limestone) + en- sis (L.) = at the place of (refers to the source of the type material). Diagnosis.—A species of Porocrinus characterized by a relatively large obovate dorsal cup, IBB about as tall as wide, column articular facet small; moderate to prominent calyx ridges with superimposed secondary ridges; well-developed goniospires, decrease in size from bottom to top of calyx, those of R-R-O corners small; R facets small; zone of proximal columnals narrow, short. Description.—Dorsal cup obovate; large for genus, height approximately 19 mm in largest specimen. Ca- lyx ridges moderate to prominently developed for ge- nus, abut across plate boundaries; with zero to four, usually three superimposed secondary ridges that dis- appear at plate centers (Pl. 11, fig. 3). Plate surfaces, particularly between calyx ridges, with fine punctae; no punctae on goniospires; plate sutures flush. Go- niospires moderately inset, at corners of adjacent cup plates, of moderate size, decreasing in size distally; those of R-R-O corners small; IB-IB-B goniospires with approximately 12 folds. IBB about as high as wide (Pl. 10, fig. 23), widest distally, much smaller than BB. Lower lateral vertices of IBB can be rounded, form notches into which attenuated proximal columnals fit; column articular facet small for genus, averaging about as wide as each IBB. BB large, as large to slightly larger than RR, as tall as wide to slightly taller than wide; CD and probably BC interray BB heptagonal, other BB hexagonal. RR large, approximately as tall as wide, R facets horeshoe-shaped, slightly elevated, laterally constricted, one-third to one-fourth as wide as R widths, RA small, quadrate, slightly wider than high, upper left shoulder articulates with anal X. Anal X small, only slightly larger than RA, slightly higher than wide, sub- pentagonal. OO poorly known, apparently large. Anal area large, covered with small, irregular, polygonal plates. Arms narrow, long unbranched, uniserial, at least two-and-one-half times longer than calyx height. Brr much wider than tall proximally, becoming nearly as tall as wide distally, lateral margins rounded, form notches between adjacent Brr along lateral margins of arms. Ambulacrals apparently of single alternating bi- series, much wider than high proximally; articulate medially along distinct zigzag suture. Column heteromorphic; proximal column region short, narrow, averaging about 3 mm long for three specimens, subpentagonal proximally, becoming round distally; of thin, expanding columnals with finely cren- ulate or possibly smooth outer surfaces. One specimen (UI X-5796) with approximately five faint longitudinal radially-disposed striae that continue for 1 to 2 cm onto distal column (Pl. 11, бр. 3). Proximal column tapers rapidly, passes transitionally to narrower me- dial/distal column. Medial/distal column known for approximately 3 cm in two specimens; transversely round to subround, slightly tapering distally; of thin straight to slightly convex-sided much wider than tall columnals, height to width ratio becoming somewhat smaller distally. Alternating thin and thick variable columnals proximally; distally of thicker, variable co- lumnals, intermittent thicker columnals with slightly convex sides. Medial column articulae cryptosymplec- tial; lumen small, pentagonal. Remarks.—The relatively small, slightly depressed goniospires, the usually prominent calyx ridges, narrow short proximal column and IBB that are as tall as wide together separate Р. lebanonensis п. sp. from other species of Porocrinus. P. lebanonensis n. sp. closely resembles P. elegans Kesling and Paul, 1968, from the early Trentonian (Rocklandian) Decorah Shale of Min- nesota but the latter species differs in having IBB av- eraging approximately 1.5 times wider than tall, a rel- atively larger column attachment facet and slightly wider proximal column region, and a smaller known crown size. Porocrinus plattinensis Shourd and Winter, 1976, from the Blackriveran Plattin Limestone, Mis- souri, is also similar to the new species but can be most easily distinguished by its low wide infrabasals aver- aging 1.7 times wider than tall and lack of secondary calyx ridges. Porocrinus fayettensis Slocum, 1924, from the Upper Ordovician Maquoketa Formation, Iowa, is only broadly similar to the new species and can be readily distinguished by its much larger goniospires. UI X-5796 differs slightly from other specimens as- signed to P. lebanonensis n. sp. in having low, wide calyx ridges with more prominent widely-separated secondary ridges. Types and occurrence.—The holotype UI X-5803 and three paratypes, UI X-5796, 5800, 5889, are from the lower member, Lebanon Limestone, at locality 7-654а. All type specimens are figured. One isolated thecal plate tentatively referred to this species, UI Х- 5890, is from Ше upper member, Lebanon Limestone, at locality Z-653. Superfamily DENDROCRINACEA Wachsmuth and Springer, 1886 Family DENDROCRINIDAE Wachsmuth and Springer, 1886 LEBANON LIMESTONE ECHINODERMS: GUENSBURG 55 Genus QUINQUECAUDEX Brower and Veinus, 1982 Type species. — Quinquecaudex glabellus Brower and Veinus, 1982, p. 134, pl. 10, figs. 1-11. Quinquecaudex species A Plate 11, figures 1, 2, 11 Description. — Crown large, more than 50 mm tall; narrow. Dorsal cup conical, steep-sided; plates smooth, sutures slightly depressed, small depressions at triple juncture points. IBB hexagonal, wider than high. BB probably hexagonal, slightly higher than wide, largest plates in cup; B below anal X truncated, heptagonal, higher than wide. RR wider than high; facets nearly as wide as (B ray) or as wide as (C ray) R plates. RA below C ray R, probably subpentagonal, much wider poste- riorly; facet articulating with C ray R tilted strongly away from anal series. Arrangement of anals within cup not discernable. Proximal anals above dorsal cup smooth, small, generally slightly wider than high; dis- tally becoming much wider than high, plicate. Arms long, slender, non-pinnulate, isotomously branching proximally; dorsal margins flattened proximally, rounded distally. IBr; axillary, IBrr much wider than high. Distal Brr nearly as high as wide. Column heteromorphic, gradually changing; long, much longer than crown length; wide, apparently near- ly as wide as cup proximally, distally tapering; pen- talobate. Columnals thin, of pentameres. Large and Small columnals typically alternate proximally; even Smaller, exceedingly thin columnals intercalated over Short column segments. Distal columnals become thicker, equisized. Concave constricted ridges abut across laterally adjacent pentameres, expand to form репїатеге midsections; single pore in depressions be- tween ridges at vertical sutures, best developed distally. Protruding midsections of pentameres flat or slightly convex-sided; articulations between pentamere circlets can be slightly depressed. Lumen pentagonal, large. Articulae symplectial at least distally; crenulae faint, arranged in semicircles adjacent to external margin of each pentamere. Remarks. — This species is assigned to Quinquecau- dex Brower and Veinus, 1982, on the basis of 1) den- drocrinine crown features such as the steeply conical Cup, upflared BB, RA below C ray R, elongate anal sac Of plicate plates, and elongate narrow non-pinnulate arms and 2) a pentalobate column of pentameres (see Brower and Veinus, 1982, p. 134). R facets are unusu- ally wide compared with most other dendrocrinids. The exact shape of dorsal cup, branching pattern of the arms, and shape and arrangement of some of the Cup plates, including the anal series, are unknown. Therefore this material cannot be assigned to species. The Lebanon species appears to be most similar to Q. glabellus Brower and Veinus, 1982, from the Pooleville Member, Bromide Formation of Oklahoma, but the latter species is considerably smaller and probably has fewer IBrr. Quinquecaudex [Dendrocrinus] springeri (Kolata, 1975) from Blackriveran and early Trentonian beds of Illinois and Canada, respectively, has a non- tapering pentagonal column. Quinquecaudex cincin- natiensis (Meek, 1873), from the Upper Ordovician of Ohio, has a more angular column and possibly a wider, more rapidly expanding dorsal cup. Quinquecaudex [Dendrocrinus] sp. A of Kolata (1975) from Blackriv- eran strata, northern Illinois, is smaller, and appears to have a less angular column, without prominent su- tural pores. Types and occurrence. — Two specimens, UI X-5700 and UI Х-5699, are assigned to this species; the former specimen is from approximately 3 m below the top of the upper member of the Lebanon Limestone at lo- cality Z-651, and the latter 15 from within 8 m of the base of the Lebanon Limestone, lower member, at lo- cality Z-658. UI X-5700 is figured. Family CUPULOCRINIDAE Moore and Laudon, 1943 Genus CUPULOCRINUS d'Orbigny, 1849b Type species. — Scyphocrinus heterocostalis Hall, 1847, p. 85, pl. 28, figs. 3a-f. Cupulocrinus species cf. C. gracilis (Hall) Plate 11, figures 6, 7 Poteriocrinus gracilis Hall, 1847, p. 84, pl. 28, figs. 2a-f. Dendrocrinus gracilis (Hall). Wachsmuth and Springer, 1879, p. 299. Non Cupulocrinus gracilis Ramsbottom, 1961, p. 13, pl. 5, figs. STE Cupulocrinus gracilis (Hall). Kolata, 1975, pp. 37, 38, pl. 7, figs. 6-8. Cupulocrinus gracilis (Hall). Brower and Veinus, 1978, p. 429, pl. 13.0025. 5573 Remarks.—Cupulocrinus gracilis (Hall) was origi- nally described from the Trenton Limestone, New York. It has since been redescribed based principally on specimens from the Blackriveran Platteville Group, northern Illinois (Kolata, 1975, pp. 37, 38) and ap- proximately equivalent-aged Platteville Limestone, St. Paul, Minnesota (Brower and Veinus, 1978, pp. 426— 430). Two Lebanon specimens, UI X-5701 and UI X-5703, are tentatively referred to this species. UI X-5701 is a nearly complete crown with proximal column and UI X-5703 is a crown and proximal column with a largely disarticulated dorsal cup; neither specimen has the CD interray exposed. The Lebanon specimens are very 56 BULLETIN 319 Table 11.—Measurements of three specimens of Cupulocrinus (in ferable to Cupulocrinus, but cannot be assigned to mm). species. The specimen is larger than specimens as- Cupulocrinus sp. cf. T signed to С! upulocrinus gracilis. It is similar in size and C. gracilis Ere feet plate configuration to specimens of Cupulocrinus UI X-5701 UI Х-5703 UI X-5702 plattevillensis Kolata, 1975, from the Blackriveran Platteville Group, northern Illinois. C height 33 — == създай 65 а BE Types and occurrence. — The single figured and mea- Cup width 6.0 = = sured specimen, UI X-5702, is from the lower mem- IB height 27 — - ber, Lebanon Limestone, at locality Z-651. | IB width 1.8 - - | В height zi = 4.7 Class PARACRINOIDEA Regnell, 1945 B width 25 a 3.8 | R height 22 21 28 Order COMAROCYSTITIDA R width 2.8 2.8 3.8 Parsley and Mintz, 1975 | Column diameter at 5 | ОШОГО) 20 20 E nn AMYGDALOCYSTITIDAE Jaekel, 1900 | | [nom. corr. Kesling, 1968, pro AMYGDALOCYSTIDAE Jaekel, 1900] similar to other specimens of Cupulocrinus gracilis but differ slightly in possessing relatively wider RR and Genus OKLAHOMACYSTIS tightly appressed rather than well-separated IIBrr, Parsley and Mintz, 1975 pairs. The Lebanon а also differ from those Type species. — Amygdalocystites tribrachiatus Bass- from northern Illinois but agree with the Minnesota ler, 1943. | specimens in possessing a slightly tapering rather than Re marks, Тһе emended generic diagnosis given by non-tapering column near the crown. These differences Parsley (1982, p. 215) is applicable to O. trigonis n. are considered minor and within possible intraspecific sp., except that thecal ornament ridge morphology in- variation range. y cludes raised flat-topped trigons as well as raised pyr- | Types and occurrence. — UI X-5701 is from the Sow- amid ridges y erbyella - Diplograptus Zone of the upper member, Lebanon Limestone, at locality Z-652 and UI X-5703 is from the upper Lebanon Limestone at locality Z-651. Both specimens are figured. Plate 11, figures 10, 12, 14-18; Oklahomacystis trigonis new species | Plate 12, figures 1-8; Text-figure 11 | Cupulocrinus species Etymology ој name.—trigonis (L.) = triangle (refers Plate 11, figures 8, 9 to the thecal plate ornament). | Remarks.—One crown fragment, UI Х-5702, is ге- Diagnosis.—Oklahomacystis with two unbranched | Text-figure 11.— Oblique view restorations of typical thecal plates in Oklahomacystis trigonis n. sp. (left) and Oklahomacystis bibrachiatus Parsley, 1982 (right); Oklahomacystis trigonis plate has partial flat-topped trigons; Oklahomacystis bibrachiatus has partial raised trigons; pore slits (PS) are demarcated in both illustrations. Much enlarged. LEBANON LIMESTONE ECHINODERMS: GUENSBURG 57 recumbent ambulacra; periproct situated to right of thecal apex on slightly elevated rim, adjacent to right ambulacra; hydropore and gonopore subapical, aligned with, continuing toward apex from left ambulacra; gonopore elevated atop moundlike plate; thecal or- nament ridge morphology varies from low to raised, nearly flat-topped trigons to raised pyramids; pore slits extend onto pyramid faces, border margins of trigons; raised pyramids concentrated near arms, apical region; stem articular facet ventralmost point of theca. Description. —Theca small- to moderate-sized, rang- ing from 14.0 through 22.5 mm high among known specimens; suboval to amygdaloid in profile; cross- sectional outline oval in small individuals, broadly oval in larger individuals, largest diameter toward dorsal end of theca; of approximately 50 to 60 plates; right Side protuberant. Areas about peristome, periproct, centers of thecal plates, ambulacra, brachioles with риз ове ornament; thecal plate centers can have sin- gle, large pustule. Three basals offset with stem articular facet to left, apparently without mutual pore slits. Stem articular facet ventralmost point on theca (theca does not extend below level of facet), suboval (?), with small central pore for lumen. Thecal plates typically hexagonal, plate centers depressed; major ornamental rays radiate from near plate centers toward plate corners (triple juncture points), expand away from plate centers. Ray mor- Phology variable, gradational among, within speci- mens. Rays meet similar rays at plate corners, together form moderately-raised nearly flat-topped trigons (Pl. 11, figs. 10, 16; Text-fig. 11) to triangular pyramids (or triacts); pyramids concentrated near arm, peristome regions (Pl. 11, fig. 15; Pl. 12, fig. 2). Pyramid faces, trigon sides bisected by plate sutures so that each triact/ trigon formed equally of parts from three plates. Pyr- amids and trigons arranged in rosettes about plate cen- ters on thecal surface. Thin skeletal sheets extend out- ward along sides of trigons, above and along pyramid faces. Pore slits in gaps between trigons/pyramids and adjacent skeletal sheets; slits from opposing pyramid faces, trigon sides form slit pairs in low areas between adjacent pyramid/trigon faces. Pore slits open to ex- terior through slits between trigons, skeletal sheets and gaps in sheets above distal faces of pyramids (Pl. 11, figs. 10, 14, 15; Pl. 12, fig. 3); openings possibly due to weathering of available material; therefore, slits pos- Sibly completely enclosed in skeletal material. Ambulacra two, unbranched, recumbent, nearly Straight, transversely arranged on theca; taper gradu- ally, distally away from peristome. Food groove an- terior in left ambulacra; posterior in right ambulacra. Rightambulacra with 13 to 15 segments, extends about two-thirds or three-fourths distance to stem facet. Left ambulacra with 13 to 16 segments, extends along an- terolateral side of theca to anterior side of stem artic- ular facet. Left ambulacra of UI X-5920 appears to touch proximal stem. Covering plates small, appar- ently of alternating biseries meeting submedially along zigzag suture. Brachials thin, shorter than thecal height in UI X-5920 (Pl. 12, fig. 3), gradually tapering, in- crease in size toward peristome. Brachiolars taller than wide, transversely evenly rounded away from food grooves; food grooves covered by tiny transversely elongate plates. Gonopore, hydropore subapically positioned, aligned with left ambulacra. Gonopore adjacent to peristome, elevated atop rounded moundlike plate. Hydropore V- or S-shaped, surrounded by elevated rim, rim appar- ently comprised of three plates. Peristome subapical, offset slightly to left. Periproct opening circular, offset to right of apex on anterior face, adjacent to left arm; bordered by four plates forming low rim, slightly to (rarely) moderately raised above theca. Periproct cov- ered by anal pyramid of seven wedge-shaped plates in UI X-5974; each plate with single prominent pustule. Stem round, homeomorphic, apparently shorter than thecal height, proximally flexed, slightly expanding dis- tally in UI X-5932 (Pl. 12, figs. 1, 2); of thin columnals. Columnals with nearly smooth, irregularly rounded, or nodose sides (Pl. 12, fig. 4). Stem in UI X-5932 articulates distally with possible low, digitate, encrust- ing holdfast. Remarks.—This species is assigned to Oklahoma- cystis because it has 1) a broadly elliptical transverse outline, and 2) thecal plate structure similar to though slightly different from previously described species of Oklahomacystis. Previously known species of Okla- homacystis are O. tribrachiatus (Bassler, 1943) and O. bibrachiatus Parsley, 1982, both from the Upper Echi- noderm Zone, Mountain Lake Member, Bromide For- mation, Oklahoma. Thecal plates of Oklahomacystis are characterized by pore slits that extend from the interior toward the thecal exterior. Plates have sharp, raised ridges ra- diating from plate centers, expanding toward corners (triple juncture points). Ridges from adjacent plates in O. bibrachiatus and O. tribrachiatus abut forming tri- angular pyramids (triacts) (Parsley and Mintz, 1975, p. 53; Parsley, 1982, p. 215). In well-preserved spec- imens of these species, thin skeletal sheets extend out- ward above and along pyramid faces; these sheets co- alesce with pyramids along pyramid ridges. Pore slits occupy the gaps between skeletal sheets above and pyr- amids below and are closed to the exterior by skeletal material (Parsley and Mintz, 1975, p. 53) (Text-fig. 58 BULLETIN 319 11). Weathered specimens of these species have the thin outer skeletal sheets partly removed, exposing the pore slits and pyramids below. Raised pyramids and accompanying skeletal sheets are arranged in rosettes on the thecal surface similarly to pyramids/trigons in Oklahomacystis trigonis n. sp. The thecal plates of Oklahomacystis trigonis п. sp. differ from those in the two previously described species in having plate rays that form pyramids to raised flat to nearly flat-topped trigons (triangles). As noted in the description, skeletal sheets covering pore slits in avail- able specimens of O. trigonis are incomplete, leaving openings (to the exterior) contiguous with pore slits. These openings result from corrosion (dissolution) or were incompletely calcified in life. Pyramids on O. trigonis greatly resemble weathered examples of O. bibrachiatus and O. tribrachiatus (compare Pl. 11, fig. 16; Pl. 12, fig. 3 with Parsley and Mintz, 1975, pl. 6, fig. 1) suggesting the former possibility is correct. How- ever, some specimens with openings display good pres- ervation including delicate pustules on thecal plates and brachiolars and tiny ambulacral coverplates. If skeletal sheets did completely enclose pore slits in O. trigonis, they must have been very thin. Of the two previously described species, Oklaho- macystis trigonis is most similar to O. bibrachiatus. Both species have two transversely arranged ambula- cra. In addition to the above thecal plate differences, O. bibrachiatus can be distinguished from O. trigonis in having 1) an apically positioned periproct mounted atop a prominent low chimney, 2) the peristome below the level of the thecal apex and canted to the left, 3) relatively thinner ambulacra, 4) the right side of the theca extending well below the level of the stem artic- ular facet in larger specimens, 5) the gonopore situated atop a small tubercle and positioned ventrally from the hydropore and ambulacra, 6) a more straight-sided, relatively less bulbous dorsal region of the theca, and 7) a right ambulacra typically moderately to strongly deflected around the anal chimney. O. tribrachiatus can be easily distinguished from the new species by its three ambulacra. UI X-5931 consists of an incomplete theca and par- tial stem buried in hard matrix. The thecal size is small and pore slits are relatively short and do not reach as far toward plate centers as they do in larger specimens (РІ. 12, fig. 8); both are features that Parsley and Mintz (1975, p. 54, pl. 6, figs. 2, 3) attributed to immaturity in Oklahomacystis tribrachiatus. Thecal shape, thecal plate shapes, distribution and position of the periproct and stratigraphic occurrence within the lower Lebanon Limestone indicate that this specimen belongs to O. trigonis n. sp. and is immature. UI X-5932 is a poorly preserved, crushed, but nearly complete individual. Thecal plate and arm morphol- ogy is identical, insofar as can be determined, to other, better preserved specimens assigned to Oklahomacys- tis trigonis. The specimen is of considerable interest because it possesses a nearly complete column, which is attached to an apparent low solid irregular holdfast (PL 12, fig. 2). The surface of the holdfast is obscured, as it was appressed against or encrusted by a bryozoan; a broken edge, however, shows characteristic echino- derm cleavage. The holdfast encrusts a trepostome bryozoan. The column is shorter than the thecal height and expands slightly toward the holdfast. Amygdalocystitid life modes have long been enig- matic. This problem is due to their unique thecal na- ture and lack of preserved columns with associated attachment devices. There is general agreement that thecal symmetry and arm distribution indicate that most if not all amygdalocystitids were rheophyllic or- ganisms with the theca canted into prevailing currents, but it was not known how this was accomplished. The holdfast and short column in Oklahomacystis trigonis (specimen UI X-5932) indicates this species was an attached low level suspension feeder. The proximal column flexure, shown best in other specimens of O. trigonis (Pl. 12, fig. 4), could have helped orient the theca upright from irregular, inclined, firm attachment sites as well as into prevailing currents. This life mode resembles that shown in a restoration by Parsley and Mintz (1975, text-fig. 6) for Amygdalocystites florealis Billings, 1854 (though their interpretation lacks an en- crusting holdfast). The occurrence of UI X-5932 in an associated thick mat of trepostome bryozoans is sim- ilar to O. tribrachiatus, which occurs along the edges of bryozoan bioherms (Fay and Стай нат, 1969, p. 39) and might indicate Oklahomacystis utilized the current baffling effects of other organisms (or irregular sub- strates) to maintain stability. Types and Occurrence.—Twenty specimens are as- signed to Oklahomacystis trigonis n. sp. The holotype UI X-5922 and paratypes UI X-5801, 5921, 5923 through 5930, 5933 through 5937, and 5974 are from the lower member of the Lebanon Limestone at locality 7-654а. Paratypes UI Х-5920, 5931, and 5932 аге from the lower member of the Lebanon Limestone at locality Z-651. Isolated thecal plates probably belong- ing to this species were observed at many exposures of the Lebanon Limestone. Figured specimens are UI X-5920, 5922, 5931, 5932, 5934, and 5936. Oklahomacystis species aff. O. trigonis new species Plate 11, figures 13, 19, 20 LEBANON LIMESTONE ECHINODERMS: GUENSBURG 59 Remarks. – Опе damaged but uncrushed partial the- ca from the upper member of the Lebanon Limestone resembles specimens of Oklahomacystis trigonis n. sp. from the lower member of that unit in the nature of the thecal plate ray and pore slit morphology, the length and disposition of the arms and pustulose ornament, and position and shape of the periproct, hydropore, gonopore and stem facet. The specimen differs, how- ever, in being about one-third larger, having a more amygdaloid profile, a more compressed transverse out- line, and greater number of thecal plates. The differ- ences may be within the range of variation for O. tri- gonis, or the specimen may represent a new, closely related species. The single incomplete specimen is con- sidered insufficient to warrant erection of a new taxon. Types and occurrence.—' The single figured speci- men, UI X-5938 is from the upper member of the Lebanon Limestone at locality Z-653. Class RHOMBIFERA Zittel, 1879 (emended Paul, 1968) Order DICHOPORATA Jaekel, 1899 Superfamily GLYPTOCYSTIDA Bather, 1899 V IV E МА А УЛТ ane WM tine RANAS NN A ji ~ УД SS; LO чудо Family GLYPTOCYSTIDAE Bather, 1899 (emend. Paul, 1972; Sprinkle, 1982b) Genus TANAOCYSTIS Sprinkle, 1982b Туре species.— Tanaocystis watkinsi Sprinkle, 1982b. Remarks.— The generic diagnosis of Sprinkle (1982b) encompasses the Lebanon species except that the new form possesses 13 rhombs and demirhombs instead of 10 to 12 and apparently rounded rather than skirted outer proximal stem columnals. These differences are considered specific taxobases and are incorporated into the generic diagnosis. Tanaocystis sprinklei new species Plate 12, figures 9-14, 16, 17; Text-figure 12 Etymology of name.—The specific name honors James Sprinkle. Diagnosis.—A species of Tanaocystis characterized by an elongate theca; B1-B2, B2-B3 sutures curved; rhombs greatly elongate, medium length dichopores with approximately 0.14 mm average spacing, IL5-L5 rhomb present; thecal plates ornamented with numer- ous prominent, wide, subcentral, radiating, discontin- uous ridges, nodes common between ridges; periproct E QS еМ ААУ `` „ә B2 Text-figure 12.— Tanaocystis sprinklei n. sp., composite partial plate diagram; based primarily on holotype UI X-5880, also on paratype Plates UI X-5877 through UI X-5879, UI X-5881 through 5885. Thecal ridges are stippled, inferred areas are dashed; areas where pore slits Or thecal ridges cannot be inferred accurately are marked by question marks. Much enlarged. subround; outer columnals of distal portion of proxi- mal stem region rounded with faint nodes. Description. — Theca moderate to large for the genus, approximate height of holotype 28 mm; apparently tapering, subcylindrical, with rounded summit; max- imum width just above base of ILL circlet. Thecal plates thin. Prominent, thick, irregular slight- ly curved ridges radiate from thecal plate centers, abut across adjacent plates; can be discontinuous, especially at plate edges. Major ridges typically radiate to mid- points of plate faces, in some to plate corners; second- ary, commonly discontinuous ridges radiate to plate edges between major ridges, many do not reach edges of plates; major chevron-shaped ridges can be along rhombs, between rhomb and adjacent plate face. Plate surfaces of ILL, LL, some BB series typically with faint to moderately prominent closely-spaced tiny pustules between ridges; nodes can be arranged in subconcentric bands perpendicular to radiating ridges (Pl. 12, fig. 13). Partial B2 only B known; distal, proximal tips miss- ing, probably hexagonal, B1-B2 and B2-B3 sutures curved. All ILL known, medium to large. ПЛ and IL2 share rhombs with B2; ПА subpentagonal, IL2 through ILS all subhexagonal, IL4 forms lower margin of peri- proct opening; IL3 and IL4 without rhombs or demi- rhombs; IL5-L5 share rhomb. L1 and L3 through L5 known, large to very large; LL circlet interrupted by R2; L1 hexagonal; L3 subpentagonal, oral margin steeply inclined; Ld and L5 subpentagonal, form sides, upper margin of periproct opening; L5 with two rhombs, L1 and L4 with one rhomb, L3 with two rhombs and one demirhomb (UI X-5899 has a very small second ?irregular demirhomb [РІ. 12, fig. 121). RR2 through 6 known, RR apparently form a closed circlet; RZ very large, elongate, other RR medium to large. R3 through R5 modified rectangular (assumes radial sinuses as one side), R2 modified hexagonal; R2 and R3 with one rhomb and two demirhombs or two rhombs and one demirhomb each, R4 and R5 apparently with two demirhombs each, R6 with at least one rhomb and one demirhomb. Ambulacral sinuses of medium length ex- tending nearly to aboral edges of R3 through R5. R6 shares ambulacrum 1 with R1. OO poorly known, ap- parently medium-sized, elongate. Periproct round or nearly so, medium-sized (for cy- lindrical rhombiferans), not more than 5 mm in di- ameter in holotype, surrounded by three thecal plates (ILA, LA, L5). Rhombs, demirhombs 13, greatly elongated, of closely-spaced medium length dichopores each sur- rounded by faint, low, narrow rim. Dichopores spaced moderately closely, from 0.12 to 0.18 mm, averaging 0.14 mm apart. BULLETIN 319 Ambulacra extend about one-third distance down theca as indicated by RR sinuses. Ambulacrum II, probably ambulacrum I, longer than other ambulacra. Ambulacral floorplates, covering plates, brachioles un- known. Only medial to distal region of proximal stem known; gradually tapering, of medium width (Pl. 12, fig. 9); stem apparently mounted in basal concavity as indi- cated by shape of B2 thecal plate. Outer columnals evenly rounded, some, at least, with faint nodes; inner columnals subpentagonal. Remarks.— Tanaocystis sprinklei n. sp. is incom- pletely known (Text-fig. 12). The holotype is a partial theca with associated partial proximal column (Pl. 12, fig. 9). Isolated thecal plates were used to determine shapes and arrangement of some plates not present in the holotype. Tanaocystis sprinklei n. sp. resembles Tanaocystis watkinsi Sprinkle, 1982b, from the upper part of the Pooleville Member, Bromide Formation, Oklahoma, but the latter species differs most conspicuously in pos- sessing fewer, thinner, thecal plate ornamenting ridges, slightly more widely spaced dichopores, no IL5-L5 rhomb and a relatively larger L1 plate. Coronocystis durandensis Kolata, 1975 is provisionally assigned to ?Tanaocystis by Sprinkle (1982b). This species differs from T. sprinklei n. sp. in apparently possessing four plates around the periproct, fewer, high, thecal plate ridges, and fewer (10 vs. 13) rhombs and demirhombs. Types and occurrence. — The holotype, UI X-5880 and all paratype isolated thecal plates UI X-5874 through 5879, 5881 through 5887 are from the lower member, Lebanon Limestone, about 14 m below the massive member at locality Z-651. UI X-5877 through 5884 are figured. Family PLEUROCYSTIDAE Neumayr, 1889 Genus PRAEPLEUROCYSTIS Paul, 1967a Type species. — Pleurocystites watkinsi Strimple, 1948. Praepleurocystis ranaformis new species Plate 12, figures 15, 18-20, 22-27; Text-figures 13, 14 Etymology of name.—rana (L.) = frog + formis (L-) = resembles (refers to the holotype theca which was mistaken for a frog by a quarry worker). Diagnosis.— А species of Praepleurocystis character- ized by a strongly-arched dorsal surface; thecal outline rounded subtriangular; large to rarely moderate-sized pectinirhombs; dorsal thecal plates ornamented with prominent, radiating, rounded ridges; B2, B3 typically contacting or nearly contacting L2, L3, respectively: LEBANON LIMESTONE ECHINODERMS: GUENSBURG 61 B1, B4, attaining periproctal margin; L1, 14 contact along narrow suture. Description.—Theca moderate-sized, 26 mm long and 21 mm wide in holotype; rounded symmetrical triangular outline as seen in dorsal or ventral views; dorsal surface strongly arched (moderate for genus); ventral surface subplanar, sides (LL, ILL area) in- curved gently with slight keel along each margin (Pl. 12, fig. 24); thecal margins surrounding periproct mod- erately wide; stem embayment shallow. Plates of dorsal surface thick, ornamented with broad, low, discontinuous, sometimes faint ridges; ridges typ- ically extend to, abut across plate corners of plate face centers; ridges can disappear toward plate centers; prominent ridges form cross at juncture of B2, ILI, IL2, L2 plates; rarely with a few faint concentric ridges on ПА or IL3 (РІ. 12, fig. 19). B2, B3 basals pentagonal, together with V-shaped Outline (as in P. watkinsi), distally contact or nearly contact L2, L3, respectively. B1, B4 lateral to, below B2, B3 on dorsal face, subtriangular in dorsal outline, articulating with B2, B4, IL1, ILS, and В1, B3, IL3, ПА, respectively; articulate sagittally on ventral sur- face to form lateral, ventral margins of stem articular facet; form proximal periproct margin. IL1 large, sub- pentagonal, borders B1, B2, L1, L2, IL5; attains per- iproctal margin along short distance because of in- clined ventral sutures with ILS (extending distally) and Li (extending proximally). IL2 large, quadrangular, borders B2, B3, L2, L3. IL3 subhexagonal, large, bor- ders ILA, B3, B4, L3, L4; with short periproctal border as in ПА because of inclined sutures with IL4, L4. ПА, ILS small, rounded subtriangular in dorsal view, form Text-figure 13.— Praepleurocystis ranaformis n. sp., plate diagram 9f dorsal face, based mainly on holotype UI X-5697. Dashed lines drawn from paratypes UI X-5698 and 5850. Plate abbreviations after Parsley, 1970, approx. x2. rounded proximal thecal angles. L2, L3 very large, as large as IL2, subhexagonal, to subheptagonal depend- ing on whether very short B2 or B3 facet present; L2 borders B2 (barely), IL1, IL2, L3, R1, R2, L1; L3 bor- ders B3 (barely), IL2, IL3, L2, 14, R2, R3. L1, 14 laterally directed on dorsal face, both subquadrangular, meet distally along short suture, form entire proximal periproct border. R1 relatively large, heptagonal; ar- ticulates dorsally with L1, L2, R2, D1, and D1’; artic- ulates ventrally with L1, O7, O1, D1’. R2 hexagonal, articulates with L2, L3, R1, R3, ОЗ, and D1; R3 ћех- agonal, articulating with L3, L4, R2, R4, O3, D2; R4 small, confined to dorsal surface, pentagonal, articu- lating with L4, R3, R5, D2, D2'; R5 pentagonal, pri- marily confined to ventral surface but forms small part of lateral thecal margin, articulates with Ld, R4, Об, D3', D2'. Distodorsal thecal border between brachioles formed by D2, O3, DI, and DI' (from left to right as seen from dorsal). Ventral distal somatic series of three plates, O7, O6, O1; O7 pentagonal, fits proximally into notch formed by L1, L4; O6 pentagonal, articulates along right distal O7 margin; O1 hexagonal, articulates along left distal O7 margin, separated from L1 by O7, R1. Narrow hydropore slit shared by O1, O6 plates; abuts across their common suture. Gonopore round, ap- proximately at triple juncture between O1, O6, O7 plates. Pectinirhombs three, distributed on, shared across B2/IL2, L1/L2, and L3/L4 plates; disjunct, el- evated, with prominent wide rims particularly on dor- sal halves (IL2, L2, L3), low rims on ventral halves. Corridor between pectinirhomb halves raised ridge bi- sected by plate sutures, plate sutures at these points convex toward dorsal. Form 11 to 16 slits per half rhomb, average is 13. Periproct incompletely known, of many (approx. 200-300) thin, small, subpolygonal plates; plates typ- ically hexagonal or pentagonal subequant away from Text-figure 14.— Praepleurocystis ranaformis n. sp. Plate diagram of distal part of ventral face (also based mainly on holotype). Dashed lines from paratype UI X-5850. Plate abbreviations after Parsley, 1970. Approx. x2. 62 BULLETIN 319 periproct margin, elongate subquadrate near periproct margin. Anal pyramid unknown. Brachioles long, apparently somewhat longer than theca, thin, gradually tapering, subround in cross-sec- tion; brachiolars biserially arranged, wider than long; coverplates biserial, wider than long, some with small intercalated wedge-shaped plates (Pl. 12, fig. 26). Stem typical for pleurocystids; long, approximately 4 ст known in UI X-5850, of proximal, distal regions; the two regions rapidly merge. Proximal region of al- ternating wide outer columnals, narrow inner colum- nals; outer columnals evenly rounded, smooth, occa- sionally with faint longitudinal ridges (Pl. 12, fig. 27); lumen large, round. Distal stem of barrel-shaped, wid- er-than-tall columnals near proximal column region, much taller than wide cylindrical columnals far dis- tally; articular surfaces with slightly depressed ring be- tween lumen and outer edge (Pl. 12, fig. 20). Remarks. — Praepleurocystis ranaformis n. sp. closely resembles P. watkinsi (Strimple, 1948) from the upper Pooleville Member, Bromide Formation, Oklahoma, particularly in plate arrangements; the latter species can be distinguished primarily by its angular thecal outline in dorsal or ventral views, relatively larger, more protruding distal ventral region, prominent sharp concentric ornamental ridges on the dorsal thecal plates; L1 and 14 plates that in ventral view are relatively wider and that mutually articulate along a relatively long suture, generally smaller pectinirhombs, and B2, ВЗ plates that usually do not articulate with ПА and IL3, respectively. Praepleurocystis nodosus Westphal, 1974a, from the Blackriveran Platteville Group, southwest Wisconsin, is poorly known but can be differentiated from the Lebanon species by its highly arched dorsal surface, nodose radiating dorsal thecal plate ridges, and the wide separation of B2 and ВЗ from IL1 and IL3, ге- spectively, in the single known specimen. Types and occurrence.—This species is represented by 19 specimens, most of which are fragmentary or buried in hard matrices. The holotype, UI X-5697, and paratypes UI X-5698, 5977 through 5979, and 5981 through 5990 are from the lower member, Leb- anon Limestone, approximately 3 m below the massive member at locality Z-652. Paratype UI X-5691 is from the Sowerbyella - Diplograptus Zone of the upper member, Lebanon Limestone, at locality Z-654 and paratype UI X-5692 is from the same horizon at lo- cality Z-666. One isolated thecal plate UI X-5851 1S from the lower member of the Lebanon Limestone at locality Z-656. Paratype UI X-5850 is from the lower member of the Lebanon Limestone at locality Z-655. Figured specimens are UI X-5697, 5698, 5850, 5983, and 5985. Praepleurocystis ranaformis n. sp. is one of the more common and widespread echinoderms in the Lebanon Limestone at or below the Sowerbyella - Dip- lograptus Zone of the upper member. No specimens have been found above the Sowerbyella - Diplograptus Zone. Praepleurocystis species cf. P. watkinsi (Strimple, 1948) Plate 12, figure 21 Pleurocystites watkinsi Strimple, 1948, pp. 761-764, pl. 1, figs. 1-3. Praepleurocystis watkinsi (Strimple). Paul, 1967a, p. 120. Pleurocystites watkinsi Strimple. Parsley, 1970, pp. 171-177, pl. 24, fig. 2, pl. 28, figs. 3-8. Praepleurocystis watkinsi (Strimple). Parsley, 1982, pp. 275-279, pl. 34, figs. 1, 5, 7, 9-11, 13-19. Remarks.—This species is represented by three the- cal plates; probable L2, L3 plates, and a fragment of uncertain derivation (its position was isolated). The (?)L2, (?)L3 plates each possess a relatively small dis- junct pectinirhomb encircled by an upraised rim. All plates are ornamented with flattened to rounded con- centric discontinuous ridges broken by prominent ra- diating ridges. The plates closely resemble those of P. watkinsi from the Pooleville Member, Bromide For- mation, Oklahoma, but plates of the latter species pos- sess somewhat sharper, more numerous concentric ridges. Incomplete material precludes definite specific assignment. Types and occurrence. — Two associated thecal plates, UI X-5893, are from the upper member, Lebanon Limestone, at locality Z-663, and one of these is fig- ured. A single thecal plate, UI X-5870, is from the upper member, Lebanon Limestone, at locality Z-653. Genus AMECYSTIS Ulrich and Kirk, 1921 Type species. —Pleurocystites laevus Raymond, 1921. Amecystis nanus new species Plate 13, figures 1-5; Text-figure 15 Etymology of name.—nanus (L.) = dwarf (refers to the small size of this species). Diagnosis.— A species of Amecystis Ulrich and Kirk, 1921, characterized by a small, flattened, evenly sag- ittate theca; proximal lobes evenly rounded, formed principally by B1, B4; thecal plates thin, smooth; dor- sal RR, OO small, compressed distally with L2 not articulating with R3; R1 and R3 small, in contact, R2 not contacting O3; outer proximal stem columnals with short spinose projections. Description.—Theca small, 9 mm long, 6 mm wide in holotype, approximately 15 mm long and 10.5 mm wide in largest paratype, evenly sagittate in outline; LEBANON LIMESTONE ECHINODERMS: GUENSBURG 63 proximal lobes rounded equidimensional, slightly ex- panded; thecal margins along LL slightly curved to nearly straight; thecal margin about periproct narrow. Dorsal surface slightly convex with shallow depression running adjacent to thecal margin (PI. 13, fig. 1); ven- tral surface nearly flat. Dorsal thecal plates thin, though thickened along thecal, periproct margin; smooth, flat Or nearly so. B2, B3 mediodorsal to B1, B4; both subpentagonal, moderate-sized; articulate with thecal plates B1, B3, Па, IL2, possibly IL5, and B2, B4, IL2, IL 3, respec- tively. B1, B4 subtriangular in dorsal view, moderate- sized; form proximal lobe angles; mutually articulate medioventrally, form proximalmost periproct margin; also articulate with B2, possibly ПА, IL5 and B3, IL3, ПА, respectively. IL1, IL3 large, elongate proximo- distally; IL1 borders dorsal thecal plates possibly B1, B2, IL2, ILS, L1 and L2; IL3 borders dorsal plates B3, B4, IL2, ПА, L3, LA. I1 2 large, hexagonal, central on dorsal surface. L1, Ld moderately large, proximodis- tally elongate; L1 articulates with dorsal plates ILI, L2, possibly R1, R2; L4 articulates with dorsal plates IL3, L3, R3, R2. L2, L3 moderate-sized, smaller than L1, L4 in holotype (UI X-5871); L2 articulates with Пал, IL2, L1, L3, R2; L3 articulates with IL2, IL3, L2, 14, R2. КК small, restricted to far distal thecal surface In dorsal view, designations uncertain. R1 small, bor- dered dorsally by L1, R2, R3, and O3. R2 small, hex- agonal, lying between L1, L4; borders L1, L2, L3, L4, R1, R3. R3 small, articulates with L4, R1, R2, and ОЗ. R1, R3 meet R2 proximally. R2 does not contact O3. O3 very small. Distoventral thecal region unknown. Periproct large, covered by many (200 +) small, thin Plates; plates typically hexagonal, arranged in poorly defined longitudinal rows (Pl. 13, fig. 3). Anal pyramid Small, subcircular, of seven wedge-shaped plates in UI X-5872. Text-figure 15.— Amecystis nanus n. sp., plate tracing of adoral Part of dorsal face (from photo of holotype UI X-5872). Some plates Overlap others slightly, approx. x6. Single brachiole known in holotype (UI X-5871); it is gradually tapering, 6.2 mm in length (slightly longer than thecal length), biserial (Pl. 13, fig. 1). Brachiolars much wider than tall proximally, becoming slightly wider than tall distally; smooth or nearly smooth. Stem longer than thecal length, typical for family, of proximal, distal regions. Proximal region of alternating outer (wider), inner columnals; outer columnals with short spinose projections. Distal region of subcylin- drical columnals, gradually become much taller than wide distally. Remarks.— Amecystis nanus n. sp. most closely re- sembles the type species, А. laevus (Raymond, 1921), from Ше Trentonian “Lower Trenton Limestone," northern Michigan, and the Kirkfield Limestone, Kirk- field, Ontario, but the latter species 15 most readily distinguished by its much larger known size, different ventral plate arrangement with L3 articulating with R3, L4 not articulating with R2, and smooth or grooved (instead of spinose) outer proximal columnals. А. woodi Broadhead and Strimple, 1975, differs in having pus- tulose dorsal thecal plate ornament as well as all the differences listed for A. laevus. A. raymondi Parsley, 1970, from the Blackriveran Shippensburg Limestone, Pennsylvania, and (?) the Benbolt Formation, Ten- nessee, differs in having angular proximal lobes, fine radiating ridges on dorsal thecal plates, and L3 con- tacting R3. Types and occurrence. — The holotype, UI X-5871, is from the lower member of the Lebanon Limestone approximately 15 m below the middle massive mem- ber at locality Z-651; paratype UI X-5873 is from the lower member, Lebanon Limestone, also at locality 7-651. Paratype UI X-5872 15 from the lower member, Lebanon Limestone, about 3 m below the massive member at locality Z-652. All specimens are figured. Class ASTEROIDEA de Blainville, 1830 Remarks.—Students of fossil sea stars have tradi- tionally assigned Paleozoic taxa to modern orders, however, affinities between the groups are not clearly established (Kesling, 1969; McKnight, 1975). None of the Lebanon asteroids, therefore, are assigned to an order at this time. Order Uncertain Suborder PUSTULOSINA Spencer, 1951 Family HUDSONASTERIDAE Schuchert, 1915 Genus HUDSONASTER Stürtz, 1900 Type species. — Palasterina rugosus Billings, 1857, p. 291; Palasterina rugosa Billings, 1858, p. 77, pl. 9, figs. 2a-c. 64 BULLETIN 319 Remarks. —Schuchert (1915, pp. 53-56) suppressed Protopaleaster Hudson, 1912, and reassigned the type species, P. narrawayi, to Hudsonaster. Schuchert ar- gued that Hudson (1912, p. 24) had misinterpreted the morphology of his specimen and that the specimen was morphologically similar to others assigned to Hud- sonaster. Subsequently, Spencer and Wright (1966, p. 050) and Branstrator (1982) resurrected Protopaleas- ter. Spencer and Wright (1966) separated the two on the basis of ray shape: those in Hudsonaster are “clearly not fused" whereas rays “tend to fuse" in Protopal- easter (Spencer and Wright, 1966, p. U50). The mean- ing of these characterizations does not appear clear, but might refer to the outline of the interbrachial an- gles. Branstrator (1982) separated the two genera on the basis of a single criterion: in adult specimens, the rays of Hudsonaster taper more rapidly than do those of Protopaleaster. Insofar as I have been able to as- certain, other characters, including ossicle shape and arrangement, are the same for both. Degree ofray taper alone is here considered insufficient for generic differ- entiation; therefore Schuchert's view that Protopal- easter be suppressed is followed. Hudsonaster species cf. H. narrawayi (Hudson) Plate 13, figure 12 Protopaleaster narrawayi Hudson, 1912, p. 25, pls. 1-3; Hudson, 1913, pp. 77-84; Spencer, 1914, p. 21, fig. 19. Hudsonaster narrawayi (Hudson). Schuchert, 1915, p. 59, pl. 1, fig. 1; pl. 2, fig. 1; pl. 4, fig. 1. Protopaleaster narrawayi Hudson. Spencer and Wright, 1966, p. USO, fig. 43a-d. Protopaleaster narrawayi Hudson. Branstrator, 1982, p. 320, pl. 42, figs. 4, 5. Remarks. — А single, small, well-preserved individ- ual, USNM 60619, was studied. The entire actinal side and parts of at least two abactinal rays are exposed. Primary measurements in mm of the specimen as pre- served are as follows: R (major radius: center of disc to tips of normal arms) = 5.2, r (mean shortest distance between disc center and thecal margin) — 2.7, Rw (arm width at the base) = 2.8. The specimen agrees well with the illustrations and drawings of the type specimen from the “Васк River" Formation at Ottawa, Ontario, except that the latter is about twice as large, has pro- portionately slightly longer, less rapidly tapering rays, and slightly more ossicles in ossicle rows. These dif- ferences are here attributed to an earlier ontogenetic stage of the Lebanon specimen. Branstrator (1982) states that many juvenile pustulosinids, such as mem- bers of the Promopaleasteridae and Mesopaleasteri- dae, could go through an early postlarval Hudsonas- ter-like stage. If true, this would cast doubt as to the identity of the Lebanon Limestone specimen; the spec- imen might be a juvenile pustulosinid. Until Bran- strator's view is tested by adequate growth series, how- ever, the assignment of the Lebanon Limestone specimen to Hudsonaster must be retained. Types and occurrence. — The single specimen, USNM 60619, is here figured; it is from an unknown horizon in the Lebanon Limestone at Shelbyville, Tennessee. The surrounding matrix suggests the specimen is from the upper member. Order Uncertain Family SCHUCHERTIIDAE Schuchert, 1915 Genus SCHUCHERTIA Gregory, 1899 Type species.— Palasterina stellata Billings, 1857, p. 290; Billings, 1858, p. 76, pl. 9, figs. la, 1b. Schuchertia darwini new species Plate 14, figures 1—3 Etymology of name.—The specific name honors Charles Darwin (1809—1882). Diagnosis. — А species of Schuchertia characterized by relatively large proximal marginals and tiny acces- sory axillaries along adradiodistal margins of axillary marginals; otherwise resembles S. stellata as far as can be compared. Description.—Size small, primary measurements of holotype in mm as preserved are: R — approximately 9.9, r — 3.9, Rw — 3.8 (see Remarks under Hudson- aster sp. cf. H. narrawayi for definitions of measure- ments). Disc moderately large with small interbrachial arcs; rays taper rapidly, more rapidly nearer disc 50 that arms appear slightly concave in abactinal view. Ambulacral grooves narrow as preserved. МАРР, Adambb, marginals, abactinals all (2) with pustules bearing short, tiny spines; spines dense, slightly larger in ambulacral grooves, oral area. Adambb, marginals, abactinals with convex rounded external surfaces; mar- ginals can have identations along sides. Ambb hidden. MAPP proximodistally elongate, with arcuate keels along adradioactinal margins. Adambb wider than long, with well-developed furrow promi- nences; equisized nearly to ray tips, then rapidly di- minishing in size. Apparent tiny accessory axillary 05- sicles at junctures of Adambb, axillary marginal, and succeeding marginal (Pl. 14, fig. 3). Single row of mar- ginals border Adambb throughout ray length except where interrupted by accessory axillaries; proximally confined to actinal surface, becoming lateral medially, extending to ray tips. Marginals decrease in size grad- ually but significantly distally; subequant in outline proximally, becoming proximodistally elongate along distal rays. Axillary marginals large; each abuts twO LEBANON LIMESTONE ECHINODERMS: GUENSBURG 65 MAPP, four Adambb, two marginals, two accessory axillaries, and abactinal; abuts abactinal along narrow Surface. Abactinal ossicles arranged in quincunx rows; of similar-sized and -shaped ossicles as far as can be Observed. Ossicle rows terminate against marginals, extend interbrachially onto actinal surface (Pl. 14, fig. 1); overlap adjacent rows abradially. Abactinals small, Outlines suboval to subrectangular, closely abutting, Not separated by significant gaps. Ratio of ossicle length to width increases, and size decreases distally along rows. Remarks. — Schuchertia darwini n. sp. is based on a single, well-preserved, slightly compressed specimen. The actinal surface is exposed showing two nearly com- plete rays, two partial rays, the disc, and the lateral Parts of the abactinal surface of two rays. It is assigned to Schuchertia based on the relatively large disc and Short rays, abactinal surface of small non-differentiated Ossicles arranged in quincunx, interbrachial arcs formed by abactinal ossicles, a single row of marginals abutting Adambb, and large axillary marginals bordering the МАРР: all are features listed in Schuchert’s emended description of the genus (1915, p. 195). The new species differs from all other species assigned to Schuchertia apparently possessing a few small accessory axillar- les between the Adambb and marginals. These acces- Sory axillaries are considered to be part of the specific diagnosis only because they are small, and do not form distinct rows that separate marginals and Adambb rows, but rather fill gaps caused by indentations along the Sides of the proximalmost marginals. 5. darwini n. sp. is similar to the type species S. Stellata (Billings, 1857) from the Trentonian at Ottawa, Ontario, Canada, but the latter species possesses rel- atively smaller proximal Inf MM, and lacks accessory Interbrachials. 5. /axata Schuchert, 1915, from the Richmondian Waynesville Formation at Waynesville, Ohio, and S. ordinaria Schuchert, 1915, from the Up- рег Ordovician Girardeau Limestone, Alexander County, Illinois, differ in having smaller more nu- Merous abactinals. Types and occurrence.— The single specimen, UI X-5844, 1s figured; it is from the lower member, Le- anon Limestone, at locality Z-656. Order Uncertain Suborder URACTININA Spencer and Wright, 1966 Family URASTERELLIDAE Schuchert, 1914 Genus SALTERASTER Stürtz, 1893 Type species. — Palaeaster coronella Salter, 1857, p. 326 [no illustr.]. Remarks.—Salteraster is distinguished from the closely-related genus Urasterella McCoy, 1851, by its more highly convex (i.e., “swollen” of Spencer and Wright, 1966, p. 071) abactinal ray surfaces and the presence of abactinals of similar size and shape be- tween the carinals and marginals (Spencer, 1950, p. 406). Abactinal ray convexity is often difficult to assess in fossil asteroids because of distortion and disarti- culation; it is therefore here considered not as useful a generic criterion as are the nature and arrangement of the abactinals. Salteraster species cf. S. grandis (Meek) Plate 13, figures 7, 10, 13 Stenaster grandis Meek, 1872, p. 258; 1873, p. 66, pl. 3, figs. 7а-с. Urasterella grandis Meek, 1873, p. 67. Palaeaster harrisi Miller, 1879, p. 117, pl. 10, figs. 2, 2a. Urasterella grandis (Meek). Schuchert, 1915, pp. 180, 181, pl. 27, figs. 6-8; pl. 28, figs. 1—2; pl. 30, fig. 104. Salteraster grandis (Meek). Spencer, 1950, p. 406. Salteraster grandis (Meek). Spencer and Wright, 1966, p. U71, fig. 64, 4c. Remarks. — Two Lebanon specimens are assigned to this species. UI X-5776 is a large, slightly flattened and distorted specimen preserving the central disc, three rays nearly complete and two partial rays (Pl. 13, fig. 13). UI X-6035 is a partial ray of a large specimen. Measurements of UI X-5775 in mm as preserved are: К = 68 (1 ray only), r= 4.5, Rw = 6.5 (see Remarks under Hudsonaster sp. cf. H. narrawayi for definitions of measurements). The Lebanon specimens are similar to the holotype of 5. (Urasterella) grandis from the upper Ordovician Richmondian stage at Richmond, Indiana, except the type specimen differs in possessing somewhat thicker paxillary shafts and less proximo- distally elongate marginals. The new occurrence significantly extends the strati- graphic range of Salteraster. Types and occurrence.— The two specimens, UI X-5775 and UI X-6035 are from the lower member of the Lebanon Limestone at locality Z-654a. UI X- 5775 1s figured. juvenile Salteraster species Plate 13, figure 9 Remarks.— This form is represented by two speci- mens. UI X-5845 is a somewhat corroded complete specimen with the disc, two complete and two partial rays of the abactinal surface exposed. UI X-5854 is an extensively disarticulated individual exposing the abactinal surface. Measurements in mm of UI X-5845 as preserved аге: R = 7.0, r= 2.0, Rw = 1.9 (see Re- marks under Hudsonaster sp. cf. H. narrawayi for def- 66 BULLETIN 319 initions of measurements). The disc is relatively large with rounded, convex rays that taper uniformly and rapidly. Abactinals typically are small; each carries a single large paxillary shaft. They are arranged in quin- cunx rows and are differentiated into carinals and lat- eral abactinals. Carinals are proximodistally elongate, slightly larger than lateral abactinals. Lateral abactinals are subequal in size. They extend along the sides of abactinal surfaces and are Y-shaped when well pre- served with pores through gaps created by excavated ossicle sides. The abactinal disc has a small non-pax- illate centrale surrounded by two circles of six, then 15 paxillate ossicles. The outer circlet contains the five proximalmost carinals. The madreporite is small, cren- ulostriate, and positioned laterally on the disc between two rays. Generic assignment of these specimens to Salteraster is based on similar shape and arrangement of abactinal ossicles, particularly the slightly enlarged carinals, which are abradially flanked by rows of abactinals having similar size and shape. The madreporite size and po- sition, and the convex arms also resemble Salteraster. The specimens described here differ principally from larger specimens of Salteraster in possessing a rela- tively larger disc, more rapidly tapering rays, and fewer rows of lateral abactinals. These differences, and the small size of the specimens, reflect ontogenetic stages. Modern asteroids show similar changes during early postlarval ontogeny. In addition, a closely-related im- mature form attributed to Ulrichaster (Urasterella) ul- richi (Schuchert, 1915) from the Blackriveran, Min- nesota, by Schuchert (1915, pl. 30, figs. 6, 7; Spencer and Wright, 1966, fig. 64, 2a, 2b) has a similar large disc, rapidly tapering rays, and reduced number of lat- eral abactinal ossicle rows. The Lebanon specimens could belong to Salteraster sp. cf. S. grandis (Meek, 1873) which also occurs in the Lebanon but incom- pletely exposed material and lack of intermediate-sized individuals precludes specific assignment. Types and occurrence.—UI X-5845 and 5854 are from the lower member, Lebanon Limestone, at lo- cality Z-651. UI X-5845 is figured. Class EDRIOASTEROIDEA Billings, 1858 Order ISOROPHIDA Bell, 1976 Isorophida species indeterminate Plate 13, figure 11 Remarks.—A single fragment consisting of a weath- ered partial peripheral rim only can be assigned to the Isorophida but no further. The rim consists of ap- proximately six circlets of plates; the plates overlap proximally and diminish rapidly in size distally. The proximalmost circlet of plates are concentrically elon- gate; the distal circlet plates are radially elongate. Types and occurrence.—The single specimen, UI X-5848, is figured; it is from the lower member of the Lebanon Limestone at locality Z-655. Suborder CYATHOCYSTINA Bell, 1975 Family CYATHOCYSTIDAE Bather, 1899 Genus CYATHOCYSTIS Schmidt, 1879 ?Cyathocystis species Plate 13, figure 6 Remarks.—A single specimen showing the distinc- tive solid canister-shaped sides and bottom of the theca is tentatively assigned to Cyathocystis. The oral surface is buried in very hard matrix and may be missing, so more definite assignment is impractical at this time. No sutures are evident and the sides and bottom of the specimen appear to behave optically as a single crystal. Type and occurrence.—The single specimen, UI X-5846, is from the lower member of the Lebanon Limestone at locality Z-651. It is figured. Order EDRIOASTERIDA Bell, 1976 Family EDRIOASTERIDAE Bather, 1899 (1898) Genus EDRIOASTER Billings, 1858 Type species. — Edrioaster bigsbyi (Billings, 1858), pp- 82, 83, pl. 8, figs. 1, la, 2, 2a. ?Edrioaster species A Plate 13, figures 8, 14 Remarks.—Two incomplete specimens are referred to ?Edrioaster sp. UI X-5847 is an extensively dam- aged theca with only distal parts of ambulacra and interambulacra preserved; UI X-5849 is a well-pre- served thecal fragment with a short ambulacral seg- ment and a few adjacent interambulacral plates. The clavate thecal shape, thick tessellate interambulacrals, and characteristic ambulacral coverplates and floor- plates easily allow assignment of the Lebanon speci- mens to the Edrioasteridae. Further subdivision within this family to the closely related Edrioaster or Edrio- phus Bell, 1976, is based on the nature of the oral frame and hydropore structure, ambulacral curvature, and details of ambulacral coverplate and floorplate тог“ phology (Bell, 1976). Only coverplate morphology was available and well-preserved in the Lebanon speci- mens (PI. 13, fig. 8); these resemble Edrioaster in being evenly curved along the exterior surface and pointed perradially, yielding a zigzag perradial suture line (see Вей, 1976, p. 296). Edriophus differs in having соу“ LEBANON LIMESTONE ECHINODERMS: GUENSBURG 67 erplates upraised along perradial margins; coverplates further have sinuous perradial margins (Bell, 1976, p. 306). The Lebanon Limestone specimens, therefore, are provisionally assigned to Edrioaster; they differ slightly from the type species Edrioaster bigsbyi in pos- sessing fine to medium reticulate plate ornament and interambulacral plates with slightly impressed sutures (РІ. 13, fig. 14). Incomplete material precludes specific assignment. The Lebanon Limestone specimens have very similar plate ornament to fragmentary edrioas- teroids from the Mountain Lake Member, Bromide Formation, Oklahoma, and referred to Edriophus sp. Cf. E. laevus (Bather, 1914) by Bell (1982, р. 297). Bell based his assignment on ambulacral curvature. The Oklahoma and Tennessee specimens extend the range Of the Edrioasteridae downward to the Blackriveran. Types and occurrence.—The two specimens, UI X-5847 and UI X-5849, are both figured. They are from the lower member of the Lebanon Limestone at locality Z-655. Class ECHINOIDEA Leske, 1778 Order BOTHRIOCIDARIDA Zittel, 1879 Family BOTHRIOCIDARIDAE Klem, 1904 Genus BOTHRIOCIDARIS Eichwald, 1859 Type species. —Bothriocidaris globulus Eichwald, 1859, p. 654. Bothriocidaris vulcani new species Plate 14, figures 4-10 Etymology of name.—The specific name honors the Vulcan Materials Company, Nashville, Tennessee, whose cooperation in this study is greatly appreciated. Diagnosis.—A species of Bothriocidaris character- ized by ambulacral plates averaging twice as wide as tall near ambitus; two perforate primary tubercles on Peripodia; ambulacrals without secondary perforate tubercles; largest interambulacral plates with one per- forate primary tubercle and up to 12 secondary per- forate tubercles. Description.—Test apparently spherical, approxi- Mately 16 mm in diameter in holotype, UI X-5842, approximately twice as wide as paratype UI X-5841. Apical system as shown in paratype dominated by five large radials. Radials wider than tall, ornamented With two to four primary tubercles, several smaller Secondary tubercles; radials barely contiguous along Narrow sutures; largest radial is the madreporite; mad- Teporite externally with crenulostriate openings. Five Small quadrate plates intercalated interradially, api- Cally in notches between radials (Pl. 14, figs. 6, 7). At least one additional circlet of small plates fills peri- proct. Ambulacra of two columns of alternating plates; ap- proximately 60 plates per ambulacrum in holotype; perradial suture a pronounced zigzag, adradial sutures with interambulacra slightly zigzag to straight. Plates much wider than tall, averaging twice as wide as tall for plates near ambitus; with two primary tubercles, one on each side of the peripodia; typically no sec- ondary tubercles, rarely can have up to four secondary tubercles (Pl. 14, fig. 9); plate surfaces with slight ir- regularities. Primary tubercles average 0.5 mm in di- ameter, separated by about 0.6 or 0.7 mm. Peripodia elliptical, low, highest apically between primary tu- bercles. Pore pairs arranged obliquely at approximately 30° to long axis of plate with perradialmost pore po- sitioned apically. Interambulacra without distinct plate rows, from one to three plates across interambulacra. Interambulacrals vary greatly in size, largest plates near ambitus, larger than adjacent ambulacrals, smallest plates much small- er than adjacent ambulacrals; shapes variable, typically subequant, quadrangular to hexagonal in outline; larger plates with subcentral primary tubercle, up to 14 sec- ondary tubercles; smaller plates typically with second- ary tubercles only. Primary spines conical, short, with longitudinal striations, secondary spines similar to primary spines but shorter, slightly narrower. Three small apparent tube feet extending from podial pores in UI X-5841 (PL 14, fig. 7); conical with concentric thin ossicles aligned perpendicular to tube foot axis; no indication of suctorial disc. Remarks. — Bothriocidaris vulcani n. sp. most closely resembles B. solemi Kolata, 1975, from the Blackriv- eran Grand Detour Formation, northern Illinois, but the latter species differs in having ambulacrals with a width-to-height ratio less than 2.0, and five to six secondary tubercles. Bothriocidaris kolatai Kier, 1982, is easily distinguishable from the new species by its up to four large primary ambulacral plate tubercles and lack of peripodial rims. B. maquoketensis Kolata, Strimple, and Levorson, 1977, from the Cincinnatian Fort Atkinson Formation is also similar to the new species but differs most conspicuously in having no secondary tubercles on the interambulacra, and radials that articulate along broader sutures. Bothriocidaris eichwaldi Mánnil, 1962, from the Upper Ordovician Pirgu Stage (F,.), Estonia, and B. pahleni Schmidt, 1864, from the Middle Ordovician Johvi D, strata, Estonia, are the only other species with two primary tubercles on the peripodial rim. These species differ in possessing ambulacrals with a much greater height-to- 68 BULLETIN 319 width ratio and a more prominent round peripodial rim; both species also differ in having no primary tu- bercles on the interambulacrals. Types and occurrence.—The holotype, UI X-5842, is from the lower member, Lebanon Limestone, at locality Z-656. Two paratypes, UI X-5841 and UI X-5839, are from the lower member of the Lebanon Limestone at locality Z-651. The holotype, and para- type UI X-5841, are partial tests; paratype UI X-5843 is an ambulacral plate. All specimens are figured. Genus UNIBOTHRIOCIDARIS Kier, 1982 Type species.— Unibothriocidaris bromidensis Kier, 1982, p. 310. Unibothriocidaris kieri new species Plate 14, figures 12-15; Text-figure 16 Etymology of name.—The specific name honors Porter M. Kier. Diagnosis.—A species of Unibothriocidaris charac- terized by up to eight columns per ambulacrum; am- bulacral plates with peripodia absent or low, faint; a single perforate tubercle perradially adjacent to single plate pore (on peripodia, if present), up to 16 pustules (12 average); interambulacrals without tubercles for spine attachment, with up to 14 pustules. Description.—Test shape, diameter unknown. Co- ronal plates thick, tightly abut along vertical to nearly vertical sutures. Ambulacra dominate corona, with eight columns of plates at ambitus; plates increase in size perradially, ambitally, lateralmost columns of plates mutually ar- ticulate along narrow sutures. Plates of each ambulacra arranged in offset rows, rows form chevron patterns (Pl. 14, fig. 13); hexagonal to pentagonal, quadrate in outermost column; largest are wider than high, smaller plates about as high as wide but variable. Large plate Text-figure 16.— Unibothriocidaris Кет! n. sp., tracing of partial peristome region in paratype, UI X-5835, x6. Radial plates (R) alternate with interradial plates (I); subinterradial plates (S) are also present. is 1.7 mm wide, 1.3 mm high. Each plate possesses a single large elliptical pore (correspond with pore pairs) that passes vertically through the test, can be con- stricted medially along major axis of ellipse; pores sit- uated toward adradial margins of plates. Peripodia ab- sent or low, faint; a single large perforate tubercle situated perradially, adjacent to pore near plate center, on peripodia (if present); plates near peristome with two closely-spaced tubercles, also near plate centers (Pl. 14, fig. 12). Pustules widely distributed on plate surfaces; larger plates with as many as 16 pustules, averaging 12. Interambulacra of a single column of hexagonal plates at least; columns originate at or close to apical region, do not reach peristome (Pl. 14, fig. 12). Plates without tubercles or spines, possess up to 14 pustules. Large interambulacral plate in holotype is 1.5 mm wide, 1.3 mm high. Peristome partly known in holotype; apparently of five small subquadrate radial plates alternating with five subequally-sized interradial plates. Radials with- out pores, articulating with two ambulacrals adapical- ly, two small (?)subinterradials actinally; small non- differentiated plates present actinally (Pl. 14, fig. 12). Remarks.—The type material of Unibothriocidaris kieri n. sp. consists of several articulated patches of plates and a few single plates that are scattered over approximately one-third square meter of a single bed- ding plane. No preserved parts are duplicated, and the sizes of the plates are comparable, strongly suggesting a single individual is present. The patches and isolated plates are each assigned a separate number, however, because it is possible that more than one individual is present. The new species closely resembles isolated plates assigned by Kolata (1975, p. 68, pl. 14, figs. 3-8) to Bothriocidaridae, genus and species unknown. The plate morphologies are remarkably similar in the two, but the latter specimens possess slightly more promi- nent peripodia. U. bromidensis Kier, 1982, differs from the Lebanon species in possessing ambulacral plates with more prominent peripodia and fewer pustules. U. kierin. sp. is the only species of Unibothriocidaris in which the peristome is known. The peristome in this species greatly resembles the one illustrated by Männil for Bothriocidaris eichwaldi, Mánnil, 1962, and reinforces the opinion of Kier (1982) that the two gen” era are closely related. Types and occurrence. — The holotype, UI Х-5832 and paratypes UI Х-5833 through 5838, and 6036 through 6044 are from the lower member of the Leb- anon Limestone at locality Z-651. UI X-5832, 5835, 5837, and 5838 are figured. LEBANON LIMESTONE ECHINODERMS: GUENSBURG 69 ?Unibothriocidaris species Plate 13, figure 15 Remarks. — А single patch of plates, UI X-5840, ex- posed from the inside only is tentatively referred to Unibothriocidaris Kier, 1982, because it possesses a single pore passing through each plate. The specimen differs slightly from others referred to this genus in possessing podial pores with distinct medial constric- tions nearly producing pore pairs, in having moder- ately inclined plate sutures, and in being larger than any other specimen referred to this genus. Without external plate surfaces exposed, the specimen cannot be assigned further. Types and occurrence.— The single specimen, UI X-5840, is from the lower member of the Lebanon Limestone at locality Z-655. It is figured. Genus NEOBOTHRIOCIDARIS Paul, 1967b ?Neobothriocidaris species Plate 14, figure 11 Remarks.— А single ambulacral plate, UI X-5839, is provisionally referred to Neobothriocidaris. The plate has a subcentral pore and one sutural pore and there- fore bordered an interambulacrum. (Ambulacrals not bordering interambulacra in this genus have one sub- central pore and two sutural pores.) A large perforate tubercle is positioned subcentrally on the peripodium. The peripodium is low but distinct. The plate is 1.1 mm high, and 1.5 mm wide. Types and occurrence. — The single plate, UI X-5839, is figured; it is from the Sowerbyella - Diplograptus Zone of the upper member, Lebanon Limestone, at locality Z-652. Class Unknown Holdfasts Remarks. — At least three kinds of holdfasts, a basal attachment plate, and a cirrus-bearing column seg- ment were collected, which could not be definitely assigned to any Lebanon echinoderm. Most holdfasts, including types B, C, and some type А specimens prob- ably belong to crinoids because associated distal co- lumnals, when present, are divided into pentameres, and the columnal lumena are stellate (Lewis, 1982, pp. 58—60). The basal attachment plate апа cirrus-bearing Column segment probably also belong to crinoids be- Cause their morphologies are similar to specimens known or thought to be crinoids. holdfast type A Plate 14, figures 17-19, 23, 24 Remarks. — This holdfast is small- to moderate-sized (approximately 4 to 12 mm in diameter) and appears to consist dorsally of a single piece of calcite. It is low to medium height and subconical with short, irregular, lobate radicles. Deeply weathered (Pl. 14, fig. 24) spec- imens showing holdfast interiors are penetrated by small tubular canals; the canals may extend beyond the hold- fast into the distal column. The column articular facet is large, averaging approximately half as wide as the holdfast; it is dish-shaped with faint, radially disposed crenulations. The axial canal as seen from the column articular facet is typically irregularly lobate; in UI X-6030, it is irregularly pentalobate. A small attached column segment in UI X-6005 is composed of thin columnals divided into pentameres with small platelets between the columnals; the axial canal is large and pentastellate. Specimens show a wide range of weathering states ranging from essentially un- weathered (UI X-6005) to nearly completely obliter- ated (UI X-6004). Identification of many holdfasts was accomplished using specimens in intermediate stages of weathering. Several small specimens tentatively assigned to this holdfast type could belong to a different echinoderm from the other specimens. These small specimens are unweathered, lack attached columns, and show only gross external morphology. They are similar to other specimens as far as can be compared. The specimens show no diagnostic features (Pl. 14, fig. 19). Holdfasts of this type, particularly larger specimens, closely match type 1A of Lewis (1982, p. 58) and may be congeneric with it. Lewis (1982, p. 59) suggests this type of holdfast was formed by Carabocrinus Billings, 1857, or Paleocrinus Billings, 1857. Paleocrinus is as yet unknown from the Lebanon. Carabocrinus thecal plates, however, are not uncommon in the Lebanon and occur at the same horizons as the holdfasts, though they have not yet been found directly associated with them. Carabocrinus, therefore, seems to be a likely candidate for the origin of at least some of these Leb- anon holdfasts. Types and occurrence.— Thirteen specimens are re- ferred to this holdfast type. UI X-6004, 6005, 6010, 6020, and 6029 are from the lower member, Lebanon Limestone, at locality Z-652; UI X-6002 through 6027, 6030, and 6031 are from the lower member of the Lebanon Limestone at locality Z-654. UI X-6009 is from the lower member of the Lebanon Limestone at locality Z-651. Figured specimens are UI X-6010, and 6029 through 6031. holdfast type B Plate 14, figures 21, 25 Remarks.—This is a moderate-sized holdfast with 70 BULLETIN 319 (?) five, large, long, branching radicles that extend into the substrate. Radicles are comprised of typically wid- er-than-tall plates that form distally alternating bi- series; opposing plates of the biseries articulated along zigzag sutures. The distal column is subpentagonal, 7 mm in diameter, and it is composed of thin straight- sided columnals divided into pentamere circlets. The lumen is pentastellate, points of the pentastellate lu- men correspond with lateral pentamere articulations. Columnal articular surfaces are symplectial with ra- dially disposed crenulae. This type of holdfast closely resembles type 1B of Lewis (1982) from the Bromide Formation, Okla- homa, who tentatively assigned his specimens to Cleio- crinus Billings, 1857. Cleiocrinus tessellatus (Troost, in Wood, 1909) from the Lebanon Limestone probably possessed a large conical holdfast and different co- lumnal articulation surfaces with flat-bottomed tri- angular notches at lateral pentamere articulation points (see ?Cleiocrinus column and holdfast descriptions). Therefore, assignment to this species appears highly unlikely. It could be that the Lebanon Limestone species Cleiocrinus laevus Springer, 1911, which has a crown more like the common Bromide species (Cleiocrinus bromidensis Kolata, 1982), is the source of this hold- fast. Both this type of holdfast and C. laevus crown fragments are rare in the Lebanon Limestone (two specimens and one specimen, respectively). However, I feel assignment to С. laevus is also unlikely. Quin- quecaudex springeri (Kolata, 1975), from the Trenton (Ottawa) Limestone, Ottawa, Ontario, has holdfast morphology similar to these Lebanon specimens (see Kolata, 1975, pl. 6, fig. 5 for comparison). The resem- blance is particularly striking in radicle morphology. Quinquecaudex or some other dendrocrinid, is, in my opinion, the most likely choice for this holdfast, given available information. Types and occurrence. — UI X-6034 is figured and is from the upper member, Lebanon Limestone, at lo- cality Z-653. It is a largely complete distal column and holdfast though much broken. Possible radicles of this type, UI X-6035, are from the lower member, Lebanon Limestone, at locality Z-651. holdfast type C (Lichenocrinus) Plate 14, figures 20, 23, 26, 27 Remarks.— These are small, discoidal, encrusting holdfasts, approximately 3 to 5 mm in diameter. The upper surface has a central depression and is polyplated with small, subpolygonal, tightly-interlocking plates. Attached column fragments are narrow, approximately 0.8 mm in diameter, and composed of pentagonal co- lumnals. The columnals are composed of pentameres; the lumen is pentastellate. The holdfasts are attached to bioclasts such as brachiopods, and arthropod frag- ments, or to firm substrates. These holdfasts are the same as those previously described as Lichenocrinus Sardeson, 1908, and type IC of Lewis (1982). Various authors (Sardeson, 1908; Warn and Strimple, 1977; Lewis, 1982; etc.) have ascribed these holdfasts to juvenile cincinnaticrinids or homocrinids. Four known cincinnaticrinid and homocrinid genera occur in the Lebanon: Columbi- crinus Ulrich, 1925, Doliocrinus Warn, 1982, Trys- socrinus n. gen., and Apodasmocrinus Warn and Strim- ple, 1977. None of the specimens of these genera were found.associated with the holdfasts described here. Adults of Columbicrinus and Tryssocrinus have larger holdfasts with relatively wider column articular facets than specimens assigned to this holdfast type but are otherwise similar (see Modes of Life under superfamily Cincinnaticrinacea). Holdfasts of Doliocrinus and Apo- dasmocrinus are unknown. Types and occurrence. —UI X-6008 and 6021 аге from the lower member, Lebanon Limestone, at lo- cality Z-651. UI X-6013 through 6019 are from the lower member, Lebanon Limestone, at locality Z-652, and UI X-6028 is from the lower member of the Leb- anon Limestone at locality Z-656. UI X-6014, 6015, and 6028 are figured. basal attachment plate Plate 14, figure 22 Remarks. — А single small subcircular basal attach- ment disc has numerous radially-disposed low septa; five enlarged septa are arranged in rough pentagonal symmetry. The specimen resembles attachment plates referred to “Podolithus” and “Disconia” (see Brower and Veinus, 1978; Westphal, 1974a, 1974b) and at- tachment plates of holdfast type 1c of Lewis (1982, p. 60, pl. 3, figs. 12, 33). Types and occurrence.—' The single specimen UI X-6032 is from the lower member, Lebanon Lime- stone, at locality Z-654 and is associated with a type A crinoid holdfast. The specimen 15 figured. column segment with cirrus attachment facets Plate 14, figure 16 Remarks.— A single column fragment has numerous variously-sized facets for attachment of cirri. The co- lumnals are nearly straight-sided and thick. Types and occurrence.— The single specimen UI X-6033 is figured, and is from the Sowerbyella - Diplo- graptus Zone of the upper member, Lebanon Lime- stone, at locality Z-652. LEBANON LIMESTONE ECHINODERMS: GUENSBURG 71 APPENDIX LOCALITY REGISTER The following 12 localities yielded the echinoderms found during this study. Approximate positions of the localities are shown on Text-figure 1. Tennessee Grid Coordinate System (T.G.C.S.)* numbers are in feet. All references made to members are to subunits of the Lebanon Limestone. Z-651: Upper 28 m of Lebanon Limestone including most of the lower member and all of the massive and upper members, and all of the Carters Limestone; Vulcan Materials Company Quarry at Una on southeast side of Nashville, Davidson County, Tennessee; T.G.C.S. 617,500 N, 1,801,900 E, An- tioch 7.5’ Quad. Z-652: Ridley Limestone, and the lower 27 m of the Lebanon Lime- stone including all of lower and massive members, lower part of upper member; M. C. West Lime Company Quarry at Turney, 6 km south of Pulaski, Giles County, Tennessee; T.G.C.S. 275,400 N, 1,684,400 Е, Pulaski 7.5’ Quad. 2-653: Upper 16 m of the Lebanon Limestone including nearly all of the upper member, and all of the Carters Limestone; Stone Man Company Quarry on south side of U.S. Highway 41 about 1.2 km east of Duck River bridge and approximately 7 km southeast of Shelbyville, Bedford County, Tennessee; T.G.C.S. 389,000 N, 1,890,000 Е, Normandy 7.5’ Quad. Z-654: Upper 27 m of the Lebanon Limestone including most of the lower member and all of the massive and upper mem- bers, and the lower Carters Limestone. Highway cut, mile 43 on Interstate 65 approximately 12 km east-southeast of Columbia, Maury County, Tennessee; T.G.C.S. 440,400 N, 1,732,000 E, Glendale 7.5' Quad. Locality Z-654a is a gray shale interbed in the lower member, approximately 3.8 m below the lower contact of the Sowerbyella - Diplograptus Zone of the upper member and approximately 3.5 m below the massive member. 27-655: Approximately 10 m of Ше lower member of the Lebanon Limestone; roadcut on Interstate 24 approximately 2.8 km north of Hoovers Gap exit (approx. 15 km southeast of Murphreesboro), Rutherford County, Tennessee; T.G.C.S. 474,000 N, 1,911,450 E, Webbs Jungle 7.5’ Quad. Approximately 8 m of the lower member of the Lebanon Limestone; roadcut on Interstate 65 approximately 0.3 km south of bridge over Duck River, Maury County, Tennessee; T.G.C.S. 426,400 N, 1,730,500 E, Glendale 7.5’ Quad. The upper part of the Ridley Limestone, and the lower 14 m of the lower member of the Lebanon Limestone; Ruth- erford County Quarry, approximately 6 km south of Mur- phreesboro on east side of Highway 231, Rutherford County, Tennessee; T.G.C.S. 499,000 N, 1,881,200 E, Murphrees- boro 7.5' Quad. Approximately 6 m of the lower member of the Lebanon Limestone; small quarry on northeast corner of intersection of Sam Ridley Parkway and U.S. 70 (= Highway 41), Ruth- erford County, Tennessee; T.G.C.S. 584,000 N, 1,842,000 E, Smyrna 7.5’ Quad. Approximately 3 m of the upper member of the Lebanon Limestone; roadcut on highway 64 2.2 km east-southeast of Shelbyville, Bedford County, Tennessee; T.G.C.S. 394,500 N, 1,877,000 E, Shelbyville 7.5’ Quad. Upper 6 m of the upper member of the Lebanon Limestone; most of the Carters Limestone; abandoned quarry of Mar- quette Cement Manufacturing Company at Martha, Wilson County, Tennessee; T.G.C.S. 666,050 N, 1,870,700 E, Mar- tha 7.5’ Quad. Approximately 4 m of the upper member of the Lebanon Limestone; roadcut at mile 54 on Interstate 65 approxi- mately 1.6 km north of county line, Williamson County, Tennessee; T.G.C.S. 497,000 N, 1,742,250 E, Bethesda 7.5" Quad. Upper 2.5 m of the lower member, all of the massive mem- ber, and the upper 2 m of the upper member of the Lebanon Limestone; roadcut on highway 231, approximately 3.0 km north of Rutherford County line, Wilson County, Tennessee; T.G.C.S. 601,050 N, 1,895,500 E, Vine 7.5" Quad. Z-656: Z-658: Z-659: Z-661: Z-662: Z-663: Z-666: REFERENCES CITED Bambach, R. K., Scotese, C. R., and Ziegler, A. M. 1980. Before Pangea: the geographies of the Paleozoic world. Am. Scientist, vol. 68, No. 1, pp. 26-38. Bassler, К. 5. 1932. 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Cletocrinus tessellatus; (Шоона. оК ЛЕ CE б AA иуи бышын + 20 | 3. Holotype, USNM 39910, ?anterior of weathered, steeply-conical dorsal cup, x 1.5. 5, 13. Hypotype, UI X-5908, partial cup: 5. ?Anterior view, showing relatively wide-flaring sides, x2; 13. Detail of inside of cup, with low, wide radial grooves flanked by pore-bearing ridges, x4. 6, 7. Hypotype, UI X-5898, x4: 6. Cup plates, showing unweathered external plate surfaces, with very fine | concentric ridges at plate centers; 7. Plate facet, showing interarticular canal (arrow) connecting with slits n from above, and articular surfaces with fine vermicular ridges and grooves. 8, 9. Hypotype UI X-5917, column fragment, x 2: 8. Weathered exterior; 9. Interior, showing attenuated areolae, and crenulate lateral pentamere articulation surfaces. 11, 15. Hypotype, UI X-5897, column fragment found associated with cup fragment in figs. 6, 7: 11. Exterior view, showing pitted columnals (?weathered), lateral pentamere articulations, x2; 15. Articular surface, showing vermiform ridges, triangular flat-bottomed notch at pentamere articulation, x4. 12. Hypotype, USNM 166816, cup fragment, becoming more deeply weathered aborally, interarticular canals exposed (arrow), x4. 14. Hypotype, UI X-5895, broken cup fragment, anterior slightly weathered, x2. 16. Hypotype, USNM 42271, cup fragment, showing transition from deeply weathered to unweathered calyx plates, x4. | 17. Нурогуре, UI Х-5900, cup fragment, showing unweathered plates, х4. | 19. Hypotype, ОЛ Х-5903, partial сир, ?anterior view, Х1.5. 20, 21. Hypotype, UI X-5902, cup fragment, x 1.5: 20. Basal view, showing CD interray bulge; 21. Anterior view. 4, 18. holdfasts of ?Cleiocrinus species ж укын куи, кзз жое. een e Ae бр nen enn e Сз... 4. Figured specimen, UI X-5914, showing subcircular outline, х1. 18. Figured specimen, UI X-5913, large specimen, showing short lobate radicles, х 1. 22-24. "tegmen of ?Cleiocrinus species ie o ir e n eea ehh hehehe e he emen 24 22-24. Figured specimen, UI X-5919: 22. Entire specimen, х 1.5; 23. Detail of small area, х 4; 24. Detail of apex | region, showing Запа! structure (arrow). Specimen photographed under xylene, х4, 24 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 PLATE 1 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 PLATE 2 Figure l-3, 6, 7. 4, 8, 9. За 10-13, 18. 14-16, 20. LEBANON LIMESTONE ECHINODERMS: GUENSBURG 79 EXPLANATION OF PLATE 2 Glelochinusaiaev us SPINE OIE Men ол К КОЕ... а о LO о аа 22 Holotype, USNM 50045, partial cup: 1. Basal view, x2; 2. Oblique view, x1.5; 3. Cup plate detail, showing fine concentric ridges, pores along sutures, х4; 6. CD interray view, x2; 7. A ray view, х2. Clerocrinus;springertmew Species mir ш o. TIEN eo; he nes eee MEE ULM Eee IE LL di 22 Holotype, UI X-5918, partial cup: 4. Cup plate detail, unweathered, х4; 8. E ray view, specimen badly corroded, x 1.5; 9. B ray view, x 1.5. Reteocrinus: polki newispeties. oo айы cx рани пе o hu А ANS а Т. e T 31 5. Paratype, UI X-5413, partial cup, showing interinfrabasal depressions (arrow), x2. 17. Paratype, UI X-5417, small crown and proximal column, anterior view, showing incipient spine-like lateral projections (arrow), small plates in interbasal gaps, x2. Reteocrinüs fenes twats new SPOe nE ли EAR E ену ONE абе ОВ ey LU asus LU d m 30 10. Paratype, UI X-5696, partial cup, proximal column, posterior view, x2. 11. Paratype, UI X-5695, partial crown, posterior view, showing Brr and RR with lateral projections (arrow), iBrr with stellate ridges, х 1.5. 12, 13, 18. Holotype UI X-5690: 12. Partial cup, column, x1; 13. Detail of cup, proximal column in posterior view, x2; 18. Detail of medial column segment, x2. Reteocrinus variabilicaulismew:species. ...... new RU SI UTIMUR EN LE 32 14. Paratype, UI X-5409, DE interray view of small crown, x3. 15. Paratype, UI X-5408, partial cup, proximal column, showing thin nodals-internodals, х 2. 16. Figured specimen, UI X-5415, crown, proximal column, in D ray view, x2. 20. Paratype, UI X-5412, CD interray view of crown, proximal column, showing supernumerary cup plates, x2. w Reteochinus-SDECIES as X D EINE cL ЖЖ noe 5» ы у с з ot Do o uM c 33 Figured specimen, UI X-5416, D ray view of partial crown, proximal column, х2. 80 Figure 1,3: 2, 4, 11. 5-10, 13. BULLETIN 319 EXPLANATION OF PLATE 3 Page Gustabilicrinus plektanikaulos new genus and species, and Reteocrinus polki new Species а empleo ea: 33, 31 1. Paratype, UI X-5791, Gustabilicrinus plektanikaulos partial crown (lower left), anterior view; and holotype, UI X-5790, Reteocrinus polki, flattened, nearly complete crown (right), DE interray view, showing distal anal series (arrow), х2. 3. Paratype, UI X-5791, Gustabilicrinus plektanikaulos, partial crown (above left); paratype, UI X-5792, Gustabilicrinus plektanikaulos, coiled distal column (below); and holotype, UI X-5790, Reteocrinus polki, nearly complete individual (except for distalmost column and holdfast), x 1. Reteoerinus polki new Species...» Ол A a OE dtque ada а E 2. Paratype, UI X-5410, partial crown, anterior view, showing distended cup plates, x1. 4, 11. Holotype, UI X-5790: 4. Detail of interbasal gap area showing small plates in gap, x4; 11. Distal column showing pentameres changing rapidly to mosaic of tiny plates, х 2s Reteocrnas variubilicaulis mew: ресіев: de ges eere onse emet tes e EST Qua бана ыж ера + IA 32 5—8, 13. Paratype, UI X-5805: 5. Nearly complete individual, x1; 6. Detail of medial column showing widely-spaced nodals and ornate ornament, x2; 7. Distal column of pentameres with holdfast, x4; 8. CD interray view of crown and proximal column, x2; 13. Interbasal gap with small plates, x4. 9, 10. Holotype, UI X-5411: 9. Nearly complete specimen (except for holdfast), A ray view, showing wide nodals, internodals compressed and not visible, x1; 10. Detail of crown, showing interinfrabasal depressions open below, x 2 31 55 . Reteocrinus species cf. R. variabilicaulis new species ..................6 nenne nennen Figured specimen, UI X-5414, C ray view of partial crown and column, showing iBrr with stellate ornament, x2. PLATE 3 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 PLATE 4 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 LEBANON LIMESTONE ECHINODERMS: GUENSBURG 81 EXPLANATION OF PLATE 4 Figure Page ДОК CLCOCHI NUS STCLIGHES SSID ps АВ ер c. ende EI 28 1. Lectotype, GSC-1525, anterior view of partial crown, showing proximal column, interbasal areas, х2. 2, 3. Paralectotype, GSC-1525b: 2. Detail of interbasal gap (photographed in water), showing small stellate plates in gap (arrow), х4; 3. Posterior view of partial crown, х2. 4 Reteocrinus-elongatus Raymond. 9999 ti ds Loc хк ши. о. le 30 Holotype, MCZ-3370, crown and proximal column, slightly corroded, showing small lateral projections, x2. 5-7, 9, 14, 15. Reteocrinus alveolatus Miller and E OY fe музды ee ст оо а И a RE... NARI ON DO QR EA Ei EE 29 5, 7. Hypotype, UC-35991 (Kopf Collection): 5. Crown, anterior view, showing proximal column, х2; 7. Detail of interbrachial area, showing pores in iBrr, x4. | 6. Holotype, FM(UC)-6045, coarsely silicified crown, x2. | 9. Hypotype, UC-34514 (Kopf Collection), proximal column with IBB, showing weathered, very thin columnals, | cryptosymplectial articulations, х4. 14, 15. Hypotype, UC-36006 (Kopf Collection): 14. Partial crown, x2; 15. Detail of partial interbasal gap, showing | plates in gap, х4. 8, 10: 1 По Gustubilicrinus:latominmnewiSpecles. ИИ A TENE 35 Holotype, UI X-5961: 8. Well-preserved ray of partial cup, showing wide, short RR and IBrr, х3; 10. Proximal column fragment with attenuated serrate latera, x 3; 11. ?Anterior view of cup fragment, x3; 16. Lateral view weathered side | of cup, showing spinose projections on tegmen plates (arrow), х3; 17. Detail of R, IBrr, showing very fine ornamenting ridges, х5. 12.213: Retepcrinus:vartabilıcaulis new DECIS еее 32 | Paratype, UI X-5694: 12. Detail of iBr plates, tegmen, x4; 13. Crown and proximal column, showing supernumerary cup plates, х2. 82 BULLETIN 319 EXPLANATION OF PLATE 5 Figure 1, 6. Gustabilicrinus latomium new species iiis eher Paratype, UI X-5962, x3: 1. C ray view, showing probable CD interray (arrow); 6. EA interray view. 2-5, 12, 13, 18, 20-22. Gustabilicrinus plektanikaulos new genus ама SPECIES! un... RED КО су to Ол ICI UE 33 2, 3. Holotype, UI X-5952, x 1.5: 2. View of flattened crown, position of CD interray unknown; 3. Reverse side of specimen. 4. Paratype, UI X-5951, damaged cup fragment showing broken BB, IBB, proximalmost column, x2. 5. Paratype, UI X-5758, partial crown, flattened, broken, lowermost cup missing, showing slightly impressed sutures, slight depressions at plate centers, х 1557 12, 18, 20. Paratype, UI X-5943, partial cup: 12. Cup, lateral view, showing large iR (lower left), x 3; 18. Anterior view, x3; 20. Detail of cup plates, showing slightly impressed sutures, faint median ray ridges, fine pitting, хб. 13. Paratype, UI X-5773, fragmentary crown with attached column segment, x1. 21, 22. Paratype, UI X-5959, coiled distal column: 21. Column wrapped around distal column of ?Trysso- crinus endotomitus n. gen. et sp., x1; 22. Detail showing nodals with radially-arranged ridges on epifacets, internodes of many thin columnals, х4. 111, 14217.19, Archacocrinus snyderi new Species ore ceeste ЙИ Ыз ныны. o e s ST 7. Paratype, UI X-5745, medial column segment, х1. 8. Paratype UI X-5744, proximal column segment, X1.5. 9, 14. Paratype, UI X-5742, partial crown: 9. Detail of free IIBrr, ШВгг, showing ray bifurcation, x3; 14. 2Anterior view, specimen partially flattened, showing faint median ray ridges, х | 10. Paratype, UI Х-5737, distal column segment, showing faint, low, node-like projections on nodals, 3175; 11, 15, 16. Holotype, UI X-5740, cup, x1: 11. A ray view; 15. CD interray view, showing two plates in primanal position; 16. Basal view, showing large basal concavity. 17. Paratype, UI X-5743, proximomedial column segment, x 119% 19. Paratype, UI X-5739, medial column segment, nodals рагйу broken away, showing symplectial articulations, х3. 36 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 PLATE 5 РГАТЕ 6 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 LEBANON LIMESTONE ECHINODERMS: GUENSBURG 83 ee nn рсе -- = ~ EXPLANATION OF PLATE 6 ) | Figure Page y 1.2, 4 Diabolocrinusspecies Ac. те АЛ E ee. en 37 | 1. Figured specimen, UI X-5767, anterior view of small crown, proximal column, х4. 2, 4. Figured specimen, UI X-5853, flattened crown, х4: 2. A ray view; 4. CD interray view. } 3; 5.16, Archacocrinus Nyder EW SPEIS па се сн ee A O m E. 36 | 3, 6. Paratype, UI X-5750, small cup, x1: 3. Basal view, showing large basal concavity; 6. Anterior view. | 5. Paratype, UI X-5741, partial cup showing CD interray of primanal followed by row of three iIBrr, x1. : 1. 105 11: Rhodocrinitid species A. СОНИН M же 37 | Figured specimen, UI X-5752, fragmentary crown, proximal column: 7. Detail of biserial free ПВгг, х4; 10. Partial cup, arms, X 1.5; 11. Partial cup, column, x 1.5. 4 SSOMBRUOGOCHBIQOKBSDOCISSEAM S И С а а AES а ee 38 | Figured specimen, UI Х-5753, much-weathered calyx: 8. Lateral view, х1; 9. Detail of well-preserved plates, showing fine i ridges and pustules, x4. | 12218 Abludoglyptochinus pregatussnew Species... ee о EE nn 39 \ 12, 14. Holotype, UI X-5964, slightly flattened calyx and proximal arms, х 2: 12. A ray view; 14. CD interray view. | 13, 18. Paratype, UI X-5969, calyx, х 1.5: 13. Basal view; 18. А ray view. \ 15. Paratype, UI X-5998, partial cup and proximalmost column, х2. ‚ 16. Paratypes ОТ Х-5996 (upper left) and UI X-5995 (lower right), partial crowns embedded in matrix, х1.5. 17. Paratype, UI X-5997, distal column segment, showing low projections (nodes) on nodals, x 2. 84 Figure 2,7 3-5, 10-13, 16. 6, 14, 15. BULLETIN 319 EXPLANATION OF PLATE 7 Cremacrinus' species: св C puncetams ШС a eins os se ne TD RUE оо 1, 2. Figured specimen, UI X-5892, partial crown, x3: 1. E ray view, showing small Е ray; 2. CD interray view. 7. Figured specimen, UI X-5891, A ray view of crown, proximal column, in matrix, showing large E ray, x2. Columbicrimis cras suc BICIS Е ee ee ee nz 3, 4. Holotype, USNM 89826, weathered cup and proximal arms. х2: 3. A ray view; 4. CD interray view. 5, 12. Hypotype, UI X-5707, crown with proximal and medial column, approximately 1 cm column segment between figures is missing, X 1.25: 5. Crown, CD interray view, showing pinnulation, proximal column; 12. Proximal to medial column, showing atypical distal taper. 10, 11. Hypotype, UI X-5705, crown and proximal column, x1.25: 10. A ray view of cup and proximal arms; 11. С ray view of crown and proximal column. 13. Hypotype, UI X-5708, EA interray view of crown and proximalmost column, showing atypical wide-flaring BB, 125. 16. Hypotype, UI X-5712, partial cup and proximal column, aboral margins of BB project slightly beyond column articular facet, cup plates with impressed sutures, X 2. Apodasmocrinus species cf. A. dauber Warn and Ѕігітріе..................... ш шын язын nn Figured specimen, UI X-5716, crown and most of column, in hard matrix: 6. Detail of crown, proximal column, DE interray view, х 3; 14. Entire specimen, x1; 15. Detail of medial column, showing striations adjacent to lateral pentamere articulations, transition in column morphology, х3. DON ocrmus montlieaulisHew.speriesern.. oot en ls ar vcre e ERN ee КК ee лата Мк. Holotype, UI Х-5716: 8. Crown and partial column, showing beadlike columnals, x2; 9. Detail of crown, A ray view, showing that iBr touches RR in AB interray, х4. 40 42 ! || | | | PLATE 7 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 РГАТЕ 8 VOLUME 86 2 BULLETINS OF AMERICAN PALEONTOLOGY ) LEBANON LIMESTONE ECHINODERMS: GUENSBURG 85 EXPLANATION OF PLATE 8 Figure Page 1. 2746-10, 17.. ColumbicrinusicrassussUlrich. s o d UNE а 44 1. Hypotype, UI X-5711, А ray view of crown, proximal column, showing narrow, steep-sided cup, impressed sutures, x2. | 2, 10. Hypotype, UI X-5713, x2: 2. Crown, CD interray view, showing narrow cup, impressed sutures; 10. A ray | view of cup, proximal arms. 4, 9. Hypotype, UI X-5704, ?gerontic individual, x 1.5: 4. D ray view of large robust cup; 9. A ray view of cup, proximal arms, and proximal column with thick and wide nodals. 6. Hypotype, UI X-5714, holdfast and distalmost column, x4. 7, 8. Hypotype, UI X-5693: 7. Distal column and attached partial holdfast, distal columnals of pentameres with faint striations (arrow), x1.25; 8. Detail of distalmost column, holdfast, x2. 17. Hypotype, UI X-5768, distal column with attached partial holdfast, x 1.25. ! 3, 5,1 1-16, 18-20. Tryssocrinus endotomitus new BENUS-ANdISDECIES We Ste o eer, RARE lios ие... 46 3. Paratype, UI X-5809, crown and proximal column, showing multiplated anal tube (arrow), supernumerary cup plates in this specimen, x2. 5. Paratype, UI X-5804, CD interray view of crown, including proximal anal tube, x2. | 11-13. Paratype, UI X-5831, cup and proximal arms, х4: 11. A ray view; 12. В ray view; 13. CD имеггау view. 14-16. Paratype, UI X-5816, proximal column, cup, and proximal arms, x4: 14. A ray view; 15. CD interray view; - 16. D ray view. | 18. Paratype, UI X-5826, pentagonal distalmost column and multiplated discoidal holdfast, х3. | 19. Paratype, UI X-5824, С ray view of cup, showing ри оп С ray iR caused by parasite, х4. ) 20. Paratype, UI X-5823, distalmost column and holdfast, х2. | | | | 86 BULLETIN 319 EXPLANATION OF PLATE 9 Figure 1, 2, 6, 7, 14, 15. Tryssocrinus endotomitus new genus and species... enn 1, 2, 6, 7. Holotype, UI X-5808: 1. Nearly complete individual except for distalmost column, holdfast, x1; 2. Closeup of crown, proximal column, E ray view, x2; 6. Detail of column segment, showing fine, faint longitudinal ridges (interval located by lines to fig. 1), x2; 7. Detail of more distal column segment (again, interval located by lines to fig. 1), x2. 14. Paratype, UI X-5819, distal column segment, showing pentameres, fine longitudinal ridges, x 2% 15. Paratype, UI X-5764, distalmost column segment, showing alternating rows of pentameres, encrusting bifoliate bryozoan, х3. 3-5, 9, 12, 13, 20. Tornatilicrinus longicaudis new genus and species ...................... nenn nenn 3-5, 12, 20. Holotype, UI X-5755, crown and most of column: 3. Detail of proximal column segment (interval located by lines to fig. 4), x4; 4. Entire specimen, impression only at distal end of preserved column, х 1; 5. Detail of crown, D ray view, proximalmost column, showing armlets оп Е ray arm bifurcation (arrows), х 4; 12. Column articular surface (position located by line to fig. 4), showing petaloid cren- ularium, small lumen, x6; 20. Detail of medial column segment, showing intercalated lens-shaped plates (arrow), х4. 9. Figured specimen, UI X-5757, ?distalmost column segment, х4. 13. Figured specimen, USNM 82273, plaster cast of “Dendrocrinus modestus” Safford (nom. nud.), showing arm branching and shape, cup shape similar to that of holotype, x2. 8, 10, 11, 16, 18, 19. Hybocrinus bilateralis new species nderi aieeaii ke eaa hehehe eher 8, 10. Holotype, UI X-5807: 8. Partial crown, column, and holdfast, CD interray view, showing coarsely pitted cup, x2; 10. Detail of column and small encrusting holdfast, x 4. 11. Paratype, UI X-5799, BC interray view of crushed crown, showing large A ray, х2. 16. Paratype, UI X-5862, C ray view of small (juvenile) specimen, showing diminutive B ray arm, mod- erate-sized C ray arm, partly-exposed large A ray arm (on right), x4. 18, 19. Paratype, UI X-5863, crown: 18. D ray view, arms folded, x2; 19. Detail of C ray arm segment, showing biserial coverplates, pitting, х6. 17. YAnomalocrinus antiquus new SPECIES... |... or Holotype, UI X-5852, basal view of partial crown, showing CD interray (arrow). See Text-fig. 7 for plate boundaries, x3. 50 51 48 | | РГАТЕ 9 VOLUME 86 > BULLETINS OF AMERICAN PALEONTOLOGY BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 PLATE 10 | LEBANON LIMESTONE ECHINODERMS: GUENSBURG 87 EXPLANATION OF PLATE 10 Figure Page 1—115113—16, 18.21, 22. Hybocrinus bilateraliinew Species. nu. пир eere e b се sR е лә. 51 1-4, 9. Paratype, UI X-5868, large calyx: 1. CD ray view, x2; 2. Oral view, х 2; 3. A ray view, x2; 4. Detail of oral area, showing coverplates, hydropore (upper center), x4; 9. Detail of cup plates showing coarse pitting, х4. 5-8, 11. Paratype, UI X-5869, calyx: 5. D ray view, x2; 6. A ray view, x2; 7. Oral view, x2; 8. CD interray view, showing fine pitting, x 2; 11. Detail of ?valvular anal structure (arrow), x6. 10, 13-16. Paratype, UI X-5867, globular calyx: 10. Detail of cup plates, showing fine pitting (compare with fig. 9), x4; 13. Oral view, x2; 14. A ray view, x2; 15. CD interray view, x2; 16. DE interray view, х 2. 18, 22. Paratype, UI X-5864: 18. Nearly complete partly flattened specimen, showing cup plates with fine pitting, x 2; 22. Detail of column and small holdfast, х4. 21. Paratype, UI X-5861, detail of proximal columnal articular surface, showing pentalobate lumen, х4, | 127197220, 24,25. Carádbocrinussspsclosuatemwt.maxmo setae ere о EU fers ens Шен e ed e e IS a 53 | 12. Figured specimen, UI X-5991, proximalmost column, and fragments of broken IBB, with thin undulatory columnals, x 2. 17. Figured specimen, UI X-5855, C ray B, showing radiating ridges, x2. 20. Figured specimen, UI X-5856, partial R, arm, x2. 24. Figured specimen, UI X-5980, column fragment of pentameres, associated with other fragments of this species, x2. А | 25. Figured specimen, UI X-5857, crown fragment, showing distorted, broken, and corroded сир | plates, х1. 19, 23. Porocrinus lebanonensis new species ......................... ira sb LL M gery y 54 | 19. Paratype, UI X-5796, anterior view of flattened crown, showing secondary ridges superimposed on primary calyx ridges, x 2. 23. Paratype, UI X-5889, IBB and proximal column, showing relatively tall IBB (for genus), х3. | | | | BULLETIN 319 EXPLANATION OF PLATE 11 Figure Page | ИФ Onpintécatdex species isn S300 pb A LM AEn. ТУО. о ЕСИ ARA. 55 | Figured specimen, UI X-5700: 1. Partial crown and column, showing columnals of pentameres, x1; 2. Detail of cup, proximal column, and proximal anal sac, x2; 11. Detail of distalmost preserved column, last preserved columnal is disarticulated, showing symplectial articulation, х2. 3-5 - BOVOCTHIBUS LEDANONENSISTIG WISE ебет A: Aii NT ePID, ЕГ АЛИКЕ ques ks И А У А acd T. o OUR 54 3. Paratype, UI X-5800, partial cup, attached column segment, showing secondary ridges superimposed on primary calyx ridges, x2. 4, 5. Holotype, UI X-5803, flattened crown, proximal column: 4. Detail of calyx, D ray view of proximalmost column, showing faint secondary cup plate ridges, x2; 5. Overall view, showing long slender non-branching arms, х І. ба биршостипе врестевс и С. gracilis (НАШЕ 22... A е 55 ) 6. Figured specimen, UI X-5701, anterior view of crown, proximal column, x2. | 7. Figured specimen, UI X-5703, partly disarticulated crown, proximal column, x2. | o S CUP UIOCFINUS: SDCCIC uma AR ое ы кг UNO пир а га у ds 56 Figured specimen, UI X-5702, crown fragment, х2: 8. Е ray view; 9. ОЕ interray view. 10; OKlanomacystisstrigonis ILE WASIC CLES”, И oe eee Red а OA EEE. а лес 56 10, 17, 18. Paratype, UI X-5936, small theca: 10. Posterior view, x2; 17. Anterior view, x2; 18. Dorsal view, showing pustulose ornament, large periproct opening, x3. | 12, 14-16. Holotype, UI X-5922, theca: 12. Dorsal view, x2; 14. Posterior view, x2; 15. Detail of dorsal area, showing | gonopore (ей arrow), hydropore (right arrow), pustulose ornament, x4; 16. Anterior view, х2. | 13; 19, 20; Oklahomacystis:species ай, O. frigonis new Species РН ee SN UE ee UE 58 Figured specimen, UI X-5938, damaged large partial theca, х2: 13. Anterior view; 19. Dorsal view; 20. Posterior view. | PLATE 11 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 PLATE 12 | | | | | LEBANON LIMESTONE ECHINODERMS: GUENSBURG 89 EXPLANATION OF PLATE 12 Figure Page = See OK авотасу STS, tritonis new Spoo rea ER Ome es me re walls cack wale we 56 1, 2. Paratype, UI X-5932: 1. Crushed, locally corroded, but nearly complete individual, x2; 2. Detail of column attached to probable holdfast, x4. 3, 4. Paratype, UI X-5920: 3. Theca with brachioles, partial column, ?posterior view, х2; 4. Closeup of column, x4. 5, 6. Paratype, UI X-5934: 5. Crushed theca, anterior view, x 2; 6. Detail of periproct area, showing covering plates, x 6. 7, 8. Paratype, UI X-5931: 7. Juvenile partial theca, column, anterior view, x2; 8. Detail of thecal plates, x4. 9S IA 6:17 Tandocpstis; sprinkle DONE E tn n eR EIER T Уз А E. 59 9, 10. Holotype, UI X-5880, x2: 9. Side view of partly disarticulated theca, showing the three plates that surround the periproct; 10. Another side view, with L1 (right center). 11. Paratype, UI X-5877, L3, lower left margin broken, x2. 12. Paratype, UI X-5879, L3, ?irregular demirhomb (arrow), showing broken lower left margin, х2. 13. Paratype, UI X-5881 (left), probable IL3, paratype UI X-5882 (right), IL2, x2. 14. Paratype, UI X-5884, R3, x2. 16. Paratype, UI X-5883, R2, x2. 17. Paratype, UI X-5878, B2, upper corner broken, x2. 15, 1852019222017 Praepleurocystis.ranaformis New SDCCICS. солото са iio choris esi user E Nt алма лар 60 15. Paratype, UI X-5983, patch of periproctal plates, x 4. 18—20, 24, 25. Holotype, UI X-5697, well-preserved theca, x2: 18. Right lateral view; 19. Left lateral view; 20. Proximal transverse view; 24. Periproctal view; 25. Rhomb view. 22. Paratype, UI X-5698, rhomb (dorsal) view, theca distorted, partly disarticulated, x2. 23, 26. Paratype, UI X-5850: 23. Partial specimen, showing nearly complete stem, x1; 26. Detail of distal periproctal face, showing gonopore and hydropore slit, x3. 27. Paratype, UI X-5985, partial theca, proximal column, showing longitudinal grooves and ridges on columnals, x2. Dl PraepIeuroGy stis SPECIES o E Pe Warsi (боре) SUE I ele ese NU OD оа, 62 Figured specimen, UI X-5893, possible L3 thecal plate, x2. 90 Figure 1-5. a TALLOS BULLETIN 319 EXPLANATION OF PLATE 13 тесуге nanus DEW SOECES sant dai ТОЛОО los dE ОА i 1, 5. Holotype, UI X-5871, nearly complete small specimen: 1. Dorsal face, one arm preserved, х3; 5. Detail of posterior theca (for plate designations, see Text-fig. 15), x6. 2. Paratype, UI X-5873, periproctal view: Proximal column showing nodose outer columnals, and proximal portion of theca; most of theca preserved as mold, x4. 3, 4. Paratype, ОЛ X-5872, partial theca, stem: 3. Periproctal face, distal stem reflexed toward theca, х3; 4. Detail, showing valvular anal structure, nodose proximal stem, x6. IA улл SEG sch eS, дч A О Figured specimen, UI Х-5846, lateral view of canister-shaped theca, x2. Salteraster species ch Si grandis (Meek) eo. „юке сипи Body на ee ее не а Figured specimen, UI X-5775: 7. Lateral ray detail, showing abactinals with long paxillary shafts, x4; 10. Detail of actinal surface, dismembered ray showing ambulacrals and adambulacrals in transverse view (arrow), X 4; 13. Entire specimen, showing primarily actinal view, x1. ‚ UE Abioaster species-A.... а Жу... ИРА mies И 8. Figured specimen, UI X-5849, thecal fragment, showing ambulacral coverplates in place, x3. 14. Figured specimen, UI X-5847, extensively damaged theca, showing coarse reticulate ornament, x 2. . juvenile овај егам е species. сос се и nop we cher ieu Br ТӨКК Ў Figured specimen, UI X-5845, abactinal view, x4. = 1soropin а зреле ве Пол тате ea. io a А u Figured specimen, UI X-5848, partial peripheral rim, x 35 . Hudsonaster species cf. Н. narrawayi (Нийвоп)......................++ emet Figured specimen, USNM 60619, actinal surface, x4. SUD O T A a e a. ae К ОО Am. casis naso обр na то шумо У А Figured specimen, UI X-5840, inner surface of ambulacral plate patch, pores constricted, almost forming pore pairs, x4. 66 65 66 65 66 64 69 | | | | | | | BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 PLATE 13 PLATE 14 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 Figure 12-15. 16. 17-19, 24. 20, 26, 27. 2253 22: Do LEBANON LIMESTONE ECHINODERMS: GUENSBURG 91 EXPLANATION OF PLATE 14 SS CHUCHE NEL A Arin ON SDE CEIA Lo RA туе ыза не ы CE E а 64 Holotype, UI X-5844: 1. Actinal view, x4; 2. Lateral view, х4; 3. Detail of oral area, showing tiny accessory axillaries (arrows), x6. : г Вотвтостайтезиш сити DEW SPECIES" na t S Ecco Lis tM Oe NE RM E и UM оу” 67 4, 5, 8, 9. Holotype, UI X-5842: 4. Partial ambulacral column, x6; 5. Partly collapsed corona, showing interambulacrals with single primary tubercle and many secondary tubercles, x4; 8. Madreporite, adjacent ambulacrals, x6; 9. Loose ambulacral plate, showing secondary tubercles on right, x8. 6, 7. Paratype, UI X-5841: 6. Small slightly corroded specimen, apical view, showing madreporite (arrow), x 4; 7. Detail, showing irregular arrangement of interambulacrals with up to three plates across interambulacra, possible tube foot (arrow), x8. 10. Paratype, UI X-5843, typical ambulacral plate from near ambitus, x8. SOHNPODOTHTIOCIQ ЕО вел ган OUR S Eoo QUIE QU ER Ee M M CK 69 Figured specimen, UI X-5839, ambulacral plate presumably from along interradial suture, x8. Omborhtibcida is Kiori DEN species ык ХШ OO cocer EO egies E ERE XE USO RU 68 12. Paratype, UI X-5835, large patch of plates, showing partial peristome, two tubercles on some plates, no inter- ambulacrals, x 6. 13. Holotype, UI X-5832, patch of plates from near ambitus, showing non-perforate interambulacrals on right, x6. 14. Paratype, UI X-5838, patch of plates from near apical region, showing interambulacrals near right edge, х6. 15. Paratype, UI X-5837, patch of plates from ambulacrum, showing perradial suture, constricted pores, x6. tolumnssegmentwith-cirtussattachment fACOtS a. ser DM oc d ОО WIE У EE 70 Figured specimen, UI X-6033, x2. holdisBtyperAt ae m d C m E lu ii. mic MIN I оС 69 17. Figured specimen, UI X-6010, deeply weathered, exposing tubular canals, x3. 18. Figured specimen, UI X-6031, unweathered, medium-sized, x3. 19. Figured specimen, UI X-6030, unweathered, small, x3. 24. Figured specimen, UI X-6029, deeply weathered, large, x3. holdfastety pe С (ИЛО пи з son тех X S. сена и We cL уугу E ARS 70 20. Figured specimen, UI X-6014, typical, small, multiplated, discoidal, with attached narrow column of pentameres, х4, 26. Figured specimen, UI X-6015, adjacent to articulated bifoliate bryozoan holdfast, х4. 27. Figured specimen, UI X-6028, attached to ostracod carapace, x4. ШОТАЕВ areas Race c MI a M MEM ON ее es sic aUe es 69 Figured specimen, UI X-6034, x3: 21. Lateral view, showing broken radicle (right), and columnals of pentameres; 25. Ar- ticular surface with pentastellate lumen, cryptosymplectial articulation. Dasaleattachmemt: А ы a око». ТЫКЕ csc се оно te M d mM a M ci. 70 Figured specimen, UI X-6032, large septa forming pentastellate pattern, x 2. noldtasttypeseasand-G odes Nu ее с Cte eens етер иде ae ee en LE E e 69, 70 Figured specimens, х 2: UI X-6005 (center), по аз! type A, well-preserved specimen with distalmost column, showing tiny platelets between columnals; UI X-6004 (left), holdfast type A, deeply weathered, broken; UI X-6006, holdfast type C (right, arrow). 3 BULLETIN 319 EXPLANATION OF PLATE 15 Figure Page 1. Interface between organic-rich dolomite clay shale interbed and poorly-sorted biocalcirudite, showing possible truncated ostracod, WIN ALLOWS) om backoroimd то show contact locality Z-654a, XIS -ceiro 2m, hne op: DU T Ui 14 2-6. Lebanon Limestone Шо астев photograpned.m. plane helt, < 15... „arme кизи екет жу» +.» Ж e шр кын a 10 2. Brachiopod, bryozoan, echinoderm, biocalcirudite (lithofacies 4), bioclasts typically well-rounded, often with micrite envelopes, mainly spar matrix, upper member at locality Z-651. . Poorly-sorted brachiopod biocalcirudite (lithofacies 4) with lutite matrix, spar-filled umbrella structure across center of photo- graph, Sowerbyella-Diplograptus Zone, upper member at locality Z-653. . Moderately well-sorted pellet biocalcarenite (lithofacies 1), massive member at locality Z-660. . Oncolite in poorly-sorted pellet biocalcirudite (mixed lithofacies 1 and 4), upper member at locality Z-651. . Laminated biocalcisilite-calcilutite (lithofacies 2). Sowerbyella-Diplograptus Zone, upper member at locality Z-654. | | BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 PLATE 15 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 86 PLATE 16 LEBANON LIMESTONE ECHINODERMS: GUENSBURG 93 EXPLANATION OF PLATE 16 Figure Page ШЕ ЕРА ОЛЕШ Св {One eX DOSULES ти заслон P M ы MM oni ыо а ис шым мо ee o 7,8,10 1. Lower member, thinly-developed massive member (arrow), locality Z-654. 2. Quarry floor exposing single bedding plane of megaripples (small scale dunes) just above massive member, crane (for scale) approximately 14 m long, locality Z-653. Prominent megaripples occur at an approximately equivalent horizon at locality Z-654. 3. Calcirudite-calcarenite lenses in nodular weathering calcilutite: note abrupt lateral margin of lens (arrow), lens cap for scale is approximately 7 cm long, locality Z-652. 4. Approximate level of Lebanon Limestone-Carters Limestone contact marked by head of hammer (arrow). BULLETIN 319 INDEX Note: Page numbers are in lightface, plate numbers are in boldface type. Abludoglyptocrinus Kolata, 1982 13,17,18,39,40 CROTILONT (ОВАН NS IS) ee ee 18,40 Тера E. o ies Cire eee aes 5,13,18,39,40 TSIEN GUUS: ROWLEY ТОО а а р bene en. TRES 40 ОЕЕО СОАТА 1982 А 40 БАУ А е 40 pustulosus (Kolata, 1975) 0) AOV SONUS арг [963 мл кеши а 40 Acanthocrama Williams, 19043 ааа. D duse ono о NM PNE M Y 55 Ава (oral corn 1980) Arne 21 alveolatus, Reteoerinus ...... een dr 19,26-27,29,31 Amecysus ат and Kirk, 19917 АИ на. 18,62,63 lievos Raymond ВОДЕН nen. en 63 MOROS MESA Goh ысыка НЕ e 5.13,62.63 PAV ONG PI AESI O ое 63 wood! Broadhead and Strimple, 1975 2 АИ, 63 Amygdalocystites КОЙСО ВАСЕ 1854. 2.0.20 0 en ен. 100 tribrachiatus Bassler, 1943 ПОША ОСОО ЫЕ Anomalocrinus Meek and Worthen, 1865 ........................ 48,49 ihemvus Meek and Worthen, 1809. сала acces oe voy ва vie 48,49 ?Anomalocrinus antiquus n. SP. .................... Ө, 5,13,48,49 Anthracocrinus Strimple and Watkins, 1953 ........................ 33 primitivus Strimple and Watkins, 1955 ANDUNI, LANOMAIOCHTNUS е Apodasmocrinus Warn and Strimple, 1977 ............. 18,40,41,70 daubei Warn and Strimple, 1977... an... 18,40,41 sp. cf. A. daubei Warn and Strimple, 1977 ....... «Ды Кыа 13,16, 40,41 Archaeocrinus Wachsmuth and Springer, 1881 ........ 13,18,36,37 DUCKJIO ensis Колата 198 2 22.2.2 ies suds инал, л: 37 14 1117120731. a eee одеа. NOT o I2 P5 бе od аре subovalis Strimple, 1953 ... ETS: GUCTIGCIINUS Sr De sors MER EC века NE oC S NUS A УВО ТА BS, ROO лке н o dip an Вашае Scotese and Ziegler (1980)... nn 6 абава астен рате san een 14. 3 70 aola onmi йде cum Парна AE ad ale ОТ. Bassler (1935) Bassler (1938) Bassler (1941) Bassler (1943) Bassler (1952) Bassler and Moodey (1943) Bather (1899) Bather (1914) vs Bell ДӘ) ls Ed E. ВЕ ыо) жле a е скн тн IO CICLO SU) A Ча ыры акы ы ача солу E е Вей (БОК rex Gas aan Вешие с (1994) A б ооо Bennedict (oral commun., 1981) Бейле and Wakor О) sans tae eae coh ehe 14 Dibrachtalus, ORIANOWUACY SUIS nennen ehren 56-58 ПРЕРАНО O be cs E rue pP Fe SUE RES 66,67 Dyoeralis, IIVDOCHMUS S rod ees ОЛО. 5113530951552. Billings (1854) Billings (1856) ... Billings (1857) ... 5,15,17,19-21,40,51,63-65,69,70 ТИ О снн ы ок и ы ы егы 5,13,52,63,64,66 Billings (1859) 5,13,15,19,21,23-28,31,36,52,53 Blackriveran (Black River) Stage ................ 5,7,16-19,21,23,27, 31,32,36,37,40,41,53-56,62,63,67 de Blainville (1825) 12,16 de Blainville (1830) = са ix UE 63 Blakes(lO 7) e edt н т QU а 19 Blake (oral commun. 19 S019 Scenes. 5 0 8 secretes cae 19 BMNH (Burpee Museum of Natural History, Rockford, IL) .. 20,31 Вобсауреошошлевроте ара зае IEEE 14,17,30 Bothriocidaris Eichwald, 1859 uu ee er t 18,67 е спите Мати 9007 gore utes vy deri е EE 67,68 SIODULUSSEICHWAI sel SOO E iu t ace ae cree Did S 67 kolari Kiet TIRI лиг cR ME E см меда T rM 67 maquoketensis Kolata, Strimple, and Levorson, 1977 ........ 67 Рао 18645 А и ул. ы ee 67 SOLEIL КОА а АКО Жоли азу Карр ы Кылык ee 18,67 ИШ Ер зоом леу ы ме АН 5,13,18,67,68 Branstrator ОВ в ИК bl tU а on TE 64 Brett ang Б айе ӨЛӨ И и си петина EROR 14,24 Broadhead ana Staple (TIIS) X EE ЛЕ лы 63 Bromide БОА ОМА Уул. ЕЯ 1751812152370 Mountain Lake Member ......... 17-19,37,40,44,46,52,53,57,67 Upper echinoderms Zone на л. TM 57 Еоовуше Member ea. bromidensis, Сеооа Unibothriocidaris Brower (1974) Brower, Lane, and Rasmussen, 1978 Brower and Venus GITA) er 5,17,18 Brower aid Venas (1978) ата tees S 5,17,41,55,70 Brower and Vemus ОЗ) о 17155 ОНССПОГПО ПУНА ТОНОВЕ ен путањи кран лы р T eat te 37 Butts II) палата ЫА. КАКША ОЛИ а СООРО. 53 Campanulites tessellatus Troost, 1849 (nom. nud.) ............... 20 Camp Nelson Formation Canada: се кеу een За QUE А со avin. enol tae орела ЛАС а крк ы И E Моне пода евра ин er ын E OD 52 Ontario .. 17-19,21,23,24,27-29,40,52,63,65,70 Kirkhieldiareae Кыды ачыса een. 5,14,17,30,40,63 нама аса а 52940520600 Caraboerinus ВШ РЕ WES Ва S 15,18,53,69 ПОПИО Боо fcr cies лн уназ пен Ре С o АИИ LS Le жас кси уа е TN По treadwelll:Sinelammal 9450, за тура eeu UTR Gr RE (CATOZA (ОЙЛ нө бл йи укны ка уа казыла EUREN Ceramoparela TaN, 51 u... een СОКЕ ООС ENS арии И (на рана ЗОдан e ite ери er charltoni, AVDINGOSINDIOCHINUS Erin е ОЛ ООЗЕ и к еы нне ө AE E cscs cau окоо ло ated Спагуаш (Chazy) Эре а LEBANON LIMESTONE ECHINODERMS: GUENSBURG 95 CINE al La ha VISIO) 112 Mr oan pm Use ины E 52 (Сана елитни ТОЛСО T T 59 (emeimngdeATohem e aici дни ан сат ENS 6 Cincinnaticrinus Warn and Strimple, 1977 .......................... 42 аалнапеп OMNGUECAUR ER T T IT S DS эш (CH SD) ВИ A uude E 20 Gleiocrinus Billings, 1807. ee 15,18,20-25,53,70 ОТООСУ Kolata, 1982 ЕЕ. 18,21,23,70 libanus Safford, 1869 (nom. nud.) ... ает у Springer TI на TI OT OUS BUNDES TESI N себе RR Y 21532 AVAL IS De EG UN EE 21 Проте Ку ЫНЧАН TITT Ж REM 2123 regius Billings, 1857 а БОЙУ SPIN POL SLOT MER 2122 Сани АЕ reer a RR Дз 23,24 SDEUISCHISD SD E E ee Dee 5,13,21-23 tessellatus (Troost, in Wood, 1909) ..... qe 12,13,20-23,70 Cobo une SUOSTHPE КЕ Л Л О UM RO 16 Columbicrinus Ulrich, 1925 ..................... 5,13,15,18,43-46,70 ASUS LEC Oa) etre tot TS 5,13,16,18,28,42–46 Sulphurensis (Frest, Strimple, and McGinnis, 1979) ...... 16,18, 44,46 column segment with cirrus attachment facets .......... ал 70 conicus, VE A аи аи рана анана 51 ПОСТ О л O Бо 58 Conotreta Walcott, 1889 12 Фопга (ка ee EN S 12 КОНТА BASE en ПАДЕ (SODIUM ыны a IN То T516 осе (OSG) utra ER UNE 7,11,12,16,17,19 Gore Ra a FOI ETH eg i etl ue ce ER. 65 Coronocystis durandensis Kolata, 1975 ............................... 60 A e ee Craig and Hallam (1963) Crassus, Columbicrinus ................ курше cartier эне аза ВЕ ce лак пина Sp. cf. C. punctatus Ulrich, 1886 ... (Сеоне NOES (LOG meen ohn ТН Tra SSA ta ee ннан 12 Gorino erniet КЛА te Sia EO 6 Cupulocrinus d'Orbigny, 18495 2 18.55.96 AQUA EM S "t “17, 18,55,56 Пеев оО о и ар 19 ПОЛОВЕ ЛАКА TOTI ОВЕ 56 SD мед A ВА EO SS еч АКЕ ы 13,56 SD Cie ОООО ОЕА ИВА) СИЕ 13,55,56 (Сре от Schmidt ТЕТО А TS 18,66 С ОУ Ср ОЛ O ВИ ата їз 13,66 Daedalocrinus Ulrich, 1925 Dalbyan Stage Dalman (1827) T ТЕ ат Пи тук МР они darwini, Schuchertia ща daubei, Apodasmocrinus ............... daubei (sp. cf.), Apodasmocrinus ... Deocrinus Hudson, ТООЛ А BERIO EN о M Ие Dendrocrinus gracilis (Hall) eec * Dendrocrinus modestus" Safford, 1869 .................... OF est: 21 EDV ESSUS ООС E I DE OSE 26,28 Diabolocrinus Wachsmuth and Springer, 1897 ............. 17,3738 oklahomensis Kolata, 1982 коте авина отоо МИЕ ET ADOLOCIINUSSDieAm ы ти Юс 13.37-38 Difficilicrinus Frest, Strimple, and McGinnis, 1979 ............... 44 Diplograptus multidens Elles and Wood, 1907 ........... 8,11.12.16 СОТА ee ee 70 DOE ОНУ COOP OS Wii a 12:15: 15 (Но sostiene fice О РА ооа ЫЕ преходни 16 оленонагех Ово ЧОВ nenn 12 НОСИ IN ALE LOO: Шеше нон Са E S 18,42,70 ШОЛ ЛООНУ ЗЫ, созсо ылу ые: E схе 5,13,42,43 ПА Ма LIED ee вина АНИНА ig o 42,43 торат оо е. 10 durandensis, Coronocystis 60 Dodo NR ió me sinet. Dybowski (1877) SEN Т т е сәен а ыды a Edgewood Limestone es де ы PU OUSE BINAS Еа 18,66,67 ПОПА ВУ Шун: Нес жаран Оша камео налога тже акы NI OE 66,67 РОАН УМ ш а осе IEORIODNUSIBELBLO On ne sp. cf. E. laevus (Bather, 1914) ............ = ЕТЕН SD ee en СТОН ИВО СУ "elo ини Узай ere керди aio (нации ШОККО КОКО хл а RA ы E ENES оса ПОШУ ee оо а ла elongatus, IRCLCOCTINUS nern. O ы ec RE ыс ыс се lectins узе о. КИШИСИ (Le УЛУШ ee ыу ыы INCAS oet pute КӨМ re E ete ceu cere Ыы: endotomitus, Tryssocrinus .............. оеро SWA TITI и en SCAT ODONE AUS АВ ра а tte eee ГОВИ EC T ЧЫ, MAG eae talons КОХ УРАП SE tee ee Fay and Graffham (1969) DIES DEA ANS AREA анына аза АР qut ide ЫЗ бурча СУНА fenestratus, Reteocrinus ... ESA emer e florealis, Amygdalocystites FM(UC) (Walker Museum Collection, Field Museum of Natural History, Chicago, IL) Gets ОУ ЕЕ ее ROBA ТА Um Ваше, TOAT toe ат аа ТОБОН Е О lewis kc, oe рр QI OUD ser ee Сш Fort Atkinson Formation CAC on ИИ ОСУИ ы Decorah Subgroup ...... Гон утен a oe eg Gaurocrinus Miller, 1883 ... SALE CAS ПОЛА es moat Ал л ы е Girvanella Nicholson and Etheridge, 1880 glabellus, Quinquecaudex Glenwoud РОДИТИ СОЛ Е ce TG 16,17 96 BULLETIN 319 ODIA EB OUT FOCI ANUS ay Nr инт пр ере elie Ee eL ee 67 Glyptocrinus СОЛГО О ТАТА OS ТИ Loon Lie ee | 39 ТАСИРИ LEII ае inate de ктык. cateo 36 gracilis, Cupulocrinus Dendrocrinus Poteriocrinus на AVISA СИРИ ОСИ rennen АМЕ cee 13:35:56 grandis, TIA ren, Aer (КИК diim ace AMI RU E 65 MELIA o deas арр NN RID RO eer Per LTS Fe 65 HESSEN уло oco Tea ч uum Жор EEN AA d 65 gandis (SD. СЕ), Salteraster..... eee eene ee 13,16,65,66 (оса И to «c cue quc N O 12 gregatus, Abludoglyptocrinus .................... Ore. es 5,13,18,39,40 EE) E ыы DONDE UM LM CIA MNA M 64 GSC (Geological Survey of Canada, Ottawa, Ontario, CA A И retire NEÓN 20,29,81 БТС Me Рей. s etre cere terree ree emen oett 5,13,15,16,33-36 VOL DAD les od DR qe CM US NU E 4552. 5,13,34-36 Ni D, SD енини rare prn SEM 5,13,33-35 BIOS A шыруы н жәй пава н MESAS E ee СЫ ER DM ЫРУ ОДА е 12 iS lu A AN ee ae 119, EE AR ыз A ES 12 PAV TBS E OE EE cable DN oo аа hc DU EE 65 E КО ee 33 ELCTCOCIIIUS Hudson, ТОШУ еее у паун ae penas cat an 33 Herpetocrinus Salter, 1873 49 Klesperortins:Schuchert And Cooper, ТЭЗ 12 heterocostalis, Scyphocrinus Еее (19638) u.a LHotstetier (1965D) „u... c err IUIS DEAS rece eere НОНИ ento pete ME bud ыа M енын ы А га holdfast type С (Lichenocrinus) Homotrypa Ulrich, 1882 ................... Ша ПОО nn an an таа Авена ea ale ES inician A Бад vem tI Me MK tas cM EID RA Је о а 0 A И boves e EE LEO LIA TU CREE Hudsonaster Stürtz, 1900 ....... narrawayi (Hudson, 1912) зр. cl. H. varrawayr (Hudson, 1902) „2... AI o ЈАПАН IERI S ОНЫ eek А а E A EU 30 РОСЕН SCO ALD эы, a EE IPEA 123, SUR ЖЫ DUTIES V SD мыз. cus ae ПА ДЕТИ 35:09:35: 515577 conicus Billings, 1857 ... S ПР S Boy път задаване споени UN FU TUS IUE: VES 57 РЕТИ БИШР, О ине НИЕ аис nera 52 punctatus (Miller and Gurley, 1899). 2... з. оюнну з 17 ODA MA 1999. css аео er МАНИ НИ ПР Уй гуа 92 Ibexocrinus Lane, 1970 аи и га Illaenus Dalman, 1827 НОТЕ nenne Wa Euler СШ сел дн Бре О S наи En ыы 65 INCHFYUS, ATIOTAAIOCIINGS. . ce desse nn sn eto horto ero ro oae into КАК Ери Indiana, Richmond Ingram (1953)... insperatus, Abludoglyptocrinus TSW enn d MK NE E Ра ба TEEN 14,16,54 Isorophida, species Indeterminate nen: 13258 15 оле РЕЋИ А ОДИ ОК po а алк а E 12 ISOLOMMOGHINUS IA CHy ПО Л Өде Дл та е 42 ЕУ О (LITO) азуы уу ет а ee Таеке! (1899) Jaekel (1900) Таеке! (1918) Jessamine Dome jewetti, Cupulocrinus ... JONA Stage coo ren en ee et ПРВА ША ee a DU А АЫ ee И аа Јотаа ео ee ULT а ae erase рд Kent арена ни ee Kesling (1968) Kesling (1969) мү Та s иа KeslingandsBaulel 968) Sr ED 54 FEUI) аа Пепа nope nen ee ges 5,17,19,67-69 kidi TAO ASAS we ТАБУ 5,13,18,68 ST SACA На 40 Kirkfieldian (Kirkfield) Stage .............. erre 16,21,30 Kalte! deletirestOlep E E SEE E 17,30,63 Кет У 0406 Ser Nescis а ОА 67 Коста ШОУ ДЗ) экеи те 5,17-19,25-28,30-32,39-41, 53,55,56,60,67,68,70 KOA Кы н шишүү етсе ТӨ 13,17,21,23,26,36–40,70 Kolarar(oraleom mu LIS ee 18 АДА BOMO AN cea ы ы een re 67 Kolata, Strimple, and Levorson (1977) .............................:: 67 lacunosus GLY DIOGIINUSU es venas е en EL E CDS 36 laevus, : AINEGC) STIS” И CO арка данъка ке ое (С О ООШ ША Уи T S ТЕ Pleurocystites EVS SO. Ch) BAOU A SE e o iu LE 67 ДАЛ ЛО ЛО ee ык а оа аа LU E E 50 UNA AS ААО УР ОСТИ 40 latomium, Gustabilicrinus с... з. ДЕ 5,13,34-36 VIANA AAA RA CA E LEE 65 ЕДО @arters Contarini Пре улс len 7 lebanonensis, Porocrinus ................... ТО. 5,12,13,18,54 AE АИ таене амалан т 12 Leske (1778) Lewis (1982) Lexington Limestone Curdsville Member or Curdsville Limestone .... 18,19,21,22,30 А ТОРИНА УЛ ШУЛЕН КОМИК КЕС ea 21 TICHeNOCHNUS:SÄtdesoms 1908 к 14 а 70 Таас үү, у, oe ани арнын aria m earn 15 ПОРТЕ ОВУ АО USOS longicaudis, Tornatilicrinus Lophospira Whitfield, 1886 Mac unes ЛО ВОИС ТД а шок ыиы хо ое Ee DEP 12 MEE den WISS) eer es M o qe I so у URL M 14 TIC ООЛО ОО AS A АН ЕП C duque 21,23 mahlburgi, ATEN Ыру у ы И e eot EE 27,28,31 TPAO PASTEUR IA IN NO ү 31 МАТИ 1962) у ee 67,68 LEBANON LIMESTONE ECHINODERMS: GUENSBURG 97 Maou oketnakonmatlone