ne as ast AGE 4 * sta pare my HARVARD UNIVERSITY e Library of the Museum of Comparative Zoology : NOR pe i ae i ‘if Gilbert Dennison Harris (1864 - 1952) Founder of the Bulletins of American Paleontology (1895) ISBN 0-877 Bee! MARCH 13, 1992 Neogene Paleontology in the northern Dominican Republic 12. The Genus Spondylus (Bivalvia: Spondylidae) by Harold E. Vokes and Emily H. Vokes 13. The Class Echinoidea (Echinodermata) by Porter M. Kier Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York, 14850 U.S.A. PALEONTOLOGICAL RESEARCH INSTITUTION Officers IRRESIDEN ia a oir tele oem heeeraeee ie et tal sae Ce ee ae HARRY A. LEFFINGWELL WIGE-PRESIDEN Tisch ivenes ooo eee oes CS ae J. THOMAS DuTRO, JR. SEGRETARIW tes cine eget ie ots Si LON CPC enc at Sei gh Pe cn cee HENRY W. THEISEN SIREASURER AGS, fc cere aatine (2 Sachin sin ear eater ee a pier JAMES C. SHOWACRE ASSISTANT AD REASURER GS One 4 atid Fs ots era ee a ee ae ROGER J. 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BROWNE WILLIAM A. OLIVER, JR. J. DAvip BUKRY SAMUEL T. PEES SysiL B. BURGER RICHARD E. PETIT Lyte D. CAMPBELL EDWARD B. Picou, JR. JOHN L. CARTER RosBertT A. POHOWSKY ANNELIESE S. CASTER JOHN PojseTA, JR. KENNETH E. CASTER JOHN K. Pope JOHN E. DUPONT ANTHONY RESO J. THOMAS DuTrRO, JR. ARTHUR W. ROCKER J. MARK ERICKSON ARNOLD Ross RICHARD J. ERICKSON WALTER E. SAGE, III Lois S. FOGELSANGER JOHN B. SAUNDERS A. EUGENE FRITSCHE JUDITH SCHIEBOUT CHRISTOPHER L. GARVIE MIRIAM W. SCHRINER ERNEST H. GILMOUR EDWARD S. SLAGLE MERRILL W. HAAS ROBERT E. SLOAN ANITA G. HARRIS RICHARD L. SQUIRES STEVEN M. HERRICK Davip H. STANSBERY CAROLE S. HICKMAN JORGE P. VALDES ROBERT C. HOERLE RAYMOND VAN HOUTTE F. D. HOLLAND, JR. CHARLES G. VENTRESS FREDERICK H. C. HOTCHKISS WILLIAM P. S. VENTRESS DAvipD JABLONSKI EmiLy H. VOKES RICHARD I. JOHNSON HAROLD E. VOKES Davip B. JONES CHRISTINE C. WAKELEY PETER JUNG THOMAS R. WALLER TOMOKI KASE ALBERT D. WARREN, JR. PATRICIA H. KELLEY Gary D. WEBSTER Davip GARRETT KERR RALPH H. WILLOUGHBY Cecit H. KINDLE ARMOUR C. WINSLOW WILLIAM F. K1osg, II THOMAS E. YANCEY Jiri Kriz Victor A. ZULLO ILUME 102, NUMBER 339 MARCH 13, 1992 Neogene Paleontology in the northern Dominican Republic 12. The Genus Spondylus (Bivalvia: Spondylidae) by Harold E. Vokes and Emily H. Vokes 13. The Class Echinoidea (Echinodermata) by Porter M. Kier Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York, 14850 U.S.A. Library of Congress Card Number: 85-63715 Printed in the United States of America Allen Press, Inc. Lawrence, KS 66044 U.S.A. References Cited FIBHOS © 5 5 GSB 6 chee PE ares Aine tee ot Seer grep rar err Bie etre ee Nd te iy Se eae Page Do oe A i Pinte hs Re eam RTC KM De hyo. oc eR Ao Ser eh teccls cyzhise drmiahe Petet and drorah doreeee Espacio CONTENTS The Genus Spondylus (Bivalvia: Spondylidae) Harold E. Vokes and Emily H. Vokes PNEDSUUea tee ke succeeds eh To ces route ene ae eis es Patt recayan sos are val avontyts eacbsiges Siena eee RES UIXLCI ET Here ro eT MEIN, SEL Shoes Acsct yah tie PNT cal CUSTOM TEN? Nellore Watieue, ake eben eaictitiasel cWsaalte seh oMpMortevotans itis Ata Alatevenautteh erable tadenareban [RATS LN TSLITOy Tt cl I me i ere en oP ney cnn eRe ee ee RICA ONS PeCon a ONS Sch Cn ore ela neta S ool (NP aONU eNO decaosuonoreebos cobubpooobe gaadnoed WedSen ner eke cos oAacbomhdetinh PaapESooabs dies ou doen Scoot Oc BIOStLaligraphiysand \haleGecOlopysmrcrevacis evevetete evarste tel eeeret eta tne lhe eet obeli eked hol saved de totede hele) SMO Mele eat er=ye tates a tel et tot tetersyatar Abbreviations of Repository Institutions .................2.---+--++ 02s Systematic Paleontology MMEGOGUGTIOM Pip) serene ae leat acceler ey atelier ach sca tose eanre eee eee Systematics Family Spondylidae Gray, 1826 Genus: Spondylusibinnacuss lS Stew. srresere cease clea areas ee ties ete cite ice esis iapee oe pete tetra st cge tec dey ole hatte ens SPONAYIUSIDOSTFYV CHILES GUPDYsglOOUaen an sect eae i etiaie Cine ee Eee ease aye ee See race eretote etre SponaylusilucastIMaUtye LODO, yaccccts serie crs cake A steccle ave cote ele terete ee Se ee Ce SS TRE al eh eae tL esrecees ens anayaa one Spondylus sumanomocon Brown andeelsbry, UO s eer rev= cts crater = clet-y= eletol ayer tebe raters seteaiel lerey torsion ele rore ele ele The Class Echinoidea (Echinodermata) Porter M. Kier PRESS CAC Crew pee Rees SIRS ess eis eh au NBT ANGNT Uo hea See lone eae Bundles ease a Meee eh Mec (eae) Feet lars ee Covicele euepetera fey onc eveve arastineres aye ac esueNeb aM et ens IROSTIDTEVGIN 2 aes de te 0 sue ee ernie pena UME Snr ERT can pack aero ne OTe ane. d CeO enn aaa An aA aD eee OM OMe. De Waiinayaltre[aYoy Net Sakae ce Perera ceorcrteres pices aio Ga chao ccna aie Rey ha aN ex cae Ackn ow led grmiemts are sccn2y 5c fezcuscoes sack sushteepetay ape ote ons cents sparen fogsasde Migs navevaveles neues Previous work ......... ET OT CORA TOT re nee RTS Cra ne ane c Henan Meech oa ee en ey ood COONS Biostratigraphy ........ BI Ae ee ot Da ance ORR eo Uno SOLO OquCS Oa aueaee oon lPalleyerarere) lela Me pe com ene a ae OU e anon tae ame Rae Oe eee Se ns ero ee sae Systematic Paleontology MTN E OD UCL OM See ersecne ec cice ead ee, chaashns a lepegb ses toasted besrene cose Pavalr een apes sk ayieresus Oicauieles ace opaiye as covanstiensijsteys ete atuch seayahas vet enststeretaieus ter Weneney ey qh tavecteys tite setee ell shokesesavave eu-tayeyshsisbaiaqenstes ste) Siejetayers ete) saber GenussSchizastersVoie ws AGASSIZ y BSD oe oan cea at execpt she es Epepate aes evehe erase fone Re vayse Pet ce nearer catietenenevehereys ehcp asheyer te ienek cheyea Merete LIST OF ILLUSTRATIONS AND TABLES 12. The Genus Spondylus (Bivalvia: Spondylidae) Harold E. Vokes and Emily H. Vokes Text-figure Page 1. Geological sketch map of the Cibao Valley, northern Dominican Republic, showing areas from which samples for this project were COWS CCEA ccc cence ste STE SPT eee R overs step cgnyegne Toa) care suis crcl e megane ate Sue et crtega clishe cyeparS chante vee ore pe ir 6 2. Right valve of Spondylus bostrychites Guppy from locality TU 1219 ........ 2.2... e eee eee eee enter enna 1 13. The Class Echinoidea (Echinodermata) Porter M. Kier Text-figure 1. Locality map for the sections measured and described by Saunders, Jung, and Biju-Duvall (1986)! see ase eee eee 14 2. Occurrence of:echinoids in the Rio\Gana Section .< ... ence ww ce een ww ocr ena oie = ola wisine) = jay eet ate ie [a a=) ll ele oaoho oko keaiee ohh netelatets 15 3. Dorsal view of the holotype of Clypeaster maoadentroensis, N. SP... - 2... 6 eee ee eee ee eee ees 18 4. Comparison of left poriferous zone of petal III of C/ypeaster maoadentroensis, n. sp. and Clypeaster caudatus Jackson ..........- 19 5. Dorsal and ventral views of Brissopsis jimenoi Cotteau ..... foals ebvwne lt. ces eltpaysee oebe 6 orarstoneanwaie ale ionene etd yes oieteseteanhe eee nee eae 21 6. Oral view of Schizaster doederleini (Chesher) ......... eee A en er re SOMA Uc GdéAoooc oo zor s 22 Table 1. Distribution of northern Dominican Republic echinoid species, by formation and age .... 2... 6.6.66 eee eee ee eee 14 2. Schizaster doederleini (Chesher): comparison of northern Dominican Republic fossils with Recent specimens from Carrie Bow Cay, 12) bd ee eee ee Repel t n e ne ien Rn PrN eae ee RCE CHAU OP ODA GUM En Ee cacoog sons DOU. 23 NEOGENE PALEONTOLOGY IN THE NORTHERN DOMINICAN REPUBLIC 12. The Genus Spondylus (Bivalvia: Spondylidae) By HAROLD E. VOKES AND EMILY H. VOKES Department of Geology Tulane University New Orleans, LA 70118, U.S.A. ABSTRACT Only three species of Spondylus have been collected in the Neogene strata of the northern Dominican Republic. These include: S. bostrychites Guppy, which is largely confined to the more shallow facies of the late Miocene-early Pliocene Gurabo Formation; and S. gumanomocon 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. /ucasi Maury from the Gurabo Formation. No specimens are known from the early Miocene Baitoa Formation and only three specimens (one of S. bostrychites and two of S. gumanomocon) have been collected in the late Miocene Cercado Formation. RESUMEN Se han colectada solo tres especies de Spondylus en los stratos Nedgenos del norte de la Republica Dominicana. Estas incluyen: S. bostrychites Guppy, por la mayor parte restringida a las facies mas someras de la Formacion Gurabo del Mioceno tardio al Plioceno principio; y S. gumanomocon Brown y Pilsbry, por la mayor parte restringida a las facies coralinas de la Formacion Gurabo, y del Miembro Mao Adentro de la Formacién Mao. La tercera especie esta representada por un solo espécimen de S? lucasi Maury de la Formacion Gurabo. No se encuentran espécimenes de Spondylus en la Formacion Baitoa del Mioceno principio y solamente tres (uno de S. bostrychites y dos de S. gumanomocon) se han colectado en la Formacion Cercado del Mioceno tardio. INTRODUCTION This study is a small contribution to the on-going project on the Neogene fossils from exposures in the Cibao Valley, northern Dominican Republic (Text-fig. 1). The geological setting and the history of the Do- minican Republic Project have been thoroughly cov- ered by several authors (see Saunders, Jung, and Biju- Duval, 1986; Jung, 1986; E. Vokes, 1989; H. Vokes, 1989). The material upon which this paper is based was collected by both the the Naturhistorisches Museum Basel team of John Saunders and Peter Jung, and by the authors, as has been documented in the aforemen- tioned papers. For information on localities, stratig- raphy, and ages, the reader is referred to the first work in this series: Saunders, Jung, and Biju-Duval (1986). 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 ma- terial 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 NMB team that collected much of the material utilized herein. We are grateful to Dr. Niles Eldredge, 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. Waller, U. S. National Museum of Natural History, and Dr. Gary Rosenberg, Academy of Natural Sci- ences, Philadelphia, for reviewing the manuscript. BIOSTRATIGRAPHY AND PALEOECOLOGY As there are but three species of Spondylidae in the Dominican Republic beds, and one of these is 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 corals or other hard substrate. The bulk of the material for S. bostrychites comes from beds that, on the basis of other molluscan species present, we think of as ‘shallow-water Gurabo”’ (see E. Vokes, 1989, p. 21), with water depths of about 20 to 50 m. From all ap- pearances the specimens of S. bostrychites were not attached but lay “‘floating”’ in the fine silty sediment. 6 BULLETIN 339 This quiet environment permitted the growth of long delicate 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 Gurabo Formation and the Mao Adentro Member of the Mao Formation. It ev- idently lived in the more normal spondylid fashion, attached to dead coral. 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 environ- ment than on time, as there is one specimen of S. bostrychites and two of S. gumanomocon from the Cer- cado beds. One of the specimens of S. gumanomocon is from locality NMB 16853, which is just upstream 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 Biyu-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 of Spondylus Linnaeus, 1758 are known from the Baitoa Formation. It is as- sumed 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 Spondylus are almost non-existent. Throughout the Caribbean, species of Spondylus occur in other early Miocene beds that are correlated with the Baitoa Formation. eS ABBREVIATIONS OF REPOSITORY INSTITUTIONS The following abbreviations for repository institu- tions are used in this paper: ANSP: Academy of Natural Sciences, Philadelphia, PA, U.S.A. BMNH: British Museum (Natural History), London, England, U.K. NMB: Naturhistorisches Museum Basel, Basel, Swit- zerland. PRI: Paleontological Research Institution, Ithaca, NY, U.S.A. TU: Tulane University, New Orleans, LA, U.S.A. USNM: United States National Museum of Natural History, Washington, DC, U.S.A. SYSTEMATIC PALEONTOLOGY INTRODUCTION The genus Spondylus Linnaeus, 1758 is known from the Jurassic to the Recent (Hertlein 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 it is growing. The shell spines are produced by extensions of the °) 10 20km 4 Rio Cana 2 Rio Gurabo 3 Rio Mao 4 Rio Amina 5 Canada Zalaya 6 Rio Yaque del Norte 7 City of Santiago 8 Arroyo Punal 9 Rio Verde Upper Cenozoic E-.] Oligocene - Early Miocene ? ESS] Mesozoic Text-figure 1.— Geological sketch map of the Cibao Valley, northern Dominican Republic, showing areas from which samples for this project were collected (after Saunders, Jung, and Biju-Duval, 1986, text-fig. 3). DOMINICAN REPUBLIC NEOGENE. 12: H. E. VOKES AND E. H. VOKES 7/. mantle beyond the shell margin and, as noted by Root (1988, p. 7), the ‘“‘feathery, fingerlike projections” of the mantle, which produce the spines, are strongly af- fected by the force of the current; large elongate spines can form in quiet waters, while those in a swift current have to be short because the projection cannot be held out long enough to permit the secretion of a large cal- careous spine. The net result of this great variability has been the assignment of numerous generic and specific names to spondylid species. The problem is also accentuated in Recent species by the tendency to have color variations ranging, for example in the eastern Pacific Spondylus princeps Broderip, 1833, from white to pink, orange or coral-red, often with spines that may vary somewhat from the color of the shell itself. Just to give an idea of the complicated taxonomy of the Recent species of Spondylus, in a recent popular work (illustrated by exquisite color photographs), Lamprell (1987) states that 224 specific names have been proposed for species of Spondylus. Of this num- ber he considers approximately 75 (even he is uncer- tain) as valid species, the remainder synonyms. In the Recent fauna of the western Atlantic there are probably six species, plus another three on the eastern Pacific coast of tropical America (Lamprell, 1989). In ee the Neogene of the Americas there are possibly 12 valid species, in addition to fossil occurrences of forms de- scribed from the Recent fauna. It is obvious that the determination of the validity of a species on shell morphology alone is impossible. There will have to be some better method of species determination, such as using electrophoresis of amino acids or proteins, before any meaningful synthesis for the living forms can be achieved. This is not going to be a great help with the fossil species. All we can do is try to compare the morphology of spine patterns, numbers and relative sizes of ribs, and overall shell shape. Thus, the determination of which name 1s cor- rect for any particular fossil is tentative at best. Time may well prove that names used herein are synonyms of some previously described taxon. Likewise, almost certainly, time will prove that there are synyonyms that might have been included in the synonymy of any particular species. We have chosen to take a conser- vative approach and only include as synonyms those for which we have no doubts. Measuring these irregular shells is also difficult. In this study the height is measured (as nearly as possible) perpendicular to the hinge, along a line from the umbo to the ventral edge. However, this is easier said than done. For example, the lectotype of S. guwmanomocon Text-figure 2.—Right valve of Spondylus bostrychites Guppy, 1867, still encased in the outcrop at locality TU 1219, on the Rio Amina, near Potrero. 8 BULLETIN 339 Brown and Pilsbry, 1913, was stated to be 175 mm in length by the original authors. Palmer (1938, expla- nation to pl. 16) says “Height 181 mm”’. If one mea- sures the maximum dimension of this shell it is 181 mm, but if one measures along a line that more or less bisects the valve then it is 175 mm. The length is here taken as the maximum distance from the anterior to the posterior margins of the shell, along a line as nearly as possible parallel to the hinge and, more importantly, at right angles to the measured height. Diameter is measured only for paired valves and is the maximum distance between two planes tangent to the outside of the right and left valves, paralleling the valve margins. SYSTEMATICS Family SPONDYLIDAE Gray, 1826 Genus SPONDYLUS Linnaeus, 1758 Spondylus Linnaeus, 1758, p. 690. Type species.— Spondylus gaederopus Linnaeus, 1758 (by subsequent designation, Schmidt, 1818); Recent, ““M. Mediterraneo” (Linnaeus, 1758); and “‘in the east- ern Atlantic from Morocco to Senegal, and in the At- lantic islands” (Dodge, 1952, p. 126). Diagnosis.—Shell pectiniform but usually deformed as a result of fixation of the right valve umbonal area to the substrate; right valve generally larger and more inflated than the left, with a higher triangular cardinal area in which the internal ligament is lodged in a pit located between two stout crural teeth that, in the mght valve, are immediately adjacent to the pit but, in the left valve, are outside of the sockets for the reception of those of the right valve; sculpture primarily radial, with ribs of primary, secondary, and tertiary size, the primary almost universally bearing strong pointed spines; smaller spines or pointed nodules often present on the secondary and tertiary radials; mght valve may be concentrically foliaceous adjacent to attachment area. Remarks.— Although Gaideropa Deshayes, 1832 is cited as an objective synonym of Spondylus Linnaeus, 1758 in the Treatise on Invertebrate Paleontology (Her- tlein and Cox, 1969, p. N378), when one examines the original reference of Deshayes (1832, p. 163)!, it is obvious he was merely giving the vernacular name of the species Spondylus gaederopus Linnaeus, 1758. '‘ What Deshayes says is [translated]: “GAIDEROPE. Gaideropa. We used to name thus, or we used to give the name of ‘donkey’s foot’, which is synonymous, to a rather common shell, that the ancients placed in the spiny oysters, and which is placed today in the genus Spondyle, under the denomination of Spondylus gaderopus [sic]. See SPONDYLE.” Spondylus bostrychites Guppy Plate 1, figures 1-3; Plate 2, figure 1; Text-figure 2 Spondylus bifrons Sowerby, 1850, p. 53. (non S. bifrons Goldfuss, 1835). Spondylus bostrychites Guppy, 1867, pp. 164 (list), 176 (nom. nov. pro Spondylus bifrons Sowerby non Goldfuss) (reprinted Harris, 1921, pp. 191, 203); Gabb, 1873, p. 257; Guppy, 1874, p. 443 (in part); Dall, 1898, p. 758 (in part, not all localities); Maury, 1917, p. 190 (354), pl. 32(58), fig. 4; Vaughan et a/., 1921, pp. 129, 145; Pilsbry, 1922, p. 413; Hanna, 1926, p. 477, pl. 24, figs. 3, 4; Palmer, 1938, p. 6(150) (in part, not references to Bowden spec- imen), pl. 1(16), fig. 2 only; pl. 2(17), figs. 1, 3, 5; pl. 3(18), figs. 1-5; Ramirez, 1956, pp. 13, 15-17; Pflug, 1961, p. 77, pl. 23, figs. 18: [2] Spondylus bostrychites Guppy. Maury, 1920, p. 22. (?] Spondylus bostrychites? Guppy. Hubbard, 1920, p. 97; Vaughan et al., 1921, p. 150. Spondylus sp. indet. Vaughan ef a/., 1921, p. 123. INCORRECT REFERENCES TO Spondylus bostrychites: Spondylus bostrychites Guppy. Guppy, 1873, p. 87 (reprinted Harris, 1921, p. 219); Dall, 1903, p. 1586; Woodring, 1925, p. 76, pl. 9, figs. 5-7; Palmer, 1938, p. 6(150) (in part, Bowden specimen only), pl. 1(16), fig. | only (Bowden Formation, Jamaica; probably = S. americanus Hermann, 1781). Spondylus bostrychites Guppy. Dall, 1915, p. 124, pl. 19, fig. 4 (Tam- pa Limestone, Florida; = S. chipolanus tampaensis Mansfield, 1937). Spondylus bostrychites Guppy. Cooke, 1919, p. 144, pl. 11, figs. 11a, 11b (Anguilla Limestone, Lesser Antilles; = S. scotti Brown and Pilsbry, 1913, fide Woodring, 1982, p. 601; however, specimens illustrated by Cooke have a strong similarity to S. chipolanus Dall, 1898). Spondylus bostrychites Guppy. Trechmann, 1930, p. 211 (Manchi- oneal Beds, Jamaica; = S. chiriquiensis Olsson, 1922, fide Palmer, 1938, p. 155). Spondylus bostrychites Guppy. Anderson, 1929, p. 158 (Tubara Group, Colombia; = S. colombiensis Weisbord, 1929). Spondylus bostrychites Guppy. Perrilliat Montoya, 1963, p. 10, pl. 2, figs. 2, 6 (Agueguexquite Formation, Mexico; = S. americanus Hermann, 1781). Spondylus bostrychites Guppy. Ferreira, 1965, p. 5, figs. 7-10 (Pir- abas Limestone, Brazil; appears to be S. chipolanus tampanensis Mansfield, 1937°). Spondylus bostrychites Guppy. H. Vokes, 1986, p. 174, text-fig. 1 (Moin Formation, Costa Rica; = S. chiriquiensis Olsson, 1922). Testa subregularis, rotundata, ventricosa, margine latiusculo, val- ide denticulato; extus radiatim costata, costis 5 ad 6 spiniferis; area cardinali alterius valvae angustissima, alterius latiori. Nearest to S. imperialis [Chenu, 1843], easily distinguishable by the area of one valve being very narrow, and that of the other being rather broader, though still narrow. (Sowerby, 1850, p. 53) Diagnosis.—Shell shape pectiniform, ornamented with alternating coarser and finer spinose ribs; valves 2In 1887, C. A. White described Spondylus pinguisculus (p. 47, pl. 9, figs. 22, 23) from the same Pirabas Limestone locality. Un- fortunately the White species is based upon an internal mold lacking any trace of the external ornamentation. The name has forty years priority over the Mansfield name and, if the Pirabas species proves to be the same as the Tampa one, would supersede it. DOMINICAN REPUBLIC NEOGENE. 12: H. E. VOKES AND E. H. VOKES 9 unusually symmetrical, rarely any trace of an attach- ment area. Description.—Shell rounded, pectiniform, subequi- valved with the lower (right) valve slightly more in- flated than the upper (left) one; attachment area on lower valve small or absent, marked occasionally by small area of raised concentric lamellae; remainder of surface of right valve and all of the left with radial ribbing consisting, on well-preserved valves, of ribs of four different strengths: the strongest, primary ribs most prominent and bearing large, elongate flattened or flut- ed spines, the ribs of secondary strength with distantly spaced, nodose spinules; the tertiary riblets occurring in groups of two to four, commonly three, separating the primary and secondaries, and often relatively smooth; quaternary riblets visible only on exception- ally well-preserved valves, submicroscopic and ex- ceedingly numerous, often being present on the sides and upper surfaces of the primary, secondary, and ter- tiary ribs. Well-preserved valves also with ribbing crossed by fine raised growth lines that form small raised spinose flutings on the rib tops. Commonly sev- en primary ribs on the lower (right) valve and six on the upper (left) one. Cardinal area of the lower valve somewhat broader than that of the narrow, often linear, upper one; both coarsely striated. Auricles moderately large, with radial ribbing that may be obscured by strong, raised growth lamellae. Adductor scar gently arched to almost straight dorsally, broadly rounded on lateral and ventral sides; ventral internal valve margins crenulated by termination of external radial ornament. Lectotype. —BMNH LL 9946 (selected by Palmer, 1938). Type locality.—Locality TU 1219 (here restricted), Gurabo Formation, Rio Amina, bluffs on east side of river immediately upstream from ford that is 2 km west of Potrero and about 3 km downstream from “La Represa”’ (= locality USGS 8516; see Saunders, Jung, and Biju-Duval, 1986, text-fig. 34). Material.—Over 100 valves, many paired; plus nu- merous immature and/or fragmentary specimens, from many localities, mostly in the shallow-water facies of the Gurabo Formation. Measurements (in mm).— diameter (paired specimen height’ length valves) locality BMNH LL 9946! 100 101.5 _— unknown?‘ BMNH LL 257067 89 81 62 unknown* BMNH LL 25707? 94 90 — unknown* BMNH LL 25708? 94 90 - unknown? BMNH LL 25709? 62 58 — unknown* BMNH LL 25710? 55 55 = unknown? USNM 450387 114 88 _— TU 1278 NMB G 16963 80 75 57 NMB 16821 ' lectotype; * paralectotypes; * Heneken Collection, “San Domingo”. Remarks.—A\though there are rare specimens of S. bostrychites Guppy, 1867, from the Cercado Forma- tion (one pair from loc. TU 1222, near Moncion) and from the Mao Adentro Member of the Mao Formation on the Rio Gurabo (locs. TU 1440, NMB 15822: see Saunders, Jung, and Biju-Duval, 1986, text-figs. 4, 5), the vast majority of the specimens have been taken in the shallow-water facies of the Gurabo Formation. In our collections and those of the Naturhistorisches Mu- seum, the species is most abundant along the Rio Gur- abo above the ford, in beds that Maury (1917, p. 437) referred to her Zones D (loc. TU 1215 = NMB 15842- 15858: see Saunders, Jung, and Biju-Duval, 1986, text- figs. 4, 5) and F (locs. TU 1277, 1296 = locs. NMB 15864-15875; see Saunders, Jung, and Biju-Duval, 1986, text-figs. 4, 5), at “Bluff 1” (of Maury, 1917, p. 426, where she noted it was ‘“‘common’’) on the Rio Mao (locs. TU 1293, NMB 16910; see Saunders, Jung, and Biju-Duval, 1986, text-fig. 29), and at the ford on the Rio Amina (locs. TU 1219, NMB 16807; see Saun- ders, Jung, and Biju-Duval, 1986, text-fig. 34). Al- though the NMB team was not able to collect the latter locality to any great degree, in the TU collections from locality TU 1219 alone we have 20 adult valves (three paired) and 40 immature valves. Inasmuch as this is certainly one of the localities where the original He- neken Collection’ was made, locality TU 1219 is here offered as a restriction of the type locality, which was only “San Domingo’”’. The species of Spondylus are all discouragingly sim- ilar and one species may be distinguished from another only with great difficulty. One of the most striking characteristics exhibited by Spondylus bostrychites is the Jack of an attachment area. Most species are marked by a relatively large area of concentric foliations in the umbonal area of the right valve. As may be seen from the various specimens figured, this is not usually de- veloped in S. bostrychites. Of all of the specimens in the TU collections only three show any trace of this attachment area. The significance of the lack of attachment area would seem to be that, unlike the majority of spondylid species, which attach to some hard substrate — corals, rocks, etc., S. bostrychites did not affix itself to any hard sur- face but “floated” on a silty bottom. Waller (written communication, April 2, 1990) has suggested that the species may be an opportunistic cementer, with settling spat cementing themselves to tiny hard objects on the silty bottom when such objects may be found. The same life-style has been documented for the large oys- ter Hyotissa haitensis (Sowerby, 1850), which occurs * The Heneken Collection was sent to the Geological Society of London in 1848 and was the basis for the molluscan descriptions of Sowerby (1850) and Guppy (1876). 10 BULLETIN 339 together with S. bostrychites at many localities (see Meeder, 1987, p. 11). Specimens from several other formations through- out the western Atlantic have been cited as S. bostry- chites. In our opinion, all of these, with the possible exception of the Puerto Rican specimens listed but not figured by Maury (1920, p. 22) and Hubbard (1920, p. 97), both of which were based on incomplete molds, prove to be other species. In particular, the examples from the Bowden Formation, Jamaica, which have been figured by Woodring (1925, pl. 9, figs. S—7) and Palmer (1938, pl. 16, fig.1), show the attachment area (see Pl. 1, fig. 4). They are here assigned to the Recent species S. americanus Hermann, 1781. Comparisons.—The Recent Caribbean species Spondylus erinaceus Reeve, 1856 (pl. 11, fig. 39), su- perfically resembles S. bostrychites in the general aspect of the surface ornament, but differs in having only five primary radial ribs (in contrast to the six or seven of S. bostrychites) adorned with palmate scales rather than spines, with five (rarely six) well-developed scaly ridges devoid of any sort of spine formation between them. The shell also tends to be attached to coralliferous surfaces and, consequently, to develop a relatively strong dorsal triangular area above the hinge region on the right valve. This latter feature seems to be the principal difference between the two forms. Neverthe- less, S. erinaceus with its globose, symmetrical shells, well shown in the illustration given by Garcia (1989, pl. 1) may be the descendant of S. bostrychites. Cer- tainly, no other species comes as close. Occurrence.— Unnamed formation: Lopez area (TU 1445). Cercado/Gurabo formations: Rio Cana area (TU 1354: NMB 16817, 16818, 16821, 16824, 16861, 16862, 16865, 16867, 16868, 16878); Rio Gurabo area (UN2ZOSEI2Z1OM12T1E 1213=12' 15912225 123i 1277; 1278, 1279, 1296, 1338, 1368, 1369, 1416; NMB 15804, 15806, 15807, 15809, 15815, 15836, 15842- 15844, 15846, 15848, 15851, 15853, 15855, 15856, 15862-15868, 15871, 16808, 16809, 16811); Rio Mao area (TU 1225, 1293; NMB 16910); Rio Amina area (TU 1219, 1220, 1248, 1370, 1371, 1411, 1412, 1455; NMB 16805, 16807); Santiago area (TU 1205, 1206, 1207, 1227A, 1250, 1380, 1404, 1405, 1449, 1453; NMB 17268). Mao Formation: Rio Gurabo (TU 1440; NMB 15822). Distribution.—Unnamed formation (Lopez area), Cercado and Gurabo formations, and Mao Adentro Member of the Mao Formation, Dominican Republic. Other occurrences in the Caribbean fossil record are uncertain, but the specimen from the Imperial For- mation (Pliocene) of California reported by Hanna (1926) may well be this species. Spondylus lucasi Maury Plate 2, figures 2, 3 Spondylus lucasi Maury, 1920, p. 23, pl. 5, fig. 1. Shell oval, oblique, small for the genus. The cardinal area is de- fective and broken, but shows traces of the isodont hinge of Spon- dylus. The sculpture consists of [?eight] stronger, low, primary, radiating, rounded threads bearing spines at intervals, and between every two of these primaries are eight to ten or twelve much more delicate radiating lines. The central one of these is slightly stronger than the rest. It does not stand out sharply but the eye can discern that it is a shade thicker. The fine radii between the primaries nearly always alternate in strength, the finest lines of all being visible only with a lens. All of the fine radii are ornamented with minute scales, giving them a beaded appearance, especially on the anterior part of the valve. (Maury, 1920, p. 23) Diagnosis.—Shell shape pectiniform, ornamented by numerous finely spinose radial ribs; approximately every seventh of these slightly stronger and with longer spines. Description.—Shell, a left valve, slightly suboval, rounded in outline; a small rounded umbo; auricles moderately short and narrow, having surface ornament of fine radial ribbing similar to that on the entire valve with microscopically small spines and raised growth lamellae. Valve ornament of seven primary radial ribs bearing relatively strong radial spines, usually six or seven in number and distantly spaced, although the anterior primary rib bears but one such spine. Between each pair of primaries, on the median surface of the valve, there are three moderately small secondary ra- dials with a slightly weaker tertiary between each of the stronger ribs. The tertiary radials tend to increase in strength with growth, becoming indistinguishable from the secondaries toward the valve margin, so that there appear to be seven secondary ribs between each pair of primaries. Surface of all secondary and tertiary radials marked by closely approximate laciniate pro- jections of growth lamellae. Interspaces between ribs narrow, flat-bottomed, with three rows of microscopic spinelike projections, one medial in position and the others on the slightly raised margins of the adjacent radials. Lectotype.—AMNH 22514 (here designated). Type locality.—Reeds’ locality 370, east shore of Guanica Harbor, Puerto Rico [see Maury, 1920, pp. 1, 7; data on label with specimen]. Material.—A single left valve from locality TU 1211. Measurements (in mm).— height length locality AMNH 22514! 332 28 see above USNM 450389 54.5 53 TU 1211 ' lectotype; ? estimated, badly broken. DOMINICAN REPUBLIC NEOGENE. 12: H. E. VOKES AND E. H. VOKES 11 Remarks.—A single left valve from locality TU 1211 (Rio Gurabo; see Saunders, Jung, and Biyu-Duval, 1986, text-fig. 5) is here identified as Spondylus lucasi Maury, 1920, originally described from the Guanica Lime- stone (= Ponce Limestone) and the Quebradillas Lime- stone of Puerto Rico. The Ponce Limestone, from the south coast, although originally described as Oligocene in age is now considered to be early Miocene; the Que- bradillas Limestone, from the north coast, is now con- sidered Pliocene. Although slightly larger than the type of S. /ucasi, the Dominican specimen agrees so com- pletely in all details of the ornamentation of that form as figured by Maury (see PI. 2, fig. 3), as to leave little doubt that it represents the second record of its oc- currence in the Caribbean fossil fauna. Comparisons.—The shell of Spondylus lucasi Mau- ry, 1920, has a much more evenly patterned ornament than S. scotti Brown and Pilsbry, 1913, as figured by Woodring (1982, pl. 97, figs. 8-10), from the early Miocene La Boca Formation of Panama, with which he tentatively [p. 601] synonymized S. /ucasi. Jung (1965, pl. 55, figs. 2, 3 and 1971, pl. 1, figs. 4, 5) has figured specimens from the early Miocene Can- taure Formation and the early middle Miocene Grand Bay Formation, respectively, that he compares to S. lucasi. Both seem better assigned to S. scotti. Spondylus lucasi and the older S. scotti are similar only to the extent that both have relatively fine radial ribs, in contrast to such species as S. bostrychites Gup- py, 1867, in which the ribs are much wider (compare Pl. 1, fig. 3b, and Pl. 2, fig. 2b). In S. scotti, there are many more major radials (17 in the type specimen) with fewer secondary radials between each pair (four to eight in the type specimen) than in S. /ucasi, which has only seven or eight primary ribs, with about seven secondary ribs between each pair of primaries. In the Recent fauna of the Indo-Pacific the species S. anacanthus Mawe, 1823, is perhaps the most nearly similar to this unusual species, which is marked by having a much more regular surface ornament than is typical for most species of Spondylus. Occurrence.—Gurabo Formation: Rio Gurabo (TU PAYS Distribution.—Gurabo Formation, Dominican Re- public. Ponce Limestone, Early Miocene; Quebradillas Limestone, Pliocene; Puerto Rico. Spondylus gumanomocon Brown and Pilsbry Plate 3, figures 1—4 Spondylus Americanus Lamarck. Gabb, 1873, p. 257 (not of La- marck, 1819, nor of Hermann, 1781). Spondylus gumanomocon Brown and Pilsbry, 1913, p. 514 (foot- note); Maury, 1917, p. 191(355); Pilsbry, 1922, p. 413, pl. 43, figs. 4 (paralectotype), 5 (lectotype); Palmer, 1938, p. 9(153), pl. 1(16), figs. 3, 4 (lectotype); pl. 2(17), figs. 2, 4 (paralectotype); Pflug, 1961, ps3: [?] Spondylus gumanomocon Brown and Pilsbry. Maury, 1920, p. 23; Hubbard, 1920, p. 97; Olsson, 1922, p. 207(379), pl. 21(24), fig. 1. Spondylus guamanomocon Pilsbry and Johnson [sic]. Vaughan et al., 1921, p. 153. Spondylus carmenensis Hodson in Hodson, Hodson, and Harris, 1927, p. 41, pl. 25, figs. 1-3. A species resembling S. varians Sowb. (S. delessertii Chenu [1845]). The upper valve is Pectiniform, orbicular, of moderate thickness, with low radial ribs, the principal ones irregularly spinose, spines short; cardinal area small and short, as in S. americanus [Her- mann, 1781]. Lower valve very ponderous, with a long, level (not receding) cardinal area, and a very long, straight (or sometimes laterally curved) beak, the cavity of which is deeply excavated in young shells, nearly solidly filled in old ones. Sculpture like the upper valve, except that it is more or less extensively foliated towards the beak .... This is the form identified by Gabb as Spondylus americanus. (Brown and Pilsbry, 1913, p. 514) Diagnosis.—Shell very massive, irregular in shape, with lower (right) valve greatly enlarged. Radial or- nament of alternating larger and smaller spinose ribs. Description.—Shell attaining large size (over 150 mm in height), attached by the umbo of the right valve, which tends to become more elongate than the rela- tively pectiniform, less-inflated left one, especially as the high, triangular cardinal area increases in height with shell growth. Hinge line straight, with narrow car- dinal area marked by a much shorter ligamental pit than that extending the full height of the area of the right valve. Sculpture of concentric, foliated ribbing strong on the upper portion of the right valve, and often extending the full length of the shell, weaker on the upper, left valve. Radial ribbing variable, com- monly three to five strong spinose primaries with two to five secondary and tertiary ribs between each pair. However, the relative strength of these ribs varies greatly, with the secondaries on some specimens be- coming as strong as the primaries and developing spines of equal strength; in contrast, the tertiaries may weaken so as to be microscopic or, occasionally, divaricate as growth proceeds to the point that the number of ter- tiary ribs on a valve may differ between each pair of primaries. Lectotype.— ANSP 2869 (selected by Pilsbry, 1922, pl. 43, fig. 5). Type locality.—Mao Formation, Samba Hills, south of Guayubin (here restricted), Dominican Republic. Material.—In addition to the type lot of five valves, five paired valves, plus seven more or less complete adults and numerous fragments and juveniles, almost all from the Mao Adentro Member of the Mao For- mation. 12 BULLETIN 339 Measurements (in mm).— diameter (paired height length valves) locality ANSP 2869! 175 115 ~ unknown® ANSP 72724A? 154 103 _ unknown® ANSP 72724B? 96.5 94 _ unknown® ANSP 72724C?? 126 97.5 — unknown® ANSP 72724D> 162 123 = unknown® USNM 450390 81 56 47 TU 1221 PRI 21986 74 51 50 Hodson 2436 PRI 21987° 80 76 43 Hodson 2436 ' lectotype; ? paralectotypes; * specimen figured by Pilsbry (1922) and Palmer (1938) as “paratype”; * holotype of S. carmenensis Hod- son; > paratype of S. carmenensis Hodson; ° Gabb Collection, “Santo Domingo”. Remarks.—The specific name S. gumanomocon was given by Brown and Pilsbry (1913) to material in the Gabb Collection, which Gabb had identified as S. americanus Lamarck, 1819. The exact locality of Gabb’s five specimens remains unknown. Although we have specimens from the Rio Gurabo and the Rio Mao, the majority of our shells were collected in the Samba Hills just south of Guayubin (locs. TU 1221, 1245), an area that Gabb almost certainly collected. In the narrative of his travels in Santo Domingo he says, concerning the Samba Hills, near Guayubin: “They are barely fifty feet high immediately adjoining the Guayubin River, though twice that a mile or two east, and they almost disappear very soon after crossing the river. The horizontal beds continue to their southern base undisturbed, and where the road first reaches ris- ing ground it climbs a few feet up the face of a sort of bluff, the exposed edges of a nearly horizontal sand- stone, full of Oysters and Spondylus” (Gabb, 1873, p. 155) This description fits our localities TU 1221 and 1245 (4.5 and 5 km south of the bridge at Guayubin, re- spectively) perfectly, for here the on/y shelled fossils are ““Ostrea”’ haitensis Sowerby, 1850, and Spondylus gumanomocon (other mollusks are represented by molds). Although we are reasonably sure this is the area that Gabb’s material came from, there is one dif- ference between his material and ours: all of his shells are single valves, gray in color, and most of ours are paired valves, yellowish in color. So Gabb probably collected near our localities but not at them. For this reason, we have not restricted the type locality to one of our locality numbers. Although the majority of specimens of S. guman- omocon come from the Mao Adentro Limestone Mem- ber of the Mao Formation, the species is not confined to those beds. Maury (1917, p. 355) cited the species as occurring at “Zone D, Rio Gurabo at Los Quema- dos” (= locs. TU 1215, NMB 15842-15858; see Saun- ders, Jung, and Biju-Duval, 1986, text-figs. 4, 5) and both we and the Naturhistorisches Museum team col- lected the species at this locality. This is not unexpected as the locality is a coral reef in the Gurabo Formation. The Naturhistorisches Museum team also collected a single valve at Canada de Zamba, in the Rio Cana drainage (loc. NMB 16818; see Saunders, Jung, and Biju-Duval, 1986, text-fig. 15) in the coralline facies of the Gurabo Formation. Furthermore, they collected one specimen from the Cercado Formation, just above the mouth of Arroyo Bellaco (loc. NMB 16853; see Saunders, Jung, and Biju-Duval, 1986, text-fig. 15) near the coral reef in that tributary, and a second specimen from the Cercado Formation just below the contact with the Gurabo Formation on the Rio Cana (loc. NMB 16842; see Saunders, Jung, and Biju-Duval, 1986, text- fig. 15). The species S. carmenensis Hodson in Hodson, Hodson, and Harris, 1927 [p. 41, pl. 25, figs. 1-3], described from the “Oligocene” of the Buchivacoa Dis- trict, State of Falcon, Venezuela, but probably coming from the Miocene La Puerta Group’, agrees in shape and ornamentation with S. gumanomocon, and was synonymized by Palmer (1938, p. 155). We see no reason to disagree. Comparisons.—Gabb (1873, p. 257) identified this species as Spondylus americanus Lamarck, 1819 (= S. americanus Hermann, 1781). Specimens of that species in the Tulane collections reveal a much more spinose ornamentation with all radials — primary, secondary, and tertiary—having spines that tend to be narrowly and elongately pointed. Given the inherent variability of species of Spondylus, and the fact that many of the molluscan species present in the fauna of the Mao Formation are still living in the western Atlantic or Caribbean area today, the two forms may ultimately prove to be the same. Certainly it seems probable that S. gumanomocon and S. americanus are part of the same lineage. Occurrence.—Cercado/Gurabo formations: Rio Cana area (NMB 16818, 16842, 16853); Rio Gurabo (TU 1231 [? = TU 1215]; NMB 15837). Mao Formation: * The type locality was given subsequently by Hodson and Hodson (1931, p. 5) as southeast of Dabajuro, and expanded by Palmer (1938, p. 154), to “about 10 miles east and 4 miles south of Dabajura [sic]”’. The type locality for the La Puerta Formation (now Group) is the La Puerta syncline, southeast of Dabajuro. The Lexico Estratigrafico de Venezuela (1970, p. 672) indicates only the La Puerta Group in the Buchivacoa District. It is the lateral equivalent of the Socorro, Urumaco and Codore formations, just to the east, and the exact stratigraphic level is uncertain. Guayubin area (TU 1221, 1245, 1281); Rio Gurabo area (TU 1208, 1366; NMB 15822, 15825); Rio Mao (hwWr1336): Distribution.—Coralline facies of the Cercado For- mation and the Gurabo Formation, Mao Adentro Limestone Member of Mao Formation; Dominican Republic. ?7La Puerta Group, Venezuela; Miocene. Un- known formation, Costa Rica; Pliocene. NEOGENE PALEONTOLOGY IN THE NORTHERN DOMINICAN REPUBLIC 13. The Class Echinoidea (Echinodermata) By PorRTER M. KIER* Department of Paleobiology United States National Museum of Natural History Smithsonian Institution Washington, DC 20560, U.S. A. ABSTRACT Nine echinoid species are reported from the Miocene and Pliocene of the Dominican Republic. Four species are still living in the Caribbean and two of the others closely resemble living species. For this reason it is suggested that living conditions in the Miocene and Pliocene were very similar to those now present in the region. One new species is described from the early Pliocene: Clypeaster maoadentroensis. RESUMEN Se describen nueve especies de equinoidos del Mioceno y Plioceno en la Reptblica Dominicana. Cuatro especies estan todavia viviendo en el mar Caribe, y dos de los otros se parecen mucho a especies actuales. Por esta razon, se cree que el ambiente natural en el Mioceno y Plioceno se parece mucho a él de hoy en la region. Se describe un especie nueva del Plioceno temprano: Clypeaster maoadentroensis. INTRODUCTION Only a few fossil echinoids have been described from the Dominican Republic (Jackson, 1922). The present collection, for the most part made by Peter Jung, John Saunders, and Emily and Harold Vokes, is important not only because of the number of specimens but also because they were collected from measured sections, making it possible to know with certainty their age relative to each other. The fauna is of particular interest because of the close similarity of the fossil echinoids to those now living in the Caribbean. This similarity makes it possible to suggest the living habits and hab- itats of these fossil species. Background information on the collecting localities has been presented in detail by Saunders, Jung, and Biju-Duval (1986), and faunal monographs on many of the faunal groups present have been published pre- viously in this series (e.g., Jung, 1986; Foster, 1986; Foster, 1987; Cairns and Wells, 1987; Logan, 1987; Bold, 1988; E. Vokes, 1989; H. Vokes, 1989; Jung and Petit, 1990). A map showing collected sections in the Cibao Valley of the northern Dominican Republic that appears in all of these publications (e.g., Saunders, Jung, and Biju-Duval, 1986, text-fig. 3) is reproduced here as Text-figure 1. ACKNOWLEDGMENTS I thank Peter Jung, John Saunders, Emily and Har- old Vokes not only for collecting these fossil echinoids but for permitting me to study them. They also read the manuscript, and I thank them for their suggestions. Additional reviews were undertaken by M. McKinney (University of Tennessee, Knoxville, TN) and Graig Shaak and Douglas Jones (Florida State Museum, Uni- versity of Florida, Gainesville, FL). Peter Hoover was of great assistance in the preparation of the manuscript for publication. Without his help the paper would not have been published. The photography was done by Arnold Powell and the artwork by Mary Parrish. PREVIOUS WORK Jackson (1922, p. 6) studied a small collection of fossil echinoids from the Dominican Republic that is now in the USNM. In this collection, he identified seven species, two of them, C/ypeaster concavus Cot- teau, 1875 and Brissopsis antillarum Cotteau, 1875, from beds he considered Oligocene. Four he consid- ered Miocene, including Cidaris sp. a of Jackson (1922), * Current address: Osprey Point, Route 784, Lottsburg, VA 22511. 14 BULLETIN 339 Table 1.—Distribution of northern Dominican Republic echinoid species, by formation and age. age unit species early Pliocene Gurabo Fm. late Miocene Cercado Fm. late early Miocene Baitoa Fm. Mao Fm., Mao Adentro Mbr. Echinometra lucunter (Linnaeus, 1758) Clypeaster maoadentroensis, new species Moira atropos (Lamarck, 1816) Brissopsis jimenoi Cotteau, 1875 Schizaster doederleini (Chesher, 1972) Clypeaster caudatus Jackson, 1922 Clypeaster cf. C. sunnilandensis Kier, 1963 Encope aberrans imperforata Kier, 1963 Mellita sp. Cidaris melitensis Wright, 1855, Echinopedina cub- ensis Cotteau, 1881, and C/ypeaster caudatus Jackson, 1922. One species, C/ypeaster dalli (Twitchell in Clark and Twitchell, 1915), he considered Miocene or Plio- cene. Jackson had five specimens from the Cevicos Lime- stone that he referred to C/ypeaster concavus, but four of them are only fragments and can not be specifically identified with any certainty. The one whole specimen is aberrant, having only four petals, and may be C. concavus, although its oral surface is not as concave as is typical in this species. No specimens of this species are present in the new collections. The specimen from the Cevicos Limestone that Jackson identified as Brissopsis antillarum is badly crushed, making positive identification impossible. It is not conspecific with the specimen in the present collections referred to Brissopsis jimenoi Cotteau, 1875. VALVERDE SANTIAGO RODRIGUEZ, ( The posterior petals in Jackson’s specimens are straight, whereas in B. jimenoi they are confluent adapically but diverge distally. The Dominican Republic specimens that Jackson referred to Cidaris sp. a and Cidaris melitensis are based only on spines. Many cidarid spines are in the present collection but are not described. The Domin- ican Republic specimen from the Gurabo Formation that Jackson referred to Echinopedina cubensis was considered to be a different species by Lambert (in Sanchez Roig, 1949, p. 39), who made it the holotype of anew species, Hebertia jacksoni Lambert in Sanchez Roig, 1949. However, this specimen is very badly weathered, with all the tuberculation removed and the peristome broken away. It cannot be generically iden- tified. Jackson’s C/ypeaster caudatus is well-represented in the present collections and one of the new specimens 10 20km 4 Rio Cana 2 Rio Gurabo 3 Rio Mao 4 Rio Amina ~) Oligocene - Early Miocene ? 5 Canada Zaloya 6 Rio Yaque del Norte 7 City of Santiago 8 Arroyo Punal 9 Rio Verde Upper Cenozoic Text-figure 1.—Locality map for the sections measured and described by Saunders, Jung, and Biju-Duval (1986). The TU collections were made in these same areas but in intervening areas also. See Appendix 4 of that work for a complete description of all TU localities. DOMINICAN REPUBLIC NEOGENE. 13: P. M. KIER 15 is illustrated (PI. 5, figs. 1, 2). The specimen he referred to Clypeaster dalli is consubspecific with specimens of that species from the Pleistocene Caloosahatchee For- mation and post-Caloosahatchee beds in Florida (Kier, 1963, p. 29). Kier considered C. da/li to be a subspecies of C. rosaceus (Linnaeus, 1758). BIOSTRATIGRAPHY Nine species of echinoids can be identified from the Neogene of the Dominican Republic (Table 1). Six occur in the Rio Cana section (Text-fig. 2): one in the Cercado Formation, four in the Gurabo Formation, and one in the Mao Adentro Limestone. The oldest species, Encope aberrans imperforata Kier, 1963, oc- curs in the Cercado Formation 150 m above the base of the Rio Cana section in beds considered of probable late Miocene age (Saunders, Jung, and Biju-Duval, 1986, text-fig. 16). This subspecies has been reported elsewhere (Kier, 1963, p. 35) from the post-Caloosa- hatchee beds and the Tamiami Formation (Bucking- ham Limestone Member) of Florida. Dubar (in Oaks and Dubar, 1974) dates the Caloosahatchee as early to medial Pleistocene and the Buckingham Limestone Member of the Tamiami as medial to late Pliocene. Four species occur near the base of the Gurabo For- mation (317-350 m above the base of the Rio Cana section) in beds considered by Saunders, Jung, and Biju-Duval (1986, text-fig. 16) to be of early Pliocene age. Brissopsis jimenoi Cotteau, 1875 is reported from beds in Cuba considered of Miocene age, but this age determination was made in 1875 and it is not known from what formation the specimens were collected. B. jimenoi is very similar and may be synonymous with Brissopsis elongata Mortensen, 1907, now living in the Caribbean. Clypeaster caudatus Jackson, 1922 was originally re- ported from the Mao Adentro Limestone or Gurabo Formation of the Dominican Republic. Sanchez Roig (1949, p. 80) reported the species as occurring in the Oligocene of Cuba, but most modern workers consider Sanchez Roig’s “‘late Oligocene” of Cuba as Miocene. I have not been able to compare the Cuban specimens with those from the Dominican Republic so I can not confirm this identification. Gordon (1963, p. 636) con- sidered C. caudatus to be a a synonym of C. cubensis Cotteau, 1875, which is known from the Miocene of Cuba and lower or middle Miocene of Puerto Rico. The petals in C. caudatus are less inflated than in the specimens Gordon referred to C. cubensis, and the two species are probably distinct. C. caudatus is also very similar to C/ypeaster rosaceus (Linnaeus, 1758), known from the Miocene to Recent of the Caribbean region. Specimens of C. caudatus also occur in the northern Dominican Republic at Arroyo Lopez. Poddubiuk (1985) argues that C. rosaceus is derived from C. cub- GE FORMATION A 800| THICKNESS IN METER 4 7+ Clypeaster maoadentroensis, new species | MAO (MAO ADENTRO LIMESTONE) FORMATION 500 600 700 L L L FORMATION EARLY PLIOCENE 4 GURABO 350 t Schizaster doederleini (Chesher) Moira atropos Lamarck Clypeaster caudatus Jackson Brissopsis jimenoi Cotteau 4 FORMATION PROBABLY LATE MIOCENE 3 Encope aberrans imperforata Kier CERCADO 100 i= Text-figure 2.—Occurrence of echinoids in the Rio Cana section. 16 BULLETIN 339 ensis and C. caudatus and discusses evolutionary changes in morphology in the lineage. Moira atropos (Lamarck, 1816) lives today in the Caribbean and has been found as fossil in the Pliocene of South Carolina and Venezuela. Schizaster doeder- leini (Chesher, 1972) now lives in the Caribbean and has never before been found as a fossil. The youngest species in the Rio Cana section, C/y- peaster maoadentroensis, n. sp., occurs low in the Mao Adentro Limestone, approximately 765 m above the base of the section. Saunders, Jung, and Biju-Duval (1986, text-fig. 16) consider these beds to be of late early Pliocene age. This species is quite distinct from C. caudatus, which occurs lower in the section in the Gurabo Formation. I know of no well-dated species of Clypeaster Lamarck, 1801, with which it has clear af- finities. Three species do not occur in the Rio Cana section. Mellita sp. is present in the Baitoa Formation, which is considered older than the Cercado Formation. Eames et al. (1962, text-fig. 5) place the Baitoa in the middle or late Burdigalian or late early Miocene. In the Baitoa, Mellita is represented only by immature specimens whose affinities with other species are uncertain. C/y- peaster cf. C. sunnilandensis Kier, 1963 occurs in the Cercado Formation (of probable late Miocene age). The specimens are only fragments but are quite similar to, though probably not conspecific with, specimens of C. sunnilandensis from the Pliocene of Florida. Echi- nometra lucunter (Linnaeus, 1758) is found in the Mao Adentro Limestone (of late early Pliocene age, accord- ing to Saunders, Jung, and Bijyu-Duval, 1986, text-fig. 16). This species lives today in the West Indies and off the west coast of Africa. It has been found previously as fossil from Jamaica, and from the Pleistocene of Angola and Florida. Fragments of sand dollars occur at many horizons in the measured sections at Rio Cana (Saunders, Jung, and Biju-Duval, 1986, text-figs. 15, 16). They are found in the Cercado Formation at localities NMB 16836, 16839, and 16842, approximately 228 to 241 m above the base of the section. They also occur in this section in the Gurabo Formation at localities NMB 16818 and 16858, 315-322 m above the base. Higher in the sec- tion, they occur in the Mao Adentro Limestone at locality NMB 16873, 690 m above the base. In the Rio Gurabo section (Saunders, Jung, and Biju-Duval, 1986, text-figs, 5, 6), fragments of sand dollars occur below the base of the Gurabo Formation at localities NMB 15904, 15906, 15907, 15910, 15911, 15914, and 15915, which are 110-132 m above the base of the section. Finally, they occur in the Rio Yaque del Norte section (Saunders, Jung, and Biju-Duval, 1986, text- fig. 21) at La Barranca at locality NMB 17268. PALEOECOLOGY Because so many of the Dominican Republic echi- noid species are still extant, it is feasible to predict how the fossil echinoids lived. Encope aberrans imperforata Kier, 1963 is very similar to Encope aberrans aberrans Martens, 1867, which according to Serafy (1979, p. 76) lives from Cape Hatteras south to the Bahama Islands and throughout the Gulf of Mexico at depths from 12- 90 m. In the Gulf of Mexico, Serafy reports that it lives on bottoms of crushed shell and quartz sand. Brissopsis jimenoi Cotteau, 1875 is very similar to, and may be synonymous with Brissopsis elongata Mor- tensen, 1907, which lives along the coasts from Ven- ezuela to Panama, and Belize and Puerto Rico at depths from 13-72 m. Kier (1975, p. 16) found this species living buried 40-100 mm in mud at depths of 12-18 m. Clypeaster caudatus Jackson, 1922 is extinct, but is very similar to C/ypeaster rosaceus (Linnaeus, 1758), now living in the Caribbean at depths from intertidal to 285 m. In the Florida Keys (Kier and Grant, 1965, pp. 26, 27), this species is abundant in 1-15 m, living on sand patches in or near fields of turtle grass. Schizaster doederleini (Chesher, 1972) and Moira atropos (Lamarck, 1816) occur in the Gurabo For- mation at the same locality. They also are found living together today in the Caribbean. Kier (1975, p. 14) reports the two species living buried 40-100 mm in mud at a depth of 12-18 m off Belize. Elsewhere M. atropos has been reported from the coast of North Carolina south to Brazil from littoral depths to 145 m. Clypeaster maoadentroensis, n. sp., 18 quite distinct from any living species. Clypeasterids today are con- fined to tropical or subtropical regions and commonly are littoral—-sublittoral, rarely occurring as deep as 500 m (Mortensen, 1948, p. 17). Among living species of Clypeaster Lamarck, 1801, Clypeaster cf. C. sunnilan- densis Kier, 1963 most resembles C. subdepressus (Gray, 1825), which according to Serafy (1979, p. 66) is known from North Carolina southward through the Greater and Lesser Antilles, the Gulf of Mexico, and southward along the coasts of Central and South Amer- ica to Brazil. It lives at depths from 5-210 m, but is most common in less than 50 m. Kier and Grant (1965, p. 28) found it most abundant between 5 and 12 m on sandy areas with little grass. Echinometra lucunter (Linnaeus, 1758) lives today from Florida to Brazil and off the west coast of Africa. It is most common in shallow water less than 3 m deep (Kier and Grant, 1965, p. 18) and commonly lives in niches in rock. In summary, seven of the Dominican Republic species are alive today or are very similar to species that live today. All of these living species occur in DOMINICAN REPUBLIC NEOGENE. 13: P. M. KIER 17 shallow water in the West Indies. Presumably, the con- ditions where the fossils lived were similar to those now in the Caribbean. The most striking aspect of the Dominican Republic echinoid faunas is the large number of Miocene and Pliocene species that are still extant or are very similar to species now living. Three of the six early Pliocene species: Echinometra lucunter (Linnaeus, 1758), Moira atropos (Lamarck, 1816), and Schizaster doederleini (Chesher, 1972), are still alive; and a fourth, Brissopsis jimenoi Cotteau, 1875, may be synonymous with a living species, B. elongata Mortensen, 1907. The early Pliocene Clypeaster caudatus Jackson, 1922 is very similar to the living C. rosaceus (Linnaeus, 1758). Only one early Pliocene species, C/ypeaster maoadentroen- sis, N. Sp., 18 clearly distinct from any living species. Of the two late Miocene species, only Encope aber- rans imperforata Kier, 1963 is still extant, although subspecifically differentiated. The second, C/ypeaster cf. C. sunnilandensis Kier, 1963 is very similar to the living C. subdepressus (Gray, 1825). The early Miocene Mellita sp. appears to be distinct from any living species. The longevity of the Dominican Republic species is in sharp contrast to the Neogene echinoid faunas of the southeastern United States. Only two of the 16 species of echinoids that can be identified with cer- tainty from the Pliocene of the eastern United States are still alive and both these species are subspecifically differentiated: Lytechinus variegatus plurituberculatus Kier, 1963 and Encope aberrans imperforata Kier, 1963. I suspect that the longevity of the Dominican Republic species is probably an indication that the environmental conditions in the region remained rel- atively the same, whereas they changed in the south- eastern United States. SYSTEMATIC PALEONTOLOGY INTRODUCTION The type specimens are housed in the Naturhisto- risches Museum, Basel, Switzerland (NMB) and the U. S. National Museum of Natural History (USNM). The localities are either those of the NMB or of Emily and Harold Vokes of Tulane University, New Orleans, LA (TU). The stratigraphic and geographic location of all the NMB localities and some of the TU localities are plotted on maps and sections in Saunders, Jung, and Biju-Duval (1986). Other abbreviations used in this section include: Museum of Comparative Zoology, Harvard University, Cambridge, MA (MCZ). Many measured taxonomic characters of echinoids can be expressed in terms of percent of test length, herein abbreviated as %L. SYSTEMATICS Genus ECHINOMETRA Gray, 1825 Echinometra lucunter (Linnaeus, 1758) Plate 4, figures 1-4 Echinus lucunter Linnaeus, 1758, p. 665. For a synonymy, see Mortensen, 1943, p. 357. Kier and Grant (1965) and McPherson (1969) de- scribe aspects of the biology of this species. Five specimens can be referred to this species. They are identical in all respects to Recent specimens of this species from the Florida Keys (see Pl. 4, figs. 1-4 for comparison of one of the fossil specimens with a Re- cent specimen). The fossil and Recent specimens share a similar number of plates relative to size, similar num- ber of pore pairs in each ambulacrum, similar tuber- culation, and tests of similar elongated shape. Type material.— Figured specimen, USNM 375449. Occurrence.—Echinometra lucunter occurs at local- ity TU 1438 in the Mao Adentro Limestone (Pliocene), in a roadcut 0.5 km south of the bridge at Guayubin, on the road to Sabaneta. Distribution.—Echinometra lucunter lives today in the West Indies from Florida to Brazil and off the west coast of Africa. Arnold and Clark (1934, p. 140) report it as a fossil from Jamaica, Dartevelle (1953, p. 38) found it in the Pleistocene of Angola, Kier (1963, p. 19) found it in the Pleistocene Caloosahatchee For- mation of Florida, where Donovan and Gordon (1989) also reported it in the Pleistocene Falmouth Formation. Genus CLYPEASTER Lamarck, 1801 Clypeaster maoadentroensis, new species Plate 4, figures 5-7; Text-figures 3, 4A Etymology.—The species is named for the formation where the holotype was collected. Material.—One moderately well-preserved speci- men, which is somewhat weathered dorsally but whose test appears to be undistorted. Shape and size.—Test, 98 mm long, 84 mm wide, and 44 mm high; greatest width central, greatest height slightly posterior to center, with test sloping more sharply posteriorly than anteriorly. Petals only very slightly inflated. Ventrally, test depressed deeply only in immediate vicinity of peristome. Apical system.—Monobasal, five genital pores, cen- ter of apical system located slightly posterior to center at distance from anterior margin equal to 53 %L. Ambulacra.—Petals long, extending almost to mar- gin of test; anterior petal III slightly longer than other petals, length = 44 %L, petal IV = 39 %L, petal V = 42 %L; width of petals = 23-25 %L. Poriferous zone at greatest width = 5.5 %L, or 39 percent of the width 18 BULLETIN 339 of interporiferous zone at its greatest width. Petals clos- ing distally with poriferous zone of same petal almost in contact; petal III with 80 pore pairs (in both zones), petal IV with 84, and petal II with 82. Distinct am- bulacral groove extending ventrally from near margin to peristome. Peristome.—Situated slightly posterior to center with center of peristome located at distance from anterior margin equal to 54 %L. Size of opening uncertain be- cause of fracturing around peristome. Periproct.—Located near posterior margin, opening wider than high with width = 5.4 %L, height = 5.1 %L. Sutures not clear enough to determine which plates enclose periproct. Diagnosis.—Species characterized by smooth dorsal surface with petals only slightly inflated, pore pairs widely separated from each other, resulting in fewer pore pairs in petals, ventral surface depressed only in vicinity of peristome. Type material.— Holotype, NMB M9746. Comparison with other species.—This species is dis- tinguished from Clypeaster caudatus Jackson, 1922, which occurs lower in the Rio Cana section by its longer petals, wider poriferous zones, and in having the pore pairs in its petals more widely separated from each other in the same poriferous zones, resulting in fewer pore pairs in each petal. For example, a specimen of C. caudatus (NMB catalogue number unknown) with a petal II that is 39 mm long has 120 pore pairs, where- as the holotype of C. maoadentroensis, having a slight- ly longer petal (43.2 mm), has only 80 pore pairs. The greatest distance between two adjacent outer pores (measured along the length of the petal) near the mid- length (Text-fig. 4A) of petal III in the holotype of C. maoadentroensis is 1.3 mm, but is only 0.8 mm in C. caudatus (Text-fig. 4B) in petal III of a specimen of similar size. Furthermore, the peristome in C. maoad- entroensis is slightly posterior to the center, whereas it is central in C. caudatus. Finally, the ventral surface is depressed only around the peristome in C. maoad- entroensis instead of over much of the ventral surface in C. caudatus. Although the relative inflation of the petals, position of the peristome, and shape of the test are characters that vary considerably within a species of Clypeaster, the distance between pore pairs in a poriferous zone is fairly constant, and therefore a re- liable character for specific differentiation. Of all the other species of C/ypeaster from the West Indian region, C. maoadentroensis most resembles Clypeaster duchassaingi Michelin, 1855 from the “‘for- mations madréporiques” of Guadeloupe. C. maoad- entroensis differs in having its apical system posterior to the center of the test, as opposed to anterior in C. duchassaingi, in having its posterior petals extending much nearer the posterior margin, a more posteriorly situated peristome, a blunter posterior margin, and a narrower anterior. Occurrence. —Early Pliocene, Mao Adentro Lime- stone, locality NMB 17022, Rio Cana section (Saun- ders, Jung, and Biyu-Duval (1986, text-fig. 15). E. Vokes (written commun., 1984) considers this formation to be middle Pliocene in age. Clypeaster caudatus Jackson, 1922 Plate 5, figures 1, 2; Text-figure 4B Clypeaster caudatus Jackson, 1922, p. 36; pl. 3, figs. 1, 2; Sanchez Roig, 1949, p. 80, pl. 10, fig. 2; ?Gordon, 1963, p. 636. Eight specimens and four fragments can be referred to this species, whose holotype is also from the Do- minican Republic. They are similar to the holotype in having only slightly inflated petals, which close distally with their poriferous zones meeting at the end of each petal. Their tests are also relatively wide (84 to 94 %L) and their ventral surfaces are deeply depressed around the peristome. Gordon (1963, p. 636) considered this species a syn- onym of Clypeaster cubensis Cotteau, 1875. I have compared the Dominican Republic specimens with the specimens that Gordon referred to C. cubensis from the Miocene Ponce Limestone of Puerto Rico. Al- though they are very similar, the Dominican Republic specimens have less inflated petals. Whether or not Text-figure 3.— C/ypeaster maoadentroensis, n. sp., dorsal view of holotype, NMB M9746, x1. DOMINICAN REPUBLIC NEOGENE. 13: P. M. KIER 19 this difference is significant will not be known until more specimens can be studied. As pointed out by Kier (1963, p. 29), the extent of inflation of the petals is very variable in C/ypeaster rosaceus (Linnaeus, 1758), a species quite similar to C. caudatus. Type material.—Holotype, USNM 328235; para- type, USNM 328236; figured specimen, NMB M9747. Occurrence.— According to Jackson (1922, p. 36), the holotype of Clypeaster caudatus came from float from the Gurabo Formation, or the Mao Adentro Limestone, Rio Gurabo, near Los Quemados, Domin- ican Republic. The new material is from locality NMB 16858 (Saunders, Jung, and Biyu-Duval, 1986, text- figs. 15, 16) in the Rio Cana section 320 m above the base of the section near the base of the Gurabo For- mation, from beds considered to be of early Pliocene age; and localities NMB 17272, 17274 (Saunders, Jung, and Biju-Duval, 1986, text-figs. 24, 26) in the Arroyo Lopez section, beds B and D. The Vokes’ material came from locality TU 1215, Gurabo Formation, Rio Gurabo, from bluffs on both sides, from the ford on the Los Quemados-Sabaneta road, upstream to ap- proximately | km above the ford (= locs. USGS 8539- 8543; Maury’s Zone D), and locality TU 1444, Gurabo Formation, Rio Yaque del Norte, east bank, at Lopez, appoximately 0.5 km upstream from the mouth of Arroyo Lopez, between the middle and lower hard limestone ledges. Distribution. —Clypeaster caudatus was reported by Sanchez Roig (1949) from El Jobo, near Puriales, \\ Len A B Text-figure 4.—Comparison of left poriferous zone of petal III of Clypeaster maoadentroensis (A) and C. caudatus (B) showing greater width separating pores of a pore pair in C. maoadentroensis and greater distances between adjacent pore pairs. A, holotype, NMB M9746; B, figured specimen, NMB M9747, x3. Guantanamo, Oriente Province, Cuba. He attributed a late Oligocene age to the specimen, but it would probably now be considered Miocene. Clypeaster cf. C. sunnilandensis Kier, 1963 Plate 6, figures 1, 2 cf. Clypeaster sunnilandensis Kier, 1963, pl. 3, fig. 3, pls. 12, 13. The three fragments recovered appear to be very similar to C. sunnilandensis from the Pliocene Tam- iami Limestone of Florida. The test of one of the frag- ments is wider more anteriorly than in specimens of C. sunnilandensis and the test appears to be lower. These specimens probably represent a new species, but with only three fragments, none of which show the ventral surface, it is not possible to make this deter- mination. Type material.— Figured specimens, USNM 375450, 375451. Occurrence.— The form here compared to C/ypeaster sunnilandensis was recovered from locality TU 1223, Cercado Formation, roadcut 5.3 km north of the plaza at Moncion, on the road to Los Quemados. Distribution.—Clypeaster sunnilandensis is known from the Pliocene Tamiami Limestone of Florida. Genus ENCOPE J. L. R. Agassiz, 1841 Encope aberrans imperforata Kier, 1963 Plate 6, figure 3 ?Encope wiedenmayeri Jeannet, 1928, p. 17. ?Encope michelini J. L. R. Agassiz. Cooke, 1961, p. 17. Encope michelini imperforata Kier, 1963, p. 33, pl. 5, fig. 1; pl. 6, figs. 3, 4, text-figs. 25-30, table 2; Phelan, 1972, pp. 117, 125, 126. One specimen showing its dorsal surface appears to belong to this subspecies, originally assigned to Encope michelini J. L. R. Agassiz, 1841. Subsequently Phelan (1972) revised the genus and showed that FE. aberrans Martens, 1867 should be maintained as a species sep- arate from E. michelini, and that E. michelini imper- forata should be referred to E. aberrans. E. aberrans imperforata was based on specimens from the Plio-Pleistocene Caloosahatchee and Tam- iami formations of Florida. This subspecies is similar in all respects to the living E. aberrans aberrans except that its posterior closed lunule is quite small or absent, whereas it is always fully developed in the living species. The specimen from the Dominican Republic also has a smaller posterior lunule. Although its posterior petals are longer than the other petals, and its test narrower than normally found in E. aberrans, I have seen a few modern specimens of EF. aberrans that are as narrow and have posterior petals longer than the anterior. The specimens from the Miocene or Pliocene of Ven- 20 BULLETIN 339 ezuela that Jeannet (1928, p. 17) referred to Encope wiedenmayeri Jeannet, 1928, and Cooke (1961, p. 17) referred to Encope michelini J. L. R. Agassiz, 1841, may be consubspecific with E. aberrans imperforata. More specimens are needed before this assignment can be made with any certainty. Type material.—Florida specimens: holotype, USNM 648167; figured specimens, USNM 648168- 648172: Dominican Republic figured specimen, NMB M9748. Occurrence.—Encope aberrans imperforata has been recovered in the present collections from locality NMB 16857 (Saunders, Jung, and Biyu-Duval, 1986, text- figs. 15, 16), late Miocene from the Rio Cana section, 150 m above the base. Distribution.— Florida: Plio-Pleistocene post-Ca- loosahatchee beds, and Caloosahatchee and Tamiami formations (see Kier, 1963, p. 35 for exact localities). Specimens questionably consubspecific with E. aber- rans imperforata have been reported from the Miocene or Pliocene of Venezuela (Jeannet, 1928; Cooke, 1961). Genus MELLITA J. L. R. Agassiz, 1841 Mellita species Plate 6, figures 4, 5 Eight small specimens can be referred tentatively to this genus. Although they have five ambulacral notches and their periprocts are outside the basicoronal plate, characters (Kier, 1963, p. 44) commonly found in Leo- dia Gray, 1852, they have more affinities with Mellita, including: (1) a posterior lunule that extends far an- teriorly between the posterior petals; (2) paired inter- ambulacra separated from the basicoronal row by one pair of ambulacral plates; (3) the first pair of post- basicoronal plates in the paired interambulacra elon- gated: and (4) lunules formed by the closing of marginal notches. These specimens appear to represent a new species but because no large specimens are present it seems inadvisable to erect one. They differ from Mellita acli- nensis Kier, 1963, from the Plio-Pleistocene of the southeastern United States, in having a wider posterior lunule and the periproct not partly within the basi- coronal interambulacral plate. They differ from Mellita quinquiesperforata (Leske, 1778), now living in the Ca- ribbean, in having five instead of four ambulacral notches, and their periproct more posterior and outside of the basicoronal plate. No small specimens of Mellita caroliniana (Ravenel, 1841) from Plio-Pleistocene beds in South Carolina are available, so it is difficult to compare these specimens to this South Carolina species. However, the periproct in M. caroliniana is within the basicoronal plate (Kier, 1972, fig. 4), whereas in the Dominican Republic specimens it is posterior to it. Type material.— Figured specimen, USNM 375452. Occurrence.— Late early Miocene, locality TU 1253, Baitoa Formation, roadcut on the west side of road from Santiago de los Caballeros to Baitoa, 1 km north of the village of Baitoa (= loc. USGS 8559). Locality TU 1362 (Saunders, Jung, and Biju-Duval 1986, text- fig. 21), Baitoa Formation, trail that leads to top of bluff, east side of Rio Yaque del Norte, just down- stream from Baitoa. Locs. NMB 16943, 17282, 17283, 17290 (Saunders, Jung, and Biju-Duval, 1986, text-fig. 25), Lopez section of Rio Yaque del Norte. Genus MOIRA A. Agassiz, 1872 Moira atropos (Lamarck, 1816) Plate 7, figures 3, 4 Spatangus atropos Lamarck, 1816, p. 32. For synonymies, see Mortensen, 1951, p. 329, and Serafy, 1979, p. 91. The single specimen recovered in this study is in- distinguishable from specimens of this species now living from the coast of North Carolina and Bermuda through the Caribbean, Gulf of Mexico, and south to Brazil. Although the specimen is distorted and partially covered with a hard matrix, enough of the test is visible that it can be referred with certainty to M. atropos. Its petals are like those in Recent specimens, deeply sunk- en but constricted dorsally where the interambulacra almost meet above them. The petals of the fossil and Recent specimens have similar length and shape, and ambulacrum III is similar. Furthermore, the shape of their tests are the same, as are the positions of their apical system and peristome. The positions of both the peripetalous and lateroanal fascioles are the same, al- though because of preservation it is not possible to determine which plates the fascioles occur on in the fossil specimen. Type material.— Figured specimen, USNM 375453. Occurrence.—Early Pliocene, Gurabo Formation, locality TU 1354, Canada de Zamba, a tributary on the west side of the Rio Cana, approximately 2.5 km east of the village of Zamba, which is 7 km north of Cruz de Santiago (Santiago Rodriguez), on the road to Guayubin; or 4.5 km (airline) below the ford at Caimito (see Saunders, Jung, and Biju-Duval, 1986, text-fig. 15). Distribution.—Moira atropos (Lamarck, 1816) has been found (Cooke, 1959, p. 74) in beds now consid- ered of Pliocene age in the Intracoastal Waterway canal in Horry County, South Carolina, one-half to 1 mi southwest of the bridge on US 17 near Nixons Cross- roads, about 15 mi northwest of Myrtle Beach (loc. USGS 18759). Cooke (1961, p. 22) reports the species from the Pliocene San Gregorio Formation, Rio Seco area, Falcon, Venezuela. DOMINICAN REPUBLIC NEOGENE. 13: P. M. KIER ZA Genus BRISSOPSIS J. L. R. Agassiz, 1840 Brissopsis jimenoi Cotteau, 1875 Plate 7, figures 1, 2; Text-figures 5A, 5B Brissopsis jimenoi Cotteau, 1875, p. 6; Cotteau, 1881, p. 33, pl. 3, figs. S—9; Cotteau, 1897, p. 79, pl. 24, figs. 5-9; Jackson, 1922, p. 81; Jeannet, 1928, p. 12, pl. 1, fig. 35; Sanchez Roig, 1949, p. 225; Kier, 1984, p. 87, pl. 46, figs. 1, 2. Material.—Only a single specimen is available. It was originally covered with a sandy silt matrix, with most of its spines still attached to the test, showing B Text-figure 5.— Brissopsis jimenoi Cotteau, A, dorsal view; B, ven- tral view, NMB M9747, x1. that the test was covered at or very soon after death. The test is flattened and weathered, obscuring the fas- cioles. Shape and size.—Test, 65 mm long, 54 mm wide (83 %L), height, 25 mm (38 %L). Apical system.—Central, located at distance from anterior margin to center of genital pores equal to 50 %L. Four genital pores, ethmolytic with genital plate 2 extending far posteriorly. Ambulacra.— Anterior ambulacrum not petaloid, in groove extending from apical system to peristome; pore pairs largest adapical to peripetalous fasciole: minute between fasciole and phyllode; 56 plates in ambula- crum. First enlarged pore pairs in plate 7. Anterior petals (II and IV) curved convexly ante- riorly with their greatest width near end of petal; de- pressed in groove. Span of ends of petals = 42.6 %L. Short, with length slightly greater than one-half dis- tance from apical system to ambitus; length = 21 %L; greatest width = 8.1 %L; 44 petaloid pore pairs in each petal; pore pairs in anterior poriferous zones reduced in size adapically. First petaloid pore pair in plate 12. Posterior petals (V and I) confluent for three-fifths distance from apical system to end of petals, sharing common groove; curving away from each other dis- tally; short, length = 25 %L; greatest width = 6.3 %L: 48 pore pairs. Petaloid pore pairs in posterior porifer- ous zones greatly reduced in size from apical system for three-fifths length of petal. First petaloid pore pair in plate 19. Peristome.—Anterior, distance from anterior edge of peristome to anterior margin = 24 %L: width of opening = 21 %L; height = 7 %L. Periproct.—Inframarginal, large, width = 15 %L, height = 12 %L, occurring within plates 5-9. Fascioles.—Only short tract of peripetalous fasciole preserved in lobe extending anteriorly in anterior am- bulacrum (III), crossing plate 7 at distance from apical system = 36 %L. Oral plate arrangement.—Labrum extending to sec- ond adjoining ambulacral plate (Text-fig. 5B), with length = 6.6 %L; sternal plate (first plate of plastron) length = 41 %L (estimated), combined width = 31 %L: episternal plate (second plate of plastron) length = 14 %L, combined width = 32 %L. Type material.—Lectotype in the Cotteau Collec- tion, Université Claude Bernard, Lyon, France. Do- minican Republic figured specimen, NMB M9749. Remarks.—This Dominican Republic specimen ap- pears to be conspecific with the holotype of B. jimenoi. The two specimens are of similar shape, have similar petals, and labrum and sternal plates of similar di- mensions. Because the Cuban holotype is a cast, de- scription of the Dominican Republic specimen herein NO i) is justified; it exhibits features not visible on the ho- lotype. Comparison with other species.—This species differs from Brissopsis aguayoi Sanchez Roig, 1952, from the Oligocene-Miocene of Cuba in its larger, lower and longer anterior petals. It is distinguished by its shorter petals from Brissopsis antillarum Cotteau, 1875 from the Miocene Anguilla Formation of Anguilla and the Miocene La Vela Formation in Venezuela. Its anterior petals extend little over one-half the distance from the apical system to the margin, but in B. antillarum these petals extend two-thirds this distance. Furthermore, the test in B. jimenoi is more elongate, and lower. B. jimenoi has a much lower test and less divergent and more curved petals than the specimen from the Miocene of Costa Rica that Durham (1961, p. 484) referred to Brissopsis, N. sp. This species is very similar and may be synonymous with Brissopsis elongata Mortensen, 1907, now living in the Caribbean. The two species are similar in the following features (the dimensions for B. elongata are from Chesher, 1968): 1. Width of the test: 83 %L in B. jimenoi; mean of 82.3 %L in B. elongata, with a range from 77-86 %L. 2. Labrum extends to the second adjoining ambu- lacral plate in both species. 3. Same number of adjoining ambulacral plates along the plastron plates with the sixth ambulacral plate oc- curring at the suture between the first and second plates of the plastron. 4. Posterior petals confluent proximally, very di- vergent distally. Although many specimens of B. elon- gata do not have their posterior petals as divergent, Chesher (1968, pl. 15, fig. a) shows a specimen with petals of similar shape to those in B. jimenoi. 5. In both species, the first petaloid pore pair in an anterior petal is in plate 12, and the first in a posterior petal is in plate 19. 6. The peristome is in a similar position. In B. ji- menoi, it is located at a distance from the anterior margin equal to 24 %L; in B. elongata it is at 25 %L with a range of 23-28 %L. 7. The periproct in B. jimenoi is enclosed by inter- ambulacral plates 5-9; in B. elongata by interambu- lacral plates 5-8. 8. The span of the ends of the anterior petals is 42.6 %L in B. jimenoi, 48.8 %L in B. elongata with a range of 39-56 %L. 9. The peripetalous fasciole crosses ambulacrum Ill at a distance from the apical system of 36 %L in B. jimenoi. In B. elongata the mean of the distance is 38.9 %L with a range of 34-43 %L. 10. The length of the first plastron plates in B. ji- menoi is 41 %L in both B. jimenoi and B. elongata. 11. Although the apical system in B. jimenoi is more BULLETIN 339 posterior than in most specimens of B. elongata, with a distance equal to 50 %L versus a mean of 38 %L in B. elongata (no range given by Chesher), some speci- mens of B. elongata have their apical system as pos- terior. In three specimens from Belize in the USNM, the distance ranges from 44 to 52 %L. Although these species appear identical, I hesitate to place them in synonymy. Nothing is known of the pedicellariae in B. jimenoi, and with only two speci- mens known of B. jimenoi (each from a different lo- cality), it is not possible to make a statistical compar- ison of populations of the two species. Furthermore, the position of most of the length of the fascioles is unknown on the fossil specimens. This feature has been shown by Chesher to be very important systematically in Brissopsis. Finally B. elongata is junior to B. ji- menoi, and it would be unfortunate to synonymize this well-known living species with a fossil species based on a holotype which is only an internal mold. Jackson (1922, p. 83) referred a Dominican Republic specimen to Brissopsis antillarum Cotteau, 1875. This specimen in the USNM (uncatalogued) is from the Cevicos Limestone, Arroyo La Mora, west of Cevicos. It is badly crushed and can not be specifically identified with any certainty. Occurrence.—Early Pliocene, near base of Gurabo Formation, locality NMB 16824 (Saunders, Jung, and Biju-Duval, 1986, text-figs. 15, 16) 317 m above base of Rio Cana section. Distribution.—Cuba, Miocene?, San Martin Cien- fuegos, Las Villas Province. Jeannet’s (1928, p. 12) specimen from the middle Miocene of Trinidad may or may not belong to this species. Genus SCHIZASTER J. L. R. Agassiz, 1835 Schizaster doederleini (Chesher, 1972) Plate 7, figure 5; Plate 8, figures 1-6; Text-figure 6 Text-figure 6. — Schizaster doederleini (Chesher), oral view, figured specimen, USNM 375456, *2. DOMINICAN REPUBLIC NEOGENE. 13: P. M. KIER 23 Paraster doederleini Chesher, 1972, pp. 10-25, figs. 1-9; Kier, 1975, p. 9, pls. 7, 8, figs. 1-5, text-figs. 7, 8. Schizaster orbignyanus A. Agassiz, 1880. Serafy, 1979, fig. 37A. Twenty-four Dominican Republic specimens can be referred to this species, which now lives in the Carib- bean off Belize and Florida, and in the Gulf of Mexico offthe Dry Tortugas. Their dimensions are very similar to those of a living population of this species from Belize (Table 2). The specimen that Serafy (1979, fig. 37A) referred to Schizaster orbignyanus A. Agassiz, 1880 is S. doederleini. The fossil specimens differ only in having slightly shorter petals with a narrower span. Furthermore, the fossil specimens are similar to the Recent specimens in having very small anterior genital pores, similar number of pore pairs in ambulacrum III within the peripetalous fasciole (a fossil specimen 35.6 mm long has 56 pore pairs, a Recent specimen 36.7 mm long has 58), and similar number of petaloid pore pairs. In both the fossil and Recent specimens the peripetalous fasciole crosses plate 4 in ambulacrum III, plate 11 or 12 in ambulacrum IV, and in both populations the labrum extends back (Text-fig. 6) to the posterior por- tion of the first adjoining ambulacral plate on the an- terior portion of the second. Finally, both have the periproct within interambulacral plates 5-8. Type material.—Holotype, USNM E11376; para- types, MCZ 8397c, 8397d; figured specimens, USNM E13735, 13749, 13743-13750, E30468. Dominican Republic fossil figured specimens, USNM 375454, 375455, 375456. Comparison with other species.—This species is eas- ily distinguished from any fossil species known from Table 2.—Schizaster doederleini (Chesher): comparison of north- ern Dominican Republic fossils with Recent specimens from Carrie Bow Cay, Belize. mean percent of length fossil Recent (min/max) (min/max) width 90.3 (87-94) 96 (92-99) height 73.1 (69-80) 73.8 (70-80) apical system to anterior margin 53 (49-59) 53 (48-57) apical system to fasciole on ambulacrum III 50. (46-53) 53 (48-56) length of anterior petal 31.7 (30-33) 33 (34-38) width of anterior petal 11.2 (10.8-12.3) 12 (9-14) length of posterior petal 15 (14.1-16.3) 16 (14-19) width of posterior petal 8.4 (7.7-10.0) 9 (7-11) peristome to anterior of test 24.4 (19.4-27.2) 20 (15-25) span of anterior petals Sil, ((47—5'5) 58 (55-62) the region. It differs from Schizaster delgadoi (Sanchez Roig, 1953) from the Miocene of Cuba by its much smaller test, with more central apical system, shorter posterior petals, wider anterior petals that distally curve anteriorly instead of the reverse, wider ambulacrum III, and more posterior peristome. It differs from Schi- zaster fernandezi Sanchez Roig, 1952, also from the Miocene of Cuba, in its more central apical system, and wider ambulacrum III, which has more enlarged pore pairs, 50 in a specimen 26 mm long as opposed to only 28 in a specimen of the same size of S. fer- nandezi. It differs from Schizaster cartagensis (Sanchez Roig, 1949) from the Oligocene—Miocene of Cuba by its much shorter and wider petals, and from Schizaster rojasi Sanchez Roig, 1952, in having wider petals, and a higher test with its greatest width more posterior. Its posterior petals are shorter and the anterior petals more curved anteriorly than in Schizaster sanctmariae Sanchez Roig, 1949 also from the Oligocene—Miocene of Cuba. Of all the Oligocene—Miocene Cuban species, S. doederleini most resembles Schizaster munozi Sanchez Roig, 1949, from which it differs in having shorter, wider posterior petals and wider interior petals. S. doederleini is easily distinguished from the Pleis- tocene Schizaster eustatii (Engel, 1961) from the Ca- ribbean island of St. Eustatius by having many more pore pairs in the dorsal part of ambulacrum III within the peripetalous fasciole. Only 32 pore pairs occur in this region in S. eustatii, whereas in a specimen of similar size of S. doederleini, 50 pore pairs are present. Furthermore, the anterior petals in S. eustatii curve posteriorly, whereas in S. doederleini they curve an- teriorly. Occurrence.—Schizaster doederleini is known as fos- sil from the early Pliocene, near the base of the Gurabo Formation, locality NMB 16862 (Saunders, Jung, and Biju-Duval, 1986, text-figs. 15, 16), Rio Cana section, 350 m above the base; locality TU 1354 (at approxi- mately same stratigraphic level in Gurabo Formation as loc. NMB 16862), Canada de Zamba, a tributary on the west side of the Rio Cana, approximately 2.5 km east of the village of Zamba, which is 7 km north of Cruz de Santiago (Santiago Rodriguez), on road to Guayubin; or 4.5 km (airline) below the ford at Caim- ito. Distribution.—This species has been reported by Chesher (1972) and Serafy (1979) living off Colombia and Florida, but Kier (1975, p. 11) suggests that Chesh- er’s Colombian specimens probably belong to another species. 24 BULLETIN 339 REFERENCES CITED Agassiz, A. 1872. Revision of the Echini, Pt. I, in Illustrated Catalogue of the Museum. Harvard University, Museum of Compar- ative Zoology. 1880. 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Republica de Venezuela, Ministerio de Minas e Hidrocarburos, Direccién de Geologia, Boletin de Geologia, Publicacion Especial No. 4, 756 pp., 2 maps, 1 correlation chart. Linnaeus, C. 1758. Systema Naturae per Regna tria Naturae, secundum Class- es, Ordines, Genera, Species, cum characteribus differentis, synonymis, locis. 10th ed., revised, 824 pp. Holmiae. Logan, A. 1987. Neogene paleontology in the northern Dominican Republic. 6. The Phylum Brachiopoda. Bulletins of American Pa- leontology, vol. 93, No. 328, pp. 44-52, pl. 12, 1 text-fig., 1 table. Mansfield, W. C. 1937. Mollusks of the Tampa and Suwannee limestones of Flor- ida. State of Florida Department of Conservation, Geo- logical Bulletin No. 15, 334 pp., pls. A-D, 1-21, 2 text- figs., 2 tables. Martens, E. C. von 1867. Uber ostasiatische Echinodermaten. Naturgeschichte, vol. 33, No. 1, pp. 106-119, pl. 3. Maury, C. J. 1917. Santo Domingo type sections and fossils, Pt. 1: Mollusca. 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Geology and Paleontology of Canal Zone and adjoining parts of Panama. Description of Tertiary mollusks (pelec- ypods: Propeamusstidae to Cuspidariidae; additions to families covered in P 306-E; additions to gastropods; ceph- alopods). United States Geological Survey, Professional Paper 306-F, pp. i-iv, 541-759, pls. 83-124, | text-fig., 1 map (labeled as pl. 125 in contents, as pl. 1 on map). Wright, T. W. 1855. On fossil echinoderms from the island of Malta; with notes on the stratigraphical distribution of the fossil organisms in the Maltese beds. Annals and Magazine of Natural His- tory, ser. 2, vol. 15, pp. 101-127, 175-196, 262-277, pls. 4-7. 28 BULLETIN 339 EXPLANATION OF PLATE 1 Figure Page 1-3. Spondylus bostrychites Guppy ................... MNS Sais a aidan iva s,eraua. a eteighevea eiviatdconea Gia Reals c Ritieto oS ee 8 1. Left valve, BMNH LL 9946 (lectotype), #4. Gurabo Formation, locality unknown. Height 100 mm, length 101.5 mm. a, exterior view; b, interior view. BMNH LL 25706 (paralectotype). Gurabo Formation, locality unknown. Height 89 mm, length 81 mm, diameter of paired valves 62 mm. a, exterior of right valve, x1; b, dorsal view of paired valves, x 2. 3. Right valve, USNM 450387 (hypotype). Gurabo Formation, Rio Gurabo area, locality TU 1278. Height 114 mm, length 88 mm. a, exterior view, x +4; b, enlargement of portion of valve, x 3. 4:oSpondylus americanus Hermann .)..02ie.crte 2% wa os 00 201 sana te ee Ta ade ee eT 10 Right valve, USNM 450388 (hypotype), x 2. Bowden Formation, Jamaica, locality TU 705. Height 31.8 mm, length 34.6 mm. N PLATE | BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 102 PLATE 2 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 102 eR, __—— eer me 5 FD pete Sas me DOMINICAN REPUBLIC NEOGENE. 12 AND 13 29 EXPLANATION OF PLATE 2 "ge Page MPS PONG VIUSTDOSUFVCRILESIGUDDYE eyereeiaies= isie tere cis aot ar staat late eats ie i 3 vey ay sce ay o-s¥auey al sha¥otlfonstaleyaverseclslaVadaserstayaie eereetess iol foieneloloror Oreo 8 NMB G 16963 (hypotype), <1. Rio Cana, Gurabo Formation, locality NMB 16821. Height 80 mm, length 75 mm, diameter of paired valves 57 mm. a, exterior view of left valve; b, exterior view of right valve; c, anterior view of paired valves; d, dorsal view of paired valves. 2=3, Grong iia NEN In, gangcbdoGen ud bonoe spon oatos Hee meee ee ome O SOMO AT aoe Aaboramenrs coho poco s ad aoone 10 2. Left valve, USNM 450389 (hypotype). Rio Gurabo, Gurabo Formation, locality TU 1211. Height 54.5 mm, length 53 mm. a, exterior view, <1'2; b, enlargement of portion of valve, x3. 3. Left valve, AMNH 22514 (lectotype), =x 1'2. Guanica Limestone (= Ponce Limestone), Puerto Rico. Height 33 mm, length 28 mm (from Maury, 1920, pl. 5, fig. 10). 30 Figure 1-4. Spondylus gumanomocon Brown and Pilsbry ... il wh BULLETIN 339 EXPLANATION OF PLATE 3 Right valve, ANSP 2869 (lectotype), x ¥. 7Mao Formation, locality unknown. Height 175 mm, length 115 mm. Left valve, ANSP 72724B (paralectotype), <¥4. 7Mao Formation, locality unknown. Height 96.5 mm, length 94 mm. Right valve, ANSP 72724C (paralectotype), x34. ?Mao Formation, locality unknown. Height 126 mm, length 97.5 mm. . USNM 450390 (hypotype), x1. Guayubin, Mao Formation, locality TU 1221. Height 81 mm, length 56 mm, diameter of paired valves 47 mm. a, anterior view of paired valves; b, exterior view of left valve; c, exterior view of right valve. BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 102 PLATE 3 PLATE 4 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 102 DOMINICAN REPUBLIC NEOGENE. 12 AND 13 31 EXPLANATION OF PLATE 4 Figure Page 14. Echinometra lucunter (Linnaeus, 1758) .......... 1. Dorsal view of Recent figured specimen (USNM 30469) from 1 m depth in holes in rock off Rodriguez Key, Florida Keys, x2. 2-4. Dorsal, ventral, and right side views of figured specimen (USNM 375449) from locality TU 1438 in the Pliocene Mao Adentro Limestone, in a roadcut 0.5 km south of the bridge at Guayubin on the road to Sabaneta, x 2. Bem CC Ly DeastersImaOddentroensis:MewsiSPECIES: seen ey rere eee ee ee RF eh eet ees spss cus st epson ehettne iss oa aps oaaeils Biacasea® 17 Left side, dorsal, ventral views of the holotype (NMB M9746) from locality NMB 17022, early Pliocene, Mao Adentro Limestone, 765 m above the base of the Rio Cana section (Saunders, Jung, and Biju-Duval, 1986, text-figs. 15-16), <1. 12 Glypeastercaudatus; Jackson; V92205r erie ice cee ere ales sere eet nese = ae ene ah ae ate aoe re ale le ee ee 18 Dorsal and right side views of figured specimen (NMB M9747) from locality NMB 17274, bed B, Arroyo Lopez section (Saunders, 32 BULLETIN 339 EXPLANATION OF PLATE 5 Figure Page Jung, and Biju-Duval, 1986, text-fig. 26), x11. | \ PLATE 5 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 102 PLATE 6 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 102 DOMINICAN REPUBLIC NEOGENE. 12 AND 13 33 EXPLANATION OF PLATE 6 Figure Page 2. 8: 4,5. Gly peaster cia Sunn ilandensisaKierli9 63s gc.