VOLUME 93, NUMBER 328 оня ЈОМЕ 24, 1987 186: OT inf АХМЕТ ZƏN Neogene Paleontology in the northern Dominican Republic 4. The Genus Stephanocoenia (Anthozoa: Scleractinia: Astrocoeniidae) by . Ann Budd Foster 5. The Suborders Caryophylliina and Dendrophylliina (Anthozoa: Scleractinia) 3 Бу Stephen р, Cairns and John W. Wells 6. The Phylum Brachiopoda by Alan Logan Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York, 14850 U.S.A. PALEONTOLOGICAL RESEARCH INSTITUTION Officers и. WILLIAM P. S. VENTRESS Ра. JAMES E. SORAUF р. HENRY W. THEISEN دی د زغ‎ ЕЛЕ аске АА IM NU LU c ы ALI er E JAMES C. SHOWACRE ASSISTANT В. Јонм L. CISNE Е НЕКИ TR KM a Cd PETER R. НООУЕВ EECA СОМЕН C Е cue ce mesi cR Ree eet HENRY W. THEISEN Trustees BRUCE M. BELL (to 6/30/90) WILLIAM A. OLIVER, JR. (to 6/30/89) RICHARD E. BYRD (to 6/30/89) EDWARD B. Рлсоџ, JR. (to 6/30/89) Јонм L. CISNE (to 6/30/88) JAMES С. SHOWACRE (to 6/30/90) J. THOMAS DUTRO, JR. (to 6/30/90) JAMES E. 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Hoover Director Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York 14850 U.S.A. 607-273-6623 VOLUME 93, NUMBER 328 JUNE 24, 1987 Neogene Paleontology in the northern Dominican Republic 4. The Genus Stephanocoenia (Anthozoa: Scleractinia: Astrocoeniidae) by Ann Budd Foster 5. The Suborders Caryophylliina and Dendrophylliina (Anthozoa: Scleractinia) by Stephen D. Cairns and John W. Wells 6. The Phylum Brachiopoda by Alan Logan 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. CONTENTS Page 4. The Genus Stephanocoenia (Anthozoa: Scleractinia: Astrocoeniidae) Ann Budd Foster ЈАКА о ር የር. o RE PM MRNA وور‎ E uu лы COE GT 5 RESUMED cemere ee eee eb iR оо. ЕЕЕ Е А, ое TE рин пануе И. POLE 5 Пади око hi Kolo) ددد‎ а WR uM ul cu Еа ПОБИВ 5 6 АСКОН. Fi ОВО VIA СКОЈ КО ОО АЙ ПА У ИО военни a MEL ፡. ድታ M M е у 77د‎ 7 BiosüraupraphysandabalegecologyA р 7 Taxonomic Method NN eR onec M pM در‎ ра cT MANEAT Un ፣ን እ... ፕ “ተ ሥፕፕ። ፕ ታፓታኃጵኃጵፖጵ።ምፖም ጋ ጋመ... 10 Матеа и Wee Че АН ен е m Me cr Md NM Да EN CIR NU IU I ካን. a 10 Саас Тете ew LM و‎ Pe CEDE Dr RP و‎ Ne رس‎ et شه د له‎ d T M E ELEM دخ‎ 11 С соает eri XM OIM DORMIT M መ ሚም NM Ue E EE ME ETE TT EE. 12 ов ка TOO а оо رک‎ FX Oe VATER ORO PP RIP E M T TC E ee UE 12 [DAS De CEU а QUE OTI И а о нат LUCUS A A የ. “ከርም” NAR E теуш ыы ..፡. ምሽ. 14 Systematic Paleontology ТООСОО СИ NN со ЕЗ derer Uwe A P Е EU RUNI MMe UU е ДАУТ ን. 18 усеше йд. кеч Kor ME CE ERI HO RE Mr E MR UN NO UM Ru MEET اس‎ UD 18 Appendix.— Means (+1 standard deviation) of all corallite characters in the two fossil species of Stephanocoenia and in three РОРШаНОнУ ОНО ОНИ У ЕРИ СО ВИ О ер МЕ ue rere CS M CEA ушШ... CA eee t qe 22 5. The Suborders Сагуорћупа and Dendrophylliina (Anthozoa: Scleractinia) | Stephen D. Cairns and John W. Wells LA መ NOME 2 Lm Me e Iu E o ои јадено QUIM AL ее 23 КЕШТЕ OS o а КО ፡.. uq um M CLA HII PE MM MAE i E ни دد‎ 23 ПАТОН EE OTN E IDE DECRE و‎ er, 28 ASRS WIE UES Te а حا‎ E شه‎ esegue os a И а: I ILS d SM 23 ALC OC COLOR ИН e c ПИ ለ NL диб هه اوه‎ чуда M RUM цент M ردو‎ Mec ولو‎ E сн 24 Systematic Paleontology ТОССО ЕСЕН E A Li ረ ы مو‎ M е ор 25 PhilosopbicalGOonsidelatlonss. а оса هو ده‎ d A M RM uc M 26 Бе о ро ПО о مت‎ oo EI WE Ws uu вте 27 Checklist of Neogene Caryophylliina and Dendrophylliina from the Dominican Republic .................................. 27 Зета о ue به‎ a A E Du RI A е qct درد‎ ۷ ٢ c: MI СОС 2 6. The Phylum Brachiopoda Alan Logan 1101111001. NNNM LU A Wes We Ide دنه کسه کم ام دمه‎ ጋ ددد‎ 44 Röse so e MeL Е بي‎ cU EE 44 Таоа Со Ко везни io: м 44 ACK OWI СС ГЕИ O cM dc للم و‎ Пее cua, о ce па е ОН 45 Bale Oe Col Oat SUR ика ААИ Се НЕТ ac he. ТИЕ а И Туын واو سه‎ гис 45 УО A Брук а ра ОК САНИ en eM ده و‎ EE ده هرمو‎ 47 Systematic Paleontology TURON OIM ооо оао л РАНЕ ME me V EE و‎ UON ET A У DR с кут ко: 47 Тао оу оловке Cons суртот ዓዓ). ጢን» о Си TE I C а Ра ај 48 Eormat Measurements, aid- Acronyms OE SpecimenwRGpOSIUOIIGS O по ጋን ንን ንጋ 48 о p وار ورو‎ ПЕ НЕ НН UE MEM پو‎ зена а SU D Со موس و نه‎ 48 Ке те жады a ара, TR ПК لودو دو ده سم‎ MIETEN ده له‎ ИРЕ та دس‎ на НЕТ СО 52 NES E а а то E А ИЕ ТАО Е Mee О A ео А 56 LIST OF ILLUSTRATIONS AND TABLES 4. The Genus Stephanocoenia (Anthozoa: Scleractinia: Astrocoeniidae) Ann Budd Foster Text-figure Page Т. Мар mdicating the location of the river sections sampled. cece a 6 2. Diagrams showing the distribution of species of Stephanocoenia within selected ШУБЕ ШОШ ИЛ res лл л ОНЕ E E ај 8 3. Variation within species in corallite characters through a composite of the Río Cana and Río Gurabo stratigraphic sections . . . 9 په‎ Гауе showing some: ofthe 0001069 mea د‎ а оа ده‎ обет رو هره‎ ИСАР ШЕ در‎ в هه هغ د ریه ار خم وم‎ E а 1 5. Cluster analysis of all colonies of Stephanocoenia in the ММВ collections. ون نن ې‎ 13 6. Canonical discriminant analysis between species of all Stephanocoenia in Ше ММВ and modern collections ................. 14 7. Means and standard deviations for six characters in the two ММВ and one modern ѕресіеѕ................................ 12 8. Intraspecific variation in the two fossil ММВ species for canonical variables-1 and 2 of the canonical discriminant analysis BINO species НУЛЕ ONOCORITG eee eet eter судиите eru ре 16 9. Canonical discriminant analysis between populations of modern Stephanocoenia intersepta from three different reef habitats near Discovery Bays) ЕЦД, e ду ке Ра а а uen Pane ана Puede dde а а ГКИС оа eu ከ 17 10. Photographs of the colony surfaces of three holotypes important in the nomenclatural problem involving Stephanocoenia EAS AS иа E Nc Со uL REMIT M رورو دواد مد‎ 19 Table 1. List and description of characters measured on corallites in transverse thin-sections of Stephanocoenia ..................... 11 2. Final canonical discriminant analysis of three fossil and modern Stephanocoenia species ..... sse 12 3. Canonical discriminant analysis of three populations of modern Stephanocoenia intersepta .................-.............. 15 5. The Suborders Caryophylliina and Dendrophylliina (Anthozoa: Scleractinia) Stephen D. Cairns and John W. Wells Text-figure Page 1. Мар сао Ше location of tiver sections ва није ва c) ар писа eee ee ም ምች ፡ጋ..፡ት ፐ de ር TE HI TL ее 24 2. Diagram of cross-section of a calice illustrating some of the morphological terms used in the text .......................... 26 3. Diagram showing the NMB records of species within the two stratigraphically well-documented river sections: Río Gurabo and Tan he о ae rue che ean mau NO سه‎ ከ. eher ን pei Кок E UE 29 Table 1. Localities at which Pourtalocyathus hispidus and Deltocyathus italicus were collected ..................................... 25 2. Species believed to occur in deep water, and depth ranges of their Recent counterparts ...... مهه نن نه نن هننن نن نن‎ 25 3. Characters of the леизеноушеврестев бї ANTOS ото они Fo oa tae cg ORE ም ۷ 77 а рој n M Rd. 38, 39 6. The Phylum Brachiopoda Alan Logan Text-figure Page 1. Geological sketch map of the Cibao Valley, showing the location of the Río Cana, Río Gurabo, and Arroyo Zalaya sections, ош обе brachiopods have Бей collected: oor EU oa. eile m EE መ: ያ. бо, 45 Table 1. Stratigraphic sections from Río Cana, Río Gurabo, and Arroyo Zalaya, showing approximate positions of brachiopod-bearing lloro: ее ru Mx »مهو سم وه همو مهد‎ E A codi cosi deba КА 46 NEOGENE PALEONTOLOGY IN THE NORTHERN DOMINICAN REPUBLIC 4. The Genus Stephanocoenia (Anthozoa: Scleractinia: Astrocoeniidae) By ANN BUDD FOSTER Geology Department The University of Iowa Iowa City, ТА 52242, U.S.A. ABSTRACT Multivariate statistical procedures are used to distinguish species in the reef-coral genus Stephanocoenia through a continuous Neogene sequence (five-million year time interval) in the Cibao Valley of the northern Dominican Republic. This genus is the only member of the family Astrocoeniidae that occurs in the sequence. The material consists of 56 colonies (17 of which are measured) from 24 localities in four river sections, the most important being Río Gurabo and Río Cana. Ten characters are measured on each of 10 corallites per colony. The data are analyzed using cluster and canonical discriminant analysis to group colonies into clusters representing species. Identical measurements on modern colonies collected near Discovery Bay, Jamaica are included for comparison. Two fossil species are defined in the analysis, one of which is new (Stephanocoenia duncani, n. sp.). Both species are significantly distinct from the single modern species (S. intersepta) that is the sole living representative of the genus. Study of collections from other reef localities shows that both fossil species occur only during Neogene time and only at a limited number of localities. Patterns within each species are traced up a composite stratigraphic section using nonparametric statistical analyses. One of the two fossil species (S. spongiformis) is found to remain stable through time, whereas the other (S. duncani) changes its morphology in a direction approaching the cluster for the modern species. Further study of patterns of variation within the one modern and two fossil clusters shows that intraspecific variation is unusually complicated in this genus. The clusters overlap, and colonies within each cluster differ widely. Variation between populations within the modern species occurs in the same characters as those which distinguish the modern species from the fossil species converging with it (S. duncani). However, these two species form a morphologic continuum that cannot be explained by environment alone. Therefore, they may represent two gradually intergrading chronospecies within one lineage. Of the two fossil species of Stephanocoenia defined, one species (S. spongiformis) exhibits an evolutionary pattern similar to that observed in the family Poritidae. In this pattern, species were found to have short durations and stable morphologies and to have become extinct during the mid- to late Pliocene through early Pleistocene mass extinction. In contrast, the second species of Stephanocoenia (S. duncani) may have evolved over a long time period, possibly forming chronospecies that survived the mass extinction. Unlike genera in the Poritidae, however, no radiation of taxa occurred in the genus after the extinction event. Since no consistent relationship has been discovered between morphology and environment in these corals with the data at hand, their paleoecologic value can only be determined after data on more taxa are collected, and their associations with other corals are studied. This study represents part of a multidisciplinary project on the stratigraphy and paleontology of the northern Dominican Republic, coordinated by P. Jung and J. B. Saunders of the Naturhistorisches Museum Basel, Switzerland. RESUMEN Se usan procedimientos estadísticos para distinguir especies de corales arrecifales en el género Stephanocoenia a travéz de una secuencia Neogéna contínua (intervalo de cinco millones de años) en el valle Cibao del norte de la República Dominicana. Este género es el único miembro de la familia Astrocoeniidae que se encuentra en la secuencia. El material consiste de 56 colonias (17 de las cuales se miden) de 24 localidades en cuatro secciones del río, siendo las mas importantes las del Río Gurabo y del Río Cana. Se miden diez caracteres en cada una de diez coralitas por colonia. Se analizan los datos usando análisis canónicos discriminatorios y de grupo para combinar las colonias en conjuntos que representen especies. Para compararlas, se incluyen colonias modernas de medidas idénticas recolectadas cerca de Discovery Bay, Jamaica. Se definen dos especies de fósiles en el análisis, una de las cuales es nueva (Stephanocoenia duncani, esp. n.). Ambas especies son significativamente distintas de la única especie moderna (5. intersepta), que es la única representante viviente del género. Estudios de colecciones de otras localidades de corales muestran que ambas especies fósiles son encontradas solo durante el tiempo Neogéno y solo en un numero limitado de localidades. Dentro de cada especie se trazan modelos de sección estratigráfica compuesta usando análisis estadísticos no paramétricos. Se encuentra que una de las dos especies fósiles (S. spongiformis) permanece estable a travéz del tiempo, mientras que la otra (S. duncani) cambia su morfología acercandose al grupo de las especies modernas. Estudios más avanzados de modelos de variación dentro de un grupo moderno y dos de fósiles, muestran que la variación intraespecífica es inusualmente complicada en este género. Los grupos se translapan, y las colonias en cada grupo se diferencian ampliamente. Dentro de las especies modernas ocurren variaciones entre las poblaciones en los mismos caracteres que en aquellos que distinguen las especies modernas de las especies de fósiles que convergen en ella (5. duncani). Sin embargo, estas dos especies forman un contínuo morfológico que no puede ser explicado solamente por el medio ambiente. Por lo tanto, pueden representar dos cronoespecies que se integran gradualmente en un linaje. De las dos especies de fósiles definidas de Stephanocoenia, una especie (S. spongiformis) exibe un 6 BULLETIN 328 modelo evolucionario similar a aquel observado en la familia Poritidae. En este modelo, se encontraron que las especies tienen duraciones cortas y morfologías estables y que se extinguieron durante el medio a tarde Plioceno a travéz de destrucciones masivas en el Pleistoceno temprano. En contraste, la segunda especie de Stephanocoenia (S. duncani) puede haber evolucionado en un periodo de tiempo largo, posiblemente formando cronoespecies que sobrevivieron la destrucción masiva. Sin embargo, en forma diferente al género de Poritidae, no ocurrió ninguna radiación de taxa en el género despues de la destrucción. Ya que con los datos a mano no se descubrió ninguna relación consistente entre la morfología y el medio ambiente, su valor paleocológico se puede determinar solamente luego de haber coleccionado mas datos en taxa, y luego de haber estudiado sus asociaciones con corales. Este estudio representa parte de un proyecto multidisciplinario en la estratigrafía y paleontología del norte de la Repüblica Dominicana, coordinado por P. Jung y J. B. Saunders del Naturhistorisches Museum Basel, Suiza. INTRODUCTION The purpose of this paper is to describe the species of one coral family (herein represented by only a single genus) in one region by quantitative study of popula- tions collected at closely-spaced stratigraphic intervals, and to trace their evolutionary patterns over a five- to ten-million year time segment, following the major coral extinction event that occurred during late Oli- gocene and early Miocene time. This paper is the sec- ond in a series of coral faunal descriptions for the mid- dle Miocene to middle Pliocene of the northern Dominican Republic. It deals with a common but lit- tle-studied coral genus, Stephanocoenia Milne-Ed- wards and Haime, 1848a, a mound-shaped coral, sometimes large in size, that represents an important reef-building component in the Caribbean from mid- dle Miocene to modern time. As in the first paper (Foster, 1986), the coral collections in this study were made stratigraphically in bulk between 1978 and 1980 by J. Geister, P. Jung, J. B. Saunders and coworkers as part of their large-scale multidisciplinary project on the paleontology and stratigraphy of the Cibao Valley of the northern Dominican Republic. The project has focussed on this region because it contains one of the richest, most continuous, and best-studied sections through Neogene fossiliferous units in the Caribbean. All collecting localities are keyed into their detailed stratigraphic sections (Saunders, Jung, and Biju-Duval, 1986). The coral faunal descriptions are part of a series of papers on these collections, planned to cover all fossil taxa occurring in the sections. Previous work on the systematics of the Neogene Dominican Republic corals has been reviewed in Foster (1986). In this work on the Dominican Republic, the genus Stephanocoenia is only rarely mentioned. Dun- can (1863, 1864, 1868) described only two species of Stephanocoenia in the Heneken collection at the Brit- ish Museum (Natural History), London, England, U.K. [BM(NH)]. They are Stephanocoenia intersepta (Esper, 1795) and Plesiastraea spongiformis Duncan, 1864. Pourtalés (1875) listed only one species, 5. intersepta 9 10 20km 4 Rio Сапа 2 Rio Gurabo GUAYUBIN Стр لا‎ Upper Cenozoic рации N А О [++] Oligocene - Early Miocene 2 5 Сапада Zalaya 6 Rio Yaque del Norte 3 и 4 Mesozoic 7 City of Santiago Ки 4 ረ 8 Arroyo Puñal ы 9 3) Е ፦ 9 Rio Verde S S RS 1 2 Ф VALVERDE ESPERANZA NAVARRETE ZAMBA MAO Eo Ко : ٣۴ الا چ‎ di e 3 ኤ Morg ee oa LOSQUEMADOS & ^ SANTIAGO 5 1 VÉ RODRIGUEZ سي‎ 4 шын QN د"‎ 60 BULLA 3 MOCA : 5 E 6 и RUP OE < BAITOA SORGE ти . Text-figure 1.—Map indicating the location of the river sections sampled. Stephanocoenia was found only in four sections: (1) Río Cana, (2) Río Gurabo, (3) Río Mao, and (4) Río Yaque del Norte (map from Saunders, Jung, and Biju-Duval, 1986). DOMINICAN REPUBLIC NEOGENE. 4: FOSTER 7i (Esper, 1795). Similarly, Vaughan (1919, and an un- dated unpublished manuscript) synonymized Dun- can's two species and indicated that only one species of Stephanocoenia [S. intersepta (Esper, 1795)] occurs in the Neogene of the Dominican Republic (see also Vaughan et al., 1921). No species of Stephanocoenia were treated in Vaughan and Hoffmeister (1925) or in any other subsequent papers on the Dominican Re- public corals. In all the Caribbean Miocene through lower Pliocene excluding the Dominican Republic, Stephanocoenia is only known in the Bowden For- mation of Jamaica (Vaughan, 1919) and at St. Croix in Trinidad [Heneken collection, 11 ACKNOWLEDGMENTS I am indebted to J. Geister (Bern, Switzerland), P. Jung (Naturhistorisches Museum, Basel, Switzerland [ММВ]), and J. B. Saunders (ММВ) for collecting the material, providing locality information, and assis- tance in sorting and curating specimens. I am especially grateful to Ms. Wen-Jian Tang (State University of Iowa, Iowa City, IA, U.S.A. [SUI]) for measuring the corals in this study and for typing the data into the computer. The thin-sections were prepared by K. Mül- ler (NMB) and T. Bahns (SUI). U. A. Dogan (SUI) assisted with scanning electron microscopy, and W. Suter (ММВ) and M. Serrette (Muséum national d’His- toire naturelle, Paris, France [MNHNP]) made many of the colony surface photographs. J. Geister provided X-radiographic equipment. R. Petrick (SUI) assisted With typing, H. Greenberg (SUI) assisted with prepa- ration of plates, and T. Druecker (SUI) translated the abstract. Ithank the following individuals and institutions for loans and assistance with museum material: R. Pan- chaud (ММВ), J. Golden (SUI), S. D. Cairns (United States National Museum of Natural History, Wash- ington, DC, U.S.A. [USNM]), В. В. Rosen [BM(NH)], J. Maréchal (MNHNP), В. L. Langenheim, Jr. (Uni- versity of Illinois, Urbana, IL, U.S.A. [UI]), D. R. Lindberg (University of California Museum of Pa- leontology, Berkeley, CA, U.S.A. [UCMP], апа М. Eldredge (American Museum of Natural History, New York, NY, U.S.A. [АММН)). J. W. Wells (Ithaca, NY, U.S.A.), M. E. Tollitt [BM(NH)] В. V. Melville [BM(NH)], and С. Klapper (SUD provided advice on Zoological nomenclature, and G. Scheer (Darmstadt, West Germany) discussed and provided photographs of coral specimens in the Esper Collection (Natur-mu- seum Senckenberg, Frankfurt, West Germany [NMS]). I thank J. C. Lang and J. W. Wells for reviewing the manuscript. J. Golden (SUI) also read the manuscript and offered helpful suggestions. Funds were provided by a grant from the U. S. National Science Foundation (BRS83-07109). ABBREVIATIONS OF REPOSITORY INSTITUTIONS AMNH: American Museum of Natural History, New York, NY, U.S.A. BM(NH): British Museum (Natural History), London, England, U.K. MNHNP: Muséum national d'Histoire naturelle, Paris, France NMB: Naturhistorisches Museum Basel, Basel, Swit- zerland NMS: Natur-museum Senckenberg, Frankfurt, West Germany ЏСМР: University of California Museum of Paleon- tology, Berkeley, CA, U.S.A. UI: University of Illinois, Department of Geology, Ur- bana, IL, U.S.A. USNM: U. S. National Museum of Natural History, Washington, DC, U.S.A. SUI: State University of Iowa, Department of Geology, Iowa City, IA, U.S.A. BIOSTRATIGRAPHY AND PALEOECOLOGY The genus Stephanocoenia Milne-Edwards and Haime, 1848a is common in two of the river sections through the Neogene of the Cibao Valley of the north- ern Dominican Republic (Río Cana and Río Gurabo) that were described by Saunders, Jung, and Biju-Duval (1986). In addition, one colony was found in each of the Вто Mao and Río Yaque del Norte sections. А total of 56 colonies was collected at 24 localities ranging from early Miocene to early Pliocene in age. The small size of these numbers contrasts with those for the more abundant coral family Poritidae, approximately 450 colonies of which were collected in 92 localities from the same river sections. The two species represented were found in relatively equal overall abundances. Thirty-two colonies of S. spongiformis (Duncan, 1864) and 24 colonies of S. duncani, n. sp. were collected. 5. spongiformis was more commonly found in Río Gurabo (29 colonies were collected here as opposed to only four of S. dun- cani). On the other hand, 5. duncani was more com- mon in Río Cana (18 colonies were collected here as opposed to only three of S. spongiformis). Both species exhibited wide ranges in Río Cana (Text-fig. 2A), whereas in Río Gurabo, S. duncani was found only at the 275 m level (Text-fig. 2B). Little can be concluded unequivocally about the bio- stratigraphic or paleoecologic implications of the in- dividual ranges of these two species. Their long du- 8 BULLETIN 328 rations (approximately ten million years) and rare occurrence in other geographic areas imply they would have only limited biostratigraphic value. Neither of the two species exists today, and the only direct mod- ern counterpart, S. intersepta (Lamarck, 1816) (the one living representative of the genus Stephanocoenia), is widely distributed both geographically across the Ca- ribbean and ecologically from backreef lagoons to deeper areas of the forereef (Goreau, 1959; Goreau and Wells, 1967). The one fossil colony found in the older, more conglomeratic Río Yaque del Norte section of the Dominican Republic belongs to the species S. dun- cani. Assuming the shallow nearshore interpretation of Saunders, Jung, and Biju-Duval (1986) for the Río Yaque del Norte section, this information together with the restricted distribution of this species lower in the Río Gurabo section suggests that 5. duncani might have been the older of the two species and/or that it might have occurred more commonly in nearshore habitats. Obviously, more data would be needed to substantiate these hypotheses. Patterns of morphologic variation within species across a composite of the Río Cana and Río Gurabo stratigraphic sections (spanning a time interval of ap- proximately five million years) show somewhat greater stability within 5. spongiformis (Text-fig. 3). Study of Spearman's rank order correlation coefficients (SAS Inst. 1982, PROC CORR) indicates that insignificant correlations occur between morphology (canonical variables 1 and 2 of the between-species analysis de- scribed in the next section) and stratigraphic position. Intraspecific comparisons between populations (SAS Inst. 1982, PROC NPARIWAY) collected in lower and upper stratigraphic levels (with the cutoff between “lower” and “upper” being defined at approximately the 500 m level [see Text-fig. 2]) demonstrate little difference in morphology between levels. However, in S. duncani, morphology appears to change upsection in a direction approaching that of modern 5. intersepta. Comparisons between populations collected at lower and upper stratigraphic levels show a significant in- crease in canonical variable 1 (a variable related to inner wall and columella thickness) in S. duncani (Епа = 12.43; р = 0.0009). Nevertheless, only insig- nificant correlations exist between morphology and stratigraphic position. The stability within the first species suggests not only a lack of evolutionary development, but a lack of re- sponse to large-scale environmental change, since the Río Cana and Río Gurabo sections of the Cibao Valley are believed to have been deposited under deepening environmental conditions (Saunders, Jung, and Biju- Duval, 1986). Such stability was also observed in the family Poritidae (Foster, 1986) and may be character- istic of corals that today exhibit complicated patterns of intraspecific variation which are not directly cor- related with environment. The directional trend within A 4 MORAN RIO CANA 5 -3 © 1000-4 X 8 | Е ፔዞ ፎ x (= 5 © = # Е |- 5004 o о 1 13 5 “E 1 4 oO سه‎ 5... 8158 2 о ке 0 r — ላ “tor се هم‎ д? so? B / ون‎ КО 06 1000 - n=29 E | Ф Е 2 S © 5 2 ቋ 500 о = E 5 п= 4 o Ф ES 15 XA 5” 55 ዐ г 1 ۱ о со" ለፍ ме 509 9 Text-figure 2.— Diagrams showing Ше distribution of species of Stephanocoenia within selected river sections. “п” = total number of localities containing each species. Numbers to the right of points along each vertical distribution line indicate the number of localities represented by each point. (А) Río Cana, (B) Río Gurabo. DOMINICAN REPUBLIC NEOGENE. 4: FOSTER 9 m m а но CEDE ще 7007 £ 1000 6004 800 1 га к Е я < di | т | 0 о 5004 600 G с : U A ~ R | ዘ Я) А А a B 400 03 Б مه‎ + 4004 393 4፡66 ሠው оо 6 መ 6. 500ኀ ፎዐዐ 200ኀ 100 ፍመ መ. ዐ ہہ‎ T T T -6 -4 -г о А CANONICAL VARIABLE 1 m m 1200 173 2 о و‎ 7004 4 1000 600 ኀ 800: SEE EEE R я n | | | I | 0 9 | 500 4 ВОЗ ^g с : U 5 : R 5] ዘ | А б : S 400 | о Е о спи dio igi 3004 200 2007 1004 o. Дои но но ን و‎ а 0 r T т T 7 ፣ j ед -г 0 Е 8 B CANONICAL VARIABLE 2 Text-figure 3.— Variation within species in corallite characters through a composite of the Río Cana and Río Gurabo stratigraphic sections. Тће composite was constructed using the Río Cana section (Saunders, Jung, and Biju-Duval, 1986, text-fig. 16) as a standard and scaling the Río Gurabo section (Saunders, J ung, and Biju-Duval, 1986, text-fig. 6) relative to it. To accomplish this, three exact elevations were correlated between the two sections: (1) the unconformity with the Tabera Group at the base of each section, (2) the boundary between 16 6 humerosa and G. margaritae foraminiferal zones, and (3) the boundary between the G. margaritae and G. miocenica zones. The position of localities within each section were determined using text-figures 4 and 15 of Saunders, Jung, and Biju-Duval (1986). The points labelled “1" and “2” on the figure represent means for all colonies within each species from each 100 meter interval along the composite section. Horizontal lines on either side of each point are one standard deviation in length. Dotted lines connect means for each Species. 1 = S. spongiformis, 2 = S. duncani. (А) Canonical variable 1 of the canonical discriminant analysis between species. As described In the text, the thickness of the inner wall and of the columella are most heavily weighted on this canonical variable. Morphological change 1$ detected in species 2, but not in species 1. (B) Canonical variable 2 of the canonical discriminant analysis between species. The spacing between corallites and development of the exothecal intercostal area are heavily weighted on this canonical variable. Morphological change cannot be detected in either species. 10 BULLETIN 328 the second species implies gradual evolution from 5. duncani into S. intersepta, signifying that the two rep- resent merely chronospecies within one lineage. Never- theless, on the basis of the current analyses, it is im- possible completely to reject the hypothesis that morphologic change upsection in 5. duncani is related to environment. This hypothesis is discussed in more detail in the section on intraspecific variation. As in the family Poritidae, the potential usefulness of Stephanocoenia as a paleoecologic indicator can only be ascertained after all the corals in the collection of Saunders, Jung, and Biju-Duval (1986) have been stud- ied taxonomically, and the patterns of intraspecifc variation are compared with patterns of diversity and abundance for the entire coral fauna. However, unless intraspecific trends appear to be clearly correlated with environment, data on the distribution and abundance of the two species of Stephanocoenia may prove more valuable in quantitative studies of species associations, especially those studies that attempt to link certain associations with particular environments (see discus- sion in Foster, 1986). TAXONOMIC METHOD PROBLEM One genus, Stephanocoenia, of the family Astro- coeniidae occurs in the Neogene of the Dominican Republic. The genus occurs today only in the Western Atlantic region. Although it represents an important component of the coral fauna in lagoon through deep forereef areas, it is relatively unstudied ecologically or morphologically and is often confused with the more dominant reef-coral Montastraea annularis (Ellis and Solander, 1786), whose colony shape and general cor- allite morphology is strongly convergent with Stepha- nocoenia intersepta (Lamarck, 1816). Stephanocoenia differs from Montastraea Blainville, 1830, in its styli- form columella, prominent pali, and its regular sub- cerioid intercostal area. Its corallites are also generally smaller than those of Montastraea. Similarly, fossil Stephanocoenia bear a striking overall resemblance to the extinct genus Astrocoenia Milne-Edwards and Haime, 1848а, a taxonomically more diverse genus that was especially common ш the Caribbean during the Eocene and the Oligocene. Stephanocoenia differs from Astrocoenia only by having prominent pali, more septa, and a cerioid colony form. Because both genera have similar styliform columellae, this second case of resemblance is more likely truly phylogenetic. Only one species of Stephanocoenia is commonly recognized in the Caribbean from Miocene (possibly Oligocene) through modern time (Frost, 1977; Zlatar- ski and Estalella, 1982). Some workers have suggested that a second modern species may also exist whose skeletal morphology differs from the first primarily by having smaller corallites and wider, more porous in- tercostal area (Goreau and Wells, 1967; J. C. Lang, oral commun., 1983). The most convincing differ- ences, however, involve tissue coloration and other live-polyp characters. This suggestion implies that the second species could overlap in skeletal morphology with the first, especially when large populations from a range of different habitats are studied. However, the validity of this second species has never been clearly demonstrated, in part because morphologic variability 1s so poorly understood in the first, more common species. Given this lack of knowledge about species of living Stephanocoenia, in the current study, species have been differentiated in the fossil material by grouping spec- imens into clusters using purely quantitative analyses of skeletal characters. To facilitate such cluster defi- nition, direct comparisons have been made with pop- ulations of living Stephanocoenia intersepta collected in three different habitats near Discovery Bay, Jamai- ca. The specimens from the living populations have been measured using the same techniques as in the fossils, and they have been statistically analyzed to- gether with the fossils following the procedures of Foster (1984). Obviously, such techniques ignore some po- tentially diagnostic soft-tissue characters. However, they do attempt to base species definitions on means and variances of known populations, two especially important parameters in the study of highly variable and possibly overlapping species. MATERIAL The material studied consists of all specimens of the genus Stephanocoenia (56 colonies) collected in the Dominican Republic by J. Geister, P. Jung, J. B. Saun- ders, and coworkers between 1978 and 1980. It is cur- rently deposited at the Naturhistorisches Museum Ba- sel. These coral collections from the Dominican Republic are termed “NMB” collections in the follow- ing discussion in order to distinguish them from ma- terial in other collections used in the analysis. All mem- bers of the family Astrocoeniidae were first sorted from the rest of the NMB collections and identified as be- longing to the genus Stephanocoenia. The specimens were then separated into two groups, one subcerioid and the other cerioid. Eight to 10 well-preserved, larger colonies were selected from each group for thin-sec- tioning and measurement. One transverse and one lon- gitudinal thin-section were prepared from near the col- ony surface on the top and side of each colony. Parameters were measured on thin-sections from a to- tal of 18 NMB colonies. DOMINICAN REPUBLIC NEOGENE. 4: FOSTER 11 Table 1.— List and description of characters measured on corallites in transverse thin-sections of Stephanocoenia. Characters 1 and 2 were measured to the nearest 0.04 mm and characters 3 through 7 were measured to the nearest 0.015 mm. character abbreviation description 1. corallite diameter CD Linear measure across corallite center between theca/corallite cavity margins; mean of greatest and smallest lengths 2. corallite spacing CS Linear measure between centers of adjacent corallites; mean of greatest and smallest lengths 3. columella width CLW Linear measure across the columellar style; mean of greatest and smallest lengths 4. tertiary septum length TSL Linear measure from septum tip to inner thecal margin; mean of longest and shortest septa 5. total theca width TT Linear measure across entire theca from the corallite cavity margin to the center of the second wall between adjacent corallites 6. inner wall thickness ATT Linear measure across solid skeleton forming the inner wall immediately surrounding the corallite 7. septum thickness (major) ST Linear measure across major septa at septum midpoint; mean of thickest septum and next septum to its right 8. number of adjacent NAC Count of corallites whose thecae abut the theca of the measured corallite corallites 9. total number of septa NS Count of all major and minor septa 10. number of major septa MJS Count of septa extending to the columella For comparison, 10 living colonies of Stephano- coenia intersepta (Lamarck, 1816) (SUI 51064- 51093) were collected from each of three environmentally- distinct reef habitats near Discovery Bay, Jamaica. The habitats consisted of: (1) a nearshore turbid lagoon, 16 m depth, (2) a forereef coral thicket, 20 m depth, and (3) a forereef sand channel, 20 m depth (see Foster, 1980 for detailed locality descriptions). Transplant ex- periments (Foster, 1979) have shown that morphologic variation between these habitats in two other species [Montastraea annularis (Ellis and Solander, 1786) and Siderastrea siderea (Ellis and Solander, 1786)] is large- ly environmental. As in the fossil material, two trans- Verse and two longitudinal thin-sections from each col- Ony ere prepared and measured. Measurements were made only on thin-sections to insure greater accuracy and consistency of data. A total Text-figure 4. — Drawing showing some of the characters mea- sured: (a) corallite diameter, CD; (b) columella width, CLW; (c) tertiary septum length, TSL; (d) total theca width, TT; (e) inner wall thickness, АТТ; (f) septum thickness, ST. of 10 corallites was measured on each colony, since previous work (Foster, 1985) has indicated that 10 corallites per colony is an adequate sample size to es- timate colony means and variances. Ten mature cor- allites were selected at regular intervals on each of the two transverse thin-sections per colony. CHARACTERS The characters analyzed consist of linear measure- ments and counts on 10 corallite features in transverse thin-sections (Table 1, Text-fig. 4). All data are avail- able on computer tape from the author. The characters were selected to include the most important diagnostic features commonly used to distinguish species in as- trocoeniid corals. They can be grouped into the fol- lowing categories: (1) size and arrangement of coral- lites, (2) development of septa, (3) thickness of septa, columellar style, and inner corallite wall, and (4) de- velopment of the intercostal area. One possibly im- portant corallite character that was not measured is the development of the pali. In Stephanocoenia, the pali are not true pali (sensu Vaughan and Wells, 1943) formed by septal substitution, but represent thicken- ings of the inner margins of the major septa. They proved difficult to measure consistently in thin-section because of their often weak development and indistinct margins; therefore, they were not included in the anal- yses. Similarly, no whole colony measurements were included in the analyses, because preliminary obser- vations suggested that colony shape was fairly uniform in the material studied. A brief summary ofthe measured characters is given below: 1. Size and arrangement of corallites. — Cursory ex- amination suggests that, after maturity, the size and shape of individual corallites remain unchanged. No clear patterns of astogeny can be deciphered. Corallites 12 BULLETIN 328 are circular or polygonal in shape, depending on the development of the theca and the intercostal area. In general, corallites appear more circular with increased development of the intercostal area. One measure, CD (the mean of the long and short corallite dimensions) describes corallite size (Table 1, Text-fig. 4). Three measures, CS, TT, and NAC, describe corallite spacing (Table 1). The first, CS, is the distance between centers of adjacent corallites. The second, TT, is the distance across the corresponding inner walls, intercostal area and outer walls of adjacent corallites (Text-fig. 4). The third character, NAC, represents a count ofthe number of adjacent corallites. 2. Development of septa. — Тһе septa are composed of six to eight vertical trabeculae that join together horizontally to form septal plates and at their outer margins to form a septothecate or parathecate wall. The septa are radially arranged into three cycles con- sisting of six primary, six secondary, and 12 tertiary septa. Three measures (Table 1) describe development of the septa: NS, the total number of septa рег corallite; MJS, the number of septa extending completely to the columella; and TSL, the length of the tertiary septa (Text-fig. 4). 3. Thickness of septa, columellar style, and inner corallite wall. — As explained previously, the 24 septa are arranged in three cycles, with pali forming on the inner margins of the first two cycles. Costae of equal thickness extend from each septum across the sep- tothecal inner wall immediately surrounding the cor- allite to a second parathecal outer wall formed at the junction between adjacent corallites (Text-fig. 4). The intercostal area between the inner and outer wall is composed of uniform, equilateral to rectangular void cells whose margins are defined by costae, inner and outer walls, and exothecal dissepiments. The double wall structure gives the coral its so-called “‘subcerioid” appearance in transverse section. The two walls may coalesce leaving the coral a cerioid appearance. The columella is formed by a single, prominent style, cir- cular to elliptical in shape. The thickness of three ver- tical elements were measured: the major septa, ST; the columellar style, CLW; and the inner corallite wall, ATT (Table 1, Text-fig. 4). 4. ‘Development of the intercostal area.—As ех- plained previously, the intercostal area consists of cel- lular void space between the inner and outer corallite walls. The cross-sectional area of the intercostal area can be estimated using a combination of two previous measurements, TT and ATT. Therefore, no additional measures were made. STATISTICAL PROCEDURES Species were distinguished within the fossil NMB Dominican Republic Stephanocoenia using multi- Table 2.—Final canonical discriminant analysis of three fossil and modern Stephanocoenia species. Total-sample correlations between the canonical variables and the original variables (COR), and stan- dardized canonical coefficients (SCC). Only values with high mag- nitudes are given. The canonical variables are labelled CV 1-2. Ab- breviations for characters are explained in Table 1. н СУ СУ2 original variable COR SCG COR SCC CD 306 not used Он د‎ not used CS .391 not used eile) not used CLW Ki 0.13 = 182 —0.46 TSL .274 (0097 —.149 220728 TT .320 0.06 .798* 1.04* ATT .840* 1.10* —.078 دن‎ ST .343 —0.45 - 092 ОШО МАС 010 not used .081 not used NS .026 17855 4120 1.297 MJS, Е= 029 —1.08* .094 —1.07* * Most important variables. variate statistical techniques similar to those outlined in Foster (1984). The programs are contained within the computer packages SAS (SAS Inst., 1982) and SPSS-X (SPSS Inc., 1983). First, the data for each mea- sured colony in the fossil NMB and modern Jamaican collections were analyzed using cluster analysis (SAS CLUSTER procedure, average linkage method) of principal component scores (SAS PRINCOMP pro- cedure) (Text-fig. 5). The fossil colonies were thereby divided into two groups using a distance of 0.8 as a cutoff. The first group (S. spongiformis) was originally composed of six colonies; the second (S. duncani) of nine colonies (Text-fig. 5). The raw data were then analyzed using a series of stepwise canonical discrim- inant analyses (SPSS-X discriminant procedure). In the first of these analyses, the two fossil clusters and the one modern species were used to form groups, and unassigned fossil colonies were left unclassified. The results were used to assign the unclassified colonies to clusters. These clusters were then modified using sub- sequent canonical discriminant analyses. In this pro- cess, cluster assignments for various colonies lying at cluster margins were changed and the analyses rerun until the highest number of correctly classified coral- lites was obtained. Three clusters (two fossil, one mod- ern) were finally distinguished (Table 2, Text-fig. 6). Means and standard deviations of all individual measured characters are given for the two fossil species and three modern populations in the Appendix. Six characters revealing significant differences between species are plotted in Text-figure 7. RESULTS AND INTERPRETATIONS As described in the previous section, three clusters (two fossil, one modern) were detected in the final discriminant analysis. Of all colony means, 87.5 per- cent were classified correctly, with 100 percent correct DOMINICAN REPUBLIC NEOGENE. 4: FOSTER 13 classification in the two fossil clusters and 80 percent correct classification in the modern cluster. The mod- ern cluster (Stephanocoenia intersepta) overlapped most significantly with the fossil cluster described herein as a new species (S. duncani). As explained previously, these two clusters could be interpreted as representing chronospecies within one lineage. However, since the present results indicate that they constitute two tight clusters whose pooled variance extends beyond the limits of common phenotypic plasticity in S. intersepta (as estimated using the three Jamaican populations from different habitats), they will be considered sep- arate until data on more populations suggest they form one truly continuous cluster. Two discriminant functions were calculated in the final discriminant analysis, the first accounting for 58.66 percent of the variation and the second accounting for 41.34 percent. The first function primarily distinguish- es the two fossil clusters from the modern cluster, whereas the second function distinguishes between the two fossil species. Seven of the 10 variables were used in the analysis (Table 2). They are in order of entry into the stepwise analysis: ATT, TT, TSL, NS, MJS, ST, CLW. NS, ATT, and MJS are most heavily load- D5772(D) 05761(5) 05771(0) 05769(0) D5681(D) 05871(0) 05766 (0) 7] D5768(D) D5765(D) D5764(D) D5870(D) D5872(D) D5773(S) D5770(S) poe 1 D5763(S) D5767(S) D5762(S) ! ገም ፕ ፐ ፲ 2.0 1.5 1.0 AVERAGE Д Џ 0.5 0.0 DISTANCE BETWEEN CLUSTERS Text-figure 5. — Cluster analysis of all colonies of Stephanocoenia in the NMB collections. Dendrogram based on distances calculated using the average linkage method of the SAS CLUSTER procedure. Each terminal branch represents one colony. Colonies are labelled using NMB catalog numbers followed by either S or D. S — colonies later identified as 5. spongiformis; D — colonies later identified as S. duncani. The Cluster analysis was performed on principal component scores computed for all colonies in the NMB and modern collections. Clusters used initially in grouping specimens into species are indicated by double vertical lines. ed on the first function; and ATT and, to a lesser de- gree, CLW are most strongly correlated with it. This result suggests that thickness of such structures as the inner wall and the columella are most important in distinguishing the fossil from the modern Stephano- coenia. NS, MJS, and TT are most heavily loaded on the second function; and TT is most strongly corre- lated. This result suggests that the spacing between the corallites and the development of the intercostal area are most important in distinguishing the two fossil species. In summary, three overlapping clusters (two fossil and one modern) exist, and seven of the measured characters are important in making the distinction. The greatest difference between the fossil and modern clusters occurs in the thickness of the inner wall and columella. The most overlap occurs between the clus- ter representing the modern species and that repre- senting the new fossil species, S. duncani. The two fossil species are distinguished by corallite spacing and the development of the exothecal intercostal area. Thus, BULLETIN 328 S. spongiformis appears more subcerioid, whereas 5. duncani appears more cerioid. INTRASPECIFIC VARIATION Because of the overlap noted between species in the previous section, variation within species has been studied in more detail at two levels to determine if the clusters could represent true species. Variation was studied first within colonies and second between pop- ulations. Variation within the fossil colonies was studied to determine if variability within colonies could be used to estimate variation within clusters. Study of such variation yielded two important results: (1) significant overlap exists between colonies within each cluster and the magnitude of this overlap appears far greater than the overlap between the two clusters (Text-fig. 8); and (2) despite the high magnitude of colony overlap, re- sults of nested multivariate analysis of variance reveal a strong colony effect indicating that colonies within each cluster are significantly different. The overall mag- له : 1 1 1 4 1 له N‏ ul Е‏ ae) |‏ m |‏ а - 1‏ پر ብዥ ВиК‏ ve] X‏ 96 о‏ 5 2 5 < d ааа 122586‏ те Чао‏ د ርን : 2 2 Крем be mz‏ = سح ته 4 መመመ‏ е‏ 2 | 4 .27 О | E 2‏ ፖረ 1 ረ 2‏ dog с 4‏ ህመ‏ دو me‏ AAA ;‏ مسبم м e‏ —— — e 4 6‏ 0 2- 4- 6- CANONICAL VARIABLE 1 Text-figure 6.— Canonical discriminant analysis between species of all Stephanocoenia in the ММВ and modern collections. Plot ofscores on the first two canonical variables showing polygons outlining the range of variation between colonies in the three species groups defined by the statistical analyses. The points represent means for each colony. 1 — fossil NMB 5. spongiformis, 2 — fossil NMB S. duncani, 3 = modern S. intersepta. In S. intersepta, dotted lines surround the lagoon population, dashed lines surround the reef population, and dotted/dashed lines surround the sand channel population. DOMINICAN REPUBLIC NEOGENE. 4: FOSTER Table 3. — Canonical discriminant analysis of three populations of modern Stephanocoenia intersepta. Total-sample correlations be- tween the canonical variables and the original variables (COR), and standardized canonical coefficients (SCC). Only values with high magnitudes are given. The canonical variables are labelled CV 1-2. Abbreviations for characters are explained in Table 1. 25 CVI CV2 original variable COR SCC COR SCC CD .002 not used .214 not used CS 080 —0.75* .189 —0.04 CLW 495* 0.36 502 1.03* TSL .032 not used е not used TT .073 not used .065 not used ATT .625* (0) o ia .282 0.24 ST 19905 0.47 .023 —0.66 NAC .150 not used .259 not used NS = 563 —0.54 .480 0.79 MJS 2209075 not used .360 not used * Most important variables. nitude of variation in the two clusters appeared equal, and results of Box's tests (Miller, 1968) show that vari- ation within colonies of each cluster appeared equal. Thus, because of differences between colonies within clusters, variation within colonies cannot be used di- rectly to estimate variation within clusters. The amount of overlap between colonies differs in degree from that between clusters. А B 135 Variation between populations of modern S. inter- septa was studied to determine if differences between populations were similar to differences between species. To do this, first, canonical discriminant analysis was performed for the three populations of the modern species (Table 3, Text-fig. 9). In this analysis, 76.7 percent of all colony means were correctly classified. This value is more than 10 percent lower than the percentage calculated in the analysis discriminating be- tween species clusters. The most significant overlap occurred between the nearby, environmentally-similar reef and sand channel localities. When the values for these two populations were pooled, the percentage of correctly-classified colonies was raised to 90 percent, a value similar to that for the analysis between species clusters. Thus, it can be concluded that the modern populations overlap to the same degree as the species clusters. However, it must be noted that the actual difference between the species clusters is far greater than that between modern populations (Text-fig. 6). Two discriminant functions were calculated in the analysis for the three modern populations, the first accounting for 78.78 percent of the variation and the second for 21.22 percent. The first function primarily distinguishes the lagoon group from the reef and sand channel groups, whereas the second function distin- C DUN (n=107) ——— DUN(nzIO7) oR DUN(n=107) EID ија INT (пе 294) کن شتو‎ INT(n=294) ла а INT(n=294) === === SPO(n=70) U مب رو‎ SPORT “፦፦- SPO(n-70) ———— 2 25 3 6 буума ага woe a CS(mm) AT T (mm) TSL (mm) D E Е DUN(nsIOT) | —— DUN(n=107) —— DUN(n=107) pron INT(n=294) -oe INT(n=294) е INT(n=294) == SPO(n=70) “፦ SPO(n=70) —— SPO(n=70) ጭ 2 4 6 D ou 3 ts 20 2 36 CLW(mm) ፐፐ(ጠጠ) NS Text-figure 7. — Means and standard deviations for six characters in the two NMB and one modern species. The midpoint of each horizontal line represents the mean, and the length of the line on either side of the midpoint is one standard deviation. Analysis of variance (or Welch's statistic when variances are unequal) shows that the species are significantly different in every case except NS. SPO = species 1, fossil ММВ S. spongiformis, DUN — species 2, fossil NMB 5. duncani; INT — species 3, modern S. intersepta. n — number of corallites measured. (A) Corallite spacing, CS, (B) inner wall thickness, ATT, (C) tertiary septum length, TSL, (D) columella width, CLW, (Е) total theca width, TT, (F) total number of septa, NS. 16 BULLETIN 328 guishes the sand channel group from the reef and la- goon groups. Five of the 10 variables were used in the analysis (Table 3). They are, in order of entry into the analysis: ATT, NS, CLW, CS, ST. CS, ATT, and NS are most heavily loaded on the first function; and ATT and ST are most strongly correlated with it. This result suggests that the thickness of the inner wall and the septa are most important in distinguishing the lagoon population from the two forereef populations. CLW, NS, and ST were most heavily loaded on the second function; and CLW and ST are most strongly corre- lated with it. This result suggests that thickness of the columella and the septa are most important in distin- guishing between the two forereef populations (Pl. 1). To determine if these two new discriminant func- tions are related to the original two functions distin- guishing the species clusters, Spearman's rank corre- lation coefficients were calculated between pairwise combinations of the four functions using all fossil and modern colony means. The results show a very high correlation between the first canonical variable of the CANONICAL VARIABLE 2 به - between-species analysis and the first canonical vari- able of the between-population analysis (г. = 0.97469; p — 0.0001). In other words, the same variables that distinguish the fossil species from the modern species also distinguish the modern lagoon population from the two modern forereef populations. This result fur- ther supports the hypothesis that 5. duncani and 5. intersepta represent parts of a morphologic continuum formed by one continuous species. However, as a com- plicating factor, the lagoon population lies interme- diate between 5. duncani and the two forereef popu- lations of S. intersepta (Text-fig. 6). Ofthe three habitats studied, the lagoon represents a high stress, environ- mental extreme, which is especially muddy with low circulation. Therefore, it should be most similar in environment to the lowest portion of the Cibao Valley sections, assuming the sequence deepens upward as interpreted by Saunders, Jung, and Biju-Duval (1986), and not to the uppermost part ofthe sequence as shown in Text-figure 6. Furthermore, colonies from the lowest portion of the Cibao Valley sections appear to have a -8 -6 -4 | Џ T Џ 4 CANONICAL VARIABLE 1 Text-figure 8.— Intraspecific variation in the two fossil ММВ species for canonical variables 1 and 2 of the canonical discriminant analysis between species of Stephanocoenia (Text-fig. 7, Table 2). Each smaller polygon surrounds the variation observed within one colony. The total area covered by polygons of S. spongiformis is lightly shaded and outlined with a dashed line. The total area covered by polygons of 5. duncani is darkly-shaded and outlined with a dotted line. The area of overlap between species is left unshaded. Considerable overlap is seen between colonies within species and, to a lesser extent, between species. DOMINICAN REPUBLIC NEOGENE. 4: FOSTER 17 skeletal morphology more extreme than that found in even the most extreme, high-stress modern environ- ment (/.е., the lagoon). Consequently, the postulated morphologic continuum cannot be explained as a re- sponse to change in environment alone. In summary, study of intraspecific variation recon- firms that the species in the genus Stephanocoenia do not form discrete clusters composed of uniform enti- ties. Variation within species is large, its magnitude cannot be estimated by variation within colonies, and colonies within species differ significantly. Overlap be- tween the three populations of the modern species oc- curs to the same degree as overlap between the three Species, although overall variation between popula- tions of the modern species is significantly lower in magnitude than variation between species. Two species (one fossil, one Recent) appear to form a morphologic continuum, suggesting that they represent chronospe- cies within one lineage. This continuum cannot be ex- plained as an ecophenotypic response to change in en- vironment alone. Comparison ofthese evolutionary patterns in Steph- anocoenia with those exhibited by species of the family Poritidae from the same sections shows similarities and differences between evolutionary patterns in the two groups and between patterns of intraspecific vari- ation in the two groups. S. spongiformis has a short duration and 15 morphologically stable as is common among the poritid species. However, it was not as abundant or widely distributed as were many poritids. In fact, Stephanocoenia itself appears restricted to the Caribbean throughout the Neogene. In contrast to 5. spongiformis, S. duncani exhibits gradual directional evolution. Like the poritids, S. spongiformis becomes extinct during the mid- to late Pliocene or early Pleis- tocene when a mass extinction occurred among many tropical marine invertebrates (Stanley, 1984). S. dun- cani may have continued its gradual evolution through the extinction interval. Unlike the poritids, Stepha- nocoenia does not experience a radiation of new taxa after the late Neogene to early Quaternary extinction. Both the poritids and Stephanocoenia exhibit large له N =‏ LJ |‏ RT 4‏ zh 5‏ нЕ ።‏ Eh‏ 61 4 ست t j pO መሚ Е ፄ R‏ IE‏ 4 < Zan nus‏ Luar AS TS R "‏ С ] 3 3‏ z | S‏ O Е 5 R S‏ Е P‏ | < ርታ ጽዳ 5‏ ورت l— — EEE TENERE ee ሸል E r E я‏ -е 0 2 4 6‏ 4- 6- CANONICAL VARIABLE 1 Text-figure 9. — Canonical discriminant analysis between populations of modern Stephanocoenia intersepta from three different reef habitats near Discovery Bay, Jamaica. Plot of scores on the first two canonical variables showing polygons outlining the range of variation between colonies within each population. The points represent colony means. L — lagoon population, R — forereef coral thicket population, S — forereef Sand channel population. amounts of intraspecific variability, which is more di- rectly related to environment in Stephanocoenia. Al- though more extensive among the poritids, overlap between species occurs in both groups. SYSTEMATIC PALEONTOLOGY INTRODUCTION Using the classification system of Wells (1956), the family Astrocoeniidae Koby, 1890 is represented by only one genus in the collection of Saunders, Jung, and Biju-Duval (1986). The family itself is described as colonial, having extratentacular budding, a phaceloid to cerioid colony form, a septothecal wall structure, beaded septal margins, and a styliform columella (if any). The septa are composed of few, simple trabec- ulae. No attempt will be made to formally describe the family herein, since considerable doubt has been ex- pressed as to whether the family represents a true monophyletic group and it therefore is much in need of revision (see Veron and Pichon, 1976). Of the 13 genera that Wells (1956) included in the family, seven are exclusively Mesozoic, three occur in Eocene to Miocene deposits (Platycoenia Vaughan, 1900, Stylo- coenia Milne-Edwards and Haime, 18488, and Astro- coenia Milne-Edwards and Haime, 18488), two range in distribution from Mesozoic to Recent (Stephano- coenia and ?Actinastraea d'Orbigny, 1849), and one (Stylocoeniella Yabe and Sugiyama, 1935) extends from Eocene to Recent. Only three (Astrocoenia, Stephan- ocoenia, and Stylocoenia), however, are commonly re- ported in the Neogene of the Caribbean (Vaughan, 1919), and three (Stylocoenia, ?Actinastraea, and ?Pla- tycoenia) in the Neogene of the Mediterranean (Chev- alier, 1961). Thus, both Astrocoenia and Stephano- coenia were restricted to the Caribbean during the Neogene, even though the distributions of both genera extended across the Atlantic during the Oligocene (Vaughan, 1919; Frost, 1981). An interesting question involving these taxa, from a Caribbean perspective, is why Astrocoenia and Stylocoenia do not occur in the Dominican Republic Neogene. The answer cannot be resolved without more rigorous study of the common Neogene species involved [4. guantanamensis Vaughan, 1919 (— A. decaturensis Vaughan, 1919 — A. portoricensis Vaughan, 1919); A. meinzeri Vaughan, 1919; and 5. pumpellyi (Vaughan, 1900)]. The formal systematic descriptions that follow are based strictly on the results of the statistical analyses presented in the previous section. As in Foster (1986), the term “diagnosis” is used only to describe higher categories such as genera, whereas the term “descrip- tion" is reserved for complete descriptions of species, which, as described in the previous section, herein rep- resent merely morphologic clusters of specimens. The terminology used in the “Description” sections is de- BULLETIN 328 fined and described in the previous section entitled "Characters". It follows the usage of Vaughan and Wells (1943) and Wells (1956). The abbreviations used for measurements are explained in Table 1 and are iden- tical to those used in the statistical analyses. The “Material” sections give an approximate esti- mate of the amount of material in the collection of Saunders, Jung, and Biju-Duval (1986) and the num- ber of specimens in this collection that were statisti- cally analyzed. Unless otherwise indicated, the locality numbers belong to the Naturhistorisches Museum Ba- sel (NMB) and material is deposited at the NMB. As- signment of formation names to individual localities is based on Saunders, Jung, and Biju-Duval (1986) (especially text-figs. 4, 6, 15, 16), except along Río Mao where formation names are used as listed by Maury (1919). Catalogue numbers have been assigned to all figured and measured material. A unique number was given to each colony. The “Remarks” sections explain synonymies and additional museum material used to estimate distri- bution patterns. Separate sections entitled “Variabil- ity" describe the variation within each cluster. Sections entitled “Occurrence” give detailed geographic and stratigraphic information within the studied areas of the Dominican Republic, whereas those entitled ““Dis- tribution" give general information on all known oc- currences throughout the world. Ages for Caribbean localities mentioned outside the Dominican Republic are based on the following sources: Jamaica, Zans et al. (1963); Trinidad, J. B. Saunders (oral commun., 1985). Genus STEPHANOCOENIA Milne-Edwards and Haime, 1848 Stephanocoenia Milne-Edwards and Haime, 1848а, p. 469. Antillastraea Duncan, 1884, p. 108. Type species. —Astrea intersepta Lamarck, 1816, p. 266. Diagnosis. — Colonies massive, plocoid to subceri- oid. Corallites budded extratentacularly, 1-3 mm in diameter. Septa usually 24 in number arranged in three cycles (six primaries, six secondaries, and 12 tertiaries); composed of six to eight trabeculae that form minute denticles along the septal margins. Costae short, equal. One crown of 12 pali (or a multiple of six) before first two septal cycles. Walls septothecal to parathecal. Col- umella styliform. Endothecal and exothecal dissepi- ments tabular with uniform spacing. Remarks. —Milne-Edwards and Haime (18488, р. 469) described the genus Stephanocoenia citing “ፈ፥- trea intersepta Lamarck" as the type species. Lamarck (1816, p. 266) described the species Astrea intersepta citing “Ап madrep. intersepta 2 Esper, suppl. I, +. 79” as the only reference. Lamarck's specimen is figured DOMINICAN REPUBLIC NEOGENE. 4: FOSTER 19 here in Text-figure 10a. According to M. E. Tollitt (written commun., 1984) and R. V. Melville (written commun., 1984) of the International Commission on Zoological Nomenclature, the question mark in La- marck's citation indicates that “ће was not satisfied that the two species were identical" and that “Lamarck was clearly describing a different species from Esper's" (letter dated 8 June, 1984, from R. V. Melville to the author). If Lamarck (1816) is interpreted not to have been describing a new species but citing Esper (1795) as the author of intersepta, multiple nomenclatural compli- cations would ensue, since re-examination of Esper's material shows that Esper's (1795, vol. 1, p. 99, pl. 79) Madrepora intersepta is clearly a member of the genus Alveopora Blainville, 1830 of the family Poritidae in the classification system of Vaughan and Wells (1943) and Wells (1956). Esper's specimen is figured here in Text-figure 10c. Accepting Esper as the author of the Lamarck intersepta would thus relegate Milne-Ed- wards and Haime's generic name “Stephanocoenia” to a junior synonym of Alveopora, leaving the name Antillastraea Duncan, 1884 as the next chronologically valid generic name for Lamarck's coral. Assuming that Lamarck's intersepta was not the same as Esper's intersepta, many Caribbean scleractinian workers (e.g., Smith, 1948; Squires, 1958; Almy and Carrión-Torres, 1963; Laborel, 1969; Cairns, 1982) have recently assigned the name Stephanocoenia mich- elini Milne-Edwards and Haime, 1848b (vol. 10, p. 301, originally spelled “michelinii” in error) to La- marck's species, since michelini has been usually syn- onymized with Lamarck's intersepta [after Gregory, 1895, рр. 276-277] and it is chronologically the next specific name that would apply. The holotype of mich- elini is figured here in Text-figure 10b. The present study of this problem, however, indicates that the species should be referred to as Stephanocoenia inter- septa (Lamarck, 1816) and not as Stephanocoenia michelini Milne-Edwards and Haime, 1848b. This тие, ч „ ጌሎ Text-figure 10. — Photographs of the colony surfaces of three ho- lotypes important in the nomenclatural problem involving Steph- anocoenia intersepta (Lamarck, 1816). A. Stephanocoenia intersepta (Lamarck, 1816), Lamarck collection, MNHNP, “Је5 mers Aus- trales", Recent, x 5 (photo by M. Serrette, ММНМР). This specimen was used by Lamarck (1816) to describe the species Astrea intersepta. It was later used by Milne-Edwards and Haime (1848a) to describe the genus Stephanocoenia. B. Stephanocoenia michelini Milne-Ed- wards and Haime, 1848b, Michelin collection, MNHNP, locality unknown, Recent, х 5 (photo by M. Serrette, ММНМР). This species has been synonymized with 5. intersepta, since Gregory (1895). Be- cause of the nomenclatural problem involving the name intersepta, several authors have used the name michelini to refer to the species. C. Alveopora intersepta (Esper, 1795), Esper Collection 68, NMS, East Indian Ocean, Recent, x9.6 (photo by G. Scheer). This spec- imen was Esper's (1795) original specimen of intersepta, which has recently been assigned to the genus Alveopora. practice follows that of the recent book by Zlatarski and Estalella (1982), although these authors were ap- parently unaware of the problem with Esper's speci- men. The genus Stephanocoenia occurs today only in the Caribbean and is represented by only one species, Stephanocoenia intersepta (Lamarck, 1816). Its mod- ern geographic distribution ranges from Bermuda throughout the Caribbean as far south as Brazil. Eco- logically, Stephanocoenia is common in many reef habitats extending from the lagoon across the reef crest (1 m depth) to the deep forereef (greater than 95 m depth). Its optimal depth range is 3-50 m (Goreau and Wells, 1967). Because various, poorly-preserved plocoid fossil corals have been assigned at some time to the genus Stephanocoenia, it is extremely difficult to determine its temporal distribution based on limited study of the literature. Stephanocoenia is clearly distinguished from other astrocoeniid and plocoid corals by: (1) its prom- inent pali, (2) its styliform columella, (3) its subcerioid growth form, and (4) its numerous septa usually ar- ranged in cycles of six. Based on these criteria, Steph- anocoenia has occurred in the Caribbean since the low- er Cretaceous (Wells, 1932; Wells, 1933). However, it may have been extremely rare during the Paleogene, since it has only been described from the Eocene of Peru (Wells, 1941). Some doubt exists as to whether Stephanocoenia occurred in the European Cretaceous (Alloiteau, 1957, рр. 56-57), although it was common in the Mediterranean during the Paleogene (Frost, 1981). It is not known from the Mediterranean Neo- gene (Chevalier, 1961). Stephanocoenia has been com- mon in the Caribbean from mid-Miocene to Recent (Vaughan, 1919; Frost, 1977). A number of Tertiary coral species that were origi- nally assigned to Stephanocoenia do not belong, in- cluding: Stephanocoenia reussi Duncan, 1867 (— Hyd- nophora Fischer, 1807), Stephanocoenia tenuis Duncan, 1863 (— Goniopora Blainville, 1830), Stephanocoenia dendroidea Milne-Edwards and Haime, 1860 (— Mad- racis Milne-Edwards and Haime, 1849), Stephano- coenia fairbanksi Vaughan, 1900 (— Solenastrea Milne- Edwards and Haime, 18488), and Stephanocoenia ? decaseptata Weisbord, 1971 (— Madracis). Only two names have been proposed for true representatives of the genus in the Caribbean during the Neogene: Steph- anocoenia intersepta (Lamarck, 1816) and 5. spongi- formis (Duncan, 1864). Gregory (1895) and Vaughan (1919) synonymized the two names. Stephanocoenia spongiformis (Duncan, 1864) Plate 2, figures 1—8; Plate 3, figures 1—6; Plate 4, figures 1—6; Text-figures 2, 3, 6, 7, 8 Plesiastraea spongiformis Duncan, 1864, p. 39, pl. 4, figs. 6a, 6b. BULLETIN 328 Description. — Colonies massive, hemispherical, small to intermediate in size (1.5—15 cm in length, 1.5— 6.5 cm in height) with smooth outer surfaces. Growth bands well developed at regular 2-3 mm intervals. Epitheca reduced, with irregular growth ridges at less than 1 mm intervals. Calices circular in shape; small in size (1.3-2.3 mm in diameter) and shallow in depth (less than 1 mm); with wide spacing (0.5-1.5 mm be- tween adjacent calices) giving the colony form a sub- cerioid appearance. Theca composed of a thin, well- defined, parathecal wall, circular in shape, surrounding each corallite. A second, equally well-defined polygo- nal wall (that may form a prominent ridge) demarcates the junctions between adjacent corallites. The two walls are separated by void space which is regularly subdi- vided by costae of equal thickness. These costae extend completely from the outer margin of each septum to the second wall. Septa approximately 24 in number, composed of six trabeculae (not including pali) that form fine surface denticles. Primary and secondary sep- ta of equal length extending completely to the colu- mella. Tertiary septa free, one-third to one-half the length of the first two cycles. Pali reduced, 12 in num- ber, formed by thickening of two to three trabeculae at the inner margins of the first two septal cycles. Col- umella forming a prominent style, 0.2-0.5 mm in di- ameter. Endothecal dissepiments thin, at 0.4 mm in- tervals. Exothecal dissepiments moderately thick (0.04 mm) at 0.3 mm intervals. Exothecal dissepiments moderately thick (0.04 mm) at 0.3 mm intervals, form- ing uniform, approximately square cellular voids. Holotype. —8M(NH) R28756 (figured here: Pl. 2, figs. 7, 8). Measurements of the holotype. — Colony length, 5.10 cm; colony width, 4.96 cm; colony height, 5.75 cm. The holotype for this species has not been thin-sec- tioned; therefore, the following characters were mea- sured on the colony surface and are not as accurate as the thin-section measurements given above in the species description. They cannot be directly compared with the thin-section measurements given in Text-fig- ure 7 and the Appendix. Each value represents the mean of 10 corallites. CD, 2.6; TT, 1.0; NS, 24.2; CLW, 0.7; ATT, 0.13; ST, 0.12; TSL, 0.4. Type locality. —“silt of the Sandstone plain" of He- neken (1853), Dominican Republic, Neogene. Material. —Thirty-two colonies from 16 localities. Seven colonies thin-sectioned and measured. Remarks. — Duncan (1864) recognized two species which belong to Stephanocoenia in the Heneken col- lection of fossil corals from Ше “silt of the Sandstone plain" of the Dominican Republic: “Stephanocoenia intersepta, var., Edwards & Haime" [p. 27, BM(NH) R28811] and “Plesiastrea spongiformis, spec. nov.” Гр. 39, BM(NH) R28756]. Re-examination of these two DOMINICAN REPUBLIC NEOGENE. 4: FOSTER 21 specimens shows the first to belong to 5. duncani as defined herein and the second to belong to 5. spongi- formis as defined herein. Duncan (1884) later desig- nated spongiformis as the type species of his new genus Antillastraea. Gregory (1895) and Vaughan (1919) syn- onymized the species spongiformis with S. intersepta and consequently the genus Antillastrea with Stepha- nocoenia, because they could not recognize consistent distinctive characteristics in the material at the BM(NH), which Duncan labelled “spongiformis”. Nevertheless, Duncan's holotype of spongiformis clearly lies within the range of one of the two fossil species defined statistically in the present study. Therefore, Duncan's name is reinstated. Variability. — Variation within this species is consis- tently high as in S. duncani. Colonies differ widely from one another (Text-fig. 8); however, the variation does not appear directly related to environment. No direc- tional morphologic trends were observed up the stud- ied sequence. From colony top to bottom, corallite diameter increases, total theca width increases, skeletal structures thicken, and pali become better developed. Comparison. —S. spongiformis is distinguished by its well-developed, porous exothecal intercostal area and by its thinner skeletal structures. It differs statis- tically from the fossil species 5. duncani by having wider corallite spacing, a smaller columella, shorter tertiary septa, a wider intercostal area, and a thinner inner wall. It differs statistically from the modern species S. intersepta by having a smaller corallite diameter, a thinner columella, shorter tertiary septa, a wider in- teroostal area, a thinner inner wall, and thinner septa (Text-fig. 7). Its pali are elongate and poorly developed, although better developed than in 5. duncani; and its colonies are small in size. Occurrence. — Río Cana: Gurabo Formation (locs. NMB 16818, 16865) and Mao Formation (loc. NMB 16884). Río Gurabo: Gurabo Formation (locs. NMB 15846, 15847, 15850, 15853, 15855, 15858, 15859, 16811, 16883, 16934) and Mao Formation (locs. ММВ 15822, 15830, 15834). Distribution. — This species is not known to occur outside the lower to middle Pliocene of the Dominican Republic. Stephanocoenia duncani, new species Plate 5, figures 1-6; Plate 6, figures 1-6; Plate 7, figures 1-6; Text-figures 2, 3, 6, 7, 8 Etymology of the name. — After P. M. Duncan, who first recognized the species as a variety. Description. —Colonies massive, hemispherical, sometimes moderately large in size (6-30 cm in length, 3-14 cm in height) with smooth outer surfaces. Growth bands well developed at regular 3-5 mm intervals. Epitheca reduced, with irregular growth ridges at 1 mm intervals. Calices polygonal in shape; small in size (1- 2.4 mm in diameter) and shallow in depth (approxi- mately 1 mm); with narrow spacing (0.2-1 mm be- tween adjacent corallites) giving the colony form a cerioid appearance. Theca composed of a thick septothecal wall separated from theca of adjacent cor- allites by little or no void space. А thin, poorly-de- veloped second wall may form between thecae of ad- jacent corallites. Septa 22 to 24 in number, composed of six trabeculae that form minute surface denticles. Primary and secondary septa of equal length extending completely to the columella. Tertiary septa free, one- half to three-quarters the length of the first two cycles. Pali weak, 12 in number, formed by thickening of two trabeculae on the inner margins of the first two septal cycles. Columella forming a prominent style, 0.3-0.6 mm in diameter. Endothecal dissepiments thin and at 0.4 mm intervals. Exothecal dissepiments thick (0.2 mm), at 0.5 mm intervals. Exothecal dissepiments forming elongate voids, separated by thin, irregular vertical trabeculae. Holotype. —NMB D5766 (figured here: Pl. 5, figs. 2, 4-6: Pl. 7, figs. 1, 2, 5). Measurements of the holotype. — Colony length, 23.8 cm; colony width, 18.7 cm; colony height, 12.8 cm. The following measurements were made on thin-sec- tions. Each value represents the mean of 10 corallites. CD, 2.06; CS, 2.48; CLW, 0.48; TSL, 0.26; TT, 0.51; ATT, 0.229; ST, 0.074; NAC, 6.4; NS, 24.0; MJS; 12.0. Type locality. — Locality ММВ 16817, Río Cana, Gurabo Formation, Dominican Republic. Lower Plio- cene (Saunders, Jung, and Biju-Duval, 1986, text-fig. 15). Paratypes. —NMB 135765, 05768, 05769, 05771, D5868, D5870, D5871 (figured here: Pl. 5, figs. 1, 3; Pl. 6, figs. 1-6; Pl. 7, figs. 3, 4). BM(NH) R28811, one of Duncan's (1864, p. 27) specimens of S. intersepta, var. Material.— Twenty-four colonies from 11 localities. Ten colonies thin-sectioned and measured. Remarks. — As explained in the Remarks section of the previous species, specimens of this species in the BM(NH) Heneken collection from the “silt of the Sandstone plain" of the Dominican Republic were la- belled “Stephanocoenia intersepta, var." by Duncan (1864, p. 27). Vaughan (1919) also placed this species within 5. intersepta. One of Duncan's specimens [BM(NH) R28811] is designated as a paratype. Variability. — Variation within this species is high, as in S. spongiformis; however, variation between col- onies may not be as extensive as in 5. spongiformis (Text-fig. 8). An increase in the thickness of skeletal 22 BULLETIN 328 structures such as the inner wall and the columella up the studied stratigraphic sequence have been detected. This increase is interpreted as evolutionary, although environmental factors could be at least partially re- sponsible. Variation within colonies is equally as high as that observed in 5. spongiformis. From colony top to bottom, corallite diameter increases, wall thickness increases, and various skeletal structures appear more heavily calcified. Comparison. — S. duncani is distinguished primarily by its poorly-developed, denser intercostal area giving the colony form a cerioid appearance; by its especially weak pali; by its predominantly septothecal wall struc- ture; and by its thicker skeletal structures. It differs statistically from the fossil species spongiformis by having closer corallite spacing, a thicker columella, shorter tertiary septa, a narrower intercostal area, and a thicker inner wall. It differs statistically from the modern species intersepta by having a smaller corallite diameter, closer corallite spacing, a thinner columella, shorter tertiary septa, a narrower intercostal area, a thinner inner wall, and thinner septa. It sometimes forms large colonies. Occurrence. —Río Cana: Gurabo Formation (locs. NMB 16815, 16817, 16818, 16819, 16881) and Mao Formation (loc. NMB 16874). Río Gurabo: Gurabo Formation (locs. NMB 15855, 16933, 16934). Río Mao: ?Gurabo Formation (loc. NMB 16911). Río Yaque del Norte: Baitoa Formation (loc. NMB 16943). Distribution. — Stephanocoenia duncani ranges in age from middle Miocene to middle Pliocene and is rare in the southern and central Caribbean. Outside the studied sequence in the Dominican Republic, it occurs in the following strata: ?Міосепе. — Rio Yaque del Sur section of the Dominican Republic, and St. Croix, Trinidad; Pliocene. — Bowden Formation of Jamaica. Appendix. Means (+1 standard deviation) of all corallite characters in the two fossil species of Stephanocoenia and in three populations of modern Stephanocoenia intersepta. “N” refers to the number of corallites measured. Ten corallites were measured in each colony. See Table 1 for explanations of character abbreviations. S. intersepta 5. intersepta S. intersepta S. spongiformis S. duncani N = 100 N = 100 N = 100 Variable N = 70 М = 110 (lagoon) (forereef) (sand channel) CD 1:91 (2.21) 1.97 (2.28) 201 (26225) 2,08 (+.24) 210) (19) cs* 2.82 (+.46) 2.42 (+.28) 2.86 (+.34) 2.94 (+.35) 2.90 (+.26) CLW* .365 (+.062) .418 (+.058) .434 (+0.64) .536 (+.098) .508 (5.081) TSL* .230 (+.088) .268 (+.061) .300 (=.091) .295 (+.079) .293 (+.072) ET .990 (+.251) .512 (+.164) .839 )±.248( .866 (+.174) .842 (+.163) ATT* .156 (+.104) .188 )±.078( .296 (+.129) .406 (+.100) .407 (+.093) ST .071 (+.017) .076 (+.019) .076 (+.014) .090 (+.018) .094 (+.016) NAC 6.48 (+.68) 6.36 (55 77) 6.41 (+.88) 0,33 (e Ma 6.44 (+.80) NS* 23.86 (+.49) 23:59" (1590) 23,215 (35/62) 23.79 (+1.04) NI7 EIS) MJS 11.91 (2.28) PII ESS) 11.94 (+.34) 11.83 (+.69) 15፡23) * Characters whose means and standard deviations are diagrammed in Text-figure 7. NEOGENE PALEONTOLOGY IN THE NORTHERN DOMINICAN REPUBLIC 5. The Suborders Caryophylliina and Dendrophylliina (Anthozoa: Scleractinia) By STEPHEN D. CAIRNS JOHN W. WELLS Smithsonian Institution Cornell University Washington, DC 20560, U.S.A. Ithaca, NY 14853, U.S.A. ABSTRACT The 20 species of Neogene Scleractinia in the suborders Caryophylliina and Dendrophylliina known from the Dominican Republic are revised and illustrated. This research was based on 1590 specimens obtained primarily from the collections of the Naturhistorisches Museum, Basel, Switzerland; National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A.; and Tulane University, New Orleans, LA, U.S.A. Eight new records are reported for the Neogene of the Dominican Republic, including four new species: Antillocyathus alatus, Trochocyathus chevalieri, T. duncani, and Paracyathus sinuosus. Special attention is given to the genus Asterosmilia, since half (five) of the known species in this genus occur in the Dominican Republic. Most species described herein are assumed to constitute a deep-water fauna by analogy to depth ranges of the same or similar species known from the Recent. Certain localities and parts of formations are inferred to represent deep-water (> 200 m) facies. These inferences may aid in the paleoecological interpretation of other fossils collected from these areas. RESUMEN Las veinte especies de Scleractinia Neogene de los subórdenes Caryophylliina y Dendrophylliina que se conocen de la Repüblica Dominicana, son revisadas e ilustradas. Esta investigación fue hecha sobre una base de 1590 especímenes obtenidos, primaria- mente, de las colecciones del Naturhistorisches Museum, Basel, Suiza; el Museo Nacional de Historia Natural, Smithsonian Institution, Washington, DC, EEUU; y Tulane University, New Orleans, LA, EEUU. Se registran ocho localidades nuevas para las especies de Neogene de la Repüblica Dominicana, incluyendo en las mismas cuatro especies nuevas: Antillocyathus alatus, Trochocyathus chevalieri, T. duncani, y Paracyathus sinuosus. Se brinda atención especial al género Asterosmilia, puesto que la mitad (cinco) de las especies conocidas, del mismo, se encuentran en la Repüblica Dominicana. Se presume que la mayoría de las especies descriptas en esta obra constituyen una fauna de agua profunda por analogía a los ¿Tangos de profundidad de las mismas o especies similares conocidas de la época Reciente. Se infiere que ciertas localidades y partes de las formaciones representan las facies de agua profunda (> 200 m). Estas inferencias pueden ayudar a la interpretación de otros fósiles colectados en estas áreas. INTRODUCTION This paper is the third in a series of descriptions of coral faunas from the Neogene of the Northern Do- minican Republic, the first two being those of Foster (1986, 1987), and is one of many coral revisions an- ticipated from the large, multidisciplinary project of Which they are part. This project surveys in depth the paleontology and stratigraphy of the Neogene of the Cibao Valley region ofthe Dominican Republic (Text- fig. 1). The specific history of Neogene coral research In the Dominican Republic has been reviewed by Foster (1986), and the background, lithology, age, and ratio- nale of the entire project have been presented by Saun- ders, Jung, and Biju-Duval (1986). This information Will not be repeated, except to note that there are few places in the Caribbean Neogene outside the Domin- 1688 Republic where the macrofauna is so diverse and Well-preserved in continuous sequence as to allow a Meaningful examination of total assemblages and their relation to paleoenvironment and time. Itis hoped that our contribution will add to the growing knowledge about this region and eventually will contribute to a better understanding of the evolution, zoogeography, and paleoecology of this region. ACKNOWLEDGMENTS We would like to thank the following people who have generously extended to us the use of their collec- tions or loaned us specimens used in this study: P. Jung (Naturhistorisches Museum, Basel, Switzerland), B. R. Rosen [British Museum (Natural History), Lon- don, England, U.K.], E. H. Vokes (Tulane University, New Orleans, LA, U.S.A.), R. Eng (Museum of Com- parative Zoology, Cambridge, MA, U.S.A.), and G. Gil (Muséum national d'Histoire naturelle, Paris, France). We are also very grateful to Nancy Foster (University of Iowa, Iowa City, IA, U.S.A.) for in- volving us in the Dominican Republic paleontological survey, doing the initial sorting of specimens, and pro- 24 viding invaluable advice regarding the lithology of the area and the proper format for this paper. PALEOECOLOGY The suborders Caryophylliina and Dendrophylliina contain primarily azooxanthellate ahermatypic species, which are generally associated with deep-water envi- ronments. For instance, of the 109 species of Recent western Atlantic Scleractinia in these two suborders (5696 of the 194 species of Scleractinia known from the western Atlantic [Cairns, 19791), only one is a her- matypic zooxanthellate [Eusmilia fastigiata (Pallas, 1766), the remainder being ahermatypic azooxan- thellates. Ninety-three of these remaining 108 species (8696) have depth ranges that exceed 200 meters (Cairns, 1979), some known from as deep as 3475 m. If simple analogy to the Recent fauna were allowed, then most of the 20 species included in this account could be assumed to be lower-shelf or upper-slope in- habitants and the stations at which they were collected could be considered to have preserved a deep-water facies. However, depth ranges of species undoubtedly have changed with time, and the ahermatypic species composition certainly has also changed. Therefore, an alternative approach to establishing paleobathymetry would be to consider the depth ranges of the four species of the Dominican Republic that have persisted from the Neogene to Recent: P. hispidus (349-1200 m), D. italicus (403-2634 m), G. annulata (28–653 m), and D. cornucopia (132-604 m). The first two species have BULLETIN 328 depth ranges that are exclusively deep-water (г.е., ex- clusively deeper than 200 m), having been collected at 13 localities (Table 1). In addition to these two species, seven! other species were collected at these 13 stations (Table 2), the implication being that these latter species probably had bathymetric ranges that were at least as deep as the shallowest depth range of P. hispidus and D. italicus (i.e., 350—400 m). Further support for this hypothesis includes a list of depth ranges of the Recent Caribbean counterparts to all nine species (Table 2), which, for the most part, have upper continental slope ranges. It was considered inadvisable to retroextrapo- late by inferring that all stations at which these addi- tional seven species occurred were also deep-water, since some ahermatypes have broad bathymetric ranges (e.g., С. annulata: 28-653 m) and, thus, to do this would probably include some shallow-water stations as well. Some of the other 11 of 20 species probably also have deep-water distributions, which may be sub- sequently inferred by correlation with other inverte- brate groups from the same stations. Nine of the 13 deep-water stations listed in Table 1 occur in the Mao Formation (570—757 m above base of section) of Río Gurabo (early to middle Pliocene), with several of them occurring at the very base of the ГА tenth “deep-water” species, А. maoensis, was also reported from deep-water station loc. USGS 8733; however, because none of the other 48 records of this species is from deep water and because it resembles a shallow-water meandrinid, it is not proposed as a deep-water species. N E 10 20km 1 Rio Cana 2 Rio Gurabo 3 Rio Mao GUAYUBIN Св | لا‎ Upper Cenozoic 4 Rio Amina A О [+ * 2] Oligocene - Early Miocene 2 5 Cañada Zalaya 3 ; 6 Rio Yaque del Norte 3 7 4 мок 7 City of Santiago Ки 4 4 8 Аггоуо Рипа! S Е 9 Rio Verde 5 4 J y 3 : 1 2 S VALVERDE ESPERANZA NAVARRETE | 3 ZAMBA (MAO Au يلا ما‎ $ SS RET BN сова LOSQUEMADOS & BU. SANTIAGO. ፳ = CRODRIGUE 4 пена TIPS, |SANTIAGO BULLA | МОСА : 8 6 ОФ ለሰር ~ ВА!ТОА ST "eg ulna dr EE نلا‎ Text-figure 1.— Мар indicating the location of river sections sampled. Specimens reported in this paper were collected in five sections: (1) Río Cana, (2) Río Gurabo, (3) Río Mao, (4) Río Amina, and (6) Río Yaque del Norte (map from Saunders, Jung, and Biju-Duval, 1986). DOMINICAN REPUBLIC NEOGENE. 5: CAIRNS AND WELLS 25 Table 1.— Localities at which P. hispidus and D. italicus were collected (therefore presumed to be deep-water environments [over 200 m]. Level above Base of Section Locality Section Formation (meters) Species Collected NMB 15823 Río Gurabo Mao 672 P. hispidus NMB 15828 Río Gurabo Mao 713 P. hispidus NMB 15829 Río Gurabo Mao 757 P. hispidus NMB 15995 Río Gurabo Mao 570 P. hispidus NMB 16016 Río Gurabo Mao 580 P. hispidus NMB 16023 Río Gurabo Mao 640 P. hispidus NMB 16036 Río Gurabo Mao 660 P. hispidus NMB 16038 Río Gurabo Mao 660 P. hispidus USGS 8735 Río Gurabo Mao 660 P. hispidus, D. cornucopia, T. duncani USGS 8702 Río Yaque del Norte ?Baitoa ? D. italicus, A. alatus USGS 8726 Río Yaque del Norte ?Baitoa E P. hispidus, A. alatus USGS 8733 Río Mao ?Gurabo " P. hispidus, D. cornucopia, Flabellum sp., A. profunda, A. exarata, A. maoensis TU 1227A Arroyo Zalaya ?Gurabo Y P. hispidus, G. sp. cf. G. annulata, A. alatus Mao Formation. Two stations (loc. USGS 8702 and loc. USGS 8726) occur in the ?Baitoa Formation of Río Yaque del Norte, which is early to mid-Miocene; one station (loc. USGS 8733) occurs in the ?Gurabo Formation of Río Mao; and one station (loc. TU 1227A) occurs in the ?Gurabo Formation of Arroyo Zalaya. Twelve of the 20 species reported in this paper are known only from the Dominican Republic but are not necessarily considered to be endemic to this island. Future collecting from the Caribbean Neogene will un- doubtedly extend the distributions of many, if not all, of these species. SYSTEMATIC PALEONTOLOGY ~ INTRODUCTION The Dominican Republic Neogene Scleractinia ex- amined in this study were obtained from four insti- tutions, here listed in decreasing order of specimens available: NMB, NMNH, TU, and BM(NH). Most of the specimens from NMNH (collections of the U. S. Geological Survey) were previously reported by Vaughan (1919), Vaughan and Woodring (1921), and Vaughan and Hoffmeister (1925), whereas the BM(NH) Specimens were reported by Duncan (1863, 1864, 1867, 1868), altogether resulting in records of 12 species in Table 2.—Species believed to occur in deep water, and depth Tanges of their Recent counterparts. Dominican Republic Neogene Species Recent Caribbean Counterparts and Depth Ranges Antillocyathus alatus none Trochocyathus duncani T. rawsonii (82-622 m) Deltocyathus italicus D. sp. cf. D. italicus (403-2634 m) Asterosmilia exarata A. marchadi (32-229 m) Asterosmilia profunda none ?Flabellum sp. F. atlanticum (357-618 m) Pourtalocyathus hispidus P. hispidus (349-1200 m) Guynia Sp. cf. С. annulata С. annulata (28-653 m) Dendrophyllia cornucopia Р). cornucopia (132-604 m) the suborders Caryophylliina and Dendrophylliina. Reports of specimens from NMB and TU have added eight new records of these taxa to the Dominican Re- public Neogene (see Checklist herein), including four new species. Altogether, new specimens were available for 15 of the 20 taxa now known from this fauna. The five species of which additional specimens were not obtained [P. henekeni (Duncan, 1863), D. italicus (Michelotti, 1838), D. dominicensis (Vaughan, 1925), A. compressa Vaughan, 1925, and A. duncani Vaughan, 1925] are nonetheless included in this paper for com- pleteness. Instead of giving a full description of these five species, we have provided diagnoses based on a reexamination of type-material and/or topotypic spec- imens, or, in one case, on specimens subsequently col- lected from an area other than the Dominican Repub- lic. The higher-level classification used follows Wells (1956). Definitions of most terms can also be found there; however, several additional terms are defined here. The ratio of the greater calicular diameter to the lesser calicular diameter, abbreviated гсалса, is used in some species accounts. The /ateral thecal edges are the two narrow thecal edges of an elliptical calice, г.е., the part ofthe theca and calice intersected by the great- er calicular axis. The /ateral thecal faces are the two broad edges of an elliptical calice, i.e., the parts of the theca and calice intersected by the lesser calicular di- ameter (Text-fig. 2). The two principal costae and septa are those aligned with the greater calicular axis along the lateral thecal edges. Cycles of septa, costae, and pali are indicated numerically following the capital letters 5, C, and P, respectively. A species or specimen with five complete cycles of septa (96 septa), the first three cycles equal in exsertness and width and the remaining two cycles progressively smaller, is indicated by the formula: S1-3>S4>S5. S4+ indicates that in addi- 26 ВОШЕТМ 328 tion to a full fourth cycle of septa, some septa of the fifth cycle are present. When symmetry is not hex- ameral, as in Antillocyathus Wells, 1937, the septa are referred to as groups of equal-sized septa, termed pri- mary, secondary, etc. The synonymies are complete for all but five species, these five species having references to more complete synonymies. А question mark preceding a synonym entry indicates a doubtful identification not verified by examination of the specimen. The convention cf. is used for two of the 20 species identified [i.e., C. duodecimcostatus (Goldfuss, 1826) and С. annulata Duncan, 1872] to imply that the Do- minican Republic specimens are most similar, if not identical, to these species, but that not enough speci- mens are available to make a definitive statement. The type-specimens of 16 ofthe 19 described species were examined. The types of C. duodecimcostatus, D. italicus, and D. dominicensis are lost; however, refer- ence specimens of these three species were examined. Inthe Material section, localities (locs.) NMB 15803- 17273 are cited. Unless otherwise indicated, specimens may be assumed to be deposited at the collecting in- stitution: four-digit catalog numbers prefaced with a D refer to the NMB collection. The number in paren- theses in the Material sections indicates the number of specimens examined; if accompanied by a museum catalog number, this museum number refers to all spec- imens represented by the number in parentheses. In- formation on localities and ages of collection sites are found in Saunders, Jung, and Biju-Duval (1986) for NMB localities; Vaughan et al. (1921) for the USGS localities; and Vokes (1979) for TU localities. In the Comparison sections of the species accounts, the Dominican Republic species are usually first com- pared to the Neogene congeners from the Dominican Republic and Caribbean; next, to other European con- geners (as reviewed by Chevalier, 1961); and finally, to Caribbean Recent congeners (as reviewed by Cairns, 1979). In most cases these related species were ex- amined and some are illustrated in the plates. PHILOSOPHICAL CONSIDERATIONS Interspecific skeletal variation has plagued coral sys- tematists for over a century (Veron and Pichon, 1976). The high phenotypic plasticity of the coralla, easily modified by environmental conditions, caused such difficulty in determining species boundaries that Ber- nard (1903) departed from Linnaean binomial no- menclature for a noncommittal geographic numbering system. Although many nonskeletal characters of Scleractinia can be used for taxonomy at the species level (see review by Lang, 1984), the identification and classification of both Recent and fossil Scleractinia de- pend almost entirely on skeletal characters, much to the delight of the paleontologist, and in most cases the skeleton provides an adequate set of characters to dif- ferentiate species and to form a higher classification. The species concept used in this paper is that of the morphospecies, first formalized for Scleractinia by Vaughan (1907, p. 4) as: ^... a group of individuals connected among themselves by intergrading charac- ters and separated by distinct lacunae from all other individuals or groups of individuals.” It is obviously advantageous to have as many specimens as possible of each species in order to elevate diagnoses and de- scriptions from the typological level to a biological or population level, including an analysis of variation. For some species, e.g., Antillocyathus maoensis (Vaughan, 1925), it was possible to study variation, but for others, e.g., Asterosmilia duncani Vaughan, 1925, and A. compressa Vaughan, 1925, the treatment was literally typological, since only the types of these species are known. Basically, our systematic philoso- phy was to: obtain as many specimens as possible (in this case, 1590) for a study of intraspecific variation; examine all extant types of species previously de- scribed from the Dominican Republic; study closely- related species in both the Neogene and Recent of the Caribbean and Mediterranean; and qualify our iden- tifications if specimens were poorly preserved or few in number. Some of the discriminating characters useful at the species level were: (1) shape and size of corallum and resultant shape of calice; (2) number of septa at a par- ticular calicular diameter; (3) relative degree of exsert- ness of septal cycles; (4) costal development; and (5) pali and paliform lobe configuration. The character of columella shape, usually consistent at the generic level, was found to be variable within species of Asterosmilia Duncan, 1867. Thin-sections of coralla and scanning electron microscopy of septal faces were prepared for some species when adequate numbers of specimens were available, but little of systematic value was ob- tained from these approaches. lateral face principal septum principal septum / lateral edge. principal costa principal costa С! 05 co Тех!-ћриге 2.— Diagram of cross-section of a calice illustrating some of the morphological terms used in the text. S = septa, C = costae, numbers indicate respective septal or costal cycle. — е + 2 | DOMINICAN REPUBLIC NEOGENE. 5: CAIRNS AND WELLS 27 ABBREVIATIONS OF REPOSITORY INSTITUTIONS BM(NH): British Museum (Natural History), London, England, U.K. Gabb: Gabb Collection, deposited at the Museum of Comparative Zoology, Harvard College, Cambridge, MA, U.S.A. MCZ: Museum of Comparative Zoology, Harvard Col- lege, Cambridge, MA, U.S.A. MNHNP: Muséum national d'Histoire naturelle, Paris, France NMB: Naturhistorisches Museum, Basel, Switzerland NMNH: United States National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A. TU: Tulane University, New Orleans, LA, U.S.A. USGS: United States Geological Survey (specimens deposited at the NMNH) USNM: United States National Museum of Natural History (specimens deposited at the NMNH) CHECKLIST OF NEOGENE CARYOPHYLLIINA AND DENDROPHYLLIINA FROM THE DOMINICAN REPUBLIC Suborder Caryophylliina Vaughan and Wells, 1943 Superfamily Caryophylliicae Dana, 1846 Family Caryophylliidae Dana, 1846 Subfamily Caryophylliinae Dana, 1846 Antillocyathus maoensis (Vaughan in Vaughan and Hoffmeister, 1925) * Antillocyathus alatus, n. sp. Antillocyathus cristatus (Vaughan in Vaughan and Hoffmeister, 1925) *Trochocyathus (Paratrochocyathus) chevali- егі, n. sp. *Trochocyathus (Paratrochocyathus) dun- cani, n. sp. Ceratotrochus (Edwardsotrochus) sp. cf. C. duodecimcostatus (Goldfuss, 1826) Paracyathus henekeni (Duncan, 1863) *Paracyathus sinuosus, n. sp. Deltocyathus italicus (Michelotti, 1838) Subfamily Turbinoliinae Milne-Edwards and Haime, 1848b *Sphenotrochus (Eusthenotrochus) senni Wells, 1945 Dominicotrochus dominicensis (Vaughan in Vaughan and Hoffmeister, 1925) Subfamily Parasmiliinae Vaughan and Wells, 1943 Asterosmilia abnormalis (Duncan, 1864) Asterosmilia exarata Duncan, 1867 Asterosmilia profunda (Duncan, 1864) Asterosmilia duncani Vaughan in Vaughan and Hoffmeister, 1925 Asterosmilia compressa Vaughan in Vaughan and Hoffmeister, 1925 Superfamily Flabellicae Bourne, 1905 Family Flabellidae Bourne, 1905 ?Flabellum sp. Family Guyniidae Hickson, 1910 *Pourtalocyathus hispidus (Pourtalés, 1878) *Guynia sp. cf. G. annulata Duncan, 1872 Suborder Dendrophylliina Vaughan and Wells, 1943 Family Dendrophylliidae Gray, 1847 * Dendrophyllia cornucopia Pourtalés, 1871 Suborder CARYOPHYLLIINA Vaughan and Wells, 1943 Superfamily CARYOPHYLLIICAE Dana, 1846 Family CARYOPHYLLIIDAE Dana, 1846 Subfamily CARYOPHYLLIINAE Dana, 1846 Genus ANTILLOCYATHUS Wells, 1937 Diagnosis. —Solitary; corallum strongly compressed, almost flabellate. Septotheca strongly costate. Pali be- fore 53 in one crown (or before penultimate cycle of septa when more than four septal cycles or when hex- ameral symmetry obscured). Columella elongate, sublamellar. Dissepiments absent or rare. Type-species. — Placocyathus maoensis Vaughan in Vaughan and Hoffmeister, 1925, by original designa- tion. Discussion. — Other species placed in Placocyathus Milne-Edwards and Haime, 1848b, by Duncan and Vaughan, other than the monocentric P. maoensis Vaughan in Vaughan and Hoffmeister, 1925, and P. cristatus Vaughan in Vaughan and Hoffmeister, 1925 (referred to as Antillocyathus in this paper), include P. barretti Duncan, 1863; P. variabilis Duncan, 1864; P. alveolus Duncan, 1863; and P. costatus Duncan, 1864. АП of these forms in their early stages resemble Ап- tillocyathus Wells, 1937, in shape but with increase in size become compressed, elongate, continuous or sin- uous polycentric calicular series. They have dissepi- ments and paliform lobes, rather than true pali of An- tillocyathus, before the second and third rank septa, and notably, continuous principal costae (“costae of symmetry"), the persistent primary directive costae. These are the characters of Meandrina Lamarck, 1801 which, however, lacks the continuous principal costae and has lateral calicular lobes. The general morphology of these polycentric species of Placocyathus is strik- ingly similar to that of species of the short-lived late Cretaceous genus Phyllosmilia Fromentel, 1862, ex- * New record for the Dominican Republic. 28 BULLETIN 328 сер! for the presence of paliform lobes. Placocyathus barretti, P. variabilis, P. costatus, and P. alveolus thus appear to represent a distinct generic group meriting further study. Distribution. — Miocene, West Indies. Antillocyathus maoensis (Vaughan, 1925) Plate 8, figures 1-7, 13 Placocyathus maoensis Vaughan in Vaughan and Hoffmeister, 1925, pp. 317—318, pl. 1, figs. 3-10; Felix, 1927, p. 444. Antillocyathus maoensis (Vaughan). Wells, 1937, p. 11; Weisbord, 1973, p. 62; ?Ricart y Menéndez, 1983, part (figs. 1, 13-15). Not Antillocyathus maoensis (Vaughan). Frost and Langenheim, 1974, pp. 300—301; Ricart y Menéndez, 1982a, pp. 129-130, figs. 1-4. ?Antillocyathus maoensis (Vaughan). Ricart y Menéndez, 19825, pp. 133-135, figs. 1-9. Description. — Corallum straight and flabellate, nar- rowing to a short, slender pedicel about 1.2 mm in diameter, which is invariably detached from substrate. From side view, corallum shaped as an isosceles tri- angle, with the variable angle, the pedicel, ranging from 60? to 80*. Largest specimen examined (USNM 64271 from loc. USGS 8545) 23 х 9 mm in calicular diameter and 25 mm in height; however, most specimens 13- 18 mm in вед. Calice elongate with parallel to slightly concave lateral faces. Ratio of gcd:lcd ranges from 5 to 2.50; this ratio increasing with increase of the gcd, implying a greater flattening of the calice with growth. The two lateral edge costae, those associated with the narrow thecal edges, often ridged and crested for basal 4—10 mm. Otherwise, costae usually low, rounded, and granulated, about 0.30 mm wide and separated by shal- low grooves. Occasionally C1 are slightly ridged. Shal- low transverse grooves about 1.75 mm wide periodi- cally encircle theca (Pl. 8, fig. 1). Septa not arranged hexamerally; instead, 19—24 large primary septa, an equal number of secondary septa, twice this number of tertiary septa, and often several pairs of quaternary septa fill the calice. Based on 24 well-preserved coralla, the most common number of primary septa is 20 (found in 16 of the coralla), fol- lowed by 21 primary septa in four coralla, 23 in two, 19 in one, and 24 in one. Number of septa per corallum ranges from 78 to 104, the one with 104 septa having 24 primary septa and four pairs of quaternary septa. Most common numbers of septa per corallum were 84 and 82, reflecting a corallum with 20 primary septa, a full complement of secondary and tertiary septa, and two (or one pair of) quaternary septa, respectively. When present, quaternary septa occur in pairs and usu- ally in half-systems directly adjacent to the two prin- cipal septa. Primary septa highly exsert, about 0.55 mm wide, with straight vertical inner edges, which fuse to columella lower in fossa. Secondary septa about half as large and much less exsert, about 0.25 mm wide. Tertiary septa smaller but as exsert as secondary septa, about 0.10 mm thick. Quaternary septa rudimentary. Inner edges of all septa simple, not bifurcate. Septal faces covered with sharp granules up to 0.13 mm in height, arranged both in rows parallel to the inner sep- tal edge and in lines perpendicular to septal edge (fol- lowing the trabeculae), the latter arrangement more apparent on distal septal edge, the former more com- mon on septal face. Fossa elongate and relatively shallow. Lower cor- allum filled with stereome; no endotheca. А distinct crown of rounded pali present, one before each sec- ondary septum and a smaller one before each tertiary septum that is flanked by quaternary septa. Occasion- ally a small paliform lobe occurs before primary septa. Pali 0.75-1.40 mm wide, separated from secondary septa by a deep notch. Inner edges of pali vertical and straight. Columella elongate (up to 7 mm long and 0.6 mm wide), solid, and lamellar. Sides of columella cov- ered with horizontal carinae of variable length. No “inner septa" (sensu Ricart y Menéndez, 19828). Diagnostic characters. — Hexameral symmetry usu- ally not apparent (often 20 primary septa); high ged: lcd (1.75—2.50); lateral corallum faces slightly concave. Type-material. — The holotype, collected at loc. USGS 7785, is deposited at the NMNH (USNM 353644). It is 14.4 x 7.8 mm in calicular diameter, 15.8 mm in height, and has 78 septa (including 19 primary septa and one pair of quaternary septa). Three paratypes are deposited at the MCZ: one, unnumbered, appears in Vaughan (1925) as pl. 1, figs. 5-6; a second, - MCZ 9267, is Vaughan's pl. 1, figs. 7-8; and a third, MCZ 9269, is Vaughan's pl. 1, figs. 9-10. Two addi- tional specimens labelled in Vaughan's handwriting as paratypes are deposited at the NMNH (USNM 68303 from loc. USGS 7776; and USNM 68304 from loc. USGS 7777) but because these were not cited by Vaughan (in Vaughan and Hoffmeister, 1925) as types, they are not considered as such. Type-locality. — Loc. USGS 7785: Rio Mao, Bluff 1 of Maury (?Cercado Formation), Dominican Republic, late Miocene. Material. — Unnumbered ММВ specimens from locs. NMB 15804 (2); 15805 (1); 15806 (26); 15807 (165); 15809 (13); 15810 (11); 15814 (4); 15836 (26); 15843 (1); 15855 (2); 15860 (3); 15863 (50); 15864 (80); 15865 (18); 15867 (2); 15869 (3); 15871 (1); 15883 (1); 15899 (1); 15964 (1); 16809 (4); 16810 (31) 16910 (12); USNM 64512 (25) from loc. NMB 15807; USNM 68303 (1) from loc. USGS 7776; USNM 68303 (2) from loc. USGS 7777; USNM 353644 (holotype) from loc. USGS 7785; USNM 64482 (11) from loc. USGS 8516; USNM 63072 (116) from loc. USGS 8519; USNM 64267 (1) from loc. USGS 8520; USNM 68424 (34) from loc. USGS 8521; USNM 64479 (7) from loc. DOMINICAN REPUBLIC NEOGENE. 5: CAIRNS AND WELLS USGS 8522; USNM 64269 (3) from loc. USGS 8528; USNM 64481 (1) from loc. USGS 8537; USNM 64483 (3) from loc. USGS 8538; USNM 64271 (67) from loc. USGS 8545; USNM 64265 (8) from loc. USGS 8546; USNM 64270 (1) from loc. USGS 8547; USNM 64272 (2) from loc. USGS 8549; USNM 64268 (1) from loc. USGS 8727; USNM 64264 (1) from loc. USGS 8733; USNM 64266 (1) from loc. USGS 8734; unnumbered TU specimens from loc. TU 1210 (40); loc. TU 1211 (11); loc. TU 1213 (1); loc. TU 1225 (1); loc. TU 1231 (33); loc. TU 1277 (9); loc. TU 1278 (12); loc. TU 1359 (5); loc. TU 1451 (1); MCZ 9267, 9269 (para- types). Remarks. — Frost and Langenheim (1974) reported a single, poorly-preserved specimen of 4. maoensis from the lower Miocene of Mexico; however, several elements of their description do not represent this species. First, their specimen had 79 septa arranged hexamerally. At the specimen size they reported, 79 septa would not be unusual, but hexameral symmetry and the relative sizes of the septa they reported (51>82>53) are inconsistent with Dominican Re- public specimens. Also, they reported paliform lobes before the first two cycles of septa but did not mention pali before the S3; this is quite different from typical specimens of A. maoensis. We seriously doubt the Mexican specimens are conspecific with А. maoensis. Ricart y Menéndez wrote three papers on Antillo- cyathus maoensis from the Dominican Republic. His third and most detailed paper (Ricart y Menéndez, 29 1983) described and illustrated ап unnamed species closely related to but different from А. maoensis. His first paper on inner septa of А. maoensis (Ricart y Menéndez, 1982а) also appears to describe this un- named species. The second paper (Ricart y Menéndez, 1982b) is problematic from the point of view of syn- onymy because only columellar structures are illus- trated; however, his figure 1 is consistent with the known columellar structure of А. maoensis. Comparisons. — Antillocyathus maoensis is similar to A. alatus, n. sp. as discussed in the account of the latter species. There are no known Recent species of Antillocyathus nor is any species of this genus known from the European Miocene. Occurrence. — Río Gurabo (Cercado and Gurabo for- mations) late Miocene to earliest Pliocene (Text-fig. 3): locs. NMB 15804, 15805, 15806, 15807, 15809, 15810, 15814, 15836, 15843, 15855, 15860, 15863, 15864, 15865, 15867, 15869, 15871, 15883, 15899, 15964, 16809, 16810, loc. USGS 7776, loc. USGS 7777, loc. USGS 8537, loc. USGS 8538, loc. USGS 8545, loc. USGS 8546, loc. USGS 8547, loc. USGS 8549, loc. USGS 8727, Пос. TU 12107 loc. TU 12112 loca TU 1213, loc. TU 1277, loc. TU 1278, loc. TU 1359. Río Mao (?Cercado Formation) late Miocene: loc. NMB 16910, loc. USGS 8519, loc. USGS 8520, loc. USGS 8521, loc. USGS 8522, loc. USGS 8528, loc. USGS 8733, loc. USGS 8734, loc. TU 1225. Río Amina: loc. USGS 8516. Arroyo Babosico (?Gurabo Formation): loc. TU 1451. —— Е RM Бинии RIO GURABO desee kid no RIO CANA in meters age tm fm age |Б 1000 4 е Ф d o n=2 n=2 = .2 92 a 2 о А - zi o E E n-8 800 = : = 1 = 2 = 1 Ф 1 о 2 о Ф jl = 5 i ፦ 600 4 a 8 -4 = a с E n=1 n=1 ш Е = = = o ш 5 та п ርን he n + ай n : мора በ2 ei ei 5 E 6 : e! 1 .2 $5 1 E ከ = 1 6 መ | ር› 1 1 በ=1 ጅጅ 2 & 8 j 1 ет ወ = 4 H 200 + о o 1 n=2 © о Ф itii 1 °2 ወ > > е 1 e! 2 zl 3 ፍ 12 5 pps o - о ፎሬ S хо O хо 5” S ќе ው کن‎ ድቻ ም Sd КЫ ም (2 ኣው Q^ NS СА o [a < پر‎ N о ኣው ው о o Ф Ка e ው > o < ҳу T e ቅ ው” S 4 < e ЖО በ ЕЗ e 3 S2>S3>S4. Inner edges of 51-3 moderately sin- uous. Degree of septal exsertness unknown. Fossa shallow. P1 small, about 1.0 mm wide, ex- tending to columella. P2 about 1.5 mm wide, pro- jecting higher in fossa, and also extending to columella. Pairs of P3 flank each P2, the P3 about same size and height as P2 but recessed from columella. Columella composed of a field of six to 10 granulated pillars ter- minating below level of P1. Diagnostic characters. — Ceratoid to trochoid coral- la; 48-54 septa; P1 P2—P3. ; Type-material. — Holotype: USNM 74684 from loc. TU 1208. Paratypes: USNM 74685 (4) from loc. TU 1208; USNM 63042 (1) from loc. USGS 8735. Type-locality. — Loc. TU 1208: Mao Adentro Lime- stone, roadcut, both sides of highway from Mao to Los Quemados, at ridge 4 km east of Los Quemados. Comparisons.— Trochocyathus (Paratrochocyathus) duncani, n. sp. is easily distinguished from 7. (P.) chev- alieri, n. sp. by its larger size, round calice, and different palar configuration. In the European Tertiary it is most similar to T. imparipartitus (Milne-Edwards and Haime, 1848b) from the upper Miocene of Italy (see Chevalier, 1961, p. 313), but can be distinguished by its round calice and lesser number of septa at a cor- responding size. Within the Caribbean, T. (P.) duncani is very similar to 7. (P.) rawsonii Pourtalés, 1874, particularly regarding calice shape and septal, palar, and columellar shapes and configuration. The only dif- ferences separating these two species are the lack of epitheca on T. (P.) duncani and its more slender, often ceratoid corallum. DOMINICAN REPUBLIC NEOGENE. 5: CAIRNS AND WELLS 33 Occurrence. —Río Gurabo (Mao Formation) early Pliocene: loc. USGS 8735. Mao Adentro Limestone: loc. TU 1208. Distribution. — Known only from the early Pliocene *Mao Adentro Limestone" of the Dominican Repub- lic, from Río Gurabo. Genus CERATOTROCHUS Milne-Edwards and Haime, 1848b Diagnosis. —Solitary; corallum trochoid; fixed or free. Septotheca costate. Paliform lobes before S1+2. Col- umella fascicular. No endotheca. Subgenus EDWARDSOTROCHUS Chevalier, 1961 Diagnosis. — Ceratotrochus in which the columella is composed of numerous flattened horizontal paliform lobes, not vertical fascicles as in the nominal subgenus. Type-species. — Turbinolia duodecimcostata Gold- fuss, 1826, by original designation. Distribution. — Miocene and Pliocene of Europe and west Africa, ?Azores; Miocene of West Indies. Ceratotrochus (Edwardsotrochus) species cf. C. (E.) duodecimcostatus (Goldfuss, 1826) Plate 9, figures 1, 2 Turbinolia duodecimcostata Goldfuss, 1826, p. 52, pl. 15, fig. 6. Ceratotrochus duodecimcostatus (Goldfuss). Duncan, 1864, p. 22; Duncan, 1868, pp. 23, 27; Felix, 1927, pp. 395-396; Felix, 1929, p. 568. Ceratotrochus duodecim-costatus (Goldfuss). Vaughan, 1919, p. 213. Ceratotrochus (Edwardsotrochus) duodecimcostatus (Goldfuss). Chevalier, 1961, pp. 359-360, text-figs. 125-128 (synonymy). Description of Dominican Republic specimens. — Corallum ceratoid to trochoid, straight to very slightly curved in plane of gcd. Largest specimen [BM(NH) 28819] 40 mm in height. Ratio of ред Лед = 1.38- 1.50. Pedicel diameter about 1.8 mm. C1-2 very prominent for length of theca, projecting up to 1.5 mm as thin ridges about 0.6 mm wide. C3-5 low ridges or flat. Septa hexamerally arranged in five incomplete cycles. Тће only specimen for which septal number could be determined (PI. 9, fig. 2, NMB D 5886 from loc. NMB 15836) has 84 septa, lacking both pairs of S5 in two half-systems and lacking one pair of S5 in two other half-systems. The poor preservation of the specimens did not allow observation of the septal sizes, fossa, pali, and columellar structures. Diagnostic characters. — C 1—2 thin exsert ridges; cor- allum slightly curved; fifth septal cycle usually incom- plete. Type-material and type-locality. — The holotype, from the Pliocene of Plaisance, Italy, is lost (Chevalier, 1961). Material. — Four unnumbered specimens from loc. NMB 15836; BM(NH) 28819 (1), Duncan's (1864) specimen from Angostina, Dominican Republic; un- numbered TU specimens from loc. TU 1225 (2); loc. TU 1277 (1). Reference specimens of C. duodecim- costatus from Italy: USNM 326663 (3) from Rio Tor- sero, Tuscany, Pliocene; USNM 156336 (3) from AI- berga, Liguria, Pliocene; USNM 156310 (2) from Astigiano, Parona, Pliocene; USNM 155251 (2) from Coroncina, Pliocene. Remarks.—' The eight Dominican Republic speci- mens are referred to C. (E.) duodecimcostatus based on their corallum size and shape, number of septa, costal structure, and external resemblance to topotypic specimens of C. duodecimcostatus (Pl. 9, figs. 3-4). Unfortunately, calicular details are lacking in all spec- imens, making a certain identification impossible. Mi- nor differences between the Dominican Republic spec- imens and the Italian specimens include the former having virtually straight coralla, whereas the Italian specimens have strongly-curved coralla. Also, the C1— 2 of the Dominican Republic specimens are more prominent than those of the Italian specimens. How- ever, Chevalier (1961) noted that this species was quite variable, particularly regarding the prominence of cos- tae. Comparisons. —Seven species of Ceratotrochus (Ed- wardsotrochus) were discussed by Chevalier (1961), all from the Miocene and Pliocene of Europe and north- west Africa. C. (Е.) duodecimcostatus is distinguished by its hexameral symmetry, curved corallum, and by having up to five complete cycles of septa. No other species of this subgenus is known from the Caribbean Tertiary or Recent. Occurrence. — Río Gurabo (Gurabo Formation) late Miocene (Text-fig. 3): loc. NMB 15836, loc. TU 1277. Rio Мао (?Gurabo Formation): loc. TU 1225. “Yellow Shale" of Angostina (Duncan, 1864, p. 22). Distribution of C. duodecimcostatus. — Miocene: northern Europe (Felix, 1927); ?late Miocene: Domin- ican Republic; Pliocene: southern Italy and northwest Africa (Chevalier, 1961; Felix, 1929); )Neogene: Azores (Chevalier, 1964). Genus PARACYATHUS Milne-Edwards and Haime, 1848b Diagnosis. —Solitary; trochoid to turbinate; fixed. Septotheca costate. Paliform lobes often bilobed or trilobed, standing before all but last cycle of septa. Columella papillose, often indistinguishable from in- ner paliform lobes. Type-species. — Paracyathus procumbens Milne-Ed- wards and Haime, 1848b, by subsequent designation (Milne-Edwards and Haime, 1850). Distribution. — Eocene to Recent, cosmopolitan. 34 BULLETIN 328 Paracyathus henekeni (Duncan, 1863) Plate 8, figures 17, 18 Brachycyathus henekeni Duncan, 1863, pp. 426—428, pl. 15, fig. 1. Paracyathus henekeni (Duncan). Duncan, 1868, pp. 16, 22, 27; Vaughan, 1919, pp. 213-214; Felix, 1927, p. 437; Frost and Lan- genheim, 1974, p. 302. ?Paracyathus henekeni (Duncan). Vaughan and Woodring, 1921, p. 152. Description of lectotype [BM(NH) 28768].— Coral- lum conical (trochoid) and free; 5.1 mm in calicular diameter and 5.7 mm in height. Alternate costae thin and ridged, dentate near calice. Forty septa hex- amerally arranged in four incomplete cycles: S1=S2>S3>S4. Fossa shallow. Paliform lobes slen- der but prominent teeth on septa of first three cycles. Columella papillose and small. Septal granules point- ed, not carinate. Diagnosis.— Costae straight, ridged, and dentate; calice round; paliform lobes discrete. Type-material.—Eleven syntypes are deposited at the BM(NH), numbered 28768—28778. Because some of the specimens are not well preserved [e.g., BM(NH) 28776, 28778], there 15 doubt about their identity, and therefore a lectotype is here designated: the figured specimen of Duncan (1863), BM(NH) 28768. Type-locality. —“Nivaje Shale” of Heneken, Do- minican Republic, Miocene. Material. – ВМ(МН) 28768 (lectotype); BM(NH) 28769—28778 (paralectotypes). Remarks. —No additional specimens of P. henekeni were collected and thus little can be added to Duncan's (1863) original description. We disagree, however, with Duncan's description of the paliform lobes as small, indistinct, and feebly-developed papillae. The pali- form lobes are, in fact, discrete, prominent structures. Also, Duncan (1863) originally described the columella as large and papillose but later (Duncan, 1868) referred to it as small. We confirm his latter description. Vaughan and Woodring's (1921) P. henekeni is not preserved well enough for identification. Comparisons. — Paracyathus henekeni is compared to P. sinuosus in the following species account. It is unlike any other species of Paracyathus from the Ca- ribbean or Tertiary of Europe. Occurrence. — Known only from the type-locality in the Dominican Republic, Miocene. Distribution. — Known only from the Miocene of the Dominican Republic. Paracyathus sinuosus, new species Plate 9, figures 5-9 Etymology. — The specific name sinuosus (Latin for “full of bendings””) refers to the sinuous course of the costae on the upper theca. Description. — Corallum ceratoid and straight, nar- rowing to a slender pedicel 1.5-1.9 mm in diameter. Pedicel expands basally up to 3 mm in diameter, form- ing the attachment. Calice of holotype (NMB D 5888) slightly elliptical, 4.5 x 4.1 mm in diameter; corallum 7.5 mm in height. Theca slightly porcelaneous, covered basally by beaded, equal costae. In upper half of cor- allum, costae ridged and often slightly sinuous (Pl. 9, fig. 8). Septa hexamerally arranged in four cycles: 5152-93-54. 54 strongly fused to adjacent S3. This fusion, along with the P3, forms a large, solid structure in each half-system. Sides of septa and pal- iform lobes covered with prominent granules and ca- rinae, the latter up to 0.1 mm in height and 0.3 mm long (Pl. 9, fig. 5). Fossa shallow. Paliform lobes irregularly-shaped pil- lars occurring before first three cycles of septa. Colu- mella papillose, composed of 10-15 irregularly-shaped rods similar in shape to paliform lobes but only about one-third the size and more deeply seated in fossa. Diagnosis. — Corallum small; costae often sinuous; S4 fused to S3. Type-material. — Holotype: NMB D 5888 from loc. NMB 15833. Paratypes: unnumbered NMB specimens from locs. NMB 15832 (8); 15833 (36); 17023 (1); 17024 (8; USNM 74686 (8) from loc. NMB 15833. Type-locality. — Loc. ММВ 15833: Río Gurabo (Mao Formation: 892 m above base of section), Dominican Republic, middle Miocene. Reference material. — Nine specimens of P. turonen- sis from Pontlevoy, France (MNHNP L 61). Remarks. — Vaughan and Woodring (1921, pp. 133, 163) listed three undetermined species of Paracyathus from the Miocene of the Dominican Republic: Para- cyathus sp. (from loc. USGS 8540 and loc. USGS 8544), Paracyathus n. sp. a (from loc. USGS 8735 and loc. USGS 8591), and Paracyathus sp. b. (from loc. USGS 8591). Only specimens of the first two taxa have been located at the NMNH, and were judged to be inade- quately preserved to identify as to genus. Comparisons. — Paracyathus sinuosus, n. sp. is dis- tinguished from P. henekeni (Duncan, 1863) by its shape, being ceratoid and attached instead of trochoid and free. Additionally, P. sinuosus can be distinguished by its sinuous costae and its inequality of S1 and S2. It 1s distinguished from Paracyathus, n. sp. of Frost and Langenheim (1974) from the lower Miocene of Mexico by its smaller size, fewer septa, and differently- ornamented costae. Unfortunately, the Mexican species was based on only one poorly-preserved specimen. Among the three European Miocene species of Para- cyathus reported by Chevalier (1961), P. sinuosus is most similar to P. turonensis Milne-Edwards and Haime, 1848b. P. turonensis, however, has a broad DOMINICAN REPUBLIC NEOGENE. 5: CAIRNS AND WELLS 35 encrusting base, an almost cylindrical corallum, and broad, straight costae. Otherwise the calicular details are very similar. The only Recent species known from the western Atlantic, P. pulchellus (Philippi, 1842), is easily dis- tinguished by its greater corallum size, larger number of septa, and broader pedicel. Occurrence. — Río Gurabo (Mao Formation) middle Pliocene (Text-fig. 3): locs. NMB 15832, 15833. Río Cana (Cercado Formation) lower Miocene (Text-fig. 3): locs. NMB 17023, 17024. Distribution. — Known only from the late Miocene to middle Pliocene of the Dominican Republic from Río Gurabo and Río Cana. Genus DELTOCYATHUS Milne-Edwards and Haime, 1848b Diagnosis.—Solitary; discoidal to patellate; free. Costae present. Pali before all but last cycle of septa; inner edges of P3 join adjacent P2 near columella, forming deltas. Columella papillose. Type-species. — Turbinolia italica Michelotti, 1838, by monotypy. Distribution. — Eocene to Recent, cosmopolitan. Deltocyathus italicus (Michelotti, 1838) Plate 9, figures 10-12 Turbinolia italica Michelotti, 1838, p. 51, pl. 1, fig. 8. Deltocyathus italicus (Michelotti). Milne-Edwards and Haime, 1848b, pp. 326, pl. 10, fig. 11; Vaughan and Woodring, 1921, p. 147; Felix, 1927, pp. 440-441; Felix, 1929, p. 581; Chevalier, 1961, pp. 328-330 (synonymy); Chevalier, 1962, pp. 40-41. ?Deltocyathus sp. cf. D. italicus (Michelotti). Cairns, 1979, pp. 95- 97, pl. 17, figs. 1-3. ?Deltocyathus conicus Zibrowius, 1980, pp. 83-85, pl. 39, figs. A-L. Description of Dominican Republic specimen (USNM 63125 from loc. USGS 8702). — Corallum patellate: 7.6 mm in calicular diameter and 4.7 mm in height (basal angle 78°). Forty-eight costae: С1+2 equal in size, moderately ridged, and bear coarse granules. These 12 costae reach center of base. СЗ +4 progressively small- er and do not attain center of base. Forty-eight hexamerally-arranged septa present but most calicular details are obscured by sediment. Only tips of some pali and columellar elements can be seen (PL 9, fig. 10). Diagnosis. — Corallum conical (patellate), usually 10— 11 mm in diameter; costae ridged, bearing coarse gran- ules. Type-material. — According to Chevalier (1961), Michelotti's holotype is lost. Type-locality. – Tortona, Italy, Miocene (see Zibro- wius, 1980, p. 85). Material. — USNM 63125 (1) from loc. USGS 8702. Reference material of European D. italicus (Pl. 9, fig. 12): MNHNP 160 (2) from Tortona, Italy. Reference material of Caribbean Recent species (Cairns, 1979). Remarks.—No additional specimens of Deltocy- athus from the Caribbean Tertiary were examined; however, the specimen listed by Vaughan and Wood- ring (1921) was found in the collections of the NMNH (USNM 63125 from loc. USGS 8702), compared to typical European specimens, and found to be conspe- cific. However, the identity of the amphi-Atlantic Re- cent species (D. sp. cf. D. italicus (Michelotti) of Cairns, 1979 [see Pl. 9, fig. 13, herein] and D. conicus Zibro- wius, 1980) is still a matter of debate. The major dif- ference between the fossil and Recent forms is that the former has a coarser costal granulation, which led Zi- browius to describe a new species for the Recent spec- imens. The crux of the problem is that hundreds of Recent specimens are available for study, showing it to be variable in many characteristics (i.e., corallum size, palar and columellar shape, and costal ornamen- tation), whereas there are relatively few Miocene spec- imens available for the study of variation. We therefore doubt whether D. italicus occurs in the Recent as well as the Neogene. Comparisons. —No other fossil species of Deltocy- athus are known from the Caribbean, and Chevalier (1961) reported only D. italicus from the Miocene of the western Mediterranean. There are, however, six Recent species known from the Caribbean (Cairns, 1979). D. italicus is distinguished from these by its strongly conical base and its ridged costae. Occurrence. —Río Yaque del Norte (?Baitoa For- mation) early Miocene: loc. USGS 8702. Distribution. — Miocene: central and southern Eu- rope (Felix, 1927; Chevalier, 1961), Morocco, ?Aus- tralia (see Zibrowius, 1980, p. 85), Dominican Repub- lic; Pliocene: Modena, Italy (Felix, 1929); ?Recent: western Atlantic, 403-2634 m (Cairns, 1979) and east- ern Atlantic, 1500-2300 m (Zibrowius, 1980 as D. conicus). Subfamily TURBINOLIINAE Milne-Edwards and Haime, 1848b Genus SPHENOTROCHUS Milne-Edwards and Haime, 1848b Diagnosis.—Solitary and small; cuneiform; free. Costae long, prominent ridges or reduced to linear granules. Columella lamellar. Pali absent. Subgenus EUSTHENOTROCHUS Wells, 1935 Diagnosis. — Sphenotrochus in which the costae have been reduced to rows of granules and/or short carinae. Type-species. — Sphenotrochus moseri Wells, 1935 (= 5. gilchristi Gardiner, 1904), by original designa- tion. 36 BULLETIN 328 Distribution. —Eocene to Recent of Europe, West In- dies, and Indian Ocean. Sphenotrochus (Eusthenotrochus) senni Wells, 1945 Plate 9, figures 14-19 Sphenotrochus n. sp. of Vaughan, 1919, p. 212; Vaughan and Hoff- meister, 1926, p. 114. Sphenotrochus (Eusthenotrochus) senni Wells, 1945, p. 17, pl. 3, figs. 25, 34-35. ?Sphenotrochus auritus var. of Pourtalés. Lindström, 1877, pp. 11- 12, pl. 2, figs. 21-22. Description of Dominican Republic specimens. — Corallum cuneiform, highly compressed with a gently rounded base. Largest of three specimens 5.1 x 3.0 mm in calicular diameter and 7.0 mm in height. Five to seven costae on center of each lateral thecal face are continuous ridges, up to 1.9 mm long, occupying mid- dle third of each thecal face. In upper third of corallum these costae become double rows of short carinae and granules 0.2-0.4 mm long. In lower third of corallum, approximately 1.8 mm above base, the longitudinally- ridged costae become slightly constricted (Pl. 9, fig. 16) and re-radiate basally into short ridges. All other costae (i.e., those on either side of the elongate costae) com- posed of short carinae and granules, much like those of distal third of ridged costae. Commonly two rows of granules per septum near calicular edge. Septa hexamerally-arranged in three cycles: 51=52> S3. Six ofthe 12 large septa, those originating from lateral faces, reach lamellar columella. Diagnostic characters. — Distinctive costal morphol- ogy, composed of ridges in center third of lateral faces and a linear arrangement of granules for upper and lower third of faces. Type-material.—USNM 68358 (holotype): Height — 8.9 mm, calicular diameter — 6.2 x 4.4 mm. Type-locality. — Martinique, Bassignac tuffs, mid- dle Miocene. Material. —Two unnumbered specimens from loc. NMB 15900; NMB D 5891 (1) from loc. NMB 16938; USNM 68358 (holotype). Reference material: USNM 68407, Sphenotrochus n. sp. of Vaughan (1919); 25. auritus var. Pourtalés of Lindstróm (1877), Naturhis- toriska Riksmuseet, Stockholm, specimen 120. Remarks. —The Dominican Republic specimens dif- fer slightly from the type specimen in having several distinctly-ridged costae on the middle third of each lateral thecal face. The holotype has only one elongate costa 1.2 mm long on one face. Wells (1945) reported three rows of granules per septum at the calicular edge. The Dominican Republic specimens have only two rows of granules per septum and a reevaluation of the holotype shows this character to be equivocal because of poor preservation. Comparisons. — Sphenotrochus (E.) senni is similar to S. (E.) brassensis Vaughan in Vaughan and Hoff- meister, 1925, the latter known from the middle Mio- cene of Trinidad. Unfortunately, the type and only known specimen of S. brassensis is too small and dam- aged to allow a proper comparison. The specimens reported by Lindstróm (1877) as Sphenotrochus auritus var. Pourtalés from the Recent of the Virgin Islands are also very similar to S. senni, particularly in corallum shape and costal ornamenta- tion. Lindstróm's specimens are not S. auritus Pour- talés, 1874, but, instead, represent either a new species or 5. (E.) senni in the Recent. The unidentified Sphenotrochus specimen alluded to by Vaughan (1919), Vaughan and Hoffmeister (1926), and Wells (1945) from the Bowden Formation of Ja- maica (USNM 68407) was reexamined and is consid- ered to be 5. (E.) senni. Occurrence. — Río Gurabo (Cercado Formation) late Miocene (Text-fig. 3): loc. NMB 15900. Río Yaque del Norte (?Baitoa Formation) early Miocene: loc. NMB 16936. Distribution. — Miocene: Martinique, Dominican Republic; late Pliocene: Jamaica (Vaughan, 1919); ?Recent: Virgin Islands (Lindstróm, 1877 as S. auritus var. Pourtalés). Genus DOMINICOTROCHUS VWells, 1937 Diagnosis. —Solitary; cuneiform; free. Septotheca costate; principal costae alate. Lacking columella, pali, and endotheca. T ype-species. —Smilotrochus dominicensis Vaughan in Vaughan and Hoffmeister, 1925. Distribution. — Miocene of West Indies. Dominicotrochus dominicensis (Vaughan, 1925) Plate 9, figures 20, 21 Smilotrochus ? dominicensis Vaughan in Vaughan and Hoffmeister, 1925, p. 316, pl. 1, figs. 1—2; Felix, 1927, p. 401. Dominocotrochus dominicensis (Vaughan). Wells, 1937, p. 14; Wells, 1945, pp. 17-18, pl. 3, figs. 26-33; Wells, 1956, p. F426; ?Frost and Langenheim, 1974, p. 304, pl. 117, figs. 5-8. Diagnosis. — Corallum cuneiform and free, with alate lateral edge costae. Coralla up to 6.5 x 10.0 mm in calicular diameter and 13 mm in height. Costae dis- tinct. Septa hexamerally arranged in five incomplete cycles: S1—S2. Fossa deep. Type-material. — MCZ 9266 (holotype) missing from the MCZ. Type-locality. — Dominican Republic, without spe- cific locality, ?Miocene. Material. —Eight specimens from Martinique re- ported by Wells (1945): USNM 68359. DOMINICAN REPUBLIC NEOGENE. 5: CAIRNS AND WELLS 37 Remarks. — Frost and Langenheim (1974) reported D. dominicensis from the lower Miocene of Mexico. Although we did not examine these three specimens, we query their identification based on the apparent poor preservation of the specimens as illustrated and described. Comparisons. — Dominicotrochus is a monotypic ge- nus within the Turbinoliinae characterized by lacking a columella. Occurrence. — The single record known from the Do- minican Republic (the holotype), was reported without specific locality. Distribution. —?Lower Miocene: Mexico (Frost and Langenheim, 1974); middle Miocene: Martinique (Wells, 1945); Miocene: Dominican Republic. Subfamily PARASMILIINAE Vaughan and Wells, 1943 Genus ASTEROSMILIA Duncan, 1867 Diagnosis. —Solitary; ceratoid, trochoid, or flabel- late; free or attached. Paliform lobes before penulti- mate and sometimes antepenultimate cycle of septa. Columella lamellar or fascicular. Endotheca abundant. Type-species.— Trochocyathus abnormalis Duncan, 1864, by subsequent designation (Vaughan, 1919). Remarks. — The species of Asterosmilia discussed here are ordered as follows: the type species, followed by the other species found in order of decreasing sim- ilarity to the type species. Distribution. — Oligocene to Recent of West Indies. Asterosmilia abnormalis (Duncan, 1864) Plate 10, figures 1-4, 6 Trochocyathus abnormalis Duncan, 1864, p. 26, pl. 2, figs. 4а-с. Asterosmilia anomala (Duncan) [sic]. Duncan, 1867, p. 653, pl. 32, figs. 3a-d; Duncan, 1868, pp. 16, 23; Pourtalés, 1875, p. 545. Asterosmilia cornuta Duncan, 1867, p. 653, pl. 32, fig. 4; Duncan, 1868, pp. 16, 23; Vaughan, 1919, pp. 213, 354. Asterosmilia coronata Duncan. Pourtalés, 1875, p. 545. Asterosmilia abnormalis (Duncan). Vaughan, 1919, pp. 213, 354; Vaughan and Woodring, 1921, p. 133; Vaughan and Hoffmeister, 1925, p. 322; Felix, 1927, p. 312; Frost and Langenheim, 1974, PUSO Description. —Corallum ceratoid, slender, and curved 10*-100? in plane of greater calicular axis. Corallum compressed, the compression most apparent in basal рагі. Largest corallum examined (USNM 546428 from loc. USGS 8516) 46 mm in height; largest calice (in largest corallum), 18.7 x 14.8 mm in diameter. Theca light reddish-brown. Pedicel very narrow, 1.0-1.3 mm in diameter. C1+2 prominent, the C1 on convex thecal edge often forming a crest up to 2 mm in height and 10 mm long on basal one-third to one-fourth of larger coralla. Less frequently the opposing СТ on concave thecal edge is also ridged. C3-5 low and smooth. Septa hexamerally arranged in five incomplete cycles. $1 +2 equal in size and moderately exsert. Higher cycle septa progressively smaller and less exsert. Largest cor- allum has 72 septa. According to Duncan (1864), paliform lobes occur before all but last cycle of septa; however, none of the specimens available to us are preserved well enough to confirm this observation. Columella commonly la- mellar but may be spongy. Lower half of corallum solidly filled with stereome. Upper half filled with obliquely-oriented dissepiments. Diagnostic characters. — Well-developed, alate costa on convex thecal edge; paliform lobes before S1-3. Type- material. — Duncan (1864) did not specify how many specimens formed the base of his description of A. abnormalis, only that he had two or more. Six spec- imens referable to this species are deposited at the BM(NH) [BM(NH) 28801-28806], all of which are considered as syntypes. Duncan's (1864, pl. 2, fig. 4) illustrated specimen is BM(NH) 28801. The holotype of A. cornuta is also deposited at the BM(NH) [BM(NH) 28807]. Type-locality. — The type-locality of both 4. abnor- malis and A. cornuta is the **Nivaje Shale” of Heneken, Dominican Republic, Miocene. Material. — UÜnnumbered ММВ specimens from locs. NMB 15806 (1); 15814 (1); 16868 (1); 16910 (19); USNM. 546428 (73) from loc. USGS 8516; USNM 546427 (44) from loc. USGS 8519; unnumbered TU specimens from loc. TU 1219 (35); loc. TU 1380 (8); loc. TU 1411 (1); BM(NH) 28801-28806 (syntypes of A. abnormalis), BM(NH) 28807 (holotype of A. сог- nuta). Reference material: USNM 65323 (1) from loc. USGS 8321 (Costa Rica). Remarks. — Ordinarily the columella type is a con- servative character at the species level and even often at the generic level; however, the columella of A. ab- normalis may be either lamellar or spongy. A similar kind of variation was noted by Cairns (1979) for the Recent species А. prolifera (Pourtalés, 1871) [see Pl. 10, fig. 5 herein]. In an examination of over 1800 spec- imens of A. prolifera, Cairns found extreme variation in columellar and palar shape; in rare cases columella and pali were missing. This range of variation found in a congener gives support to identifying specimens that have either a lamellar or spongy columella as A. abnormalis. The lateral-edge costal crests of Duncan's illustrated Specimen (1864, pl. 2, fig. 4a) are grossly exaggerated. There is, in fact, no costal ridge on the concave thecal edge, and the convex edge crest is much smaller than illustrated (Pl. 10, figs. 2-3). Comparisons. — The long, curving ceratoid corallum with an alate costa on the convex thecal edge distin- guishes А. abnormalis from the other nine species of 38 BULLETIN 328 Table 3.— Characters of the 10 known species of Asterosmilia. pedicel septal corallum shape and size nature of costae diameter cycles? A. abnormalis (Duncan, 1864) ceratoid, curved; medium lateral costae alate (esp. narrow 54+ concave side) A. exarata Duncan, 1867 ceratoid, curved; small C1+2>C3+4 narrow S4 A. marchadi (Chevalier, 1966) ceratoid; small С1+2>С3+4 narrow S4* A. profunda (Duncan, 1864) trochoid, curved; large С1+2 slightly ridged thick S5 A. duncani! Vaughan in Vaughan and calice constricted in center, equal, all slightly ridged narrow 55" Hoffmeister, 1925 straight; medium A. compressa? Vaughan in Vaughan flabellate, straight; medium alate edge costae narrow S5* and Hoffmeister, 1925 A. machapooriensis Hoffmeister in ceratoid, curved; medium C1-3 slightly ridged narrow S4* Vaughan and Hoffmeister, 1926 A. trinitati? Vaughan in Vaughan and flabellate, straight; medium alate edge costae thick S4* Hoffmeister, 1926 A. pourtalesi* Duncan, 1873 ceratoid to sub-cylindrical, C1-3>C4 unknown S4* curved; medium A. prolifera (Pourtalés, 1871) ceratoid to trochoid, curved; equal, low narrow 54 small to medium " Based only on the holotype; ? Based only on the four syntypes; ? Based on few poorly preserved specimens (the types). Generic attribution doubtful; * Not seen; 5 A plus sign means an incomplete cycle of the next higher numbered septal cycle. Asterosmilia (Table 3). It is morphologically very sim- ilar to A. prolifera, particularly with regard to corallum size and shape. A. prolifera differs in having equal and nonalate costae, dissepiments extending to the base of long coralla, and a less compressed corallum. Occurrence. — Río Gurabo (Gurabo Formation) late Miocene to early Pliocene (Text-fig. 3): locs. NMB 15806, 15814. Río Cana (Gurabo Formation) early Pliocene (Text-fig. 3): loc. NMB 16868. Río Mao QGurabo Formation) late Miocene: loc. NMB 16910, loc. USGS 8519. Río Amina: loc. USGS 8516, loc. TU 1219. “Nivaje Shale”, South of Santiago de los Ca- balleros (?Gurabo Formation): loc. TU 1380. Río Gua- najuma (?Gurabo Formation): loc. TU 1411. Distribution. — Known only from the late Miocene to early Pliocene of the Dominican Republic from Río Gurabo, Río Mao, Río Amina, “Nivaje Shale", and Río Guanajuma. Asterosmilia exarata Duncan, 1867 Plate 10, figures 7-9, 11 Asterosmilia exarata Duncan, 1867, p. 653, pl. 32, fig. 5; Duncan, 1868, pp. 16, 23; Pourtalés, 1875, p. 545; Vaughan, 1919, pp. 213, 354; Vaughan and Hoffmeister, 1925, p. 322; Felix, 1927, p. 302, Asterosmilia hilli Vaughan, 1919, pp. 212, 228-229, 355-356, pl. 80, figs. 4—6; Vaughan and Woodring, 1921, pp. 99, 133; Vaughan and Hoffmeister, 1925, p. 322; ?Felix, 1927, p. 213; ?Frost and Langenheim, 1974, pp. 304—305, pl. 117, figs. 1-2. ?Asterosmilia exarata var. Vaughan, 1919, рр. 200, 218. Not Asterosmilia exarata Duncan. Vaughan and Woodring, 1921, р. 133 (= 4. profunda and an unidentified species of Asterosmilia). ?Asterosmilia exarata var. robusta Vaughan and Hoffmeister, 1925, p. 324; Felix, 1927, p. 312. ?Asterosmilia sp. cf. А. hilli Vaughan. Wells, 1934, р. 106. Description. — Corallum ceratoid, slender, and curved 45°-90° in plane of greater calicular axis. Corallum slightly compressed, almost round in cross-section. Holotype [BM(NH) 28944], one of the largest coralla of this species examined, 18.1 mm in height and 9.6 x 8.8 mm in calicular diameter. Pedicel narrow, about 1.4 mm in diameter. С1-2 slightly more prominent than C3-4, but all costae distinct, convex, granulated ridges. No alate costae. Septa hexamerally arranged in four cycles, the fourth cycle complete at a ged of about 9.5 mm. S1+2 equal in size and exsertness, the higher cycle septa progres- sively smaller. A prominent paliform lobe occurs be- fore each S3, forming a crown of 12 elements in large, well-preserved specimens. Paliform lobes broad (about 1.45 mm wide) and separated from their adjacent S3 by a deep and wide notch. Columella usually lamellar, often with two or three deep indentations; however, some specimens have trabecular columellas, showing the same variation found in A. abnormalis and A. pro- lifera. Endothecal dissepiments present (Pl. 10, fig. 11) but sparse. Diagnostic characters. — Corallum small; never more than four cycles of septa. Type-material. — BM(NH) 28944 (holotype of A. ex- arata), USNM 324815 (7 syntypes of A. hilli), USNM 324816 (3 syntypes of A. hilli). Type-locality. — A. exarata: “Nivaje Shale", Domin- ican Republic, Miocene. А. hilli: loc. USGS 2580, Ja- maica (Bowden Formation) late Pliocene. Material. -USNM 546422 (17) from loc. USGS 8446; USNM 546411 (7) from loc. USGS 8733, USNM 324815 (7 syntypes of A. ЛИШ), USNM 324816 (3 syn- types of A. hilli); BM(NH) 28944 (holotype of A. ex- DOMINICAN REPUBLIC NEOGENE. 5: CAIRNS AND WELLS 39 Table 3. — Continued. paliform lobe cycles nature of columella endothecal dissepiments distribution and age P1-3? lamellar and spongy abundant, oblique, stereome present Dominican Republic (late Miocene to early Pliocene) P3 lamellar and trabecular sparse ?Barbados (Eocene) ?Antigua (late Oligocene) ?Mexico (Miocene) Costa Rica, Dominican Republic (Miocene) Jamaica (Pliocene) P3 crispate abundant Atlantic (Recent: 32-229 m) P3+4 lamellar and fascicular abundant, no stereome Jamaica (Miocene) Dominican Republic (late Miocene to middle Pliocene) P4? linear—papillose abundant, stereome present Dominican Republic (?Miocene) P4 lamellar sparse Dominican Republic (?Miocene) P1-2 spongy abundant Trinidad (Miocene) P3 or P4 lamellar sparse Trinidad (Miocene) Y lamellar present St. Barthelemy (Oligocene) P3 papillose and lamellar abundant, stereome present Atlantic (Recent: 32-311 т) arata); USNM 44303 [А. sp. cf. A. hilli of Wells (1934). Reference specimens: USNM 63329 (43) from loc. USGS 2580 (Jamaica); USNM 63129 (14) from loc. USGS 20468 (Costa Rica); USNM 324814 (14) from Limon, Costa Rica, Pittier 4618; USNM 63611 (1) [A. exarata var. “robusta”] from loc. USGS 6881 (upper Oligocene, Antigua). Remarks. — The specimen reported by Wells (1934) as Asterosmilia sp. cf. A. hilli from the Eocene of Bar- bados (Pl. 10, fig. 10) is a poorly-preserved fragment missing its calicular features. Its size and costae are consistent with 4. exarata, but a definitive identifi- cation is impossible. The single, poorly-preserved, immature specimen of A. hilli reported by Frost and Langenheim (1974) from the lower Miocene of Mexico is also highly suspect on the basis of its poor preservation and the presence of an epitheca; however, this specimen was not examined by either author. Vaughan and Hoffmeister (1925) alluded to a variety of А. exarata that Vaughan called robusta, but that variety was never described or figured and therefore must be considered a nomen nudum. 'This is undoubt- edly the unnamed variety of A. exarata referred to by Vaughan (1919, pp. 200, 218) from Antigua and the Dominican Republic. According to Vaughan and Hoff- meister (1925), the form occurred together with А. dun- cani Vaughan, 1925 in the blue clay of the Dominican Republic. A specimen labelled as such in Vaughan's handwriting is deposited at the NMNH (USNM 63611, from loc. USGS 6811, Willoughby Bay, Antigua). It is not conspecific with А. exarata and is too poorly pre- served to properly identify. No specimen of variety robusta was found in the NMNH collections from the Dominican Republic. Comparisons. — Asterosmilia exarata is extremely similar to the Recent species A. marchadi (Chevalier, 1966) (Table 3). The only differences noted are that А. exarata sometimes has a lamellar columella and has equal-sized S1--2, whereas А. marchadi has а crispate columella and its S1 are distinctly larger than its S2. A poorly-preserved specimen, especially one lacking calicular details, could easily be confused with Troch- ocyathus chevalieri, n. sp., based on similarity of size, shape, number of septa and costae. Nonetheless, on closer examination one would find that T. chevalieri has twice as many pali (Р 1 +2 as well as P3), a papillose columella, and no endothecal dissepiments. Occurrence. — Río Mao (?Cercado Formation) late Miocene: loc. USGS 8733. Río Gurabo: loc. USGS 8446. Distribution. —?Eocene: Barbados (Wells, 1934, as A. Вий); ?аррег Oligocene: Antigua (Vaughan, 1919, рр. 200, 218 as А. exarata var.); ?early Miocene: Mex- ico (Frost and Langenheim, 1974); Miocene: Costa Rica (Vaughan, 1919, as A. hilli), late Miocene: Dominican Republic; late Pliocene: Jamaica (Vaughan, 1919, as A. hilli). Asterosmilia profunda (Duncan, 1864) Plate 10, figures 12-18 Trochocyathus profundus Duncan, 1864, p. 26, pl. 5, figs. За-с. Asterosmilia exarata Duncan. Vaughan and Woodring, 1921, p. 133 (in part: USNM 546415, 546416, 546417, 546424). Asterosmilia profunda (Duncan). ?Vaughan and Woodring, 1921, pp. 99, 133; Vaughan and Hoffmeister, 1925, p. 322; Vaughan and Hoffmeister, 1926, p. 117; Felix, 1927, pp. 312-313. Description. — Corallum trochoid, massive, and often irregularly curved in basal region. Largest corallum examined (USNM 63327 from loc. USGS 2580) 90.0 40 BULLETIN 328 mm long and 27 x 22 mm in calicular diameter. Theca light reddish-brown. Pedicel thick, sometimes attached to substrate, 3.3-5.8 mm in diameter. С1-2 often slightly ridged but no alate costae present. C3-5 flat and equal in width, covered by small granules. Theca smooth and sometimes porcelaneous. Septa hexamerally arranged in five cycles: S1=S2>S3>S4>S5. Fifth cycle usually complete at a gcd of about 25 mm. Paliform lobes before penultimate (S4) and ante- penultimate (S3) cycles of septa. As in most other species of Asterosmilia, the columella is variable: either fascicular (e.g., Bowden specimens) or lamellar (e.g., most of the Dominican Republic specimens). Fossa deep. Vesicular dissepiments abundant, extending to base of corallum. Diagnostic characters. — Large, robust, trochoid cor- allum; thick, attached pedicel. Type-material. —BM(NH) 40413 (holotype): height — 38.8 mm, 25.7 x 19.8 mm in calicular diameter. Type-locality. — Jamaican “Miocene”. Material. — Unnumbered ММВ specimens from locs. NMB 15832 (1); 15833 (7); USNM 546417 (3) from loc. USGS 8527; USNM 546424 (2) from loc. USGS 8528; USNM 546415 (1) from loc. USGS 8556; USNM 546416 (3) from loc. USGS 8733; unnumbered TU specimens from loc. TU 1208 (2); loc. TU 1225 (7); loc. TU 1352 (14); loc. TU 1366 (1); loc. TU 1451 (2); loc. TU 1453 (1); BM(NH) 40413 (holotype of 7. pro- fundus). Reference specimens: USNM 63327 (10) from loc. USGS 2580 (Jamaica); USNM 64213 (3) from loc. USGS 8421, Eocene of Panama. Remarks. — The specimen reported by Vaughan and Woodring (1921, p. 133) as A. profunda from loc. USGS 8733 is present at the NMNH (USNM 546412) but is too small to identify. Other specimens from the same station identified by Vaughan and Woodring (1921, p. 133) as A. exarata Duncan, 1867 are A. profunda. Comparisons. — The massive trochoid corallum and relatively thick pedicel serve to distinguish A. profunda from its congeners. Although Vaughan and Hoffmeis- ter (1926) compared it to А. machapooriensis Ноћ- meister in Vaughan and Hoffmeister, 1926, we feel that, of all the species of Asterosmilia, 11 most resem- bles 4. duncani Vaughan, 1925, but can be distin- guished by a number of characters (Table 3). Occurrence. —Río Gurabo (Mao and Gurabo for- mations) early to middle Pliocene (Text-fig. 3): locs. NMB 15832, 15833, locs. USGS 8550, 8556,1ocs. TU 1352, 1366. Río Mao (?Gurabo Formation) late Mio- cene: locs. USGS 8527, 8528, 8733, loc. TU 1453. “Nivaje Shale”. Arroyo Babosico (?Gurabo Forma- tion): loc. TU 1451. Distribution. — Eocene: Panama (reported herein); Miocene: Jamaica (Duncan, 1864); late Miocene to middle Pliocene: Dominican Republic. Asterosmilia duncani Vaughan, 1925 Plate 10, figures 19, 20 Asterosmilia n. sp. of Vaughan, 1919, p. 354 (one of two "additional species"). Asterosmilia duncani Vaughan in Vaughan and Hoffmeister, 1925, pp. 323-324, pl. 3, figs. 1-2; Felix, 1927, p. 312. Description based on holotype (MCZ 9277). — Сог- allum compressed, straight, and free; 26.1 x 19.0 mm in calicular diameter and 33.5 mm in height. Calice slightly constricted in center, about 18 mm in lcd. Lat- eral calicular edges rounded; no alate costae. Costae thin and ridged. Ninety-two costae and septa present but more detailed observations of symmetry and num- ber of cycles not possible because of poor preservation. According to Vaughan and Hoffmeister (1925), pali- form lobes present before penultimate septal cycle (usually S4). Columella well developed, composed of a row of interconnected papillae. Dissepiments abun- dant; stereome in base of corallum. Type-material. —MCZ 9277 (holotype). Type-locality. — Dominican Republic (Gabb Collec- tion), lower Miocene. Material. —We have examined the holotype (MCZ 9277), which is the only known specimen of this species. Remarks. —There was no evidence of paliform lobes in the holotype. Comparisons. — Asterosmilia duncani is distin- guished from other species of Asterosmilia by its com- pressed corallum (constricted in center) and relatively high number of septa. However, comparisons based on a single damaged specimen are inadequate. Occurrence. — Known only from the type-locality of “Dominican Republic”, ?lower Miocene. Distribution. — Known only from the ?lower Mio- cene of the Dominican Republic. Asterosmilia compressa Vaughan, 1925 Plate 10, figures 21-24 Asterosmilia n. sp. of Vaughan, 1919, p. 354 (one of two “additional species”). Asterosmilia compressa Vaughan in Vaughan and Hoffmeister, 1925, pp. 322-323, pl. 3, figs. 3-4; Vaughan and Hoffmeister, 1926, p. 117; Felix, 1927, p. 312. Diagnosis based on the syntypes. — Corallum strongly compressed, straight, and free. Pedicel 2-3 mm in di- ameter. Largest of four syntypes (specimen 4 of Vaughan and Hoffmeister, 1925) 10.8 x 25.3 mm in calicular diameter and 35.4 mm in height. Costae low and distinct, every fourth costa (i.e., C1-3) slightly more prominent; two principal costae alate. Septa hex- amerally arranged in five complete cycles plus pairs of DOMINICAN REPUBLIC NEOGENE. 5: CAIRNS AND WELLS 41 56 even at a вед of 20 mm; largest specimen with 135 septa. Small paliform lobes before penultimate septal cycle (usually S4). Columella lamellar. Dissepiments sparse. Type-material. — MCZ 9273-9276 (four syntypes). Type-locality. — Dominican Republic (Gabb Collec- tion), ?lower Miocene. Material. —We have examined the four syntypes (MCZ 9273-9276); no additional specimens are known of this species. Comparisons. — Asterosmilia compressa is similar to A. trinitatis Vaughan in Vaughan and Hoffmeister, 1926, in size, shape, and costal morphology, as pre- viously noted by Vaughan and Hoffmeister (1926). It 15 distinguished by having a narrower pedicel and а , greater number of septa at a corresponding gcd (Table 3). Occurrence. — Known only from the type-locality of “Dominican Republic", lower Miocene. Distribution. — Known only from the ?lower Mio- cene of the Dominican Republic. Superfamily FLABELLICAE Bourne, 1905 Family FLABELLIDAE Bourne, 1905 Genus FLABELLUM Lesson, 1831 Diagnosis.—Solitary; cuneiform, compressed-tur- binate, or ceratoid; free. Wall epithecal. Base not thick- ened by stereome; roots absent. Pali absent. Columella rudimentary or absent. Type-species. — Flabellum pavoninum Lesson, 1831, by monotypy. Distribution. — Eocene to Recent, cosmopolitan. ?Flabellum species Plate 11, figures 1, 2 Flabellum b Vaughan and Woodring, 1921, p. 133. Diagnosis. — Corallum flabellate, largest specimen (USNM 64202 from loc. USGS 8733) 20.8 x 13.9 mm in calicular diameter, 29.1 mm in height, and 1.5 mm in pedicel diameter. C1—2 highly ridged, especially two principal costae, which project up to 4 mm. С1 extend to pedicel; C2 about three-quarters that dis- tance. Corallum epithecate, covered by transverse ep- ithecal bands. Septa hexamerally arranged in four cycles; some pairs of S5 in lateral edge half-systems. Details of septa and fossa not preserved. Material. — USNM 64202 (2) from loc. USGS 8733; 2 unnumbered TU specimens from loc. TU 1225. Ref- erence material: USNM 155290 and 155288 (8), F. roissyanum Milne-Edwards and Haime, 1848b from Miocene of Austria. Remarks. —'This taxon is quite different from any other species described from the Dominican Republic but because its preservation does not allow an exam- ination of the calicular details, the generic attribution 15 tentative. Five species of Flabellum have been reported from the Dominican Republic: F. dubium Duncan, 1863; F. exaratum Duncan and Wall, 1865 (by Duncan, 1868, p. 16); Flabellum, n. sp. sensu Duncan (1863, pp. 411, 430); and Flabellum a and b sensu Vaughan and Woodring (1921). Flabellum dubium was transferred to Antillophyllia Vaughan, 1932 by Vaughan (1932, p. 509). After examination of the type of F. exaratum, we concur with Vaughan (1919, p. 213) that the spec- imen does not belong to Flabellum, but probably to a group of species ascribed to the genus Placocyathus (see discussion of Antillocyathus). Flabellum, n. sp. of Duncan (1863) was based on casts that do not allow a definitive determination. Flabellum a of Vaughan and Woodring (1921) was determined to be Pourtalocy- athus hispidus (Pourtalés, 1878) [this paper]. Finally, Vaughan and Woodring's Flabellum b, the specimen figured and diagnosed in this account, is also of doubt- ful generic identity. Thus, ofthe five records, no species of Flabellum are definitely known to occur in the Do- minican Republic. In corallum shape and costal characteristics the four Dominican Republic specimens resemble F. roissya- num, known from the Miocene to Pliocene of southern Europe (see Kojumdgieva and Strachimirov, 1960). Occurrence. —Río Mao (?Gurabo Formation) late Miocene: loc. USGS 8733. Río Mao (?Gurabo For- mation): loc. TU 1225. Family GUYNIIDAE Hickson, 1910 Genus POURTALOCYATHUS Cairns, 1979 Diagnosis. —Solitary and small; ceratoid; free. Wall epithecal, often with one row of mural spots (differ- ential calcification) in every interseptal space. Three cycles of septa; small paliform lobes before S2. Col- umella papillose. Туре-зрес!ез.- Ceratotrochus hispidus Pourtalés, 1878, by original designation. Distribution. — Miocene to Recent, West Indies. Pourtalocyathus hispidus (Pourtalés, 1878) Plate 11, figures 3-7, 10-11 Ceratotrochus hispidus Pourtalés, 1878, p. 202, pl. 1, figs. 19-20. Flabellum a of Vaughan and Woodring, 1921, p. 133. Pourtalocyathus hispidus (Pourtalés). Cairns, 1979, pp. 171-172, pl. 33, figs. 3-8 (includes synonymy). Description. — Corallum ceratoid, straight to slightly curved, and small. Typical corallum 8-9 mm in height and about 4.2 mm in calicular diameter; calice round. Coralla often attached by a relatively thick pedicel about 1.2 mm in diameter; base of pedicel reveals six primary 42 BULLETIN 328 septa. Costae equal, slightly convex, and smooth. Fine concentric epithecal bands encircle the theca. In some coralla, rows of round to elongate mural pores about 0.25 mm in diameter occur in every interseptal space (PI. 11, figs. 10, 11). Septa hexamerally arranged in three complete cycles (24 septa). S1 highly exsert and very thick (up to 0.26 mm). S2-3 much less exsert and progressively smaller in size and thickness (about 0.07 mm thick). Only S1 reach columella. Granules up to 68 um in height (Pl. 11, fig. 6) occur on all septal faces. Fossa shallow. Small, papillose paliform lobes often before S2. Columella well-developed and papillose, composed of several solid, basally-fused rods. Diagnostic characters. — Mural spots or pores often present; three cycles of septa; S1 very exsert; corallum small, rarely over 5 mm in calicular diameter. Type-material. —MCZ 5583 (holotype). Type-locality. —U. S. Coast Survey Steamer Blake sta. 19: 23*02'N, 83?10"W (western Straits of Florida), 56 m; Recent. Material. —Unnumbered NMB specimens from locs. NMB 15823 (10); 15828 (19); 15829 (3); 15995 (2); 16016 (4); 16023 (1); 16036 (8); 16038 (4); USNM 63040 (2) from loc. USGS 8726; USNM 64200 (2) from loc. USGS 8733; USNM 64201 (5) from loc. USGS 8735; 89 unnumbered TU specimens from loc. TU 1227A; MCZ 5583 (holotype). Reference material: 123 Recent specimens listed by Cairns (1979). Remarks. — The rows of white mural spots corre- sponding to each interseptal space, reported by Cairns (1979) for about one-third of the Recent specimens, are also found in the Dominican Republic fossils with about the same frequency. These spots, which when the specimen is alive have no surface relief, probably represent a periodic differential calcification of the the- ca because in many of the fossil specimens these spots are now pores penetratng the theca. Some Recent specimens have granulated costae, his- pid spines, or imbricate epithecal bands. Dominican Republic specimens have consistently smooth epithe- ca. This is the first fossil record of P. hispidus. The Do- minican Republic Miocene fossils are indistinguish- able from Recent specimens. Comparisons. — Pourtalocyathus is a monotypic ge- nus within the Guyniidae characterized by its papillose columella, large P2, and three cycles of septa. Occurrence. — Rio Gurabo (Mao Formation) early to middle Pliocene (Text-fig. 3): locs. NMB 15823, 15828, 15829, 15995, 16016, 16023, 16036, 16038, loc. USGS 8735. Río Mao (?Gurabo Formation) late Miocene: loc. USGS 8733. Río Yaque del Norte (?Baitoa For- mation) loc. USGS 8726. Arroyo Zalaya (?Gurabo For- mation): loc. TU 1227А. Distribution. — Late Miocene to middle Pliocene: Dominican Republic; Recent: Antilles and off eastern Florida, 349-1200 m (Cairns, 1979). Genus GUYNIA Duncan, 1872 Diagnosis. —Solitary; ceratoid to cylindrical; free or laterally attached. Wall epithecal, often with rows of mural pores between septa. Pali absent. Columella composed of one twisted ribbon. Type-species. — Guynia annulata Duncan, 1872, by monotypy. Distribution. — Eocene to Recent, cosmopolitan. Guynia species cf. G. annulata Duncan, 1872 Plate 11, figures 8, 9, 12, 13 Guynia annulata Duncan, 1872, p. 32, pl. 1, figs. 1-8; Wells, 1973, pp. 59-63, figs. 1-3; Cairns, 1979, pp. 164-165, pl. 32, figs. 1-3 (synonymy); Zibrowius, 1980, pp. 161-163, pl. 83, figs. A-Q; Cairns, 1984, p. 23, pl. 5, figs. А-В. Pyrophyllia inflata Hickson, 1910, pp. 1-7, text-figs. 1—4. Description of Dominican Republic specimens. — Corallum straight and cylindrical; largest specimen (unnumbered TU specimen from loc. TU 1227A) 3.25 mm long and 0.9 mm in calicular diameter. Coralla show no sign of lateral attachment. Concentric epithe- cal growth bands occur about every 0.12-0.15 mm. Longitudinal costae (C1) also present. Prominent spines up to 0.10 mm in height occur at the intersections of costae and epithecal bands. In two specimens mural pores about 0.10 mm in diameter are present. Eight S1. Inner septal edges very sinuous; septal faces smooth. S2, if present, are much lower in fossa than S1 and therefore obscured from view. Columella also obscured. Diagnostic characters. — Very small corallum; octameral symmetry; two septal cycles; no septal gran- ules. Type-material. —BM(NH) 1883.12.10.110-112 (18 syntypes). Type-locality. — Adventure Bank, Mediterranean, 168 m, Recent. Material.—Two unnumbered TU specimens from loc. TU 1227A; USNM 74689 (1) from loc. TU 1227A. Reference material: Recent specimens from the west- ern Atlantic and Hawaii (Cairns, 1979, 1984); BM(NH) 1883.12.10.110-112 (18 syntypes of G. annulata). Remarks. — The Dominican Republic specimens dif- fer from Recent specimens in having spines at the in- tersections of the costae and epithecal bands. But, giv- en the range of variation of the theca described by DOMINICAN REPUBLIC NEOGENE. 5: CAIRNS AND WELLS 43 Cairns (1979, p. 164), this is not considered to be a differentiating character. Comparisons. — Only one other species of Guynia is known: С. adherens (Tenison-Woods, 1878) from Ше Oligocene of Aldinga, Australia. It differs from G. an- nulata in having three cycles of hexamerally-arranged exsert septa, a columella composed of two elements, and a much greater calicular diameter. Occurrence. — Arroyo Zalaya (?Gurabo Formation): loc. TU 1227А. Distribution. — Miocene: Dominican Republic; Re- cent: Caribbean, Gulf of Mexico, Bermuda, Madeira, Azores, Persian Gulf, Hawaiian Islands, 28-653 m (Cairns, 1979, 1984). Suborder DENDROPHYLLIINA Vaughan and Wells, 1943 Family DENDROPHYLLIIDAE Gray, 1847 Genus DENDROPHYLLIA Blainville, 1830 Diagnosis. — Colonial; dendroid or bushy colonies formed by extratentacular budding; free or attached. Synapticulothecate, with well-developed costae. Septa arranged in Pourtalés Plan (see Wells, 1956, p. F341). Columella spongy; pali often present. Endothecal dis- sepiments common. Type-species. — Madrepora ramea Linnaeus, 1758, by subsequent designation (Milne-Edwards and Haime, 1850). Distribution. — Eocene to Recent, cosmopolitan. Dendrophyllia cornucopia Pourtalés, 1871 Plate 11, figures 14-17 Dendrophyllia cornucopia Pourtalés, 1871, p. 45, pl. 5, figs. 7-8; Cairns, 1979, pp. 179-181, pl. 36, figs. 1-4 (synonymy); Zibro- wius, 1980, pp. 175-176, pl. 88, figs. A-L; Hubbard and Wells, 1986, p. 139, figs. 33-35. Dendrophyllia, n. sp. A of Vaughan and Woodring, 1921, p. 135. Balanophyllia sp. of Vaughan and Woodring, 1921, p. 135. Description. — Corallum cylindrical and often slight- ly curved: up to 55 mm long and 14.2 mm in calicular diameter. Synapticulotheca covered by epitheca, which Obscures porosity of corallum. Costae equal in width (about 0.98 mm wide) and flat, separated by shallow grooves about 0.20 mm in width. Costae covered with numerous fine granules, each about 0.10 mm in di- ameter. No buds or bud scars present on any fossil specimens. Septa hexamerally arranged in four cycles, some- times with pairs of S5 in half-systems adjacent to prin- cipal septa. Higher cycle septa arranged in а Pourtalés Plan (see Wells, 1956, p. F341). Ра lacking. Columella massive, composed of an elliptical spongy mass of tra- beculae. Thin horizontal dissepiments, each traversing one to four interseptal loci, are common. Diagnostic characters. — Recumbent, subcylindrical corallum; no buds. Type-material. —-MCZ 5442, 2752, syntypes from U. S. Coast Survey Steamer Bibb stations 135 and 173, respectively; unnumbered BM(NH) syntype from Bibb sta. 173. Type-locality. — Off Key West, Florida, 220-229 m, Recent. Material. — Unnumbered NMB specimens from locs. NMB 15806 (3); 15807 (2); 15815 (2); 15836 (2); USNM 64227 (5) from loc. USGS 8528; USNM 64225 (1) from loc. USGS 8545; USNM 64226 (2) from loc. USGS 8733; USNM 63041 (2) from loc. USGS 8735; 2 unnumbered specimens from loc. TU 1210. Refer- ence specimens: specimens listed by Cairns (1979); MCZ 5442, 2752 (syntypes). Remarks. —The only difference noted between the Recent specimens and the fossils from the Dominican Republic is that the latter lack evidence of buds on all specimens examined. Buds and(or) bud scars are com- mon on almost all Recent specimens. Vaughan and Woodring's (1921, p. 135) Dendro- phyllia n. sp. A (USNM 63047 from loc. USGS 8735; USNM 64226 from loc. USGS 8733) and Balano- phyllia sp. (USNM 63041 from loc. USGS 8735) were examined and identified as D. cornucopia, but Vaughan and Woodring's Dendrophyllia n. sp. B from loc. USGS 8546 could not be found in the collections of the NMNH. Comparisons. — Dendrophyllia cornucopia is the only species of Dendrophyllia thus far reported from the Caribbean Miocene. А species of a related genus, Ва/- anophyllia pittieri Vaughan, 1919, is known from the Miocene of Costa Rica. There are two other Recent species of Dendrophyllia from the Caribbean: D. gad- itana (Duncan, 1873) and D. alternata Pourtalés, 1880. They are easily distinguished by their branching pat- terns (Cairns, 1979). Occurrence. —Río Gurabo (Gurabo and Mao for- mations) late Miocene to early Pliocene (Text-fig. 3): locs. NMB 15806, 15807, 15815, 15836, locs. USGS 8545, 8735, loc. TU 1210. Río Mao (?Gurabo For- mation) late Miocene: locs. USGS 8528, 8733. Distribution. — Late Miocene to early Pliocene: Do- minican Republic; Recent: Atlantic Ocean, 132-960 m (Cairns, 1979). NEOGENE PALEONTOLOGY IN THE NORTHERN DOMINICAN REPUBLIC 6. The Phylum Brachiopoda By ALAN LOGAN Department of Geology University of New Brunswick Saint John, New Brunswick, E2L 4L5 CANADA ABSTRACT Four species of articulate brachiopods: Tichosina? lecta (Guppy, 1866); Terebratulina cailleti Crosse, 1865; Argyrotheca johnsoni Cooper, 1934; and Lacazella caribbeanensis Cooper, 1977 have been described and illustrated from the Río Gurabo, Río Cana and Arroyo Zalaya sections of the Neogene of the Dominican Republic. Т.? lecta and Г. caribbeanensis have previously been recorded from the Tertiary of the Caribbean area; A. johnsoni, T. cailleti and L. caribbeanensis from the Recent of the same region. Brachiopod occurrences in the Dominican Republic Neogene generally support the paleoecological reconstruction of Saunders, Jung, and Biju-Duval (1986) of increasingly deeper-water conditions proceeding upsection and downstream in both the Río Gurabo and Río Cana valleys. RESUMEN Cuatro especies de braquiópodos articulados: Tichosina? lecta (Guppy, 1866); Terebratulina cailleti Crosse, 1865; Argyrotheca johnsoni Cooper, 1934; y Lacazella caribbeanensis Cooper, 1977 han sido descritos e illustrados de los secciones del Río Gurabo, Río Cana y Arroyo Zalaya del Neógeno de la República Dominicana. Т.? lecta y Г. caribbeanensis han sido identificados previamente del Terciario del área del Caribe; A. johnsoni, T. cailleti y L. caribbeanensis del Reciente de la misma región. El hallazgo de braquiópodos en el Neógeno de la República Dominicana por lo general soporta las reconstrucciones paleo- ecológicas de Saunders, Jung, y Biju-Duval (1986) de condiciones de aguas más profundas procediendo sección arriba y río abajo en los valles de los ríos Gurabo y Cana. INTRODUCTION Tertiary brachiopods are relatively rare in the Ca- ribbean region compared to the comparative abun- dance and diversity of this group in modern seas in the West Indies (Cooper, 1979). The collections from the Neogene of the Dominican Republic reflect this paucity of fossil taxa, resulting from normal tapho- nomic information loss, as well as a general reduction in articulate brachiopod diversity following the episode of mass extinction at the end of the Cretaceous. Specimens collected by the Natural History Mu- seum, Basel [NMB] expeditions to the Dominican Re- public between 1978 and 1980 came mainly from the Río Gurabo section (locs. NMB 15823, 15827 [716084], 15828 [716095], 15829 [716104], 15838, 15840, 15851, 15853, 15942 [715807], and 16811), with small collections from the Río Cana section (locs. NMB 16884 and 17023). In addition Tulane Univer- sity material was available from the Río Gurabo sec- tion (locs. TU 1210, 1211, 1215, and 1300) and the Arroyo Zalaya section (loc. TU 1227A). The location of these sections is shown in Text-figure 1 and the approximate positions of the brachiopod-bearing ho- rizons in the stratigraphic sections from these three river valleys are shown in Table 1. More detailed maps and columnar sections of these valleys, with locality numbers, may be found in Saunders, Jung, and Biju- Duval (1986, text-figs. 4-6, 15, 16, and 36), while a detailed register of both NMB and TU macrofossil collecting localities is shown in appendices 3 and 4 of the same publication. Systematic paleontological studies of Caribbean Ter- tiary brachiopods have been made mainly by Cooper (1955, 1959, 1979), while Recent brachiopods from the same region have been described by several au- thors, most notably Cooper (1977), in which reference to earlier studies is made. Since many Tertiary forms are extant today, such studies are of particular rele- vance to Tertiary brachiopod paleontology. The stan- dard for Cenozoic brachiopod systematics has been set by G. A. Cooper of the National Museum of Natural History, Smithsonian Institution, Washington, DC, U.S.A. and the present work has relied extensively on his publications on Tertiary (Cooper, 1979) and Recent (Cooper, 1977) brachiopods. ERRATA The measured апа(ог/ figured hypotypes from the Dominican Repub- lic that appear in "Neogene Paleontology in the northern Dominican Repub- lic. 6. The Phylum Brachiopoda", by Alan Logan (Bulletins of American Paleontology, volume 93, Number 328, pp. 44-52, plate 12), are the pro- perty of the Naturhistorisches Museum Basel and are deposited in that institution. By error, catalogue numbers of the United States National Museum of Natural History (USNM) have been assigned to them. The following list shows the correlation of USNM and NMB catalogue numbers: Tichosina ? lecta Guppy USNM 410560 - NMB L 9587 USNM 410561 - NMB L 9588 USNM 410562 - NMB L 9589 USNM 410563 - NMB L 9590 USNM 410564 - NMB L 9591 Terebratulina cailleti Crosse USNM 410565 - NMB L 9592 USNM 410566 - NMB L 9593 USNM 410567 - NMB L 9594 USNM 410568 - NMB L 9595 USNM 410569 - NMB L 9596 Argyrotheca johnsoni Cooper USNM 410572 - NMB L 9597 USNM 410573 - NMB L 9598 USNM 410574 - NMB L 9599 USNM 410575 - NMB L 9600 USNM 410576 - NMB L 9601 USNM 410577 - NMB L 9602 USNM 410578 - NMB L 9603 USNM 410579 - NMB L 9604 USNM 410580 - NMB L 9605 Lacazella caribbeanensis Cooper USNM 410583 - NMB L 9606 USNM 410584 USNM 410585 NMB L 9607 NMB L 9608 W^ tu DOMINICAN REPUBLIC NEOGENE. 6: LOGAN 45 ACKNOWLEDGEMENTS I am indebted to Dr. Peter Jung of the Natural His- tory Museum in Basel, Switzerland, for the opportunity to examine the Dominican Republic brachiopod ma- terial, and to Dr. Emily Vokes of Tulane University, New Orleans, LA, U.S.A., for providing additional brachiopods from the same region. Dr. G. A. Cooper of the National Museum of Natural History, Smith- sonian Institution, Washington, DC, U.S.A. kindly sent specimens of Argyrotheca johnsoni Cooper, 1934 from the type locality and Dr. Michael Risk of McMaster University, Hamilton, Ontario, Canada, and Dr. Jean Vacelet of the Station Marine d'Endoume, Marseille, France, sent specimens of Lacazella caribbeanensis Cooper, 1977 from the Bahamas. The scanning elec- tron micrographs were taken using the University of New Brunswick's Cambridge S4-10 Scanning Electron Microscope. Mr. Wilfred Morris, Research Technical Officer at the University of New Brunswick, aided in the preparation of photographs, and financial support was provided by a Natural Sciences and Engineering Research Council of Canada Operating Grant (A4331) to the author. PALEOECOLOGY Present-day articulate brachiopods are always ma- rine and benthic and there is no reason to suppose that fossil species, particularly in the Tertiary, were oth- erwise. The presence of pediculate and cementing forms usually implies the initial availability of some kind of hard substrate, either rock or pebbles, for attachment. However, studies made to date suggest that modern forms are generally not sensitive indicators of other environmental variables, such as depth, salinity and water clarity. Species of Pelagodiscus Dall, 1908, and Abyssothyris Thomson, 1927, for example, seem to be restricted to waters deeper than 500 m, while most other brachiopods, particularly diminutive forms, pre- fer depths between 0 and 500 m (Zezina, 1970) but, as Cooper (1977) points out, the relatively broad depth ranges of modern species make them unreliable bathy- metric indicators where the same or related species occur in the fossil record. The Dominican Republic collections contain species belonging to the genera Argyrotheca Dall, 1900, La- cazella Munier-Chalmas, 1881, Terebratulina d'Or- bigny, 1847, and (?) Tichosina Cooper, 1977. Of the 10 named species of Argyrotheca recognized by Cooper (1977) from modern seas in the Caribbean area, nine are restricted to the depth range 0—500 m (with four less than 100 m) and the tenth, A. barrettiana (David- son, 1866), is usually found in waters less than 200 m in depth, although occasional specimens may range down to 1473 m (Cooper, 1977). Similarly, all three species of Argyrotheca from the Mediterranean Sea UAYUBIN 9 10 20km 1 Rio Сапа 2 Rio Gurabo 3 Rio Mao 4 Rio Amina 5 Cañada Zalaya 6 Rio Yaque del Norte Upper Cenozoic [72] Oligocene - Early Miocene 2 Vid Mesozoic 7 City of Santiago 8 Arroyo Рипа! 9 Rio Verde d x iw пСО ፈር мы ее JANICO ит د‎ Text-figure 1. — Geological sketch map of the Cibao Valley, showing the location of Ше Río Cana, Río Gurabo, and Arroyo Zalaya sections from which brachiopods have been collected. (More detailed maps of these sections, with locality numbers, appear in Saunders, Jung, and Biju-Duval, 1986, text-figs. 15, 4-5, and 36, respectively.) 46 BULLETIN 328 prefer depths of less than 100 m (Logan, 1979). The nensis Cooper, 1977, has so far been found in less than presence of this genus in untransported sediments thus 100 m depth in the Caribbean (Cooper, 1977), while usually indicates relatively shallow water. Similarly, the closely-related L. mediterranea (Risso, 1826) from species of the cementing form Lacazella usually occur the Mediterranean and eastern Atlantic (Logan, 1979, in shallow waters in modern seas. Lacazella caribbea- 1983), although ranging down to 600 m, is commonest Cana Gurabo Zalaya 0 | = °С NN 16 Fa 9 5 Е | دت‎ 3 ж E 58 Ae | sds weg ህን ОТЕ NNI5| _ лос | Ist 333 ‚0 5 о мо = Е DEG 2 4 5 | = | 515 Y ш Е 5| f= шоу 2 5 5 ога # о ММ 14 ۰ سک‎ а о 0 5 9 Джа lign О z 5 & congls E а 5 NN13 2 с [ч ‚О .: .ቿ | ረ.አ 8 5 © Е 3 2 6 6 9 NNI2| ‘о » 8 =<) ھ‎ "ጋ g 5 P ۳ о О |225 n ل‎ O б = 5 де, =, 5 5 У ሠ " للا‎ ooo Z У 9 вка: للا‎ — Ф للا‎ о 2 E ርጋ 5 $ ወ = ) = 2 не = ? ጩ | Ы 1 Ош SS Qu УМУ 8, 9 5 ES congls 7 OU OV WEST EAST Table 1.—Stratigraphic sections from Rio Cana, Rio Gurabo, and Arroyo Zalaya, showing approximate positions of brachiopod-bearing horizons (stippled). Approximate stratigraphic correlations after Saunders, Jung, and Biju-Duval (1986). More detailed columnar sections appear in their text-figures 4, 6 (Rio Gurabo); and 15, 16 (Rio Cana). DOMINICAN REPUBLIC NEOGENE. 6: LOGAN 47 in depths of less than 200 m. The commonest living species of Terebratulina [T. retusa (Linnaeus, 1758) from the eastern Atlantic and Mediterranean, 7. sep- tentrionalis (Couthouy, 1838) from the western Atlan- tic and 7. cailleti Crosse, 1865 from the Caribbean] all range down below 500 m (Cooper, 1977; Logan, 1979, 1983), although 7. septentrionalis occurs in great abundance above 50 m in the Bay of Fundy (Noble, Logan, and Webb, 1976), and all species are com- monest at depths below 300 m. Species of Tichosina range down to 1000 m in the Caribbean and are rarely found in depths of less than 100 m (Cooper, 1977), suggesting deeper-water conditions where found in situ. The paleoecological reconstructions proposed by Saunders, Jung, and Biju-Duval (1986) for Neogene strata in the Dominican Republic can now be exam- ined in the light of the above observations on the depth preferences of modern brachiopods, bearing in mind that most of the fossil brachiopods show little evidence of extensive wear-and-tear or disarticulation normally associated with transported assemblages. The base of the Gurabo Formation is about 150 m above the base of the Río Gurabo section and calcareous silts between 250 and 400 m above the base of the section have yielded Argyrotheca johnsoni Cooper, 1934 and La- cazella caribbeanensis Cooper, 1977, indicating rela- tively shallow water conditions. This is compatible with the paleoecological evidence from associated fos- 5115, such as foraminifera and slumped blocks of sed- iment-enclosed reef corals belonging to Goniopora Blainville, 1830, and Montastraea Blainville, 1830. These occurrences, presumably derived from nearby reefs, suggest depths greater than 40 m and open ocean circulation, according to Saunders, Jung, and Biju-Du- val (1986). Cryptic habitats within the corals may have acted as living sites for these brachiopods, as in modern environments (Jackson, Goreau, and Hartman, 1971; Meile and Pajaud, 1971; Logan, 1977, 1981), where sclerosponges are usually an important part of this community. Mousa (1974) has found a similar asso- ciation of Argyrotheca, Lacazella and reef-associated foraminifera in Oligocene-Miocene claystones from Puerto Rico, which have also yielded sclerosponge-like spicules in wash residues. Such spicules should be sought in the sediment samples from the Dominican Republic as further evidence of derivation from nearby reefs. The upper part of the Río Gurabo section comprises the Mao Formation, which has yielded Argyrotheca johnsoni Cooper, 1934, and Tichosina? lecta (Guppy, 1866). Saunders, Jung, and Biju-Duval (1986) refer to this part of the section as follows (рр. 16-17): “From about 650 to 750 m on the column a Limopsis-Pterop- oda mollusc assemblage predominates. This assem- blage also includes Propeamussium, terebratulid bra- chiopods, and small solitary corals, and thus points to considerable water depths." The terebratulids referred to are undoubtedly those here questionably assigned to Tichosina, a form previously acknowledged to be indicative of deeper-water conditions. Furthermore, both Tichosina? lecta and Terebratulina cailleti Crosse, 1865 are found in association with Argyrotheca john- soni in alternating calcareous siltstones and coral lime- stones at a slightly higher level in the Mao Formation from the Río Cana section (Table 1). Mainly on the basis of foraminifera, Saunders, Jung, and Biju-Duval (1986) suggest water depths of at least 100 m. The brachiopod evidence of Т.? /ecta and T. cailleti is not inconsistent with this reconstruction, although the deepest recorded occurrence of А. johnsoni from the present day is 72 m (Logan, 1977). Thus in both the Río Gurabo and Río Cana sections the sedimentary and fossil evidence indicates increas- ingly deeper water conditions due to increasing dis- tance from the paleo-shoreline as one proceeds down- river and upsection, notwithstanding the different lithologies in the two sections. The Arroya Zalaya sec- tion has only a single brachiopod-bearing locality, which has yielded Argyrotheca johnsoni Cooper, 1934, in- dicative at present of water of less than 72 m depth (Logan, 1977). 'The associated foraminifera and other evidence suggest open marine conditions, moderate water depths and relatively low water-energy levels, according to Saunders, Jung, and Biju-Duval (1986). BIOSTRATIGRAPHY Brachiopods are generally intolerant of a wide va- riety of substrate types and, since they evolve slowly, are generally regarded as of limited value in biostra- tigraphy. Lacazella caribbeanensis Cooper, 1977 ranges from the Miocene to the present, while Tichosina? lecta (Guppy, 1866) is recorded from the Eocene of Cuba by Cooper (1979). Both Argyrotheca johnsoni Cooper, 1934, and Terebratulina cailleti Crosse, 1865, are com- mon in modern seas off the West Indies but have not previously been recorded from the Tertiary. SYSTEMATIC PALEONTOLOGY INTRODUCTION Taxonomic studies on both Recent and fossil bra- chiopods have traditionally been carried on mainly by paleontologists, since this phylum was much more di- verse in the past than it is at the present time. This has resulted in a consistent approach to the classifi- cation of the group as a whole and the description of taxa at various levels, with emphasis on shell char- acteristics rather than soft parts, as one might expect in a predominantly fossil group. The classification of Williams and Rowell (1965) has been adopted here; this volume of the Treatise also contains detailed de- 48 BULLETIN 328 scriptions of all supra-specific categories and a glossary of terms, an abbreviated version of which can be found in Logan (1979). PHILOSOPHICAL CONSIDERATIONS Cooper (1970, 1977) has outlined the main char- acters used in the recognition of species and genera of brachiopods, noting the emphasis on external shell fea- tures at these lower levels, in contrast to the importance of internal shell characters at family and higher taxo- nomic levels. The ornament and shell shape, together with the nature of the brachial skeleton, are the main generic characters used, with size, ornament, shell shape and shell coloration (mainly with Recent species) as the most important specific characters. The density of punctae varies greatly with the size of the individual and the position of the area sampled on the shell; con- sequently it has not proved to be a reliable specific characteristic, except in certain cases (Foster, 1974). While there 15 general agreement among brachiopod taxonomists on the nature of the characters to be used for distinguishing taxa, there is less conformity of opin- 1on on how narrowly brachiopod genera should be de- fined or on what is the limit of permissible variation within a species. Those taxonomists conversant with Recent species are well aware of the variation in size and shape associated with ontogeny and usually ap- pend scatter diagrams to species descriptions (e.g., Lo- gan, 1979). This is less easy to do with fossil species, where the number of actual specimens from a locality may be small and in these cases differences in practice arise among paleontologists. My own philosophy is to take the broad view of a species, based on natural variation, and to avoid the proliferation, whenever possible, of large numbers of species based on small samples. FORMAT, MEASUREMENTS, AND ACRONYMS OF SPECIMEN REPOSITORIES Citations included in the synonymy of each species are those considered to have identity with the de- scribed species. Measurements of figured specimens are in millimeters and include the length of each valve, maximum width and thickness, where able to be mea- sured; these and other dimensions can also be obtained from the figures. All figured specimens have been de- posited in the U. S. National Museum of Natural His- tory, Smithsonian Institution, Washington, DC, U.S.A. (formerly the United States National Museum) and bear USNM repository numbers. All other specimens examined have been returned to the Natural History Museum, Basel, Switzerland [NMB], including some Tulane University [TU] material given to the author by Dr. Emily Vokes. Class ARTICULATA Huxley, 1869 Order TEREBRATULIDA Waagen, 1883 Suborder TEREBRATULIDINA Waagen, 1883 Superfamily TEREBRATULACEA Gray, 1840 Family TEREBRATULIDAE Gray, 1840 Subfamily TEREBRATULINAE Gray, 1840 Genus TICHOSINA Cooper, 1977 Type-species. — Tichosina floridensis Cooper, 1977, Recent, West Indies. Tichosina? lecta (Guppy) Plate 12, figures 1-13 Terebratula lecta Guppy, 1866, p. 296, pl. 19, fig. 3. Tichosina? lecta (Guppy). Cooper, 1979, p. 11, pl. 2, figs. 1-5. Diagnosis. — Anteriorly-tapering Tichosina? with a narrow anterior fold. Type material. — Cooper (1979) states that this species 15 based on three specimens, all of them poorly pre- served. The lectotype (USNM 1 1 56468) is measured and figured (his pl. 2, figs. 1—5, where it is incorrectly referred to as the holotype), while a syntype (USNM 115646b) is also measured. Horizon and type locality. — Eocene, USNM locality 8180, San Fernando, Trinidad. Material. — Fragmentary material from localities NMB 15828 (14 fragments) and 15829 (about 30 frag- ments) from the Río Gurabo section, locality NMB 16884 (nine fragments) from the Río Cana section and locality TU 1300 (three relatively complete specimens and eight fragments) from the Río Gurabo section, the complete specimens showing some post-mortem dis- tortion and fracturing. Measurements (in mm).— pedicle brachial maxi- apical valve valve mum thick- angle length | length width ness (degrees) USNM 410560 29.7 25.8 23.9 15.0 82 USNM 410561 27.0 ЗЛЕ 23.9 237 71 USNM 410562 219 26.4 2558 16.9 100 Remarks and comparisons.— The relatively poor preservation of the Dominican Republic material and the absence of a complete loop in any of the specimens examined preclude positive identification with the ge- nus Tichosina, although one specimen (Pl. 12, fig. 13) shows the flat-bladed crural bases typical of the genus. The external shape, size and labiate permesothyridid foramen also suggest Tichosina. The general dimen- sions are closest to Т.? lecta from the Eocene of Trin- idad, the type material of which was recently re-figured by Cooper (1979). While the uniplicate anterior com- missure varies in intensity of development of the fold, DOMINICAN REPUBLIC NEOGENE. 6: LOGAN 49 post-mortem distortion may account for some of these differences. Thus one example from the Dominican Republic (USNM 410561) is narrower and more glo- розе than typical 7.2 lecta and has a more developed uniplicate anterior margin, resembling that of 7.2 bart- letti (Dall, 1920) from the Recent of Barbados (Cooper, 1977, 1979). None of the other Recent species of Ti- chosina described by Cooper (1977) is closely com- parable with 77? lecta; most are more globose and have a more exaggerated uniplicate anterior margin. Occurrence and distribution. — The occurrence of this species in the Dominican Republic has already been documented (see Material above). The type material 15 from the Eocene of Trinidad and the species has not previously been recorded from any other area. Family CANCELLOTHYRIDIDAE Thomson, 1926 Subfamily CANCELLOTHYRIDINAE Thomson, 1926 Genus TEREBRATULINA d'Orbigny, 1847 Type-species. – Anomia retusa Linnaeus, 1758, Re- cent, off Norway. Terebratulina cailleti Crosse Plate 12, figures 14—22 Terebratulina cailleti Crosse, 1865, p. 27, pl. 1, figs. 1-3. Terebratulina cailleti Crosse. Davidson, 1886, p. 26, pl. 5, figs. 41- 42. Terebratulina cailleti Crosse. Cooper, 1977, p. 99, pl. 25, figs. 1-16; pl. 28, figs. 4-27. Description. —Shell small, rarely exceeding 10 mm in length in adult forms, ovate to sub-pentagonal, lon- ger than wide, slightly auriculate at hinge line, partic- ularly in juvenile stages, apical angle of pedicle valve less than 90*; biconvex, brown in living specimens, punctate, costellate, with eight to 10 primary costellae, secondary costellae intercalated anteriorly. Costellae strongly beaded, particularly in juveniles. Anterior commissure uniplicate, with shallow ventral sulcus and dorsal fold. Foramen large, mesothyridid, with pedicle collar; deltidial plates disjunct. Brachial skeleton extends to almost half length of brachial valve; hinge plates absent, socket ridges and crural bases fused to form prominent ridges. Crura convergent, crural processes uniting to form a ring-like loop with ventrally-arched transverse ribbon. Cardinal process small, transverse; hinge teeth grooved on inner face. Small oval-shaped muscle impressions in both valves, interiors of both valves with branching mantle canals; lophophore plectolo- phous, spicules abundant. Diagnosis. — Small elongate-oval Terebratulina with beaded costellae. Type material. — The whereabouts of Crosse's type material is unknown. Horizon and type locality. — Recent, Guadeloupe, West Indies. Material. — А complete brachial valve from locality NMB 16884 and a complete bivalved specimen and three separate pedicle valves from locality NMB 17023, all juveniles and from the Río Cana section. Measurements.* — pedicle brachial maxi- apical valve valve mum thick- angle length length width ness (degrees) USNM 410565 — 31 4.1 — 105 USNM 410566 5.4 — 4.0 — 72 USNM 410567 5.0 4.5 3.8 2.0 74 USNM 410568 ፦ 3.0 2.3 - 60 USNM 410569 — 2.9 2.0 — 61 USNM 410570 6.6 6.0 4.7 2 55 (97) USNM 410571 7.0 6.2 6.0 2.6 71 (93) Remarks and comparisons. — Although попе of Ше Dominican Republic specimens show the diagnostic loop, the external features of both valves and the in- ternal features of the pedicle valve are typical of the genus Terebratulina rather than Eucalathis Fischer and Oehlert, 1890, or Chlidonophora Рай, 1903. The fossil material is identified with the Recent species 7. cailleti Crosse, which is common in the waters around the West Indies. This species was recently extensively re- illustrated by Cooper (1977) and specimens dredged from 125 to 150 m depth off Paynes Bay, Barbados have been figured here for comparison with the fossil forms. АП the fossil specimens appear to be juveniles but show the typical elongate-oval shape and beaded costellae typical of the species. As Cooper (1977) has remarked, 7. cailleti is generally smaller than its more northern congeners, 7. septentrionalis (Couthouy, 1838), and T. retusa (Linnaeus, 1758), the largest spec- imen of T. cailleti in the collection from Barbados measuring 13.0 mm in pedicle valve length and 11.2 mm in maximum width. Terebratulina retusa was re- cently re-described by Logan (1979) and compared in detail to 7. septentrionalis. T. cailleti differs from both these species and from T. /atifrons Dall, 1920 in being much smaller when fully grown, more elongate and with strongly-beaded costellae (Cooper, 1977). In ad- dition, 7. latifrons is sub-triangular in outline, with a stronger fold and sulcus, and is similar in these respects to the Miocene species 7.2 palmeri described by Coo- per (1979) from Cuba. Occurrence and distribution. — The occurrence of this species in the Neogene of the Dominican Republic is * The number in parentheses under apical angle is that of the brachial valve: the other number in such pairs is that of the pedicle valve. 50 BULLETIN 328 limited to the Río Cana section. The species is pre- viously known only from the Recent, where it is com- mon in the Caribbean region. Suborder TEREBRATELLIDINA Muir-Wood, 1955 Superfamily TEREBRATELLACEA King, 1850 Family MEGATHYRIDIDAE Dall, 1870 Genus ARGYROTHECA Пай, 1900 Type-species. — Terebratula cuneata Risso, 1826, Recent, Mediterranean. Argyrotheca johnsoni Cooper Plate 12, figures 23-41 Argyrotheca johnsoni Cooper, 1934, p. 2, pl. 1, fig. 10, pl. 2, figs. 1- 12. Argyrotheca johnsoni Cooper. Cooper, 1977, p. 111, pl. 1, figs. 2-7. Description. — Adult shell wider than long, rarely ex- ceeding 8 mm in width at its widest point, variable in outline from alate to semicircular, cardinal extremities occasionally mucronate; biconvex to almost plano- convex, pedicle valve more convex than brachial valve; anterior commissure shallowly sulcate; ventral inter- area broadly triangular, flat, catacline to weakly ap- sacline, dorsal interarea very narrow, anacline. Beak short, subtruncate, usually abraded, foramen large, hy- pothyridid, with pedicle collar, delthyrium flanked by rudimentary narrow deltidial plates. Ornament of low, gently-rounded costae, usually four to seven in num- ber, with occasional costae intercalated in sulcus-fold region, interspaces narrow, cream to pale yellow in living forms, in contrast to pink tinges at extremities of ribs. Shell coarsely punctate. Interior of pedicle valve shows widely-separated teeth along denticulated hinge area, pedicle collar supported by median septum extending to between about half and four-fifths length of valve, triangular in profile, low and thin. Brachial valve with short, transversely- elongated cardinal process; hinge line with elongate sockets; long median septum reaching almost to an- terior margin, apex at mid-valve, triangular in profile, occasionally with four to five serrations sloping steeply to anterior margin. Crura widely separated, crural pro- cesses pointed, horizontal; brachial skeleton compris- ing two arcuate lamellae attached to base of crura, valve floor and slightly posterior of anterior extremity of median septum; muscle scars distinct, lophophore schizolophous. Diagnosis. — Alate to semicircular Argyrotheca with catacline ventral interarea and subdued costae. Type material. — Holotype: USNM 431003. Para- types: USNM 4310038--ር (Cooper, 1934, 1977). Horizon and type locality. —Recent, Johnson- Smithsonian Deep Sea Expedition Station 52, Latitude 1910 25", Longitude 69?20'55"W to Latitude 19*10'05"N, Longitude 69?21'25"W, attached to worm tubes in association with Thecidellina barretti (David- son, 1864) at 25-40 m depth off Cape Samaná, Do- minican Republic (Cooper, 1934). Material. — Four complete specimens from locality NMB 15823, one complete specimen from locality NMB 15827, two brachial valves from locality NMB 15828, nine pedicle and eight brachial valves from locality NMB 15829, 14 complete specimens and some fragments from locality NMB 15838, two complete specimens and two pedicle valves from locality NMB 15840, a single incomplete brachial valve from locality NMB 15851, а single complete specimen from locality NMB 15853, a complete specimen and two brachial valves from locality ММВ 15942 (- 15807), а pedicle valve and two brachial valves from locality NMB 16811; all from the Río Gurabo section. One complete specimen and three fragmentary pedicle valves from locality NMB 16884, and a single complete specimen from locality NMB 17023; Río Cana section. Also a single complete example from locality TU 1215 (Río Gurabo section), and three separate pedicle valves from locality TU 1227A (Arroyo Zalaya section). Measurements (in mm).— pedicle brachial valve valve maximum length length width thickness USNM 410572 ጋ ፦ DU — USNM 410573 1.9 1.8 3.8 1.9 USNM 410574 1.8 1.8 3.0 1.6 USNM 410575 14. 1:3] 3.0 13 USNM 410576 1:5) 15 2:2 1.3 USNM 410577 epu 3.0 5.6 1.3 USNM 410578 2.9 29 3.8 11 USNM 410579 1.7 1.6 4.0 1.9 USNM 410580 2.0 240) 870 ۸12 USNM 410581 4.0 Bo 8.0 2.8 USNM 410582 4.8 4.1 7.0 2.8 Remarks and comparisons.— Although many Re- cent and fossil specimens of Argyrotheca have been described from the Caribbean region, very few are strongly alate and have a catacline ventral interarea. The Dominican Republic material 15 here identified with the modern species А. johnsoni Cooper, 1934, notwithstanding the smaller size of the fossil speci- mens, which are assumed to be immature. Many of the modern species of Argyrotheca are distinguished on the basis of shell coloration, clearly inappropriate for the fossil forms, which have lost any coloration they might once have possessed. Two specimens of modern A. johnsoni from the type locality are figured (Pl. 12, figs. 36-41) to show the range of variation in shape and number of costae, and for comparison with the figured fossil specimens. Ofthe other extant species DOMINICAN REPUBLIC NEOGENE. 6: LOGAN 51 of Argyrotheca, only A. rubrocostata Cooper, 1977, A. schrammi (Crosse and Fischer, 1866), and A. rubro- tincta (Dall, 1871) show similarities in outline of shell to A. johnsoni. The ribbing in A. rubrocostata is gen- erally finer and the ventral sulcus narrower than in 4. johnsoni, while A. schrammi has fewer but more ex- aggerated ribs, giving a strongly-scalloped anterior margin. This species, as Cooper (1977) has indicated, is poorly known. 4. rubrotincta has perhaps the strong- est resemblance to the fossil forms from the Dominican Republic, but lacks the strongly-alate outline charac- teristic of this species. None of the Tertiary species described by Cooper (1979) from Cuba and the Carib- bean is similar to the species described here. Occurrence and distribution. — This species occurs in the Neogene of the Dominican Republic in the Río Gurabo and Arroyo Zalaya sections but is otherwise unknown from the fossil record, although relatively common in modern seas in the Caribbean, where it has been recorded from the Dominican Republic [Johnson-Smithsonian Station 52, Pillsbury Station P 1153], north-east of Turks Island, Bahamas [Pillsbury Station P 1421] (Cooper, 1977), off Tongue of Ocean, Bahamas (Logan, 1977), and off the north coast of Jamaica (Jackson, Goreau, and Hartman, 1971). Order THECIDEIDA Pajaud, 1970 Suborder THECIDEIDINA Elliot, 1958 Superfamily THECIDEACEA Gray, 1840 Family THECIDEIDAE Gray, 1840 Genus LACAZELLA Munier-Chalmas, 1881 Type-species. — Thecidea mediterranea Risso, 1826, Recent, Mediterranean. Lacazella caribbeanensis Cooper Plate 12, figures 42—49 Thecidium mediterraneum (Risso). Davidson, 1864, p. 21, pl. 2, fig. 5. Thecidium mediterraneum (Risso). Davidson, 1887, p. 158. Lacazella mediterranea (Risso). Meile and Pajaud, 1971, p. 470, pl. 1, figs. 1-4. Lacazella caribbeanensis Cooper, 1977, p. 132, pl. 4, figs. 12-19. Lacazella caribbeanensis Cooper. Logan, 1979, р. 75, pl. 10, figs. 9- 14. Lacazella caribbeanensis Cooper. Cooper, 1979, p. 28, pl. 1, figs. 2-5. Description. — This species was recently described and illustrated in detail by Cooper (1977), to which the reader is referred. Diagnosis. —Small Lacazella with hemispondylium attached to the valve floor and the ascending apparatus narrow anteriorly and with smooth margins (Cooper, 1977). Type material. — Holotype: USNM 549449a. Para- types: 549448a, 549449b (Cooper, 1977). Horizon and type locality. —Recent, Johnson- Smithsonian Deep Sea Expedition Station 52, Latitude 19?10'25"N, Longitude 69?20'55"W to Latitude 19?10'05"N, Longitude 69?21'25"W, in association with Argyrotheca johnsoni Cooper, 1934 at 25-40 m depth off Cape Samaná, Dominican Republic (Cooper, 1977). This species may be the “Thecidellina barretti (Da- vidson)" referred to in Cooper (1934). Material. — Only three specimens of this species are present in the collections: two single brachial valves (from loc. NMB 15942 and loc. TU 1211) and a single pedicle valve (from loc. TU 1215), all from the Río Gurabo section. Measurements (in mm).— pedicle brachial valve valve maximum length length width USNM 410583 — 135 22 USNM 410584 — 1.8 2.6 USNM 410585 2.0 - 2.1 USNM 410586 2.6 — 2 USNM 250982 3.6 23 3.4 USNM 250983 3.6 — 3 USNM 250984 — 3.0 3.4 Remarks and comparisons. —In spite of the limited material available, there is no doubt of the identity of the Dominican Republic specimens with Lacazella caribbeanensis Cooper, 1977. The distinctive internal features of the brachial valve, mainly the ascending and descending apparatus, clearly indicate that the specimens belong to Lacazella rather than Thecidel- lina, the only other modern cemented articulate bra- chiopod genus. There are three known species of La- cazella: L. mediterranea (Risso, 1826) from the Mediterranean and eastern Atlantic (Logan, 1979, 1983), L. mauritiana Dall, 1920 from the Indian Ocean (Cooper, 1973) and L. caribbeanensis Cooper, 1977 from the Caribbean region. The main differences be- tween these three species have been discussed by Coo- per (1977, 1979) and Logan (1979). L. caribbeanensis 15 distinguished from its congeners by its smaller size, differently-shaped hemispondylium (two separate plates, rather than a bilobed plate), smoother margins to the ascending apparatus and intervening median ridge of the brachial valve, and more subdued internal markings in both valves. Modern Bahamian material belonging to Г. caribbeanensis (Pl. 12, figs. 46-49) has been figured for comparison with the fossil forms. Occurrence and distribution. — This species has been recorded from the Miocene of Cuba (Cooper, 1979); this and the present report of its occurrence in the Neogene of the Dominican Republic are the only fossil 52 BULLETIN 328 examples so far known. The species has been recorded from modern seas around the Bahamas (Meile and Pajaud, 1971; Logan, 1979) and has recently been iden- tified by the present author attached to a stalagmite at the bottom of the blue hole off South Bight, Andros Island, Bahamas, at a depth of 45 m (in Gascoyne ef al., 1979). Elsewhere it has been taken off Jamaica (Davidson, 1864, 1887; Jackson, Goreau, and Hart- man, 1971) and the Dominican Republic (Cooper, 1977). REFERENCES CITED Alloiteau, J. 1957. Contribution à la systématique des madréporaires fossiles. Centre nat. Rech. sci., Paris, 2 vols., 462 pp. 1958. Monographie des madréporaires fossiles de Madagascar. Ann. Géol. Madagascar, vol. 25, 218 pp., 38 pls. Айту, C. C., Jr., and Carrión-Torres, С. 1963. 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The columellar style and septa are thickest in the forereef and thinnest in the lagoon population. The inner wall is also especially thin in the lagoon, and the entire theca is more porous. The pali and septal teeth appear best-developed in the sand channel population. Figure i Figured specimen. SUI 51065, forereef sand channel (20 m depth), SEM of calical surface. 2. Figured specimen. SUI 51073, forereef sand channel (20 m depth), transverse thin-section. . Figured specimen. SUI 51080, forereef (20 m depth), SEM of calical surface. . Figured specimen. SUI 51082, forereef (20 m depth), transverse thin-section. . Figured specimen. SUI 51085, lagoon (16 m depth), SEM of calical surface. . Figured specimen. SUI 51085, lagoon (16 m depth), transverse thin-section. С\ л څل‎ UY PLATE 1 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 93 PLATE 2 DOMINICAN REPUBLIC NEOGENE. 4, 5, AND 6 57 EXPLANATION OF PLATE 2 Stephanocoenits pone: JO LMC е КОЕ M ш 8ዛንይ፡ፕ. MU E CT око i cM 20 Whole colonies, closeups of calical surfaces, and X-radiographs. Colonies are generally small to intermediate in size and well-rounded with smooth outer surfaces. The calices are small and circular, and the exothecal intercostal area well-developed. Figure 17 21 3 \ 4. Figured specimen. NMB D5763. Upper Miocene, locality NMB 15847, Rio Gurabo, Gurabo Formation, Dominican Republic. Upper surface of colony, х и. Figured specimen. NMB D5867. Upper Miocene, locality NMB 15858, Rio Gurabo, Gurabo Formation, Dominican Republic. Upper surface of colony, х 1. Figured specimen. NMB D5866. Upper Miocene, locality NMB 15859, Rio Gurabo, Gurabo Formation, Dominican Republic. Upper surface of colony, x1. Figured specimen. NMB D5793. Upper Miocene, locality NMB 15850, Río Gurabo, Gurabo Formation, Dominican Republic. Upper surface of colony, х1. . Figured specimen. NMB D5762. Upper Miocene, locality NMB 15846, Río Gurabo, Gurabo Formation, Dominican Republic. X-radiograph, х1. . Figured specimen. NMB D5792. Upper Miocene, locality NMB 15846. Río Gurabo, Gurabo Formation, Dominican Republic. Upper surface of colony, x1. . Holotype. BM(NH) R28756. Neogene, “silt of the sandstone plain, San Domingo." Upper surface of colony, x1. . Holotype. BM(NH) R28756. Same specimen as figure 7 above. Closeup of calical surface, x 5. 58 BULLETIN 328 EXPLANATION OF PLATE 3 STEDRUNOCOCNEMSPONET OVS (Duocan) ie Е а ереси es vnc hus de gee Ee lcu 20 Closeups of calical surfaces and transverse thin-sections. The exothecal intercostal area has a distinctive appearance with a well-developed second outer wall formed at the junction between calices. The pali are weak and 12 in number. They appear better-developed on the primary septa. The columella forms a prominent style. The tertiary septa are relatively thin and long. Figure 1. 2. ON tn 4 یں‎ Figured specimen. NMB D5866. Same specimen as Plate 2, figure 3. Calical surface, x 5. Figured specimen. NMB D5761. Upper Miocene, locality NMB 15846, Río Gurabo, Gurabo Formation, Dominican Republic. Transverse thin-section, x 10. . Figured specimen. ММВ D5793. Same specimen as Plate 2, figure 4. Calical surface, x 5. . Figured specimen. NMB D5762. Same specimen as Plate 2, figure 5. Transverse thin-section, x 10. . Figured specimen. NMB D5867. Same specimen as Plate 2, figure 2. Calical surface, x 5. . Figured specimen. NMB D5763. Same specimen as Plate 2, figure 1. Transverse thin-section, x 10. BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 93 PLATE 3 Tx NT | TA РГАТЕ 4 „э. аш а” > ed: «+»: መ - ጠጋ BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 93 Stephanocoenia spongiformis (DUCA) ое ав ER у: оро а LEUR M M аат оо ان‎ В ЕЦ, DOMINICAN REPUBLIC NEOGENE. 4, 5, AND 6 59 EXPLANATION OF PLATE 4 SEM photographs of calices and a broken longitudinal surface, and a photograph of a longitudinal thin-section. Vertical and horizontal skeletal structures are equally poorly developed. Septa are composed of six trabeculae that form moderate-sized surface denticles. The pali are elongate and consist of approximately three thickened trabeculae between the septa and columella. Endothecal dissepiments are especially weak, whereas exothecal dissepiments are better-developed with distinctly uniform spacing. Figure le 2; 3 4 . Figured specimen. ММВ 135761. Same specimen as figure 1 above showing a longitudinal break through a calice, x 20. ON м . Figured specimen x 10. Figured specimen. ММВ D5761. Same specimen as Plate 3, figure 2. Calical surface, x 20. Figured specimen. NMB D5761. Same calice as figure 1 (left), showing columella, pali, and septa, x 100. Figured specimen. NMB D5869. Upper Miocene, locality NMB 15850, Río Gurabo, Gurabo Formation, Dominican Republic. Calical surface, x 20. Figured specimen. NMB D5869. Same calice as figure 3 (left) showing columella, pali, and septa, x 100. . NMB D5879. Upper Miocene, locality NMB 15850, Río Gurabo, Dominican Republic. Longitudinal thin-section, 60 BULLETIN 328 EXPLANATION OF PLATE 5 ИДИ Ic SDCOIeS ር А በወ ".፡.፡.: UNO Мике куи PEL ከሠ... O У 21 Whole colonies, closeups of calical surfaces, and X-radiographs. Some colonies are large with hemispherical shapes and a smooth outer surface. The calices are relatively large and polygonal, and the exothecal intercostal area weakly developed, if at all. Figure 1. Paratype. NMB D5768. Lower Pliocene, locality NMB 16818, Río Cana, Gurabo Formation, Dominican Republic. Whole colony, x Y, 2. Holotype. NMB D5766. Lower Pliocene, locality NMB 16817, Río Cana, Gurabo Formation, Dominican Republic. Whole colony, x Ya, 3. Paratype. NMB D5868. Upper Miocene, locality NMB 16933, Río Gurabo, Gurabo Formation, Dominican Republic. Whole col- опу, хд4. 4. Holotype. ММВ D5766. Same specimen as figure 2 above. X-radiograph, х и. 5. Holotype. ММВ 05766. Same specimen as figure 2 above. Calical surface, х 5. 6. Holotype. ММВ 05766. Same specimen as figure 2 above. Calical surface, х 5. PLATE 5 Ми \ | wl y ; х | Ў | едра 1 ARPA 3 x $e ETA 1 ٨ 1 0 W NUM EN TET. Meet WM РТАТЕ 6 VOLUME 93 И BULLETINS OF AMERICAN PALEONTOLOGY DOMINICAN REPUBLIC NEOGENE. 4, 5, AND 6 61 EXPLANATION OF PLATE 6 Stephanocoenia duncanisnew.specles- азына С сше па и мари ала ን.6ጋፓ.ፓ.ጋንኪፓፓፕፓፕፓ ሸ6ባ››ሽባሽ፡፡፡ሽ፡ሽ፡›፡ሽ፡ ፡ ምሎ ምሺፀደቢፀቢዐቢኪቢኪቢ6ቧቢቧቢዷቢጊ0ኪ ኽሜጫሜጫሌ:ዚ፡፡ሽ: ፡፡ .-. 21 Closeups of calical surfaces and transverse thin-sections. The corallites are roughly polygonal and closely-spaced and are separated by little or no pore space. The corallite walls are thick and dense. The pali are especially weak and the columella thick. Tertiary septa are relatively thick and short. Figure 1. Paratype. NMB D5768. Same specimen as Plate 5, figure 1. Calical surface, х 5. 2. Paratype. NMB D5765. Lower Pliocene, locality NMB 16817, Río Cana, Gurabo Formation, Dominican Republic. Transverse thin- section, x 10. . Paratype. NMB D5769. Lower Pliocene, locality NMB 16817, Río Cana, Gurabo Formation, Dominican Republic. Calical sur- face х5. 4. Paratype. ММВ D5871. Upper Miocene, locality ММВ 16934, Río Gurabo, Gurabo Formation, Dominican Republic. Transverse thin-section, x 10. 5. Paratype. NMB D5868. Same specimen as Plate 5, figure 3. Calical surface, x5. 6. Paratype. NMB D5771. Lower Pliocene, locality NMB 16881, Río Cana, Gurabo Formation, Dominican Republic. Transverse thin- section, x 10. > 38 تت‎ የ ሀ SA е те BULLETIN 328 EXPLANATION OF PLATE 7 SEM photographs of calices and a broken longitudinal surface, and a photograph of a longitudinal thin-section. Vertical skeletal structures are generally less developed than horizontal ones. Septa are composed of five to six trabeculae that form minute surface denticles. The pali consist of two to three thickened trabeculae between the septa and columella. Endo- and exothecal dissepiments are thin and fairly regularly spaced. Figure [፣ 2 3. 4. Holotype. NMB D5766. Same specimen as Plate 5, figures 2, 4-6. Calical surface, x 20. Holotype. NMB D5766. Same calice as figure 1 (above), showing columella, pali, and septa, x 100. Paratype. NMB D5870. Upper Miocene, locality NMB 16933, Río Gurabo, Gurabo Formation, Dominican Republic. Calical surface, x 20. Paratype. NMB D5870. Same calice as figure 3 (above), showing columella and pali, x 100. . Holotype. NMB D5766. Same specimen as figure 1 (above), showing a longitudinal break through a calice, x 20. . Paratype. NMB D5871. Same specimen as Plate 6, figure 4. Longitudinal thin-section, x 10. E. ind са ውጋ ጩ = ~ = О. > € 9 © =] О Е ፖ ወ] ш لم‎ < es ፖሪ < S = ጩ = < بت‎ о u ደ E щ کو‎ © =) ea BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 93 ኘ > | Г EZ Sree ЕЕ Yt y, y тъга PLATE 8 Ефиге 1-7, 13. 14-16. 17, 18. 19-21. pom. DOMINICAN REPUBLIC МЕОСЕМЕ. 4, 5, AND 6 63 EXPLANATION OF PLATE $8 zintillocyaihusemuoensis«(Vaughan9jl925) oos SUR н оаа 28 1, 2. Side and calicular views of holotype (USNM 353644), x2.0, x2.1, respectively. Loc. USGS 7785. 3. Calice (NMB D5873) showing well-preserved pali and columella, x2.2. Loc. NMB 15863. 4. Damaged calice (ММВ D5874) showing well-preserved pali and columella, x 2.3. Loc. ММВ 15864. 5. SEM of columella and pali (NMB D5875), x9.8. Loc. NMB 15836. 6. Side view of upper calice (ММВ D5876) showing degree of septal exsertness of primary septa, x 2.2. Loc. ММВ 15863. 7. Aberrant paratype (MCZ 9267) with 58 septa, х 3.6. Loc. unknown. 13. Transverse thin-section through calice (ММВ D5877) showing two septa joined to palus at right, x19. Loc. ММВ 15807. comrillocyathusialütus; new. Зе 30 8, 9. Side and calicular views of holotype (ММВ D5878), х2.1, x2.2, respectively. Loc. NMB 15871. 10. Paratype (NMB D5879) showing well-developed lateral costae, x 2.0. Гос. ММВ 16857. 11. SEM of septal granules (NMB D5880), x38. Loc. NMB 15962. 12. Transverse thin-section through calice (NMB D5881) showing theca and two septa, х 54. Loc. NMB 15863. Antillocyattusicrisiatus. Vaughan оо) Cc ሲን HM i d MM 30 14. Lateral face of large specimen (ММВ D5882) showing lateral edge costae, x 0.85. Гос. ММВ 15809. 15, 16. Side and calicular views of holotype (USNM 353643), x1.4, x1.6, respectively. Loc. USGS 7785. Paracvarhus: henekeni Duncan TSO)... O E a a a a 34 Side and calicular views of lectotype [BM(NH) 28768], x6.3, x6.1, respectively. “Nivaje Shale”. Trochocyathus (Paratrochocyathus) chevalieri, new species نن نن‎ о а تت ت‎ een 31 19. Damaged calice of paratype (ММВ 135883) showing pali and columella, х 2.0. Гос. ММВ 15816. 20. Stereo view of calice of holotype (NMB D5884), x3.4. Loc. NMB 15836. 21. Transverse thin-section of septal base (NMB D5885), x86. Loc. NMB 15809. Trochocyathus (Paratrochocyathus) duncani, new species تت تت تن نن‎ hene 32 Side and calicular (stereo) views of holotype (USNM 74684), x1.35, x1.6, respectively. Loc. TU 1208. 64 Figure bea 3, 4. 5-9. 10-12. 14-19. 2U 2T. BULLETIN 328 EXPLANATION OF PLATE 9 Page Ceratotrochus (Edwardsotrochus) species cf. C. (E.) duodecimcostatus (Goldfuss, 1826) .................................... 33 Side and calicular views of specimen from Loc. ММВ 15836 (ММВ D5886), х 1.35, x1.5, respectively. የ ያያ CE. J duodeeuncostatus(Gioldiusss: 926)... сърбин и ን. осо бре ре con к T ER ыы e 33 Side and calicular views of specimen from Rio Torsero, Italy (Pliocene) (USNM 326663), x 1.3, x 1.4, respectively. VA AOS АНА ነና Бресте ване RR ET PUE ts nr ee مهس وغم‎ л у л у eani ከም... 34 5. SEM of damaged calice of paratype (ММВ D5887) showing septal granules and paliform lobes, х 8.9. Loc. NMB 15833. 6, 7. Side and calicular views of holotype (NMB D5888), x5.0, x6.9, respectively. Loc. NMB 15833. 8. SEM of upper theca (ММВ D5887) showing sinuous costae, х 20.3. Loc. ММВ 15833. 9. Transverse thin-section of calice (NMB D5890) showing wall and two septa, x67. Loc. NMB 15833. IL НД ВУИ LSI lc ос A E ИНИ du er ne T Ue 35 10, 11. Calicular and basal views of single specimen Кот Dominican Republic (USNM 63125), х 3.9. Loc. USGS 8702. 12. Basal view of specimen from the Miocene of Tortona, Italy (MNHNP L 60), x3.35. < ያያ ያር በያት speciesic Di Halens (Michelotti 1838) ረ... .ሱ.ም.....ስርስክ...ቢ፡.... S T 35 Basal view of Recent specimen (USNM 36443), x3.3, U.S. Fish commission Steamer Albatross sta. 2751: 22°41'N, 7 772 шъ. 21 Jan. 1885, Sphenotrochus (Emsthenotrochus) Seni Wells; 1247:፡፡፡.፡..፡........፡........፡..፡...።..ሥ.ም.ጵስ.ቢ..... መ... 36 14-17. SEM of specimen (ММВ 135891) from Loc. ММВ 16938. 14, 15. Side and calicular views, х 8.6, x 10.9, respectively. 16. Elongate costae in lower third of lateral face of corallum, x25.3. 17. Costal granulation near upper thecal edge, each costa composed of two rows of granules, x 23.5. 18. Holotype (USNM 68358) from Martinique, x3.9. 19. Lateral face of specimen (NMB D5892) from Loc. NMB 15900, x4.4. DION COLO CHUSCOS SEN апай LIZI) O o а АД 36 Side and calicular views of a specimen (USNM 68359) reported by Wells (1945) from Martinique, х 3.1, x3.7, respectively. PLATE 9 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 93 ነ BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 93 PLATE 10 DOMINICAN REPUBLIC NEOGENE. 4, 5, AND 6 65 EXPLANATION OF PLATE 10 Figure Page 1-456 Asterosmilia anormalis ЧӘ Сай ISO) Soo NU ፡..:..:.....:.፡፡፡፡፡.፡....052ያሆ656ዓ9 . ار د‎ 37 1. Side view of specimen (USNM 546427) showing alate costa on convex lateral edge, x 1.3. Loc. USGS 85109. 2, 3. Duncan's illustrated syntype [BM(NH) 28801], both faces figured, x 1.1. 4. Damaged calice of specimen (USNM 546424) from Loc. USGS 8519, x3.2. 6. Transverse thin-section of calice (NMB D5893) showing theca and three septa, x 20. Loc. NMB 16910. 5 Asterosmiluuproluera:(Boustales eoa) ее 37 Recent specimen (USNM 74687) from R/V Silver Bay sta. 2445: 24°08'N, 80*08"W, 252 m, 3 Nov. 1960, x 1.0. 7-9, 1o የ ያና የየየ ያ Duncan 1907 m SLES us ic GN TS уулду eo Aa Ао TT a C със 38 7, 8. Side and calicular views of largest syntype (USNM 324816) of A. hilli Vaughan, 1919, х 1.6, х 2.0, respectively. Loc. USGS 2580. 9. Holotype [BM(NH) 828944] of A. exarata, х 2.2. Specimen figured by Duncan, 1867, pl. 32, fig. 5. 11. Longitudinal fracture of specimen (USNM 63329) from Loc. USGS 2580 (Jamaica) showing dissepiments, х 2.4. TOL Asterosmiliaespecles CP A. ለ1... ее و‎ ник due куйе ጨመር ሚም ሚር፡ UD دت‎ 39 Specimen (USNM 44303) reported by Wells (1934) from the Scotland Beds of Barbados, х 3.4. 12-18. -Asterosmilia profunda Duncan Ss OA) ie eared ets pes, s RE E D C NALE ጋን ሚር ኸኸ. 39 12. SEM of longitudinal fracture through specimen (USNM 546424) showing two dissepiments, x 12.4. Loc. USGS 8528. 13. Elongate corallum (USNM 63327) from Loc. USGS 2580 (Jamaica), x0.5. 14, 15. Side and calicular views of holotype [BM(NH) 40413], х 0.88, x 1.4, respectively. 16. Longitudinal cut of specimen (NMB D5894) from Loc. NMB 15833 revealing dissepiments, x 1.3. 17. Transverse cut of specimen (USNM 63327) from Loc. USGS 2580 (Jamaica) revealing abundant dissepiments, x 2.8. 18. Transverse thin section through two septa and wall of specimen (NMB D5895), x12. Loc. NMB 15833. 19:20. 2A steyosmutu duncani Vauchan 92 Iu E m c nua s Reed Ре 40 Side and calicular views of holotype (MCZ 9277), х 1.1, х 1.4, respectively. 21-24 S M ОРОШ сорте sas ALON AD LIAS RENTUR ое о ም ፡ች፡በዎርሜማ:፡ и 21, 22. Side and calicular views of syntype (MCZ 9275), x 1.4. 23. Side view of syntype (MCZ 9276), x2.2. 24. Calice of syntype (MCZ 9273), x1.6. 66 BULLETIN 328 EXPLANATION OF PLATE 11 Figure Page БЕРРИ SDECIOS O чалан vote e e rero он го REED De a E WERL VONT OIM 41 Side and calicular views of specimen (USNM 64202) from Loc. USGS 8733, x1.4. 41 5—7 ТОТ POUT IOC OS Hispidus (Polrtales ШЕ ЛӨ} co UR о ehe OC CN 3. SEM of well-preserved calice (USNM 74688) showing three cycles of septa, columella, and pali, x 12.5. Loc. TU 12274. 4. SEM side view of upper theca of specimen (USNM 74688) showing smooth concentric banding of ерићеса (see also Pl. 11, fig. 3), x 16.4. Loc. TU 1227A. 5. Transverse thin-section through three septa of calice (USNM 61928), x62. R/V Caroline sta. 67: 18°32'18"М, 65?46'12"W, 329—512 m, Recent. 6. SEM of septal granules in specimen (USNM 74688), x145. Loc. TU 1227A. 7. Stereo SEM of well-preserved corallum (USNM 74688), x9.4. Loc. TU 1227A. 10, 11. Rows of mural pores in specimen (ММВ 05896), x 15.3, x65, respectively. Loc. ММВ 16038. 8,9, 12, 5: Guynia Species ct-6zannuldta Duncan, ое e er. rr лл л оллоо. у. A. 42 8, 12, 13. Theca (USNM 74689) showing costal spines, x 33, x65, x21, respectively. Loc. TU 1227A. 9. Calice of specimen (USNM 74689) showing octameral symmetry (see also Pl. 11, figs. 8, 12, 13), x38. 43 DII EDOnUrophylha.cormucopiaw:ouciales Г. 14. Recent specimen (USNM 46685) from Straits of Florida: 24?18.7'N, 81°56.9'\\, 220 m, х 0.64. 15-17. Side and calicular views of specimens (USNM 64227) from Loc. USGS 8528, x1.3, x2.4, x2.4, respectively. BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 93 PLATE 11 BULLETINS OF AMERICAN PALEONTOLOGY, VOLUME 93 Ефиге 1-13. 1-5. esp 33, 34. BS 36-39. 40, 41. 42—49. 42. 43. 44, 45. 46. 47. 48. 49. DOMINICAN REPUBLIC МЕОСЕМЕ. 4, 5, AND 6 EXPLANATION OF PLATE 12 Тзскоѕтна? ваа (еру) رکون د‎ А пое пе MEM دلو هخه عفد‎ Pedicle valve, brachial valve, posterior margin, anterior margin and side views of a complete specimen, x1, hypotype USNM 410560, Mao Formation, Río Gurabo section, locality TU 1300. . Pedicle valve, brachial valve, anterior margin and side views of a distorted complete specimen, showing resemblance to Т.? bartletti (Dall), x 1, hypotype USNM 410561, Mao Formation, Río Gurabo section, locality TU 1300. . Pedicle and brachial valves of a partially-crushed example, x 1, hypotype USNM 410562, Mao Formation, Río Gurabo section, locality TU 1300. . Posterior margin, showing labiate foramen, x 1, hypotype USNM 410563, x 1, Mao Formation, Río Cana section, locality NMB 16884. . Enlargement of crural bases of brachial valve, x 5, hypotype USNM 410564, Mao Formation, Río Gurabo section, locality NMB 15828. spel CF EDV ALULING: COLLLELE Gkossea woe هرد‎ о ፡..፡ሽ6.፡፡፡8ሀ6፡).7፡፡፡ амне . Pedicle valve interior, x 4, hypotype USNM 410566, Mao Formation, Río Cana section, locality NMB 17023. . Brachial valve view of complete specimen, x4, hypotype USNM 410567, Mao Formation, Río Cana section, locality NMB 170287 - Brachial valve exterior, х4, hypotype USNM 410565, Mao Formation, Río Cana section, locality ММВ 16884. . Brachial valve exteriors, x4, hypotypes USNM 410568 and 410569, respectively, Mao Formation, Rio Cana section, locality NMB 17023. . Pedicle and brachial valve views of a complete specimen, x4, hypotype USNM 410570, Recent, off Paynes Bay, Barbados, dredged from 125-150 m. . Pedicle and brachial valve interiors of a complete specimen, х 4, hypotype USNM 410571, Recent, off Paynes Bay, Barbados, dredged from 125-150 m. 5 ANB LORCA ЈОВО COOBO acim, И وش‎ i ы а . Pedicle valve, x5, hypotype USNM 410572, Mao Formation, Rio Cana section, locality NMB 17023. . Posterior view of complete specimen, x4, hypotype USNM 410573, Gurabo Formation, Rio Gurabo section, locality NMB 15840. . Posterior margin of complete specimen, x5, hypotype USNM 410574, Gurabo Formation, Rio Gurabo section, locality NMB 15838. . Anterior margin of complete specimen, hypotype USNM 410575, x5, Gurabo Formation, Rio Gurabo section, locality NMB 15823. . Side view of complete specimen, hypotype USNM 410576, x5, Gurabo Formation, Rio Gurabo section, locality NMB 15823. 28-30. Pedicle and brachial valve interiors, x6, and SEM profile of median septum of brachial valve, x 10, hypotype USNM 410577, Gurabo Formation, Rio Gurabo section, locality NMB 15942. Pedicle and brachial valve interiors of a complete specimen, х 6, hypotype USNM 410578, Mao Formation, Rio Cana section, locality NMB 16884. Pedicle and brachial valves of an alate complete specimen, x6, hypotype USNM 410579, Gurabo Formation, Rio Gurabo section, locality TU 1215. Brachial valve interior, x 12, hypotpye USNM 410580, Gurabo Formation, Rio Gurabo section, locality NMB 15853. External and internal views of a complete alate specimen, х 6, hypotpye USNM 410581 (out of USNM lot No. 549947), Recent, off Cape Samana, Dominican Republic, dredged from 75 m depth. Pedicle and brachial valves of a complete specimen, x5, hypotype USNM 410582 (out of USNM lot No. 549947), Recent, off Cape Samana, Dominican Republic, dredged from 75 m depth. Жасахейшсинрвейнейзт СООН ӨК ти. ie ое ое. Brachial valve interior of partially-worn specimen, x8, hypotype USNM 410583, Gurabo Formation, Río Gurabo section, locality NMB 15942. Brachial valve exterior, hypotype USNM 410584, Gurabo Formation, Río Gurabo section, locality TU 1211. Pedicle valve interior, x 8, and SEM enlargement of hemispondylium, x 15, hypotype USNM 410585, Gurabo Formation, Río Gurabo section, locality TU 1215. Pedicle valve interior, x 8, hypotype USNM 410586, Recent, attached to stalagmite in blue hole off South Bight, Andros Island, Bahamas, at depth of 45 m. Brachial valve of complete specimen, x 5, hypotype USNM 250952, Recent, Grand Bahama Island, Bahamas, 20 m depth. Pedicle valve interior, tilted to show hemispondylium, x6, hypotpye USNM 250983, Recent, Grand Bahama Island, Bahamas, 20 m depth. Brachial valve interior, x 5, hypotype USNM 250984, Recent, Grand Bahama Island, Bahamas, 20 m depth. 67 51 68 BULLETIN 328 INDEX Note: Page numbers are in light face, plate numbers are in bold face type; principal discussion pages are in italics. abnormalis, ДЕНОВИ та а es AA а осо рота иык Ра O E Abyssothyris Thomson, 1927 ........... Actinastraea d'Orbigny, 1849 .... adherens, GUYA መ ሠ ን о ПОЕ ОКВ а орла зас сено LIN EN К CLO DO ык но Же M D E БИ а И Ајту and Carrión-Torres (1963) aterra РЕПО e rrr IM иан йы ООО АСОСИ ЕТЕР AlveoporaBlamville 1830 imer ep a (ES retro ie ana dani АММН [American Museum of Natural History, NEVADAS USA а 7 “Ап. madrep. intersepta ? Esper, suppl. І, t. 79” ................... 18 ОАК ИОАН Еа EA AO 10,11 Е A T E E 25,26,42,43 annulata Spec) GUYNIA srnce وو وروی هس‎ Е 24,25,27,42 апота!а, Asterosmilia Anomia reinsa LACUS, 79S ووو وو‎ инве а 49 E Е а неее наи еН нн 39 УОН BIY ccc rM еН 39 Amtilastraca: Duncan, ПВА 18,19,21 И ОЈ iso corti c cce Me M а н АН شه ووهه‎ 42 nO attis Wells. о 26,27,29,41 ICES pico tu E e HM e e የ ee 23.23, 21,29 30,31 cristatus (Vaughan in Vaughan and Hoffmeister, 1925) ............. О M E 8.205 maoensis (Vaughan т Vaughan and Hoffmeister, 1925) ........... Ware pur lU T ые $... 22/2909 Annlophyila ۷ 231181399 1932 cece irdenas ioien a 41 DAV COMBE а То EA AD такое е КИ ЖО т ያ үне к 46 Argyrotheca Dall, 19005... 45,47,50 barrettiana (Davidson, 1866) ее 45 Johanson Coopt, ПОЗА ое и هغاه‎ 12:577 44,45,47,50,51 УНОГОВО На GOOD OP ረረ aas ER ታች ፡ A ET E 51 ПИРОТ теле В Ра По 51 schrammi (Crosse and Fischer, 1866) .............................. 51 Articulata Hunley, 1869 а روم هغ‎ ee دوا‎ ٧ er ም”ጥጉ”ጥ”ጥ” 48 Asterosmilia Duncan, 1867 ............. 23,26,31,37,38,40 abnormalis (Duncan, 18064) оно кое его dere 229: ИСТ: anormala неа 37 compressa Vaughan in Vaughan and Hoffmeister, 1925 ............ ПЕЕ И А 10 ...... 25-27,38,40,41 COMMU DUNCAN По 97 СО ее 37 duncani Vaughan in Vaughan and Hoffmeister, 1925 ............... Mo caede oa MEN IET 10 ...... 25-27,38,39,40 exarata Duncan, 1867 .................... wW 25,27,38,39,40 DOLI AES ан exarata var. robusta hilli Vaughan, 1919 Bp. oF A hihi of Wens, IIIA мл ove سوه ونو‎ 39 machapooriensis Hoffmeister in Vaughan and Hoffmeister, 1926 ........................... 38,40 marchadi (Chevalier, TIGO) conver odore deoa PEERS ERIS 25,38,39 ро ана ОВ е 38 profunda (Duncan, 1864) ............ 10575 25,27,29,38,39,40 profeta Vouriales TSI) CE MUN 37,38 trinitatis Vaughan in Vaughan and Hoffmeister, 1926 .... 38,41 ?Asterosmilia exarata VAC رر‎ e MM E 38 exarata var. robusta Vaughan and Hoffmeister, 1925 ......... 38 SPC ነ 4 ....ያፕፕፕ፡›፡ ::፡:፡:: Astrea intersepia Lamarck, ПВО o UT uua нает Astrocoenia Milne-Edwards and Haime, 1848a ... decaturensis Vaughan, WITO ec ec LE guantanamensis Vaughan, 1919 ۸١٠ тетией Масова оо 01101167515 Маарап IIS a Atlantic. OCEA E m па ES ќи АНА Ена ATT [inner wall thickness] QUII SSSDHOHOLLOOLUSS S TE auritus var., ISDICHOLLOCITSR ЛО ОЛУУ УК ОШО СОК: DSDLOHOLNLOGCHASQ ...ፕፕቡፕተ፡ፕ.፣፡.፡..፡፡፡፡፡ ር Australia ша ЗА ране VERA AUS кы ра А ро EN STE E RUD Id ur E RZ OVCS s OR REESE ENSE ር ርር ከ. вара ва сл и EUM NNUS TS E Andros Island, off South Bight of Tongue of Occ О. а 1 Baoa FOMA errar е MOA Е Оа о e EAE A ОТ Balanophyllia j pittieri Vaughan; 1919. sp. of Vaughan and Woodring (1921) BdrdadO о е ee Р Paynes Ва barretti, СОС 27,28 LMA E укн Л оо E HUE 50,51 ያያ Are yrotheca ROSE 45 ገረ ያ о 49 Bassignac tuffs .............. Bay of Fundy .... Bermuda sr. Bernardi1908)8 ое Blainville (1830) 10,19,20,43,47 BM(NH) [British Museum (Natural History) London, England, КЕЕ 6,7,20,21,23,25,27,32-34,37,38,40,42,43 Bounie5909) e rrin ያ... бен. Bowdon оао соо ert err Ie ER ETE В TII e eode ничка ን... EIE Brachycyathus henekeni Duncan, 1863 Pe brassensis, Sphenotrochus (Eusthenotrochus) ............ ии 36 Pra A АЕ 20 G [costal Су ee 25,26,28,33,35,37,38,40—42 DOMINICAN REPUBLIC NEOGENE. 4, 5, AND 6 69 GAEL FEED SIU e ae | fw 44,47,49 Cairns (1979) Cairns (1982) (cmd AR E E E (ваши он ПО о. Cambridge S4-10 Scanning Electron Microscope .... Cancellothyrididae Thomson, 1926 ...................... Gancellothyridinaeshhonmsonel92695. очна сис E Сар саш и и со 6-8,10,18,20,22--26,32,34,35,43--47,49--51 Car ያ 12/1. аса ን ека 125 н 44—47,51 Caryophyllia (Ceratocyathus) Seguenza, 1864 ....................... 32 Caryophyllidae Dana, 1846 Caryophylliicae Dana, 1846 Caryophylliina Vaughan and Wells, 1943 Caryophylliinae Dana, 1846 CD COANE тате TM Ceratotrochus Milne-Edwards and Haime, 1848b ................. 315 duodecim- costatus (в оа) я 58 duodecimcostatus (Goldfuss, 1826) .............................. 26,33 ри Мода А NETO сери er ES MN 41 Ceratotrochus (Edwardsotrochus) Chevalier, 1961 ................. 55 sp. cf. C. (Е.) duodecimcostatus (Goldfuss, 1826) ..................... አ ቄን IIS PE in EE Ори 8027020186 Cercado Бома полни вина ceri RN 8,29,35,36,46 TGereado:Eormallote не а а прсти М 28-32,39 Chevalier (1961) 18,20,26,32-35 CheValier:(19 62) Pir II Килен RUN CHEN 35 Ghevalier об). OP HO UIT SEND 33 Chevalier (1966) ለ n) IA MEC НИНА re 38,39 Ghevalter*Jean-Bierze- EE коскена Мк CRM пеене 31,32 chevalieri, Trochocyathus (Paratrochocyathus) mra .. ‹................... Il RN ыу кс 8. ЭР Ghiidonophora Dall ТО eei Ue КЕ 49 CEW [Columella wiati] ence aA 11,12-16,20-22 ОШ ЕТО ን и и 31 compressa, Куќа ке КИРЕ T0975. 25-27,38,40,41 conicus, Deltocyathus ?Deltocyathus Cooper (1934) ......... Cooper GOSS cae COOPER) он AAR У. EN Cooper (1970) SOOO о) tt газ a е ER УМ Cooper (1977) 44-52 Cooper (1979) 44,47–49,51 COPEN GTA и МР ИМ de СВОЕ 44,45 COR [tota sample correlations] о. eee ele ARTS cornucopia, DERN ያ T Ne rene Trochocyathus ?Trochocyathus ВОЙ а ООШ эме еа UR ад HERRERA MERERI Е CORORATA АШ АЦО ЦИЦА посно за а ан РИ AE TR Ca Раја ال‎ Со оре ПОН оноо И costatus, Placocyathus " (ОШ ОНУ SSS sans hen ede enue ከው ОНЕ cristatus, UTA ING, canker а CO AUS a и GROSSE SOS) тен M СО aR Crosse and Fischer (1866) CS оба тераса ји о ео: Dall (1870) Dall (1871) Dall (1900) Dall (1903) Dall (1908) Dall (1920) ааа IDA Со (ОБ s co stel NENNEN EE 10 маке еа US uere РАСИ ок ОИ Ов ENNIO oU c LS M DEE (ОБС) crue УНЕ MEL DN Палма сети Gl SSH) ae FEL п LL LL menant decaseptata, Stephancoenia ? .... GECALUTENSIS ASMOCOCM renoier AE INEO CEA EM Deltocyathus Milne-Edwards and Haime, 1848b ................... 35 COTLICUSSZADLOWAUS T I Ed А nu pu 35 italicus (Michelotti ИЗ) омаи 2427,35 ЗБСП (Michelotti) eee. 25,35 ?Deltocyathus conicus Zibrowius, 1980 sp. cf. D. italicus (Michelotti) .... dendroidea, Stephanocoenia Dendrophyllia. Blainville 1830; Sse OS alternata Pourtalès, 1880 cornucopia Pourtalés, 1871 .............. радиата (Duncan в) И E n. sp. А of Vaughan and Woodring (1921) sp. В. of Vaughan and Woodring (1921) ........................... 160160111111096 (531:15../1(65.፡፡፡.፡.......: Репдгорћу па Vaughan and Wells, 1943 DOLAN АТ ومع هوو‎ TET 7 Dominican Republic, АВ OS CLITA лан ددد ول و مهه وو‎ 33 2100 О 66 eerie ан ас 29,30,40 Gibao&Vvalleya У а экес кшм и мы ки ы КУ. 6,7,16,23,45 Стао Valley, Arroyo Рода нь ее ое 6,24,45 Arroyo Zalaya Cañada Zalaya 6,24,45 Ciy of Santiago xa вон аи нане د وا‎ 6,24,45 А 6,24,29,30,38,45 Rio- Сапа ии 5-9,21,22,24,29-31,35,38,44-47,49,50 Río Gurabo ....... 5-9,21,22,24,25,29-36,38-40,42-48,50,51 Вло Мао و‎ 6,7,18,22,24,25,28-31,33,38-43,45 ВАЗА ОЕ Мао 28,31 Возете та ото e : 6,24,45 Río Yaque del Norte 6–8,22,24,25,30–32,35,36,42,45 Santiago delos Caballeros ее. 31,32 Rio уб е А ы ак у а оо Е 22 (ов Quemados. ماو‎ EOE 32 МАО банани Виз ET csset oid das Jia лалы 32 (59111 Gape ата па т os. ccc NG seinen aS 50,51 Rio Guanajuma 38 South of Santiago de los Caballeros ................................. 38 dominicensis, Dominicotrochus а дори АЗ ከው ee ees ЗОО Е 5... 5.5. ን. DDOMIMICOLLOCHUS WIE 1937... add cil 36,37 dominicensis (Vaughan in Vaughan and Hoffmeister, 1925) ...... Locus aM MM E ን ሚወ ውጣም TURIN T E 9... 25298697 [Drucken вана UR са сыны ннн ees ټمر‎ ПИК $ 70 BULLETIN 328 Сара ЈАДИ SR SD هره‎ co OO due eod лана кан edid 41 DUN [species 2, fossil ММВ Stephanocoenia duncani] .......... 15 Dancan ОСО i tete ou deis tory 6,20,25,27,31,32,34,41 na ВОД), دمه وو‎ 6,20,21,25,27,31,33,37-39,40 ТОНА СВОДЕ ее 20,25-27,37,38,40 Donean 906). Lr እይ مومسم‎ ctn rr ም ክን 6,25,31–34,37,38,41 Duncan (1872) Duncan (1873) Duncan (1884) S airo Mall CLBOS A ው а 41 пасат CET ОЛАТИ A አው о 21532 duncani, ОМОТИ ARI ው ones rye ps ТО 25-27,38,39,40 Stephanocoenia ........... а-о 5,7-10,12-14,16,17,21,22 Trochocyathus (Paratrochocyathus) ......... Sates: 28125327 00. ипопеситесозгатиз COERQLOLPOGHMS posos moss AEE лм RAE 33 UROUPCHTCOSIQI, 121 ያያ፡፡፡፡ o ZELUS 33 00066772 و7‎ sass wu ve N RE sO e E 26,33 duodecimcostatus (sp. cf.), Ceratotrochus (Edwardsotrochus) ......... نه لصو سو‎ E О et ae. 0123 A O ووو غاد ول ولاو هواک و‎ 19 Eastern АА region iiio E potete 35,46,47,51 121215. S Mi ОИ СОС а 7 ШОС UNUS QU LS VE зау a e Ellis and Solander (1786) ... ER яры щч re MA К ео NE јин E ራማ loo A е га ции A ጋ ደይ ve о ОНЕ ШӘ FE Ени ЛАВЕ رواو‎ РЕНО E سه سا‎ осна Fischer aud OQehlert, О 5.67 2 AREE cores 49 2፡15 E O SOL АЦЕ ЕСЕТ САДЫ тасаг 33–36 ВТА ومو سه سم‎ e obiil cr RR 4 ፣ጸያ። 35 (ior serere ае edens SOR eite v УЧИ eS аЙ 35,41 BIS азота Pallas, 1766) وو‎ ker T DE EO 24 CRO QA, ASTEVOSIMUA. оз я ЩЙ; 25,27,38,39,40 exarata var., PASTOS II то со ив оа паноа нае toute о АИТ 39 2 ШЕ ва ано КЫМ таз и жедна рыу e ብ 38 exarata var. robusta, Asterosmilia ?Asterosmilia ESA ADE nr ክናን ከን ዋሉ ሙሻ ይክ 41 JOR OPES, SCD INANOCOCING sb ር... ни vr condone dee cea te ive ENEE 20 fasciatus, Trochocyathus (Paratrochocyathus) нн 32 ور‎ БШШШ eat ыы 24 وي ^947 و‎ HIE н СИН ео 28,30,33-40 и پد‎ MEE 53:935 PEO OO ано огои E Fischer and Oehlert (1890) 11216111686 Bourne, 1905. оное ао اوه‎ та Tie ደጸ voa 27,41 1191011 Bonne; 1905 ove و و‎ ал 27,41 “ОРЕ КОЕ o وو ووو وو‎ OER 41 atlanticum e СООО ТОСА LOS A НН 41 exaratum Doncan and Wally TE6S و‎ ла. 41 pavoninum Певвош Sd LGR OAS 41 roissyanum Milne-Edwards and Haime, 184860 ... 41 а sensu Vaughan and Woodring, ፕሳ ማሻያያ፡ርሙች፡፡፡ ee 41 bsensu Vaughan and Woodring, لا‎ o MERERI 41 I op- sensn Dinmont (DOOD)! о В ВИНТ 41 DRDO DANE о ве ES a قر‎ M ux 252/41 Florida, кс. ыт к oL eed ueber SE avv usa aevi ን ም እ. 42 (ОПАК VENAE I ER E аа UT UM АА 43 TOT S RAILS RO АЕ cc A ከ be des eh TR Eo 42 fossulus, Trochocyathus (Paratrochocyathus) |............... esses 32 Е ООО) и ES SO ое УО ОБОО) на а елы DE ОНА ика TEE E оС Foster (1980) Foster (1984) КОЗЕ (1989). A O oa EOS ORO et E ени Foster (ISO) Е Ae Бове Зее Foster, A. B. (Nancy) Eran Ces OM lV ул оо |:7:(63471:4111(:11:1(110፡4፡ 2792227922 РН E ее orar (0 рази Frost (1981)............ a" Frost and Langenheim (1974) 18,20 Gabb [Gabb Collection, Museum of Comparative Zoology, Cam- bridge МА, US. AJ ан 27,31,40 ርጋ ጋ IDLO HALO ШОН пениса NAR Me 43 Gardiner (1904) tee Gere A ако бокс Cantate аа 35 Gascoyne or dL. 11910. xc цени ridere eee io M ten tes 52 gcd [greater calicular diameter] 25,30,33,41 Geistes R екы Cv КО ERE COE DARET: 6,7,10 ИСПУНЕ ያ ያ ን de а сена сено EO thee 35 а 28 Globigerinoides trilobus fistulosus foraminiferal zone ............. 46 Globordialla exilis оташе zone ое 46 Globorotalia humerosa foraminiferal zone ............................. 9 Globorotalia margaritae foraminiferal zone ....................... 9.46 Globorotalia miocenica foraminiferal zone ......................... 9,46 EE em EE N e ያንን СА РА а 7 Goldfuss (1826) ... COMODO BILO Hle: ПОЗА. а ите SR UE 20,47 ОТВАР A Goreau and Wells (1967) Gray (1840) до Grays (USA не ከ... Овај руы ED ИРА СЕ РЕ m C а (ВАРОШ ВИКИ а сан dU ФЕ О Я ru guantanamensis, Astrocoenia m on Gili Ole УИ о и ПОРУ CLO OG) EG TEUER TQ ICONE OI Бн ИА Gurabo Formation .......... . 8,21,22,29-33,38,40,43,46,47 ?Gurabo Formation ......... 25,29-33,38,40-43 GIGA Wd LOT АА هه‎ а 42,43 adherens (Tenison=Woods; 1878) ска ninas obse 7. 43 ANNUAL DUNCAN LSTA о 25,26,42,43 sp. cf. С. annulata Duncan, 1872 .......... Ш 24,25,27,42 (Ed ТОЛО еа а 27,41,42 АА مره‎ РА НА е МАВАРА ЛИН ро ay nl RRO ССОО 42 11:1: о ДЕ АЙСЫ a O оно соо са во а о 43 Еее (5) dhe( ره وه مش اه‎ и па و‎ E ро A а 20 НеКо Colection ор лети ter er etr ране OM 2021 henekeni, Brachycyathus POPC OUD ec ки кы EA RR VPM ተ ae Hickson (1910) ............ hilli, Asterosmilia hilli (sp. сЕ), O И ረ ሴንን о О O 39 DOMINICAN REPUBLIC NEOGENE. 4, 5, AND 6 71 ЖОО Slides RAR О ЕСЕГЕ яя 38 hispidus, (GERALOW OC SA ute EURO NUT - 41 Јаја по ዎች SS ዳ..::..... Ilo 24,25,27,29,41,42 Hubbard and Wells озо) ERE 43 МЧК БУ (SGD) АН ማይ. ህወ Be Re айныш due: 48 Tiyanonhotaabuaschers08075 SCENE SRO ER 20 ICZN [International Commission on Zoological Nomenclature] ... imparipartitus, Trochocyathus .. 11111 Фесаще d I ندید هوه لصو‎ WGta ТОРА ҮТЕ жа በክር La ae INT [species 3, modern Stephanocoenia intersepta] .............. 15 intersepta, оротат AS Ar а ር ው E 19 “АШК суулоо уко o о E EOS E 18,19 ГООО а ER اور‎ ን M он HO 19 Stephanocoenia ..... Ie 5--7,8,10,11,13,14,16,17, / 9,20--22 пе 1272427777 ያፕ нае INE 35 ANG SIDON ER DANE 24-27,35 italicus (sp. cf.), ОСТИ о A УЕ ыл A ደ iet 25:35 DELICIAS A л УЛ УЛ Лл RN IS 35 Дана OA DO БА ARES есенен, 31552 COLON а sere re лг а пали NR е 33 A A reece eae 33 1.1 Која О ао е 35 Paroni Си (а по SN PES re ديد که سوه‎ ies 33 РТАТЗАПСВЕ UR ورد‎ еы ያችን rience ያ ከመ бре 33 Иели ој ЫЛАН» ЛЕ eredi ም ደ) Ови saws hats 35 TUSCANY KIO LOLSOLO AR. A ж ee A ره‎ 33 Jackson, Goreau, and Hartman (1971) ........................ 47,51,52 Jamadar ለ1... Pee iam Е ers 7,13,18,22,36,38—40,52 ID1ScOVenysBays Ro А 5510511517 ofthe ШОШ бав о Eee INTERNO PEE 51 Johnson-Smithsonian Deep Sea Expedition ..................... 50,51 Johnsoni, ANEVUOLREO ees e eee see eene rient | Ip — 44,45,47,50,51 JIE PRS II са гнасни КО а аө а. 5-7,10,23,45 EAN S USO 50 NO E o lc E A EAE EU UAE THE NM 7 NR E ee c ER Weder Жогол 18 Kojumdgieva and Strachmirov (1960) ................................ 41 L [lagoon population of modern Stephanocoenia intersepta] ... 17 Баба IK conu. e 19 Lacazella Munier-Chalmas, 1881 .............................. 45- 47,51 caribbeanensis Cooper, 1977 ................... Bohas 44-47,51 VIRO) aues eim rem О e ው оао 31 ТОДОР ПО 550: 1820) نس‎ ПОО HS 46,51 Ја АНОМ سو‎ оа 7 2 ИК ШОО) о к 8,10,11,18–20 amarok e olle ПОШ нае о 19 Саве ADAM te Se AS Ic سایس‎ УЫ Er 26 Itang ከወ በክን ላከን ን ከ ከወ... md br dro 7,10 апреле Re Eio diras eu lateral thecal edges ....... lateral thecal faces .......... latifrons, Terebratulina Icdiflessencaliculanciameter].-... oerte erm 25,30,33,40 lecta, PARAE a s Coe К К О ОО ОО LL OE ыо: 48 Тол (КОЗИ) аа ss pet om ree ما‎ За uen et 41 Limopsis-Pteropoda mollusc assemblage ............................. 47 ТОББ ИОВ: eM ел 7 Fema stro fnis и OR dass fiot S seda todas 36 Linnaeus (1758) ..... e AA e 43,47,49 ТЕСЕ QUOTA) ri ERE Se BE: 47,51 о а (КОЈА) оо سه‎ sees Se 46-49,51,52 Говап (1981) Говап (1983) PUROS Sa Мадаразса е а it ንን роба аа SU i ule ac ረረ LU Madracis Milne-Edwards and Haime, 1849 Madrepora ПОН SIUC NE cue зора caeca ан» сы RUD rp sanus DA 19 ПОВРАТАТ ЕНА iii a са РТ и shal gee 43 MaorAdonto АОЛ ба свои а паса cose CHER Слана 32.33 Mao Formation ............. 8,21,22,24,25,29,33,34,40,42,43,46,47 maoensis, Antillocyathus 8 xu 25-27,28,29-31 ROO ce Pa cater o со оа аси as AA: 27,28 marchadi, Asterosmilia Е. Мапе ва ТЕ А са ОИ deut tige LED e مس‎ 7 УГО MEE o E e e med 36,37 mauritiana, Lacazella ጄሪ о دون‎ 51 МАО рүн cce тей а е ም ዓን do es 18 McMaster University, Hamilton, Ontario, Canada ............... 45 MCZ [Museum of Comparative Zoology, Cambridge, MA, U. S. A.] ук EAE. erc دلا‎ 23,27,28,29,31,36,40-43 ሰ пиле NS Ol ооо وهن‎ АВЕ 27 mediterranea, Hog Dd ce d ЦЕ, НП NS 46,51 ШӨ АЕ E سمش‎ c ЕУ ዩር ње 31 piedüemaneunt PUCCIO ie ORE TE 51 Mecathyndidas а 1870r rri cd. یو خو‎ ahe te ee 50 Meile and Pajaud (1971) meinzeri, Astrocoenia ... Melville; В. V. sus ИКО аб مه‎ caules e d NM E michelini, Stephanocoenia michelinii, Stephanocoenia Michelotti e Ти Е (OG ВИ ран د وتن‎ а ve مده مه وشن‎ Milne-Edwards and Haime (1848а) Milne-Edwards and Haime (1848b) Milne-Edwards and Haime (1849) سه‎ ЛАДА Milne-Edwards and Haime (1850) .................................. Milne-Bdwards and Haime (1960) Саса itunes МЈ5 [number of major septa] ......................... 11,12-15,21,22 MNHNP [Museum national d'Histoire naturelle, Paris, France] ... аборта qu Me лень. des ን И ት e РИ 19.23.27. 35. 35 ДОЛУ ава Вашу ше 1 SO. ИН РРА У ПРИ tees 10,47 annularis (Ellis and Solander, 1786) ............................ 10,11 Morocco 35 Morris, Wilfred де 45 ПОЕН SDHGHOIVOGHUS e e тен оном ра ን 39 INTO ИЕ) ccm و لور‎ mee سم وص‎ ае 47 MIES ООШ (1ህ556))፲52..::.:::...:.:...፡... መሥመር... 50 Ме А o M EM с en مرل‎ f 12 BULLETIN 328 МАС [number of adjacent corallites] .................. LUIS 211. 22 Natural Sciences and Engineering Research Council of Canada ..... vieil nd RD ODE UE ы HM К 45 Naturhistoriska Riksmuseet, Stockholm, Sweden .................. 36 ТОВ 11:11 О 7 ВИ а ШШЕ oa al a ... 6 ааа Ва 34,37,38,40 ММВ [Naturhistorisches Museum Basel, Basel, Switzerland]......... 5,7,10,12-14,16,18,21-23,25,26,27,29-38,40,42,43-45,48-51 NMNH [United States National Museum of Natural History, Smith- sonian Institution, Washington, DC, U.S. A] ..................... В E 25,27,28,34,35,39,40,43-45 NMS [Natur-Museum Senckenberg, Frankfurt, West Germany] .... ПРАВИ asm RAUM O MEI dU ERAT E РА 7,19 Noble; Logam, and Webb 01976) ሻም dd ገ 47 صن و چا‎ 2177. а а A O MEETS TOES 49 NS [total number of septa] 722.2... 11,12-16,20—22 d'Orbigny (1847) d'Orbigny (1849) DA пао Б E Идеја COM ር O ERIN ERAS REE Pallas (1766) рано LL 49 O ti ti MR و‎ ታጋ алт OE 40 E EE E EE CO SRS CU ዯረ... 7 Paracyathus Milne-Edwards and Haime, 1848b ............... 33,4 Henekent (DUICAR ¡ISO oe SUEM 25,27,34 procumbens Milne-Edwards and Haime, 18486 ................. 33 DUICHOHTISGPIITDDIS LOAD) o ር ЕА ЭЭ ТРИО НОУ O ODE A ога A 23,27,29,34 turonensis Milne-Edwards and Haime, 1848b ................... 34 ОЕ c А РАТОВА ИАЕА A И ПРО E T N ПС ПОН D M i gee ОНА а Pd в а A FINES у ао دځ‎ ИСЕ тРетавушттњнетекен (Duncan) ла Parasmiliinae Vaughan and Wells, 1943 MEET TurammothocyarhusAlloneau 1958... 1:2: 31,32 collingnoni Alloiteau, 1958 ООН A TR Це дара тога ува ДОД а A SOE РС КО НЕ СЕ dodo ETT Persian Gulf О О Е зл а re: Phyllosmilia Fromentel, 1862 РОГУ Бресе о е РИСТО ЈОНА Eodem ОИ Ee ОРЕ 121772 ያያ ያ съ ооо ОЛГА ИРА Placocyathus Milne-Edwards and Haime, 18485 Н; неохота ЧӨӨ Аи او‎ ЫЛ НЫМАЛ ЛЕ DT OE DA 93 дылган а ОС СОНИ АА СОРИ из ODEO, 1604 enero И cristatus Vaughan in Vaughan and Hoffmeister, 1925 .... 27,30 maoensis Vaughan in Vaughan and Hoffmeister, 1925 ... 27,28 О Dunen, бер 17/2007 Машай ОИ нию Plesiastraea spongiformis Duncan, 1864 “ Plesiastrea spongiformis, spec. nov.” portoricensis, Astrocoenia .................... TOUT ТА АЕ SERRE ER ECO 27,37,38,43 PORE А ИО 1 የ ን о s 32,36 PO AES (LOTTIE ODER о Ie RE ОМЕТА. 6,37,38 Јој ЕЕН (ӨЛӨ) ርን ውን МИРАН r UD ተ. 27,41 РОК ШЕ SON оо МА ሮ፡፡፡...፡‹.. 43 Оа ЕТИШЕ ያ SMO OTO የ ገ ያየም ን پو‎ 43 PONAR RAS TENOR tinted reete ор 38 OUTTQIOGV ANUS: Сатов ЛОТО ate а 41,42 hispidus (Pourtalés, 1878) ........... Li 24,25,27,29,41,42 рошошакќеовіае апаверіаа n сево ог oo ፡፡ АБМЕ ЕЕ 23 ID ADU TD ТУ EAN AG! OTUS ОЕ A Е 93 profunda, Asterosmilia ................... 105254. 25,27,29,38,39,40 ТОТА ያ ዘ ያ ረ)... met Ui РА ند‎ 39,40 роет ASTCVOSITUN o rt АТ ОИ Propeamussium Ded RICO Rls pulchellus, Paracyathus pumpellyi, Stylocoenia РОО EKSO LIFO E E ТИНИ 42 R [forereef coral thicket population of modern Stephanocoenia CATS IN AL uA. ША LOCO VIAOLE DON Um Мо ыл T E 43 rawsonii, РОС ОСУ ИН К مالس هخ مره ووا د سم د‎ D مه سوه‎ p 27 ааа 32 retusa, Уол Пе завои cocti له د‎ ise UMEN а. Анди ао eb ipae 49 I OOA E E EA aE AA o EA E EE 47,49 TAS SVE PNAN ጋሪ22፡.....፡...፡.፡፡፣፡.፡..፡....፡. 20 СЗУ Кико боза се (VIZA رس‎ еи 28,29 Каса И пена (ОКОЛО) شه‎ а 28,29 Касас o (ЕО SE 28,29 Сау 1071011901. E Ои 29 КИУА O TB OUI NE EMEL EEE OES 45 О AA ЕВА T E Со, 46,50,51 TOISSW CLT ARANA ОИ ОЕ 41 ASL LL A O N DO موه ره وره‎ Ree ን Ж КОРООСУ ያር IVO OOE E те ከክ соо АЛА: 51 HUDHOUMCLO AI OV NOUN ያ... а 51 S [forereef sand channel population of modern Stephanocoenia MSO oeste ee Dee Se URS ИА isx[septalieyeles]! EROS у oli ón 25,26,27,29,31--34,36--43 SAS CLUSTER procedure, average linkage method .......... 12,13 A 19፡11. ШШЕ ын С HTC TR 8,12 SAS PRINCOME procedure 92) 8526892202. inme ES 12 Зап ера ои вени овим ооо ... 5-7,10,18 Saunders, Jung, and Biju-Duyal. (1986). Mae Ге TES LII 6-10,16,18,21,23,24,26,44--47 SCC [standardized canonical coefficients] ........................ TES SCHEGI (Os VE LO Dass os PA O оа 7 БОЛТИ АЙДОШ COU о ое A SOARES RO 51 БОРЕ ОСА) ор оса 32 senni, Sphenotrochus (Eusthenotrochus) .... 27,29,36 septentrionalis, Terebratulina .................. 47,49 ВОШЕЙ MU ооа مهنهسمهمموه‎ TT 7,19 Siderastrea siderea (Ellis and Solander, 1786) ...................... 11 Суарес ТАС По Еда по 2. A ጋ... 11 За об Ен STOOPID АИ оао ИИО ИТИ. 20,21 СИА Parae AUS и are ora 9. 2:6 23,27,29,34 Smilotrochus dominicensis Vaughan in Vaughan and Hoffmeister, 192599 ለ сле нис pert aE ERR ረም. 36 Smilotrochus ? dominicensis Vaughan in Vaughan and Hoffmeister, ROD SE TA ен Ог дине АСОИ RRA OSE 36 SHIH РОВ уллы Nerei bo оо ЛЕ И СТ ДАЛ с ር 19 DOMINICAN REPUBLIC NEOGENE. 4, 5, AND 6 73 Solenastrea Milne-Edwards and Haime, 1848a .................... 20 Spearman’s rank order correlation coefficient ......................... 8 Sphenotrochus Milne-Edwards and Haime, 1848b ............ 35,36 auritus Pourtalés, 1874 36 auritus var. Pourtalés ....... ха Hae а ки دن‎ gilchristi Gardiner, 1904 55 mosen Welse озо н сна ме 35 ТЇ, Spr OF Vaughan hol orem ое 36 Sphenotrochus (Eusthenotrochus) Wells, 1935 ...................... 35 brassensis Vaughan in Vaughan and Hoffmeister, 1925 ...... 36 Зет A A е Оо 27,29,36 ?Sphenotrochus auritus var. of Pourtalés .............................. 36 SPO [species 1, fossil ММВ Stephanocoenia spongiformis] ..... Ша spongiformis, Аа EN AS ЕРОН ТОЕ 6,20 Stephanocoenia ............. 2,84. 5,7-9,12-14,17,20,21,22 SPSS ING (1983) qus rere ዱሙ ር اک‎ е 12 1 ኤች discriminant procedure ................................ wes 1.2) Squifes (1958) .... ም ች ودند‎ 19 ST [septum thickness (major)] ................. 11,12,13,15,16,20-22 Sie ВАО O ае по ке века و دورس‎ 39 Stanley ТОВА) остри a ene ee ret Ноа Е зи Station Marine d'Endoume, Marseille, France Stephanocoenia Milne-Edwards and Haime, 1848a dove em UE ME 5-8,10-14,16,17,18,19,20,21 dendroidea Milne-Edwards and Haime, 1860 .................... 20 duncani. RSD a eA SOT une 5,7-10,12-14,16,17,21,22 fairbanksi Vaughan, 1900 ¡NTE EPIA АРЕНИ EL EE UE intersepta (Lamarck, 1816) En CEU SER у, 1 ያያ መያ Vale, Вааа «Наше 20:21 michelini Milne-Edwards and Haime, 1848b 19 michelinii Milne-Edwards and Haime, 1848b 19 réussi Duncan; ОЛ о. 20 SDONZION MIS: (IDC A Hh Der USE 2,3,4 ...... 5,7-9,12-14,17,20,21,22 ТЕЙЕШ СА По пи сине карта ыш а ра 20 Stephanocoenia ? decaseptata Weisbord, 1971 ...................... 20 Stylocoenia Milne-Edwards and Haime, 1848a ..................... 18 pumpelyi N aughan LIO. нь. 18 Stylocoeniella Yabe and Sugiyama, 1935 ............................. 18 SUI [State University of Iowa, Iowa City, ІА, U. S. АЈ .. 7,11,23 SUIS ON PERO COCA SEC E TIE 7 Switzerland: Веке a аал как: Rd IM TU I NE 7 1311 о A е 9 апр Уеа Аца е и 7 Os on оо (919/65) пика КЕШ уз NIS 43 ТООТПОС AE еа омо ETE. rM MU 20 Mo Топ Ве 50 TerobratelldmaMurs ул ОВ ла са ги есери i eee eet 50 Tierebratulaslecta Guppy SUO Сен e ጋፕ .ጢ..ሖጊ. ር 48 кесе рта Шасеа ета ПАО UN Жш... пуни ፡፡.:፡: Шетергалишчав ударена OSS аса ое Terebratulidae Gray, 1840 ............ Terebratulidina Waagen, 1883 Terebratulina d'Orbigny, 1847 caen око OOS ел ои а оке след RISAS TILO e a ro CERE ак уа cue Tation D ООО а retusa (няне 39) e SOA septentrionalis (СО Шоу; 1з 47,49 шегсогагиитат pamer Фоорев, 12/2..፡.፡፡፡፡፡፡፡፡..፡ 49 Зегеогатеай тава Gray; T340 ос ob te RI TUER. Бев задай Thecidea mediterranea Risso, 1826 си Sa Тес Час тау ISAO а A DEREN fa solene BUNU መቻች ች2::: НА NOE дшш Thecideidae Gray, 1840 .......... et СЕ 5. TibeordeidinasBllo51958..- иа а аиа 7 ረ. ፣.....፡፡፡ ፡፡፡.ዩ. око лас barrani Davidson, LIG eae aaa Thecidium mediterraneum (Risso, 1826) Thomson (1926) Thomson (1927) VE O EEO E ЛО Т T EEE EE E ee Tichosina? рае ай 49 lesta (Guppy TIBOR Е ta. 44,47,48,49 ARO My MO e aaa 7,19 URGING AG sarees аска eo E сад 18,36,39,48,49 SAM еј дип аа oe tA Mem MM EC ILIA 48 Бато усу дый M NM مه‎ EO 2 2 Taa а О с ы л a иа 38,41 Trochocyathus Milne-Edwards and Haime, 1848b ................ 31 ORO IIS DIREN SOE EL 37 cornucopia (Michelotti, 1838) ix © imparipartitus (Milne-Edwards and Haime, 18485) ............ 32 DLOf Maus Duncan, BOA Se. е 39,40 ДРАМА OL DO Uta CS Пеј West e сон, E А АДИ оа С и due 25 Trochocyathus (Paratrochocyathus) СИЕ о ИЕ E OURO DE SPO O ам в 29,2923 32 JAS CITO GAINS QD лу o бе 32 TMS aims O a 32 ТООТ РОА БЕ КОО al 32 ?Trochocyathus cornucopia (Michelotti) ............................... 31 TSL [tertiary septum length] ........................ 11,12,13,15,20-22 ET [total theca ащ a 11,12-15,20-22 TU [Tulane University, New Orleans, ГА, U S. A] ................... КЫМ cues аз ска ксы وو‎ 23,25,26,27,29-33,37,38,40-45,48,50,51 Turbinolia duodecimoostata ር)... По 33 папса успео ПВ 35 Turbinoliinae Milne-Edwards and Haime, 1848b ......... 27,35,37 ПОЛЕ Рата SERERE e ан سم‎ 34 Wr 5. Coast Survey Steamer ВАО. 43 WS. Coast Survey Steamer BURE <5 tugs Сеа as 42 UCMP [University of California Museum of Paleontology, Berkeley, GASES MAGIC on n. هسررو د‎ a መወ ሃሪ UI [University-of Illinois, Urbana; IL, U. S. АДАД 7 United States National Science Foundation ........................... T University of New BEUDSWIGR Lissi еы نند‎ opere ን 45 USGS [United States Geological Survey (specimens deposited at the INIVIINUED) cd ы. 24-26,27,28-32,34,35,37-43 USNM [United States National Museum of Natural History, Wash- DELO ОКО ШС: АЕ NR 7,27,28-43,48-51 Маут o ера TS а ds 45 VULGI Ва. 27,28 Vaughan (1900) Vaughan (1907) Muir ШОШО) cm E 7,18,20,21,25,33,34,36-41,43 Vaughan (1925 [in Vaughan and Hoffmeister]) ........................... Мана напа ЕЕ до cs a ey a Vaughan (date unknown) 74 BULLETIN 328 Vaughan and Hoffmeister (1925) ................... 7,25-28,31,36-40 Vaughan and Hoffmeister (1926) ............................... 36,38-41 Vaughan and Wells. (1943)... 11,18,19,27,37,43 Vaughan and Woodring (1921) .................... 25,34,35,37-41,43 Vaughan et al. (1921) Vero add Pichon (1976)... 297/5595”) ን k hen i nage 665.3 ር ee eee o OA ORR SR 48 a ПО A ое оа осо 20 Weisbord (1973) Wells (1932) A ece Пен а МА SES e ee e E MN E Wells (1935) Wells (1937) Wells (1941) Wells (1945) Wells (1956) Wells (1973) о о 7 West Africa West Germany, Darmstadt Wes i OO И начи 28,31,33,36,37,41,44,47,49 Westen Atlantic TOBON o arrini i E eir ene 10,24,35,42,47 Wiliams ай RO Well) :/ነነ”፡፡፡፡ 47 vade AA S Ugly Anda: (LISS hre ei i O не pcs 18 CLOW eS Mal ees о овоза жн от 33 Vans c ОВ ја г о E саа 18 ОВА Шо е ጋን ንን. 45 TADO TOS LO Оре ን ር lps 35,42,43 7 NAVAUS а а Бета Је а (1992) oes ው седеа 10,20 PREPARATION OF MANUSCRIPTS Bulletins of American Paleontology usually comprises two or more sep- arate monographs in two volumes each year. This series is a publication outlet for significant longer paleontological monographs for which high quality photo- graphic illustrations and the large quarto format are a requisite. 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