3LUME ^uiOtins of "fSMcrican yakontowqs^ 102, NUMBER 339 MARCH Neogene Paleontology 12 in the northern The Genus Spondylus 13, 1992 Dominican Republic (Bivalvia: Spondylidae) by Harold E Yokes and Emily H Yokes 13 The Class Echinoidea (Echinodermata) by Porter M Kier Paleontologicai Research Institution 1259 Trumansburg Road New York, 14850 U.S.A Ithaca, J PALEONTOLOGICAL RESEARCH INSTITUTION Officers Harry A Leffingwell J Thomas Dutro, Jr Henry W Theisen President Vice-President Secretary Treasurer Assistant Treasurer Director Legal Counsel James C Showacre Roger Howley J Peter R Hoover Henry W Theisen Trustees Edward Bruce M Bell (to 6/30/93) Carlton E Brett (to 6/30/92) William L Crepet (to 6/30/94) J Thomas Dutro, Jr (to 6/30/93) Harry A Leffingwell (to 6/30/93) Robert M Linsley (to 6/30/92) Samuel T Pees (to 6/30/92) Willl\m p B Picou, Jr (to 6/30/92) Constance A Sancetta (to 6/30/94) James C Showacre (to 6/30/93) James E Sorauf (to 6/30/94) John Steinmetz (to 6/30/94) Henry W 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LIBRARY APR 1992 HARVARD UNIVERSITY The Paleontological Research Institution acknowledges with special thanks the contributions of the following individuals and institutions PATRONS ($1000 or more E Allen (1967) American Oil Company (1976) Atlantic Richfield Company (1978) Christina L Balk (1970, 1982, 1983) Hans M Bolli(1984) James Ruth G Browne (1986) Mr & Mrs Kenneth E Caster (1967) Chevron Oil Company (1978, 1982) Exxon Company (1977 to date) Lois S Fogelsanger (1966) Gulf Oil Corporation (1978) Merrill W Haas (1975) at the discretion of the contributor) Robert C Hoerle (1974-1977) Richard Johnson (1967 1986) J M McDonald Foundation (1972, 1978) Mobil Oil Corporation (1977 to date) Samuel T Pees (1981) Richard E Petit (1983) Robert A Pohowsky (1982) Texaco, Inc (1987 to date) Union Oil of California (1982 to date) United States Steel Foundation (1976) Charles G Ventress (1983 to date) Christine C Wakeley (1976-1984) LIFE MEMBERS ($200) Tucker Abbott James E Allen R Elizabeth A Balcells-Baldwin Christina L Balk Bruce M Bell Robert A Black Richard S Boardman Hans Bolli David John Bottjer Ruth G Browne David Bukry J Sybil B Burger Lyle D Campbell John L Carter Antmeliese S Caster Kenneth E Caster John E DuPont J Thomas Dutro, Jr J Mark Erickson Richard J Erickson Lois S Fogelsanger A Eugene Fritsche Christopher L Garvie Ernest H Gilmour Merrill W Haas Anita G Harris Steven M Herrick Carole S Hickman Robert C Hoerle F D Holland, Jr Frederick H C Hotchkiss David Jablonski Richard \ Johnson David B Jones Peter Jung Tomokj Kase Patricia H Kelley David Garrett Kerr Cecil H Kindle William F Klose, 11 JiRi Kriz Ralph Harry L Langenheim, Jr A Leffingwell Egbert G Leigh, Gerard A Jr Lenhard Louie N Marincovich, Donald Shuji R Moore Niko Hiroshi Noda Sakae O'Hara WiLLL\M A Oliver, Samuel T Pees Richard E Petit Edward B Jr Picou, Jr Robert A Pohowsky John Pojeta Jr John K Pope Anthony Reso Arthur W Rocker Arnold Ross Walter E Sage, John B III Saunders Judith Schiebout Miriam W Schriner Edward S Slagle Robert E Sloan Richard L Squires David H Stansbery Jorge P Valdes Raymond Van Houtte Charles G Ventress William P S Ventress Emily H Vokes Harold E, Vokes Christine C Wakeley Thomas R Waller Albert D Warren, Jr Gary D Webster Ralph H Willoughby Armour C Winslow Thomas E Yancey Victor A Zullo Jr tAmcrxcan toxcqs) 'LUME 102, MARCH NUMBER 339 Neogene Paleontology 12 in the northern The Genus Spondylus Dominican Republic (Bivalvia: Spondylidae) by Harold 13 The E Yokes and Emily H Vokes Class Echinoidea (Echinodermata) by Porter M Kier Paleontological Research Institution 1259 Trumansburg Road New York, 14850 U.S.A Ithaca, 13, 1992 Library of Congress Card Number: 85-637 Printed in the United States of Amenca Allen Press Inc Lawrence, KS 66044 U.S.A CONTENTS Page The Genus Spondylus (Bivalvia: Spondylidae) Harold E lakes and Eiuilv H I'okes 12 Abstract Resumen Introduction Acknowledgments Biostratigraphy and Paleoecology Abbreviations of Repository Institutions Systematic Paleontology 6 Introduction Systemalics Family Spondylidae Gray, 1826 Genus Spondylus Linnaeus, 1758 Spondylus hostrychites Guppy, 1867 Spondylus lucasi Maury, 920 Spondylus gumanomocon Brown and Pilsbry, 1913 8 10 11 The Class Echinoidea (Echinodermata) Porlcr M Kier 13 Abstract 13 Resumen 13 Introduction 13 Acknowledgments Previous work 13 Biostratigraphy 15 Paleoecology 16 13 Systematic Paleontology 17 Introduction Systematics Genus Genus Genus Genus Genus Genus Genus References Cited Plates Index Echmomclra Gray, 1825 17 Clypeasier Lamarck, 1801 17 S L R Agassiz, 1841 19 \fellila J L R Agassiz, 1841 20 20 Encope Moira A Agassiz, 1872 Brissopsis Schizasler J J L R Agassiz, L R Agassiz, 840 21 1835 22 24 28 37 LIST The Genus Spondylus 12 Harold E I 'okes OF ILLUSTRATIONS AND TABLES (Bivalvia: Spondylidae) and Emily H I okes ^^^^ Text-figure Geological sketch map of the Cibao Valley, nonhem Dominican Republic, showing areas from which samples for this project were collected Right valve of Spondylus bostrychites Guppy from locality TU 1219 The Class Echinoidea (Echinodermata) Porter M Kier 3, Text-figure Locality map for the sections measured and descnbed by Saunders Jung, and Biju-Duval (1986) Occurrence of echinoids in the Rio Cana section Dorsal view of the holotype of Clypeasler maoadentroensis, n sp Comparison of left poriferous zone of petal III oi Clypeasler maoadentroensis Dorsal and ventral views of Bnssopsis jimenoi Cotteau Oral view of Schizastcr doederleim (Cheshcr) 14 '5 '8 n sp and Clypeasler caudalus Jackson 19 21 22 Table Dominican Republic echinoid Distribution of northern Schizaster doedcrlemi (Chesher): comparison of northern Belize by formation and age Dominican Republic fossils with Recent specimens from Carrie Bow Cay, species, 2, 14 23 IN THE NORTHERN DOMINICAN REPUBLIC The Genus Spondylus (Bivalvia: Spondylidae) NEOGENE PALEONTOLOGY 12 By Harold New E Yokes and Emily H Yokes Department of Geology Tulane University Orleans, LA 701 18, U S A ABSTRACT Only three species of Spondylus have been collected in the Neogene strata of the northern Dominican Republic These include: Guppy which is largely confined to the more shallow facies of the late Miocene-early Pliocene Gurabo Formation; and giimanomocon Brown and Pilsbry, which is largely confined to the coralline facies of the Gurabo Formation and the Mao Adentro Member of the Mao Formation The third species is represented by a single specimen of S lucasi Maury from the Gurabo Formation No specimens are known from the early Miocene Baitoa Formation and only three specimens (one of bostiychiles and two of giimanomocon) have been collected in the late Miocene Cercado Formation hostryclutes RESUMEN Se han colectada solo tres especies de Spondylus en los stratos Neogenos del norte de la Republica Dominicana Estas incluyen: mayor parte restringida a las facies mas someras de la Formacion Gurabo del Mioceno tardio al Plioceno principle; y S giimanomocon Brown y Pilsbry, por la mayor parte restringida a las facies coralinas de la Formacion Gurabo, y del Miembro Mao Adentro de la Formacion Mao La lercera especie esta representada por un solo especimen de lucasi Maury de la Formacion Gurabo No se encuentran especimenes de Spondylus en la Formacion Baitoa del Mioceno principio y solamente Ires (uno de S boslrychiles y dos de gumanomocon) se han colectado en la Formacion Cercado del Mioceno S bostrychnes Guppy, por la tardio INTRODUCTION This study is of the material utilized herein a small contribution to the on-going project on the Neogene fossils from exposures in the Cibao Yalley, northern Dominican Republic (Text-fig 1) The geological setting and the history of the Dominican Republic Project have been thoroughly covered by several authors (see Saunders, Jung, and Biju- Duval, 1986; Jung, 1986; E Yokes, 1989; H Yokes, We are grateful to Dr American Museum of Natural History, New York, and Dr George M Davis and Ms Elena Benamy, Academy of Natural Sciences, Philadelphia, for the loan of type material, and to Dr Thomas R Niles Eldredge, Waller, U S National Museum of Natural History, and Dr Gary Rosenberg, Academy of Natural Sci- ences, Philadelphia, for reviewing the manuscript 1989) The material upon which this paper is based was collected by both the the Naturhistorisches Museum team of John Saunders and Peter Jung, and by the authors, as has been documented in the aforementioned papers For information on localities, stratigraphy, and ages, the reader is referred to the first work in this series: Saunders, Jung, and Biju-Duval (1986) Basel ACKNOWLEDGMENTS This study owes an immense debt of gratitude to Peter Jung, of the Naturhistorisches Museum Basel, who photographed and measured all of the type material for the species of Spondylus Linnaeus, 1758 that are involved His labors made the remainder of the work much less time-consuming In addition, he and John Saunders led the team that collected much NMB BIOSTRATIGRAPHY AND PALEOECOLOGY As there are but three species of Spondylidae Dominican Republic beds, and one of these is in the repre- sented by a single specimen, not a great deal of infor- mation can gleaned from their occurrence The most abundant species is S bostrychites Guppy, 1867, and from all evidence it followed a life-style unlike the vast majority of living spondylids, which attach to dead The bulk of the material comes from beds that, on the basis of other molluscan species present, we think of as "shallow-water Gurabo" (see E Yokes, 1989, p 21), with water depths of about 20 to 50 m From all ap- corals or other hard substrate for 5' bostrychites pearances the specimens of S bostrychites were not attached but lay "floating" in the fine silty sediment Bulletin 339 This quiet environment permitted the growth of long dehcate spines (see Text-fig 2) The other species for which we have enough material to consider is the more massive S gumanomocon Brown and Pilsbry, 1913, which is found primarily in the coralline facies of the Mao Adentro Member idently lived in the Gurabo Formation and of the Mao Formation more normal spondylid It the ev- fashion, attached to dead coral ABBREVIATIONS OF REPOSITORY INSTITUTIONS The following abbreviations for repository institu- tions are used in this paper: ANSP: Academy of Natural BMNH: British Museum (Natural History), London, England, U.K Most of the records of these two species are from the Gurabo Formation and the Mao Formation, but the occurrence of both is more dependent on environment than on time, as there is one specimen of S bostrychites and two of S gumanomocon from the Cer- NMB: cado beds One of the specimens of gumanomocon 16853, which is just upstream is from locality from the mouth of Arroyo Bellaco, in the Rio Cana drainage, where a very large coral reef is located (loc TU 1422; see Saunders, Jung, and Biju-Duval, 1986, text-fig 15 [loc TU 1282 is at the mouth of Arroyo Bellaco and TU 1422 is off the map]) No specimens of any species ofSpondylus Linnaeus, 1758 are known from the Baitoa Formation It is assumed that this is due to the very shallow-water nature of this formation The facies of the Baitoa is extremely similar to the shallow-water Cercado Formation, where specimens of Spondyhts are almost non-existent Throughout the Caribbean, species of Spondylus occur in other early Miocene beds that are correlated with the Baitoa Formation USNM: NMB Sciences, Philadelphia, PA, U.S.A Naturhistorisches Museum Basel, Basel, Swit- zerland PRI: Paleontological Research Institution, Ithaca, U.S.A NY, University, New Orleans, LA, U.S.A United States National Museum of Natural History, Washington, DC, U.S.A TU: Tulane SYSTEMATIC PALEONTOLOGY Introduction The genus Spondyliis Linnaeus, the Jurassic to the Recent (Hertlein 758 is known from and Cox, 1969, p N378) and today is found in all tropical and subtropical marine waters The shell is highly variable in shape and ornamentation to the extent that no two specimens of a species are entirely alike The variation in shape is due to the nature of the surface to which the right valve becomes attached, but the surface ornament is influenced by the environment in which The shell spines are it is growing produced by extensions of the Bulletins of American Paleontology, Volume 02 Plate Dominican Republic Neocene 12 and 13 31 Explanation of Plate Figure 1-4 Page Echinometra lucunter (Linnaeus, 758) Dorsal view of Recent figured specimen 17 I (USNM 30469) from m depth in holes in rock off Rodriguez Key, Florida Keys, x2 2-4 Dorsal, ventral, Limestone, 5-7 in and right side a roadcut 0.5 Clypeaster maoadentroensis, new views of figured specimen km m 375449) from locality TU 1438 in the Pliocene Mao Adentro species 17 (NMB M9746) NMB from locality 17022, early Pliocene, Mao Adentro Limestone, above the base of the Rio Cana section (Saunders, Jung, and Biju-Duval, 1986, text-figs 15-16), x Left side, dorsal, ventral views of the holotype 765 (USNM south of the bridge at Guayubin on the road to Sabaneta, x2 Bulletin 339 32 Explanation of Plate Page Figure , Clypeaster caudalus Jackson, 1922 '° Dorsal and nght side views of figured specimen Jung, and Biju-Duval 1986 text-fig 26) x 1 (NMB M9747) from locality NMB 17274, bed B, Arroyo Lopez section (Saunders, Bulletins OF American Paleontology, Volume 102 Plate Bulletins of American Paleontology, Volume 102 Plate Dominican Republic Neogene Explanation of Plate 12 and 33 13 Page Figure 1,2 Clypeaster C sunnilandensis Kier, 963 19 375450) from the late Miocene Cercado Formation at locality TU 1233, roadcut 5.3 km north of plaza at Moncion, on road to Los Quemados, x Dorsal view of figured specimen (LISNM 375451) from same locality as above, x Encope aberrans imperforata Kier, 1963 Dorsal view of figured specimen (NMB 9748) from locality 16857, late Miocene from the Rio Cana section (Saunders, Jung, and Biju-Duval, 1986, text-figs 15, 16), xi cf Dorsal view of figured specimen (USNM 19 NMB 4, 20 Mellita species Dorsal and ventral views of figured specimen (USNM 375452) from early Miocene, Baitoa Formation, locality Jung, and Biju-Duval, 1986, text-fig 21), x2 TU 362 (Saunders, Bulletin 339 34 Explanation of Plate Page TT- Figure , f Brissopsis jimenoi Cotteau, 1875 Gurabo Formation, at locality Dorsal and ventral views of figured specimen (NMB M9749) from the early Pliocene, near base of the above the base of the Rio Cana section, x 16824 (Saunders, Jung, and Biju-Duval, 1986, text-figs 15, 16) 317 m NMB 3, Moira atropos (Lamarck, 1816) Dorsal and ventral views of figured specimen (USNM 375453) from Jung, and Biju-Duval, 1986, text-fig 15), x locality TU 1354, early Pliocene, Gurabo Formation (Saunders, 22 Schizaster doederleini (Chesher, 1972) Dorsal view of figured specimen Biju-Duval, 1986, te.\t-fig 15) (USNM x3 375454) from locality TU 1354, early Pliocene Gurabo Formation (Saunders, Jung, and Bulletins of American Paleontology, Volume 102 Plate Bulletins of American Paleontology, Volume 102 Plate Dominican Republic Neocene Explanation of Plate and 35 Figure Page 1-6 Schizaster doederleini (Chesher, 1, 972) 3-5 Dorsal, ventral, and front views of figured (Saunders, Jung, and Biju-Duval, 1986, 22 Dorsal and ventral views of figured specimen Belize (Kier, 1975 fig 1), x2 View of peristome of figured specimen (USNM specimen text-fig (USNM 30468), Recent, from depth of (USNM 15), 375455) from locality TU locality TU mud west of Water Cay, 354, early Pliocene, Gurabo Formation x3 375456) from m fields I 1354, x4,5 in Dominican Republic Neogene and 12 37 13 INDEX Note: Page numbers arc in light face; plate numbers are in bold face type; the page numbers on which principal discussions occur are in Encope aberrans 19 aberrans aberrans Encope 16,19 aberrans imperforata Encope 14-17 79,20, 33 20 aclinensis Mellila Caribbean region 6,10-12,15 Sea 10,12,13,15-17,20,22,23 carmenensis, Spondylus Angola 16,17 Morocco caudatus Clypeasler Central west coast 16,17 Agassiz, A (1872) Agassiz, A (1880) R (1835) Agassiz, J L R (1840) 21 Agassiz, J L R (1841) 19,20 22 agiiayoi Brissopsis Agueguexquite Formation America, tropical Pacific coast americamis Spondylus [American Museum 8,10-12,28 of Natural History, New NY] York, 5,10,29 anacanlhus Spondylus 11 Anderson (1929) Anguilla Limestone Formation ANSP [Academy 8,22 of Natural Sciences, Philadelphia, PA] amillarium, Brissopsis 3, Arnold and Clark (1934) Atlantic Ocean, 4,22 17 western 7,10,12 atropos Moira Spatangus 14-17,20,34 Baitoa Formation Bed B of Saunders, Jung, and Biju-Duval (1986, Bold (1988) 1,2 Durham (1961) Caloosahatchee Formation Cantaure Formation Spondylus chiriquiensis Cidaris Whghl 1855 14 of Jackson (1922) Clark and Twitchell (1915) sp a 13,14 14 Clypeasler Lamarck, 1801 16,77,18 14-17,75,19,32 13,14 cubensis Coneau 1875 15.18 Twitchell ;>; Clark and Twitchell, 1915 14,15 duchassai ngi Michehn, 1855 18 16,17 sunnilandensis Kier, 1963 n sp cC C sunnilandensis Kier \963 text-fig 26) text-fig 26) 19 13-16,77,18,19,31 16,19 14,16.17.79.33 Codore Formation 12 colombiensis, Spondylus concavus Clypeasler 13.14 Cooke (1919) Cooke (1959) Cooke (1961) 13 Cotteau (1875) Cotteau(1881) 5-7,5,9-11,28,29 Cotteau (1897) 22 CO ncavus Coneau, 1875 20 19.20 13-18,21,22,34 14,21 21 Cotteau Collection [Universite Claude Bernard, Lyon, FRANCE] 21 cubensis 13,14,22 Clypeasler 15-17,22 Echinopedina 15.18 14 14-17,27,22,34 22 Dall (1898) Dall (1903) 6-8.1 1.12 Dall (1915) dalli, Cairns and Wells (1987) chipolanus tampaensis Spondylus 27,22 } L of 22 14,16,17,22,23,34,35 5,6,14,16,20,33 8,10,28 agi