pea Prete ? tes a OF Ony ad Anth : an mn ma 9 Tran llins 125 = . el S. on = Sito Soe : oh ees 1 ae oe) = A i-4 = = =) Z. b PALEONTOLOGICAL RESEARCH INSTITUTION Officers PPRBSUOENE ceric dee tere even na} SAS UTE EN et al ehevled iat atelarbatte alr Sealy peice elena SHIRLEY K. EGAN BURST) WIGE=PRESUDENE 5% ia isis ana G oud E rehcer at seta ley chia, ed anemone tara neiae JouHN C. STEINMETZ SECOND) VIGE-PRESIDENY elie lalla eo oi2ul Aiauey pias ere teeter ie vaca tal ea THOMAS E. WHITELEY SECRETAR OH i ate lis eke AME MRE a LAC Nk ao ads a eee Gane nea Henry W. THEISEN PU RRIZASUIRE RE Se Ae ee ire) is ea sh bareuiclnes tas eet habiccevie sah ala Howarp P. HARTNETT ERECTOR) Gites Shah ei ercelates ieee eae whales chal at rte vie cab an aueeobette oma |e Roh eeatreees WaArREN D. ALLMON Trustees CARLTON E. BRETT MEGAN D. SHAY WILLIAM L. CREPET Mary M. SHUFORD J. THomas Dutro, Jr. ConsTANCE M. SoJA SHIRLEY K. EGAN JouHN C. STEINMETZ Howarp P. HARTNETT PETER B. STIFEL Harry G. LEE Henry W. THEISEN Amy R. McCuNnE Tuomas E. WHITELEY PHILLIP PROUJANSKY Trustees Emeritus Harry A. LEFFINGWELL RosBert M. LINsLEY SAMUEL T. PEES Epwarbp B. Picou, Jr. JOHN PoseTa, JR. JAMES E. SORAUF RAYMOND VAN HOUTTE WILLIAM P. S. VENTRESS BULLETINS OF AMERICAN PALEONTOLOGY and PALAEONTOGRAPHICA AMERICANA WSR REBUN ATETINION G54 02 Stevan td avila a) pois inva ial aa te Landi eet ahec ea eel eels eee euehioe atc Epitor A list of titles in both series, and available numbers and volumes may be had on request. Volumes 1—23 of Bulletins of American Paleontology are available from Periodicals Service Company, 11 Main St., Germantown, New York 12526 USA. Volume 1 of Palaeontographica Americana has been reprinted by Johnson Reprint Corporation, 111 Fifth Ave., New York, NY 10003 USA. Subscriptions to Bulletins of American Paleontology are available for US $150 per year (individual or institution) plus postage. Issues are available and priced in- dividually. Numbers of Palaeontographica Americana are priced individually. for additional information, write or call: Paleontological Research Institution 1259 Trumansburg Road Ithaca, NY 14850 USA (607) 273-6623 FAX (607) 273-6620 www.englib.cornell.edu/pri This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). Begun in 1895 NUMBER 357 DECEMBER 31, 1999 A Paleobiotic Survey of Caribbean Faunas from the Neogene of the Isthmus of Panama edited by Laurel S. Collins and Anthony G. Coates Panama Paleontology Project Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York, 14850 U.S.A. ISSN 0007-5779 ISBN 0-87710-449-2 Library of Congress Catalog Card Number: 99-76998 This publication is supported in part by a Corporate Membership from Exxon Exploration Company This publication is contribution number 9 to The Program in Tropical Biology at Florida International University. Note: Beginning with issue number 356, Bulletins of American Paleontology is no longer designating volumes. The journal will continue to publish approximately 2—4 issues per year, each of which will continue to be individually numbered. Printed in the United States of America Allen Press, Inc. Lawrence, KS 66044 U.S.A. CONTENTS Page Introduction aurelesaGollinsiandtAnthonyi Gy, Coates ye stacsye cou ress reer rca Pos ass) 2 Pees ta Feo eee a ence aan oor secu erte wae) abe Rene iss @ eigen) Bie TSr 5 PART 1. STRATIGRAPHY AND PALEOENVIRONMENT Chapter 1 Lithostratigraphy of the Neogene strata of the Caribbean coast from Limon, Costa Rica, to Colon, Panama SELON Crew CO ALES west etre et ewcn weer sein Ronee ay ite se ise ytepheg oe che Rts aayst ohn cease Ne Te tea aa ROE ERS ON MOM Seecteg ew Stole: don/nsued ai uch/aecvie ici erate 17 Appendix to Chapter 1 Newest Biostratigraphy Mane-bictre-AubiyxangdawialliameAs See Oren. rac sauhie er craier cite eee etek ere rete © lone meor a toue ete abe Tatas, MGM Ai ny SPoNen a paTAcnG Renters No cate 38 Chapter 2 Neogene calcareous nannofossil biostratigraphy of the Caribbean coast of Panama and Costa Rica [EDEN NY LS hate) I SESS ewe ae Ie Came een eee eoi es tr uusus brs 6:0 AS ie eee, beh oeeLs. b. a cain yc Gea CHOI ALOR Nree EEE Aces orc or Dade maEMEre rae 41 Chapter 3 Neogene planktic foraminiferal biochronology of the southern Central American isthmus NiatheweA AC Otome ya terre tadstic che eet hes Sate cee eet Cees etc cen Su eEN le ettee, Se EA 3 ke ee Sa ee eee 61 Chapter 4 A paleoenvironmental analysis of the Neogene of Caribbean Panama and Costa Rica using several phyla aurclnses Collis OranveleAcuilerayeamelay. Bomeand StephensDGaims ase cits peers cle) ieee ten eee aie ete sneer en enn enters 81 PART 2. PALEOBIOTIC SURVEY Chapter 5 The Miocene to Recent diversity of Caribbean benthic foraminifera from the Central American isthmus AIT CIE Sen OMIM S ght trace ick eon oy arses eye Te he Soc ay Ae ygee Suey GL EMS Ee! Fever ss GLE es Vee Fl ee aR SES UR ROA Scere 91 Chapter 6 Stratigraphic distribution of Neogene Caribbean azooxanthellate corals (Scleractinia and Stylasteridae) DtEPHeng Py NC AMS i oon re, ces to usta Sa pee Gover ivey ok avs, es siexet suspen stsi sboldie cuties shame Mach lent tt ceees Rieter ins, She ecLon ee aCe ee attests 109 Chapter 7 Pliocene to Pleistocene reef coral assemblages in the Limon Group of Costa Rica Ann EeBudG. Kenneth G Johnson, Dhomas A. Stemann and) Bridget HL Tompkins) sss ene: .) seins oo ene ore = = Gene eee 119 Chapter 8 Neogene cheilostome Bryozoa of tropical America: Comparison and contrast between the Central American isthmus (Panama, Costa Rica) and the North-Central Caribbean (Dominican Republic) Alans @heethams jeremy b.G. Jackson JoAnn sannen and, Yara» Ventocillatys: saacpny- aes eins oy st cise sl oictics helo ste) aitaseeremaiemee es cae 159 Chapter 9 Diversity and assemblages of Neogene Caribbean Mollusca of lower Central America Jeremy 3.C. Jackson, Jonathan) Aw Todds Helena\Fortunato)and|\Peter Jung. 420.52 sh ee ee ee ee ie ee ee 193 Chapter 10 Neogene-Quaternary Ostracoda and paleoenvironments of the Limon Basin, Costa Rica, and Bocas del Toro Basin, Panama Eanes Borne-slhomasy iy Croninuand J osephib Hazel eyes: 22. -veasgeiewadess ie le eceea sy one iouea sh egcisucuucierials, weirs, Musici aue euler euer ie ere 231 Chapter 11 Bathymetric distribution of Miocene to Pleistocene Caribbean teleostean fishes from the coast of Panama and Costa Rica WranvcleApiierdsangel (onc yROdM PUES) de PAS UILGLaya trea tet aren cae tis soa. a) Se yas See ee anne ay eBid AL GRITS Une LUNE coiaa aes 251 Chapter 12 A data model for the Panama Paleontology Project ISBT YG LLU CTO DE TAT ec Bree A sSoney tape o ier ty OL RMENoN eee ORCI Oct ee mI Chee REIEA RCT nG tcinG os REP en net Goria CR keen eee ee aE 271 Appendix A Maps NTMI INY (Ge CORTES Ha SB aie Bes Sin OSRie ONO CANO O le Chote la Cece Ieee chraict SVN CT aan Oi OnC no NG AS enn ae ener tien eae oa 287 Appendix B Stratigraphic sections PAEICRIOMLY gm Crem ORLESMeMea eta aey celeb sing Raye liane tesoten fete eee ey such ale eas ells Gc kel cu anne ce PereancRe mae Gree stele nee ala so an Me eee ke 299 ETC 5S RM SEER tcl ae A eccin aies arene SI cya cach eral oh eh crak er-S Gh ame ameticas ales dala! osu Casiaicaderzolictres Sah sire ldgé ules tie aness er 8 oheerei erate’ 349 vw tarot : ~ P= - 2 J ’ i) a’ & i! mesyiteaes 4d.) @mdiy . ovlty lingtme, endl 7 =. CO? tae Gre well jane? ary rca sia) 1 Ages “ tarot ee eros a ony eve Papi dole sim connie ee ; . ‘ae bapewis — a! 7 i + Saale aaa mn cnc sre m. Paste cp totn ete i Be 2c. niaienmamaresenanen hadi lf a : - INTRODUCTION LAUREL S. COLLINS Department of Earth Sciences Florida International University Miami, Florida 33199, U.S.A. AND ANTHONY G. COATES Smithsonian Institution Smithsonian Tropical Research Institute Washington, D.C. 20560-0580, U.S.A. A fundamental question in biology concerns the ex- tent to which populations and communities are affect- ed by geographic isolation and environmental change, a full comprehension of which must include under- standing environmental conditions and biodiversity of the past. The main research goal of the project that produced this volume is an assessment of patterns of changing marine invertebrate faunas of tropical Amer- ica over the last ~10 million years, for the purpose of determining the impacts of environmental change and genetic isolation on large-scale evolution and ecologic systems. This multitaxonomic paleobiotic survey takes advantage of a “natural experiment,” the Miocene constriction of the Caribbean-Pacific seaway and the Pliocene emergence of the Isthmus of Panama, which resulted in biotic isolation and changes in oceanic con- ditions on opposite sides. We initially concentrated on southern Central America because the bulk of evi- dence indicates that this is where final isolation of the tropical Atlantic and Pacific occurred. In this region, the biological effects are likely to have been most pro- nounced and directly relatable to the physical, sedi- mentary record of isthmian emergence. A remarkably complete record of these events is preserved in Neo- gene sediments of the region, including abundant, di- verse and well-preserved macrofaunas and microfau- nas. In addition, the Recent lies at the end of this time range, providing extant collections for comparative an- atomical and molecular studies. The Panama Paleontology Project (PPP), was initi- ated to make the systematic, regional fossil collections and fine-scaled chronologic framework necessary for these investigations. All geographic, stratigraphic, and taxonomic data are integrated in the PPP Database. From these and and other data, paleontologists are documenting biodiversity, biogeographic change, and the origination and extinction of tropical American or- ganisms, and relating these to patterns of environmen- tal and tectonic changes. THE PANAMA PALEONTOLOGY PROJECT The PPP is a geographically, chronologically and logistically large-scaled endeavor that has taken con- siderable time and effort to develop. The advantage of a coordinated project is that it can take a multitaxon- omic, integrated approach to investigating evolution- ary and environmental processes. The project currently involves 35 scientists from 20 institutions in 7 coun- tries (see the PPP internet site at http://www.fiu.edu/ “collinsl/), although many more have participated dur- ing its existence (Table 1). The PPP organizes expe- ditions to collect fossils and measure geologic sec- tions; prepares and curates macrofossils and microfossils from standardized, random samples; as- signs ages using microfossils, paleomagnetics and ra- diometric dating; and reconstructs paleoenvironments based on microfossil and macrofossil assemblages, sedimentology, and stable isotopes. The maintenance and development of the PPP Database and the exten- sive collections support longer term taxonomic, sys- tematic, ecologic and evolutionary studies. Below we describe the organization of the project. This formal collaboration began in 1986 with a re- connaissance survey of the Neogene geology of Pan- ama by Jeremy Jackson and Anthony Coates. The ob- jective was to determine whether the fossils were suf- ficiently abundant, both stratigraphically and geo- graphically, for research on the evolutionary and ecological consequences of the rise of the Isthmus of Panama. In 1987, Peter Jung and Laurel Collins joined the project, which became known as the Panama Pa- leontology Project. This group, with the addition of Ann Budd in 1993, formed a steering committee to plan collecting expeditions, seek funds, devise guide- BULLETIN 357 Table 1.—Members*, field participants and assistants in the Pan- ama Paleontology Project, 1986-1999. Ann Budd* Anthony Coates* Laurel Collins* Jeremy Jackson* Peter Jung* Teresita Aguilar Orangel Aguilera* Laurie Anderson* Marie-Pierre Aubry* Guillermo Barbosa Peter Baumgartner William Berggren* Pamela Borne* Laurel Bybell* Alan Cheetham* Stephen Cairns* Mathew Cotton* Timothy Collins* Thomas Cronin* John Dawson* Stephen Donovan* Harry Dowsett* Helena Fortunato* Andrew Gale Dana Geary* Thor Hansen* Antoine Heitz Nelson Jimenez Kenneth Johnson* Karl Kaufmann* Patricia Kelley* Susan Kidwell Michael Kunk Lorena Lanza Peter Marko* Donald McNeill* Jorge Mideros Daniel Miller* Simon Mitchell Richard Mooi* Galo Montenegro Steering Committee Hermatypic corals, taxono- my database Stratigraphy PPP Database, benthic fora- minfera, stable isotopes Scientific coordination, bryozoans, mollusks Mollusks Scientists Mollusks Teleost fishes Corbulid bivalves Calcareous nannofossils, biochronology Regional geology Tectonics Planktic foraminifera, biochronology Ostracodes Calcareous nannofossils, biochronology Cheilostome bryozoans Ahermatypic corals Planktic foraminifera, biochronology Gastropods, molecular biology Ostracodes Ahermatypic corals Echinoids Planktic foraminifera, paleoceanography Strombiniid gastropods, taxonomy database Facies analysis Strombid gastopods, stable isotopes Mollusks Mollusk curation and taxonomy Calcareous nannofossils Hermatypic corals, data analysis PPP Database Mollusks Stratigraphy Radiometric dating (Ar39/ 40) Regional geology Arcid bivalves, molecular biology Magnetostratigraphy Petroleum geologist Muricid gastropods, mollusk taxonomy Stratigraphy and sedimentology Clypeasteroid echinoderms Petroleum geologist U.S.A. U.S.A. U.S.A. U.S.A. Switzerland Costa Rica Venezuela U.S.A. France Costa Rica Switzerland U.S.A. U.S.A. U.S.A. U.S.A. U.S.A. U.S.A. U.S.A. U.S.A. U.S.A. England U.S.A. Panama England U.S.A. U.S.A. France Ecuador U.S.A. U.S.A. U.S.A. U.S.A. U.S.A. Nicaragua U.S.A. U.S.A. Ecuador U.S.A. Jamaica U.S.A. Ecuador Table 1.—Continued. Ross Nehm* Marginellid gastropods U.S.A. Florin Neumann Dinoflagellates Romania Hiroshi Noda Mollusks Japan Jorge Obando* Regional sedimentation Costa Rica Luis Obando Regional stratigraphy and Costa Rica geology Marta Ordonez Foraminifera, biostratigraphy Ecuador Dawn Peterson* Ostracodes U.S.A. Stephen Schellenberg Sr isotopes of reef corals U.S.A. Jay Schneider* Cardiid bivalves U.S.A. John Sutter Radiometric dating (Ar39/ U.S.A. 40) Paul Taylor* Cyclostome bryozoans England Jane Terranes* Stable isotopes of mollusks U.S.A. Jon Todd* Polystirid gastropods England Pascal Tschudin* Glycymerid bivalves Switzerland Italo Zambrano Palynology Ecuador Jijun Zhang* Planktic foraminifera, Canada biochronology Research Assistants Dione R. de Aguilera Sample processing for fishes Venezuela Raul Brito Student assistant Ecuador Eric Brown Field assistant Panama Martin Brunner Student assistant Switzerland Magnolia Calderon Sample processing Panama Rogelio Cansari Field guide Panama Daniel Castaneda Field guide Panama Sebastian Castillo Boatman Panama Janet Coates Field logistics U.S.A. John-Mark Coates Field assistant U.S.A. Chena Cooke Field logistics Panama Luis Cruz Field assistant Panama James Diaz Student assistant U.S.A. Beatrice Ferrenbach Field logistics Panama Lucien Ferrenbach Field logistics Panama Xenia Guerra Research assistant Panama Karl Hansen Photographer U.S.A. Huichan Lin Nannofossil processing U.S.A. Dorotheo Machado Field assistant Panama Claudia Mora Field assistant Costa Rica Angelica Munoz Field guide Nicaragua Agustin Paladines Student assistant Ecuador Rene Panchaud Field assistant, collections Switzerland manager Betzabeth Rios Sample processing Panama Fabricio Sierra Student assistant Ecuador Omar Sugasti Field assistant Panama Bridget Tompkins Student assistant U.S.A. Sophia Velotti Sample processing Panama Yira Ventocilla Sample processing Panama Jamie Wineberg Student assistant U.S.A. David West Research vessel, Captain Panama lines for preparing collections, coordinate studies of taxonomic groups, and organize joint publications. U.S. Geological Survey paleontologists Bybell (cal- careous nannoplankton), Dowsett (planktic foraminif- era) and Cronin (ostracodes), together with graduate student Cotton (planktic foraminifera), contributed the PPP’s biostratigraphic foundation. The question of an INTRODUCTION: COLLINS AND COATES Tl adequate fossil record was answered affirmatively by Coates et al. in 1992. Fossil collections are most useful to researchers when they reside at centralized locations. An agree- ment was signed between the Smithsonian Tropical Research Institute (STRI, the home institution of Jack- son and Coates) and the Naturhistorisches Museum Basel (Jung’s institution) that all of the mollusks and less abundant groups (crustaceans, echinoderms, bra- chiopods) would be prepared and accessioned in Basel, and that all bryozoans, corals, foraminifera, calcareous nannofossils, and ostracodes would be permanently housed, after study by the appropriate specialists, at the U.S. National Museum of Natural History, Wash- ington D.C. By 1990, growth in the volume and completeness of the collections required new procedures and a broadened taxonomic expertise. To census the macro- fauna, full-time “factories” at STRI and Basel were established for processing bulk sediment samples tak- en at shell-rich sites. The data on locality, stratigraphy, age, sample processing, and identified taxa began to be tracked in the PPP Database designed by Kaufmann (Chapter 12) and Collins. New colleagues began to study PPP bryozoans (Cheetham), mollusks (Geary, Anderson, Schneider), corals (Cairns), and regional sedimentation (Obando). Since 1993, the PPP has developed into two (over- lapping) research groups, a division reflected in the two parts of this volume. The first group establishes a chronologic and paleoenvironmental framework for each region, and the second builds its paleobiological studies upon this framework. The stratigraphic part of the first group consists of Coates, Aubry (calcareous nannofossil biostratigraphy), Berggren and Zhang (planktic foraminiferal biostratigraphy), and McNeill (paleomagnetics). Within the constraints of the physi- cal stratigraphy, the biostratigraphers use the evolu- tionary and paleoceanographic history of microfossils to establish a high-resolution chronology for strati- graphic sections. The ages of many sections are further refined by applying the chronology of paleomagnetic reversals. Paleoenvironmental determinations (Chapter 4) are based on the modern ecology of primarily ben- thic foraminifera (Collins), but also ostracodes (Borne, Cronin, Peterson), otoliths (Aguilera), ahermatypic corals (Cairns), and sedimentology. The second PPP research group includes members conducting macrofossil and microfossil studies of evo- lution, biogeography and ecology. For the mollusks, by far the most diverse group, Jackson coordinates the analysis of faunal lists of genera and subgenera that have been taxonomically standardized by Heitz, Jung and Todd. Several molluscan clades with modern tran- sisthmian distributions are being studied with morpho- metric and/or molecular techniques by Anderson, For- tunato, Jackson, Marko, Miller, Nehm, Schneider, Tshudin and Todd. Additional paleobiological research includes that of Aguilera (otoliths), Borne and Peter- son (ostracodes), Donovan and Mooi (echinoderms), Budd, Johnson, and Stemann (reef corals), Cairns and Dawson (ahermatypic corals), Cheetham, Jackson and Taylor (bryozoans), and Collins (benthic foraminifera). In addition to the PPP Database of information about locality, stratigraphy, age, and taxon occurrence (Kauf- mann and Collins), Budd designed and implemented a taxonomic database (Nmita) that contains information such as photographic and scanning electron micro- graph images on PPP and other tropical American pa- leontological collections. To obtain comparative Caribbean and Eastern Pa- cific collections, expeditions were undertaken more or less equally to both sides of the southern Central American isthmus during the first five years of the PPP (Table 2). In the next six years, most expeditions fo- cused on the relatively complete and fossiliferous Ca- ribbean sections of the Limon region of Costa Rica, and the Bocas del Toro and Colon regions of Panama. Many new formations were described, dated, and col- lected in detail to yield unparalleled collections of fos- sils from different stratigraphic levels and facies. The Caribbean stratigraphy and collections form the focus of this volume. In contrast to the Caribbean coast, the Pacific coast from the Darien (eastern Panama) to Nicaragua has yielded sequences that are less continuous chronolog- ically and not comparable in age, environment, and taxonomic diversity with the Caribbean faunas. The most important Pacific sequences are in the Darien (Middle-Upper Miocene) and the Burica Peninsula (Pliocene-Lower Pleistocene). Recently, to compensate for this inadequate record, the PPP began fieldwork in Ecuador, where richly fossiliferous, Neogene sequenc- es extend from the coast to the Andean foothills of the Borbon and Manabi basins. We hope to summarize all these sequences in a companion volume on the Neo- gene of the Pacific coast. BIODIVERSITY AND SCALE The large scale of the Panama Paleontology Project is the main characteristic that differentiates it from oth- er field-based paleontological projects. To study the diversity and distribution of taxa within a tropical ocean basin over ~10 million years at a relatively fine chronological scale requires years of constructing a re- gional stratigraphy and collecting and identifying fos- sils. For most taxonomic groups, collections have only recently become sufficient to calculate biodiversity re- Table 2.—PPP expeditions, 1986-1999. wh 26 27 28 29 30 31 32, 33 34 Begun 1/13/86 1/16/86 1/18/86 3/21/86 2/9/87 2/19/87 8/4/87 8/7/87 8/18/87 3/13/88 3/27/88 5/30/88 6/12/88 1/12/89 4/2/89 3/16/90 7/12/90 1/15/91 1/4/92 1/8/92 1/12/92 11/28/92 12/1/92 12/3/92 5/4/93 6/13/93 7/13/93 8/1/93 3/6/94 3/20/94 3/29/94 3/31/94 4/7/94 4/8/94 11/3/94 2/22/95 3/25/95 9/9/95 9/12/95 12/6/95 12/11/95 12/14/95 4/8/96 3/24/96 6/29/96 1/5/97 1/10/97 1/13/97 1/14/97 1/10/97 1/8/98 1/16/98 10/9/98 10/17/98 6/9/99 6/17/99 Country Panama Panama Panama Panama Panama Panama Panama Panama Panama Costa Rica Costa Rica Panama Panama Costa Rica Costa Rica Costa Rica Panama Panama Panama Costa Rica Costa Rica Panama Costa Rica Nicaragua Panama Panama Costa Rica Panama Panama Panama Costa Rica Panama Panama Panama Panama Costa Rica Panama Panama Panama Venezuela Venezuela Venezuela Panama Trinidad Costa Rica Venezuela Venezuela Venezuela Venezuela Panama Panama Panama Ecuador Ecuador Ecuador Ecuador Region Colon Burica Peninsula Bocas del Toro Burica Peninsula Lake Bayano Burica Peninsula North coast Bocas del Toro Colon Burica Peninsula, Nicoya Peninsula Limon Bocas del Toro Colon Burica Peninsula Limon Osa Peninsula, Nicoya Peninsula Darien Darien Colon Limon Burica Peninsula Colon San Carlos Managua Darien Colon Limon Bocas del Toro Darien Colon Golfo Dulce Burica Peninsula Bocas del Toro Burica Peninsula Bocas del Toro Limon Darien Colon Bocas del Toro Falcon Araya Peninsula Isla de Margarita Darien Manzanilla Limon Falcon Araya Peninsula Cumana Isla de Margarita Bocas del Toro Bocas del Toro Colon Borbon Manabi Borbon Manabi BULLETIN 357 Mode truck truck truck, canoe truck truck, boat truck STRI R/V truck STRI R/V truck STRI R/V truck foot, horses truck foot, truck, horses truck, canoe truck, canoe truck truck truck truck truck truck canoe truck car motor boat canoe truck truck truck motor boat truck motor boat truck canoe truck STRI R/V truck truck truck canoe car truck truck truck truck truck STRI R/V motor boat truck truck, canoe truck truck, canoe truck sults, most of which appear in this volume. The geo- graphic scale, or spatial resolution, of the research varies with taxonomic group. At one extreme, higher- level taxa that are quite diverse and widely distributed in the Caribbean (e.g., mollusks) require an enormous sampling effort for results that are meaningful at the scale of an ocean basin. At the other extreme, higher- level taxa that are less diverse and also more restricted environmentally, such as reef corals (~175 Neogene- Recent Caribbean species), require less sampling. Most taxa in this project fall between the two ex- tremes, e.g., benthic foraminifera are moderately di- verse and normally distributed across the entire Carib- bean (some globally), and cheilostome bryozoans are moderately diverse but more endemic. Differences in spatial and temporal distributions of taxa studied by the PPP are reflected by the approaches to evaluating diversity. Cheetham ef al. (Chapter 8, cheilostome bryozoans) and Jackson et al. (Chapter 9, mollusks) address the adequacy of PPP collections for calculating total diversity by plotting cumulative num- bers of the species recovered as a function of the num- bers of collections examined in each area. Whereas collections are still inadequate to determine total Ca- ribbean molluscan generic diversity per time interval, relative molluscan diversity for successive time inter- vals has been compared. The cumulative curves for bryozoan species show a slight flattening which sug- gests that total diversity is being approached, and Cheetham et al. identify trends in diversity for ade- quately sampled growth forms of cheilostomes. Col- lins (Chapter 5, benthic foraminifera) and Jackson both use Fisher’s alpha to measure local diversity, an ap- proach that avoids the enormous task of calculating total regional diversity. Cairns (Chapter 6, azooxan- thellate corals) combines PPP data with other data to address Caribbean species richness and evolution from Neogene to Recent time. AGES OF FORMATIONS EXAMINED BY THE PPP Ages have evolved over the course of the project and continue to do so. Time scales change (Berggren et al., 1985; Berggren et al., 1995), new exposures that represent older or younger parts of previously dated formations are discovered, the evolutionary and paleo- ceanographic history of microfossils becomes better known, and resampling previously dated sections and new paleomagnetic studies sometimes result in age re- finement. In this volume, biostratigraphy completed before 1993 (Bybell, Chapter 2; Cotton, Chapter 3) used an older time scale, as do most of this volume’s chapters, while current biostratigraphic research (Au- bry and Berggren, Appendix 1 of Chapter 1) has begun INTRODUCTION: COLLINS AND COATES 9 Table 3.—Ages of Caribbean stratigraphic units examined by the PPP, based on the time scale of Berggren ef al. (1995). Formation Member/facies/section Age (Ma) Cayo Agua Formation 5.0-3.4 Chagres Formation 8.6—5.3 Escudo de Veraguas 3.7-1.9 Formation Gatun Formation Lower part 11.8-11.4 Gatun Formation Middle part 9.4-8.6 Gatun Formation Upper part 9.4-8.6 Moin Formation 2.1-1.5 Moin Formation Empalme member 2.1-1.5 Moin Formation Lomas del Mar member 1.9-1.5 Nancy Point Formation 7.2-5.6 Quebrada Chocolate 3.7-2.6 Formation Quebrada Chocolate Buenos Aires member 3.3-2.6 Formation Rio Banano Formation* 3.8-3.0 Rio Banano Formation? Brazo Seco section** 5.2-4.3** Shark Hole Point 5.6-3.6 Formation Swan Cay Formation 1.8-0.8 Tobabe Formation 7.2-5.3 Uscari Formation Uppermost part 8.1-5.6 * Aubry and Berggren (App. 1, Chapter 1) and McNeill er al. (in press) include an extra section above that examined by Bybell (Chapter 2) and Cotton (Chapter 3). ** From McNeill ef al. (in press), this age (using Sr isotopes) is highly uncertain given the absence of age-diagnostic microfossils and anomalous results of Sr ages from samples in other Limon sections. to use the newer one. Most biostratigraphy (including microfossil zonation) based on older time scales can be transfered to newer ones, so that sample ages re- corded in the PPP Database use a single time scale. Table 3 summarizes current age estimates for the strati- graphic units surveyed by the PPP, based on the time scale of Berggren et al. (1995). ANALYSIS OF THE PALEOBIOTIC SURVEY DATA The paleobiotic survey was carried out as a series of separate surveys encompassing algae, protists, in- vertebrates and vertebrates. The major taxa, each rep- resented by a chapter in this book, are: calcareous nan- noplankton, planktic foraminifera, benthic foraminif- era, azooxanthellate corals, reef corals, cheilostome bryozoans, mollusks, ostracodes and teleost fishes. The records of nannoplankton and planktic foraminifera are not censuses of all species present because species are selectively identified from whole assemblages for bio- Stratigraphy. For the remaining taxonomic groups, all species or genera are recorded for each site, which has a unique PPP number. In this analysis, we combine the separate censuses into one data set, conduct cluster analyses of the assemblages in stratigraphic units, and explore the relative influence of age versus environ- ment, which helps in differentiating evolutionary and ecological changes. When combining paleontologists’ separate data sets, all occurrence data are necessarily standardized by the coarsest sampling and recording method. Because some taxa (e.g., bryozoans) are recorded only as pre- sent or absent, the other relative abundance data must be converted to presence/absence. Similarly, some taxa (e.g., azooxanthellate corals) are typically sparse at in- dividual sites or sections, so their sites are combined within each stratigraphic unit to address sampling bi- ases. The resulting data set records the presence or absence in stratigraphic units for benthic foraminifera (333 species), azooxanthellate corals (17 species), reef corals (89 species), cheilostome bryozoans (200 spe- cies), mollusks (1022 genera to subgenera!), ostra- codes (79 species) and teleost fishes (82 genera). The eleven stratigraphic units we analyzed are the: Gatun Formation, Chagres Formation, Nancy Point Formation, Shark Hole Point Formation, Cayo Agua Formation, Fish Hole section of Bastimentos Island, Escudo de Veraguas Formation, Swan Cay Formation, Uscari Formation, Rio Banano Formation, and the Lo- mas del Mar Member of the Moin Formation. Never- theless, there are missing data for taxonomic groups in many of these units. For example, reef corals are absent from most units, ostracodes are not censused in the Panama Canal Basin, and azooxanthellate corals are not yet recorded from the deeper-water units or Panama Canal Basin. Therefore, separate analyses are performed on various combinations of taxonomic groups and stratigraphic units. The cluster analyses use six different algorithms (Ward’s method and single, complete, centroid, average and median linkage) with only slightly different results; a typical result is figured below for three combinations of taxa and stratigraphic units. 1. In Text-figure 1 are the results of the only analysis that includes reef corals, as well as mollusks and fish. Bastimentos and Swan Cay, both in the Bocas del Toro Basin, are the most similar, and Lomas del Mar, in the Limon Basin, is the most different. This result (which holds with or without the corals) con- firms interbasinal differences noted previously (Collins et al., 1995), and appears to confound pre- dictions based on the Late Pliocene—Early Pleisto- cene turnovers in molluscan and reef coral taxa (Jackson et al., 1993; Budd et al., 1996). On the basis of age, Swan Cay and Lomas del Mar, de- posited after the turnovers, should be most similar: Bastimentos is 2.6—2.4 Ma, Swan Cay is 1.8—0.8 10 BULLETIN 357 Gatun Bastimentos Swan Cay Lomas del Mar | paca Knee Sl lana ee ee aca Sama | 00 0.1 02 03 04 05 06 0.7 Distances Text-figure 1.—Cluster analysis (complete linkage method) of the presence/absence of species of reef corals, genera of teleost fishes, and genera to subgenera of mollusks in the Gatun Formation (Pan- ama Canal Basin), the Fish Hole section of Bastimentos Island (Bo- cas del Toro Basin), the Swan Cay Formation (Bocas del Toro Ba- sin), and the Lomas del Mar Member of the Moin Formation (Limon Basin). Distances are Euclidean. Assemblages from the same basin are more similar than assemblages from the same age or bathymetry. Ma, and Lomas del Mar is 1.9—1.5 Ma. Units do not cluster by paleobathymetry, either: Bastimentos and Lomas del Mar are middle neritic, and Swan Cay is shallowest outer neritic with transported middle neritic material. 2. Text-figure 2 shows the similarity of five strati- graphic units using all taxa except reef corals. Age and environment affect the similarity of the units’ faunal assemblages about equally. Although the Rio Banano and Cayo Agua formations are most alike in environment (inner-middle neritic), the former is linked first to an outer neritic unit of a comparable, Early-middle Pliocene age, the Shark Hole Point Formation. However, the Escudo de Veraguas For- mation, which is Late Pliocene and mixed middle to outer neritic, is most similar to the first two units, suggesting that environment has a stronger influ- ence in this grouping than age. The unit that is most different in both environment and age, the reefal, latest Pliocene to earliest Pleistocene Lomas del Mar Member, has the most different faunal assem- blage. 3. Text-figure 3 is an analysis of ten stratigraphic units using only benthic foraminifera, fish and mollusks. It suggests that environment influences the similar- ity of assemblages more than age, although age and environment are somewhat correlated because of the progressive Neogene uplift of the Bocas del Toro and Limon basins. There are two main clus- ters. In the upper one, the Late Miocene, bathyal Uscari, Nancy Point and Chagres formations are most similar. The next shallowest units, the outer Cayo Agua Escudo de Veraguas Rio Banano Shark Hole Point Lomas del Mar a Sc i Lia. oe 1 0.0 01 0.2 03 04 05 06 0.7 Distances Text-figure 2.—Cluster analysis (complete linkage method) of the presence/absence of taxa in the Cayo Agua, Escudo de Veraguas, and Shark Hole Point formations of the Bocas del Toro Basin, and the Rio Banano Formation and Lomas del Mar Member of the Moin Formation in the Limon Basin. Included in the analysis are species of benthic foraminifera, azooxanthellate corals, cheilostome bryo- zoans, and ostracodes, genera of teleost fishes, and genera to sub- genera of mollusks. Distances are Euclidean. Age and environment have approximately equal influences on the similarity of assem- blages. neritic, Early Pliocene and Pleistocene Shark Hole Point and Swan Cay formations, are most similar to the bathyal units. The older, shallower Gatun Formation falls between the old, deep units in the upper cluster and the lower cluster of shallower- water, Pliocene—earliest Pleistocene units. Gatun Shark Hole Point Uscari Nancy Point Chagres Swan Cay Lomas del Mar Rio Banano Escudo de Veraguas Cayo Agua i Pa ia alae aaa | 0.0 0.1 02 03 04 05 06 0.7 Distances Text-figure 3.—Cluster analysis (complete linkage method) of the presence/absence of taxa in the Gatun and Chagres formations of the Panama Canal Basin; the Shark Hole Point, Nancy Point, Swan Cay, Escudo de Veraguas, and Cayo Agua formations of the Bocas del Toro Basin; and the Uscari Formation, Lomas del Mar Member of the Moin Formation, and Rio Banano Formation of the Limon Basin. Included in the analysis are species of benthic foraminifera, genera of teleost fishes, and genera to subgenera of mollusks. Dis- tances are Euclidean. Environment seems to influence the similarity of assemblages more than age. INTRODUCTION: COLLINS AND COATES iil We conclude from these exploratory analyses that there are strong age, paleoenvironment, and basin ef- fects on the similarity of PPP assemblages. Age re- flects evolutionary changes but is somewhat correlated with paleoenvironment because of regional tectonic uplift through time. Similarities of assemblages from the same paleoenvironments result from ecological as- sociations, and the basinal effect reflects more local- ized conditions. Analyses of this sort begin to disen- tangle evolutionary and ecological faunal changes for the ultimate purpose of isolating evolutionary events. These analyses are of multiple, higher-level taxa re- corded as presence/absence in stratigraphic units, and future analyses using relative abundances and a finer- scaled chronology will undoubtedly reveal other trends in evolution and ecology. CONTENT OF CHAPTERS The volume is divided into two parts. Part 1, Stra- tigraphy and Paleoenvironment, consists of four chap- ters on the formal lithostratigraphy, biostratigraphy, geochronology, and paleoenvironments of sediments from the Panama Canal, Bocas del Toro, and Limon basins. The chapters are summarized as follows: Chapter 1. Coates places the physical stratigraphy of Neogene sediments of Caribbean Panama and Costa Rica within a regional tectonic framework, incor- porating the units defined in Coates et al. (1992) and creating several new ones. In Appendix 1 of Chapter 1, Aubry and Berggren give the latest bio- stratigraphic data and chronological correlation of the new sections. Chapter 2. Bybell presents calcareous nannofossil data collected until 1991, and discusses their application to the geochronology of the formations described by Coates et al. (1992). Her research laid the founda- tion upon which the later biochronology was built. Chapter 3. Cotton presents data from planktic fora- minifera collected before 1993 and combines it with the nannofossil data for refined age estimates of for- mations. He correlates the Central American for- mations with other tropical to subtropical American formations from southeastern Virginia to Ecuador. Chapter 4. Collins, Aguilera, Borne and Cairns com- bine environmental assignments from four different phyla (benthic foraminifera, teleost fishes, ostra- codes, and ahermatypic corals, respectively) for most stratigraphic units. The results among taxa are remarkably congruent, considering the different life modes of the organisms (e.g., benthic versus nek- tonic; feeding at versus above the subtratum), as well as variations in technical and analytical ap- proach. The paleoenvironments for individual sec- tions are combined for an overview of larger-scale environmental change set within the region’s tecton- ic history. Part 2, Paleobiotic Survey, includes seven chapters that report the distribution of species or genera at PPP sites, and address topics such as faunal and paleoen- vironmental change through time. Some of the conclu- sions of Chapters 5 to 12 are summarized as follows: Chapter 5. Collins combines fossil and modern distri- butions of species of Caribbean benthic foraminifera from Panama and Costa Rica and shows that their diversity has doubled from the Late Miocene to Re- cent, through the time of seaway constriction, com- plete closure, and afterward. The proportion of taxa associated with carbonate shoals and reefs increased during this time, which agrees with the trend of in- creasing speciation in these ecologically restricted taxa. The largest faunal changes apparently occurred in the Pleistocene to Recent rather than the middle Pliocene, suggesting that complete seaway closure had little effect. Chapter 6. Cairns reports the stratigraphic ranges of 142 Caribbean azooxanthellate coral species, 101 of which are extant. The data suggest that the highest origination rate occurred in the Middle to Late Mio- cene and the highest extinction rate occurred in the Late Pliocene. Neither of these evolutionary pulses occurred near the time of complete seaway closure. Chapter 7. Budd, Johnson, Stemann, and Tompkins de- scribe the distribution of reef coral species from the Limon Basin, and identify different periods of fau- nal change that occurred at various Caribbean lo- calities during the Late Pliocene to Pleistocene. Chapter 8. Cheetham, Jackson, Sanner, and Ventocilla contrast assemblages of cheilostome bryozoans from both sides of the Central American isthmus with those of the Dominican Republic in an analyt- ical comparison of Caribbean and Pacific faunas. An unexpected result is that the complete closure of the isthmian seaway apparently had relatively little evo- lutionary effect. The authors also find a Middle Miocene to Pleistocene decline in the diversity of erect species, possibly associated with their growth on decreasingly available substrata such as seagrass. Chapter 9. Jackson, Todd, Fortunato and Jung control for sampling and taxonomic biases in an enormous dataset of Neogene molluscan genera to subgenera from Caribbean Panama and Costa Rica. Local mol- luscan diversity varied more than six-fold, and ei- ther increased or remained constant from the Mio- cene to Recent. Previous studies which identified a decline in Pliocene Caribbean molluscan diversity 12 BULLETIN 357 and associated it with seaway closure were based on inadequate sampling of the faunas. Chapter 10. Borne, Cronin and Hazel use assemblages of ostracodes from the Limon and Bocas del Toro basins to identify lagoon, carbonate platform, re- stricted nearshore, and outer shelf to upper slope facies. The distributions and morphology of several species suggest that cold, upwelling currents im- pinged on the Late Pliocene to Early Pleistocene Central American shelf. Chapter 11. Aguilera and Aguilera describe teleost fish assemblages at the genus level from otoliths, and infer bathymetries by comparison to living repre- sentatives of the genera. Several genera, found liv- ing only in the Indo- or Western Pacific, show that relict elements of Tethyan faunas persisted in the Caribbean until at least the Late Pliocene. Chapter 12. Kaufmann presents a data model that ex- plains the way in which the elements of the PPP function as a whole, as well as the working of the database of information on stratigraphy, geography, chronology, paleoenvironment, and faunal occur- rence. For complex projects, data models help clar- ify the relationships of the many, diverse parts. Appendices. Coates locates all PPP collecting sites used in this volume, plotting them on maps in Ap- pendix A. Each site is represented by a unique PPP number. In Appendix B, he places the sites strati- graphically in a series of 39 detailed sections. SUMMARY For the Panama Paleontology Project, the whole is much greater than the sum of the parts. Basing studies of multiple, higher-level taxa on the same, well-dated set of samples has provided many possibilities for in- tegrated research. A few of the conclusions the PPP can make thus far, based on the Caribbean collections, are the following: 1. Stratigraphy. There exists along the Caribbean coast of Panama and Costa Rica a series of expo- sures of richly fossiliferous, Neogene, shallow-wa- ter sediments which, when placed in stratigraphic succession, cover the late Middle Miocene to Early Pleistocene interval. 2. Chronology. The fossil collections can be dated biostratigraphically and paleomagnetically with a precision that is relatively fine for land-based for- mations, with age ranges for single samples varying between approximately 100,000 years and 1.5 mil- lion years. 3. Environments. The Panama Canal Basin was a shal- low Middle Miocene basin until deepening ~6 Ma caused an inflow of deep, Pacific water. The Bocas del Toro and Limon basins differed in sediment source, isobathyal microfaunas, and stable isotopes. They were similar in their back-arc setting, histories of uplift, and sedimentary sequences of bathyal Miocene mudstones, neritic Pliocene siltstones/ sandstones, and lower Pleistocene coral reefs. 4. Seaway closure. To date, research on PPP collec- tions shows no strong evolutionary response to the complete closure ~3.5 Ma of the Central American isthmian seaway, although an evolutionary turnover in reef corals did occur sometime between 4 and 1 Ma. Whereas the largest pulses of origination in the Neogene occurred for azooxanthellate corals and benthic foraminifera in the Middle and Late Mio- cene, during seaway constriction, they occurred in the latest Pliocene to Early Pleistocene for the mol- lusks, perhaps because of increased northern hemi- sphere glaciation. Complete seaway closure appar- ently had relatively little evolutionary effect on cheilostome bryozoans. 5. Biodiversity. From Late Miocene to Recent time, the diversity of molluscan genera either increased or remained constant and that of species of benthic foraminifera increased. From the Middle Miocene to the Pleistocene, the diversity of erect cheilostome bryozoans declined. ACKNOWLEDGMENTS Grants from the Biotic Surveys and Inventories Pro- gram of the National Science Foundation (grant num- bers BSR90-06523, DEB-9300905, DEB-9696123, DEB-9705289) provided the means to build the PPP Database, prepare large numbers of samples, track fos- sil collections, and assign ages and paleoenvironments. The National Geographic Society has consistently funded PPP fieldwork in Panama, Costa Rica, Nica- ragua and Ecuador. The Smithsonian Institution, STRI, Swiss National Science Foundation, and Naturhisto- risches Museum Basel have also funded fieldwork and the preparation of collections. This is contribution number 10 to The Program in Tropical Biology at Florida International University. INTRODUCTION: COLLINS AND COATES 13 REFERENCES CITED Berggren, W.A., Kent, D.V., Flynn, J.J., and Van Couvering, J.A. 1985. Cenozoic geochronology. Geological Society of America Bulletin, vol. 96, pp. 1407-1418. Berggren, W.A., Kent, D.V., Swisher, C.C., and Aubry, M.-P. 1995. A revised Cenozoic geochronology and chronostratigra- phy. SEPM Special Publication, no. 54, pp. 129-212. Coates, A.G., Jackson, J.B.C., Collins, L.S., Cronin, T.M., Dow- sett, H.J., Bybell, L., Jung, P., and Obando, J.A. 1996. Closure of the Isthmus of Panama: the near-shore marine record of western Panama and Costa Rica. Geological So- ciety of America Bulletin, vol. 104, pp. 814-828. Collins, L.S., Coates, A.G., Jackson, J.B.C., and Obando, J.A. 1995. Timing and rates of emergence of the Limon and Bocas del Toro basins: Caribbean effects of Cocos Ridge sub- duction? in Geologic and tectonic development of the Ca- ribbean Plate Boundary in southern Central America. Geological Society of America Special Paper, no. 295. P. Mann, ed., pp. 263-289. Jackson, J.B.C., Jung, P., Coates, A.G., and Collins, L.S. 1992. Diversity and extinction of tropical American mollusks and emergence of the Isthmus of Panama. Science, vol. 260, pp. 1624-1626. MeNeill, D.F., Coates, A.G., Budd, A.F., and Borne, P.F. 1999. Integrated paleontological and paleomagnetic stratigraphy of the upper Neogene deposits around Limon, Costa Rica: A coastal emergence record of the Central American Isth- mus. Geological Society of America Bulletin (in press). sete ‘ i F 2 é e ry Th a 1 * eel he © i : le |. qi Void 314 = Ce) «fw peerice ed be aA DP, (ied y rl > & wage! Gl ~~ < ied bow) ae ie ‘ sh Picheweviadi Wyityees, 0M let pat. Beking a4 y tah =! bia fmeenad i " ne os” oe a j ee . vend iwi ‘ Ps iu igh4 i ion, 7 cp y ii m of cui hey € Plaudatiede th | if notated iy tia Min ( omplicty MyMaWway = ely firthe eveiucemury- LitMAho 7 = ‘ap Peuly Lab Aen, be ey al neNvaren nemeta® aed px eens nad idea vf ators Wwrerdetl). Chan bs Puree ee hi ceanlity’« . te comcamneme, Serkipindah a Ae PART 1 STRATIGRAPHY AND PALEOENVIRONMENT | THAT TVG ADAIVPHIOSLTAS GMA YHIASOITAATS CHAPTER 1 LITHOSTRATIGRAPHY OF THE NEOGENE STRATA OF THE CARIBBEAN COAST FROM LIMON, COSTA RICA, TO COLON, PANAMA ANTHONY G. COATES Smithsonian Institution Smithsonian Tropical Research Institute Washington, D.C. 20560-0580, U.S.A. INTRODUCTION The Central American isthmus lies at the intersec- tion of six tectonic plates (Text-fig. 1; Burke ef al., 1984; Mann et al., 1990). The North and South Amer- ican Plates, with relative westerly and west-north- westerly motions, respectively, override two large oce- anic Pacific plates, the Cocos and Nazca, with north- easterly or easterly relative motions, respectively. The collision of these two sets of plates has formed, since the Cretaceous, a major zone of subduction along the western margin of the Americas, a segment of which constitutes the Central American volcanic arc. The southern Central American isthmus consists pri- marily of igneous and sedimentary rocks of oceanic crustal composition, generated by the Central Ameri- can volcanic arc. In this chapter, I focus on the stra- tigraphy of three important sedimentary basins that flank the magmatic arc on the Caribbean side (Text- fig. 2), namely, the southern Limon Basin in Costa Rica, and the Bocas del Toro and Panama Canal basins in Panama. These basins are dominated by volcani- clastic sediments, commonly with foraminiferal and nannofossil microfaunas. They also contain important Miocene through Pleistocene coral reefs, as well as a series of rich and diverse molluscan, bryozoan, fish (otoliths), and coral assemblages at many stratigraphic levels. In this chapter, I present a revision of the formal stratigraphy of the sediments in which the faunas oc- cur, including the definition of several new formations and their biochronology (Appendix 1 this chapter). Also included are 11 maps and detailed insets, show- ing the location of all samples (Appendix A, this vol- ume), and the computer-drawn logs of 39 sections measured across the three basins that show the strati- graphic relations of all the samples (Appendix B, this volume). The locations of the measured sections are shown in Text-figure 2. The Isthmus of Panama was the last portion of the Central American isthmus to emerge (Coates et al., 1992; Coates and Obando, 1996), closing the marine connections between the Caribbean and the Pacific about 3 Ma (Kaneps, 1970; Berggren and Hollister, 1973, 1974; Keigwin, 1978, 1982). The Panama Pa- leontology Project (PPP) set out to look for extensive upper Neogene fossiliferous sedimentary sequences in this region on the assumption that the sedimentary rec- ord here would track most closely the marine environ- mental and ecological changes caused by the emer- gence of the Isthmus. The stratigraphic sections and faunal samples ana- lyzed in this volume are located in back-arc basins (e.g., southern Limon Basin) or in marginal aprons (e.g., Bocas del Toro and Panama Canal Basins) de- rived from the Caribbean side of the Central American volcanic arc, the structure of which is shown in cross section in Text-figure 3. Although we originally un- dertook field expeditions to both Pacific fore-arc and Caribbean back-arc basins, the Caribbean sequences yielded more complete stratigraphic sections and more abundant and diverse faunal assemblages. This is largely due to erosion of many younger sequences on the tectonically active Pacific coast. Older sediments have been subducted or obducted onto the overlying plate and are either highly deformed or lost (Text-fig. 3). By contrast, on the passive Caribbean margin, the southern Limon, Bocas del Toro, and Panama Canal basins (Text-fig. 2) have yielded numerous diverse and abundant faunas. These sections are less deformed, and span a greater time interval than the Pacific sections. For example, the Pacific Burica Peninsula fore-arc ba- sin (Corrigan et al., 1990; Coates et al., 1992) has more than 4000 m of sediments, ranging from about 3.5 to <1.6 Ma, whereas the Caribbean Bocas del Toro Group has about 1000 m, ranging from 8.5 to about 1.5 Ma. In the region of the Talamanca Range (Text- fig. 1) in Costa Rica, subduction of the Cocos Ridge has elevated and structurally deformed both the inner fore-arc Terraba Basin and the now inverted back-arc southern Limon Basin (Kolarsky et al., 1995), as is shown in Text-figure 3. From 1986 to 1992, the PPP undertook a series of reconnaissance field expeditions to explore a number 18 BULLETIN 357 Mexico Caribbean Sea Honduras CHORTIS TERRANE CHOROTEGA TERRANE Middle eee . Nicaragua 23 A CHOCO TERRANE ,~ Talamanca Range North American Plate Lesser Antilles PANAMA MICROPLATE Panama Microplate jf South American Plate Nazca Plate > Text-figure 1.—Map of Central America showing location of the older Maya, Chortis, Chorotega, and Choco geological terranes, and the younger Panama microplate, picked out in thick dashed lines and gray stipple. The northern border of the Panama Microplate is called the North Panama Deformed Belt. Also shown is the Talamanca Range (dash-dot stipple) and the volcanoes of the central magmatic are (black triangles = active, open triangles = inactive). we tka! AY Km N 12° ) EE "| Caribbean Sea Oye Lees Panama Canal Basin \ Seay \4. | | rs’ Pacific Ocean EAMES Peninsula Ny ow f Colombia 78" 77° 83° Text-figure 2.—Map of Costa Rica and Panama showing location of the southern Limon, Bocas del Toro, and Panama Canal Basins (dark gray stipple). Numbers correspond to sections referred to in text and in Appendix B. LITHOSTRATIGRAPHY: COATES 19 Fore Arc Inner Outer is Fore Arc Basin Py Ridge | Osa Pacific Peninsula West Shoreline -. Subducted, Ay Deformed Sediments IS AIS, IRN INS ES Rs Ne Nae NAA Nem NmeN a Ren ae Neamt EN EXTRUSIVE OR INTRUSIVE Fore Arc Terraba Trough Magmatic At ap Talamanca Range Caribbean southern Limon Basin “AN INE IN INSTA AS Anes NaN SEDIMENTS IGNEOUS ROCKS pre fin 2) Tertiary lavas =z7] Miocene Granodiorite Pa Cretaceous volcanics [—] Eocene/ = Paleocene [7 Eocene (GA Miocene =] Oligocene/ 362) Miscene pag Oligocene Text-figure 3—Schematic geologic cross section from the Osa Peninsula on the Pacific coast to the southern Limon Basin on the Caribbean coast (see line on Text-fig. 2). Modified from “Hydrocarbon potential of Costa Rica’’, Ministry of Environment and Energy, Government of Costa Rica, 1996. of the Neogene sedimentary fore- and back-arc basins associated with the volcanic arc in southern Central America. Basins were surveyed on the Pacific coast, from the Nicoya Peninsula, northwestern Costa Rica, to Darien, eastern Panama, and on the Caribbean coast, from the northern part of the Limon Basin, Costa Rica, to the Panama Canal Basin, Panama (Text-fig. 2). Ina preliminary review of the litho- and bio-stratigraphy, Coates et al. (1992) established that a well-preserved and diverse marine fossil record existed on both coasts, containing nannofossils and planktic foraminifera ca- pable of yielding a precise geochronology for the late Neogene sediments. In 1993, with a view to more detailed comparisons of geologic history and evolutionary and ecological patterns, the PPP began a more extensive series of field expeditions. These focused particularly on the com- plete and richly fossiliferous sections of the Caribbean coast, specifically in the southern Limon, Bocas del Toro, and Panama Canal basins (Text-fig. 2) described here. The northern part of the Limon Basin is not treat- ed in this chapter because it is extensively covered by Pleistocene volcanic deposits and did not yield abun- dantly fossiliferous sections. Because the Miocene to Pleistocene sediments of the southern Limon Basin are relatively elevated and Structurally complex, the physical stratigraphy of this basin has been difficult to reconstruct (Text-fig. 3). The stratigraphic sequence has been studied mostly along rivers draining the foothills and coastal plain northeast of the Talamanca Range in the area around Limon (Map 11) and, to a lesser extent, further south as far as the Panamanian border (Map 10). Many new, ex- tensive and very fossiliferous Plio-Pleistocene sections were exposed during our field work, often only tem- porarily, by housing construction in the hills of the western part of Limon and by commercial construction along Route 32 between Buenos Aires and Limon (In- sets A, B, Map 11). In contrast to the sediments of the southern Limon Basin, the Miocene to Pleistocene deposits of the Bo- cas del Toro Basin are mostly exposed along the coast, and are generally only gently folded. In the Bocas del Toro Basin (Text-fig. 2), flat-lying sediments are ex- tensively exposed along coastal sections of the islands and peninsulas of the archipelago. Access to these sec- tions is by sea and many can only be studied in rela- tively calm weather. Geological mapping and section measuring were done using a 22-foot boat, but large- scale bulk fossil sampling was carried out by PPP ex- peditions of 6 to 12 persons using the research vessels Benjamin and Urraca of the Smithsonian Tropical Re- search Institute. The Upper Miocene Panama Canal Basin (Text-fig. 2) sediments are observed in roadside exposures from Sabanita to Colon, and in coastal exposures from Co- lon to Gobea, about 40 km to the west. (Map 1). The Gatun Formation was studied in numerous, often tem- porary, construction sites along or near Route 3, be- tween Sabanita and Colon, along the road to the Pay- ardi Oil Refinery, between Gatun and Margarita, and around the Gatun Dam (Map 1). Also included in this study are a few localities at the mouths of rivers along the north coast of Panama, between the Valiente Pen- insula, Bocas del Toro, and Gobea (Maps 2,3). The following account of the lithostratigraphy of the southern Limon, Bocas del Toro, and Panama Canal basins revises that of Coates et al. (1992) and Bottazzi et al. (1994), adding new information obtained in sub- sequent field work, including new formal stratigraphic 20 BULLETIN 357 Toro Member 7 Wéddn sadesosoonucybupestepemeannsnaskaakunaa Bf fee a =) pea aol a = 9 patbateasiaedael fl. # | bwwncuen ashe cvebenash aauneeubswnaa=esues abe rih z ° = < = 35 10 3 = nt, [bossrcrenotaseeco cedars = z =) E < o Tesi sj isamaacaaa acca Margate: Gatun Volcanics Aa Cretaceous} "7 7 Sabanita- Payardi NORTH COAST PANAMA (Undated Upper Miocene) (6) DON (8) eee ae ae | eal Miguel dela Calzones Bocade Bocade | Borda River Concepcion Concepcion East West Text-figure 4.—Correlation of measured sections from the Panama Canal Basin and along the north coast of Panama. Numbers in parentheses refer to sections in Appendix B, this volume. Lithologic patterns correspond to the lithologic key in Appendix B. units. The temporal range and formal nomenclature of the stratigraphic sequences in the three basins are sum- marized in Text-figures 4, 5, and 6. The bases for the age assignments are discussed by Aubry and Berggren Appendix 1, (this chapter). Five stratigraphic sections were measured in the Panama Canal Basin: 4 along the north coast of Pan- ama, 17 in the Bocas del Toro Basin, and 13 in the Limon Basin (Appendix B, this volume). Fossil col- lections are located by their PPP number on the maps in Appendix A, (this volume), and stratigraphically on each section in Appendix B, (this volume). These PPP numbers also link all files in the PPP Database (Kauf- mann, this volume) which is also available at the in- ternet site http://www.fiu.edu/‘collinsl/. Currently, a paleomagnetic sampling project is being completed on the stratotypes of the Limon (McNeill et al., in press) and Bocas del Toro groups that will be integrated with the litho- and biostratigraphy to refine the geochro- nology. ACKNOWLEDGMENTS Many people have helped me in the field since the PPP started in 1986. Jeremy Jackson, Laurie Collins, Peter Jung, and Ann Budd, my colleagues on the Steering Committee of the PPP, have been my most LITHOSTRATIGRAPHY: COATES DDI VALIENTE PENINSULA ESCUDO DE South VERAGUAS BASTIMENTOS Valiente ISLAND ISLAND (15) (22/23) : Reef Member ew PY snort cut Bed the GriAeORN: Huei MaeSolaris: Cay abr eas S| oo Valiente--*)-° | Coast Coast 24] Nispero xx Point | South Shark Hole Point Fm. Shark Hole PointFm. North Point- Piedra Roja ingen Tiburon Point Point West Island (14) CAYO AGUA ISLAND Feats cy PointFm. Nan El. yu = Sip lok: Job =|! 1 oO}. Je a|- {= |. Tobabe Sst Valiente Volcanics Text-figure 5—Correlation of measured sections in the Bocas del Toro Basin. Numbers in parentheses refer to sections in Appendix B, this volume. Lithologic patterns as for Text-figure 4. 22 BULLETIN 357 LIMON (35) Pueblo Nuevo Lomas del Mar Mbr. Empalme Mbr. Lomas del Mar East Buenos Aires Mbr. Chocolate River Rio Banano Fm Chocolat Quitaria ge GO) ee a Vizcaya Bananito Santa River River Rita od ue} cf. Sf. @| =) Carbon Sandbox Dos Rive Text-figure 6.—Correlation of measured sections in the southern Limon Basin. Numbers in parentheses refer to sections in Appendix B, this volume. Lithologic patterns as for Text-figure 4. constant field companions. Marie-Pierre Aubry, Bill Berggren, and Don McNeill, identifying calcareous nannofossils, planktic foraminifera, and taking and processing paleomagnetic samples, respectively, have also spent much time in the field and even more in the laboratory, integrating the litho-, bio-, and magneto- stratigraphy. To them all I owe a special debt of grat- itude. I thank Dana Geary, Tom Cronin, Susan Kid- well, John Sutter, Claudia Johnson, Erle Kauffman, Si- mon Mitchell, Orangel Aguilera, Jorge Obando, and Jon Todd, all of whom provided valuable insights into field interpretations on various occasions. Valuable as- sistance in measuring sections and collecting samples was also provided by Laurie Andersen, Mairi Best, Pam Borne, John-Mark Coates, Tim Collins, Mat Cot- ton, Helena Fortunato, Antoine Herz, Ken Johnson, Rene Panchaud, Stephen Rhodes, Jay Schneider and Tom Stemann. My thanks are also due to Sebastian Castillo, who guided my boat for many years, and to Beatrice and Lucien Ferrenbach for constant logistical LITHOSTRATIGRAPHY: COATES 23 help and wonderful hospitality throughout this project. I am also very grateful for very helpful reviews from Paul Mann and Joe Hartman. This chapter could not have been completed without the extraordinary con- tribution of my research assistant Xenia Guerra. She is responsible for the locality maps and the conversion of my field notes into computer-drawn sections that comprise Appendices A and B, and for drawing all the text-figures. She undertook the task of checking that 2531 samples were properly recorded on all sections and maps. She has accompanied me in the field to process samples, edited the manuscript, and ensured that I was always equipped with the appropriate field maps, for all of which I am especially grateful. Lastly, my profound thanks to Janet Coates, my wife, who acted as my field assistant on numerous occasions, tol- erated long absences on other occasions, helped or- ganize many expeditions, and generally supported PPP activities in so many ways. GEOLOGICAL SETTING The Central American isthmus forms the western margin of the Caribbean and lies at the center of a complex intersection of the Pacific, Cocos, and Nazca plates with the Caribbean Plate and the small Panama Microplate (Text-fig. 1). The dominantly oceanic Ca- ribbean Plate lies between the North and South Amer- ican plates. Its relative eastward motion, with respect to the North and South American plates is accommo- dated by strike-slip faults to the north and in part to the south (but now confounded by compression from the west-northwestward-moving South American Plate). In the east, it is bounded by the subduction zone of the Lesser Antilles. The western margin of the Ca- ribbean Plate is more complex; in the northern part of the western margin, the Cocos Plate is in contact with the Caribbean Plate. In the southern portion of the western margin, a triple junction brings the Cocos and Nazca Plates in contact with the small Panama Micro- plate (Text-fig. 1). The Panama Microplate appears to have formed by northward escape from compression of the South American and Cocos/Nazca plates, which created its northern border, the North Panama De- formed Belt. Much of Central America lies either on the trailing western edge of the Caribbean Plate or on the Panama Microplate but a portion lies on the south- western corner of the North American Plate (Text-fig. 1). Since their formation in the Miocene, the two Pa- cific plates have impinged on Central America with different motions. The Cocos Plate, with relative northeasterly motion, is subducting vigorously under Central America as far south as the Costa Rica-Pana- ma border so that this region is a zone of active vol- canism and seismicity. In contrast, the Nazca and Pan- ama Microplate border south of Panama, is relatively aseismic and without active volcanoes (Text-fig. 1). The oceanic crust of the Panama Microplate is typical of the widespread basalt plateau that underlies much of the rest of the Caribbean Plate (Burke, 1988; Burke et al., 1978; Case et al., 1990). This is in striking con- trast to northern Central America, where much of the crust is older continental material (Donnelly et al., 1990). Younger volcanic deposits are found only along the western margin of the isthmus adjacent to the vol- canic arc associated with the subduction of the Cocos Plate. Northern Central America consists of 1) the Maya Terrane, underlying Chiapas and Yucatan in southern Mexico, Belize, and Guatemala north of the Motagua River; and 2) the Chortis Terrane, which forms the rest of Guatemala, El Salvador, Honduras, and Nicaragua (Text-fig. 1). In the Maya and Chortis terranes, the isthmus is broad, formed of continental crust, and has a geologic history extending back to the early Paleo- zoic. For reviews of the geological history of northern Central America, see Dengo (1985), Donnelly (1992), Donnelly et al. (1990), Coates (1997), and references therein. Southern Central America, the focus of this book, includes the Panama Microplate (see papers in Mann, 1995, for recent reviews), which encompasses most of Costa Rica and Panama but also includes part of north- western Colombia (Text-fig. 1). The geology of north- western Colombia has been reviewed by Duque-Caro (1990a, b), and the rest of the region by Escalante (1990), Mann (1995), Seyfried and Hellmann (1994), and Coates and Obando (1996). Three major tectonic movements dominated the late Neogene tectonic evolution of the southern Central American isthmus (Kolarsky et al., 1995; Coates and Obando, 1996). The first was convergent tectonics of the eastern Pacific subduction zone, the primary driv- ing force creating the southern isthmus by forming a volcanic arc with associated fore- and back-arc basins (Astorga et al., 1991). During the Miocene the arc manifested itself as an extensive archipelago stretching to South America. The second tectonic movement, initiated about 4—3 Ma, was the shallow subduction of the Cocos Ridge, a lighter and relatively thick welt of oceanic crust trail- ing from the Cocos hot spot (Meschede, oral commun., 1998). This hard-to-subduct ridge rapidly elevated the Talamanca Range in particular (Text-fig. 1), and the southern isthmus in general (de Boer et al., 1988; Cor- rigan et al., 1990; Collins et al., 1995; Kolarsky et al., 1995). The elevation of the Talamanca Range probably 24 BULLETIN 357 Chagres Fn ——————_> < ‘aimito Fm ——><— Bohio Fm > se [TT toro Mbr. Bohio Lake Peninsula << Gatun > Text-figure 7.—Schematic geologic cross section of the Panama Canal Basin from the Bohio Peninsuia to the Caribbean coast west of Toro Point. also substantially reduced the number of marine con- nections between the Pacific and Caribbean. The third tectonic influence on southern Central America was the convergence of the South American and Caribbean plates (Text-fig. 1), which increasingly compressed the southern Caribbean Plate margin throughout the Neogene (Silver ef al., 1990; Kellogg and Vega, 1995; Mann and Kolarsky, 1995). This up- lifted eastern Panama and the outer Andes of north- western Colombia and, at about 3 Ma (Keigwin, 1982), finally severed all marine connections between the Pacific and Caribbean. LITHOSTRATIGRAPHY THE PANAMA CANAL BASIN The Panama Canal Basin (Text-fig. 2) is located at the junction of the Chorotega and Choco terranes (Dengo, 1985; Escalante, 1990), which is manifested by a major contrast in gravity (Case, 1974) and a series of north-south basement faults (de Boer et al., 1988; Mann and Corrigan, 1990; Mann and Kolarsky, 1995). The stratigraphy of the complex series of laterally varying Cenozoic deposits across the Panama Canal Basin is well known because many were excellently exposed during the construction of the Panama Canal. Reviews of the Cenozoic sequence were given by Woodring (1957, 1970, 1977, 1982) and Escalante (1990). In this chapter, I am concerned only with the Neogene Gatun and Chagres formations. These two formations crop out only at the northern end of the Panama Canal Basin, along the Caribbean coast, im- mediately to the west and east of Colon (Map 1), and their stratigraphic relations are shown in Text-figure 7. Collins et al. (1996) provided strong evidence to suggest that the southern Central American archipel- ago was an almost complete ecological barrier between the Pacific and the Caribbean, at the time of the de- position of the Gatun Formation (Late Miocene, about 8 Ma). The elongated outcrop pattern (Text-fig. 2), par- allel to the isthmus, and the shallow marine deposi- tional environment (20—40 m, Collins et al., this vol- ume) indicates that the Gatun Formation sediments formed as an apron of volcaniclastics flanking the isth- mian arc with no marine connection to the Pacific side. However, Collins et al. (1996) also show that the Pan- ama Canal Basin became temporarily a marine strait again, at about 6 Ma, during the time of deposition of the Chagres Formation, because it contains abundant bathyal benthic foraminifera of dominantly Pacific af- finity. The Gatun Formation Origin of the name.—The Gatun Formation was named by Howe (1907) after the village of Gatun which lies at the northern margin of Gatun Lake, 12 km southwest of Colon (Map 1). This name has come to supersede other earlier names for this unit, such as Monkey Hill and Mindi Beds (Hill, 1898). Location of stratotype.—Section 1 includes the stra- totype of the Gatun Formation defined by Coates et al. (1992). It runs from Sabanita on the main transisth- mian highway 12 km east of Colon, to 0.7 km west of the junction with Route 77 (the turnoff for Porto- belo). Reference sections.—Four reference sections have been measured (Sections 2—5) that include both the Gatun and the Chagres formations and reflect the lat- eral changes that the formations undergo from Gobea, 40 km west of Colon, to Sabanita, 12 km east of Colon (Map 1). Stratal relations.—The Gatun Formation rests un- conformably on formations of different ages in differ- LITHOSTRATIGRAPHY: COATES ent parts of the Panama Canal Basin. To the east of Colon, the Gatun Formation rests nonconformably on the unnamed Cretaceous volcanics (Text-fig. 4). To the west of Colon, including several islands in Lake Ga- tun, the Gatun Formation rests with angular unconfor- mity on the upper Oligocene Caimito Formation (Text- fig. 7). Westwards, the Gatun Formation can be traced as far as Gobea. Lithology.—The lower five m of the lower Gatun Formation (Text-fig. 4) consists of volcanic conglom- erate, with 1—5-cm clasts and a tuffaceous, arkosic ma- trix, cross-bedded, laminated, tuffaceous siltstone, and alternating laminated sandstone and siltstone, mostly deeply weathered. The overlying 40 m (Section 1) consists of massive, grey-green, clayey siltstone, with minor claystone and fine sandstone units. Zones of densely packed large concretions, pervasive bioturba- tion, and extensive thalassinoid burrow systems, as well as simple vertical and lateral hash-filled burrows, are typical of this part of the section. Shell hash of varying grain size and density is almost ubiquitous, as are units packed with diverse, whole mollusks. The middle Gatun Formation (Text-fig. 4) is de- scribed in Section 2 and covers the composite section from Gatun to Margarita. The middle Gatun Formation is about 350 m thick and consists of alternating silt- stone and sandstone with occasional 4—5-m units of interbedded sandstone and conglomerate. Concretion zones like those of the lower Gatun Formation are largely absent. Shell hash and diverse molluscan as- semblages are somewhat less abundant than in the lower Gatun Formation, but pervasive bioturbation is still very extensive. Bentonitic horizons and a higher wood fragment content are also typical of the middle Gatun Formation. The upper Gatun Formation (Text-fig. 4) is exposed around Mount Hope (upper part of Section 2) and more extensively on the western side of the canal, along and adjacent to the road to Pifia (Section 4), and is about 40 m thick. The lithology is more consistently volcaniclastic sandstone or fine conglomerate, with minor mudstone and siltstone. Thin bentonite horizons and shell hash are common but diverse, whole mol- lusks are relatively rare. A distinctive horizon is ex- posed below the overflow dam on the Chagres River west of Gatun Locks (Map 1) and has conglomeratic, tuffaceous sandstone beds with extensive thalassinoid burrows, wood, and scattered coral colonies up to 50 cm in diameter. Armored mudballs, 6—10 cm in di- ameter, are also abundant at one horizon that has nu- merous pockets filled with conglomerate. The Chagres Formation Origin of the name.—The Chagres Formation (Text- fig. 4) was named by MacDonald (1915) after the Cha- gres River. NO Nn Location of stratotype.—The stratotype is dia- grammed in Section 3. It is exposed between Toro Point and Naranjitos Point (Map 1). A distinctive lat- eral facies of the Chagres Formation is exposed at Boca del Rio Indio (Map 1) on the north coast of Pan- ama, approximately 40 km west of Colon. The Rio Indio facies is diagrammed in Section 5. Reference sections.—Sections 3, 4, and 5 include the Chagres Formation, which has been recently re- vised by Collins et al. (1996). The formation also crops out along the coast as prominent cliffs between Toro Point, at Colon, and the mouth of the Chagres River (Map 1). Stratal relations.—The Chagres Formation sits dis- conformably on the Gatun Formation; a marked change in lithology and a temporal hiatus characterizes the disconformity. Lithology.—The Chagres Formation consists of in- durated, conglomeratic, coarse-grained, volcanic sand- stone with quartz, feldspar, and lithic grains. It is per- vasively bioturbated and is relatively poor in macro- fossils. Toward the west, in the region of the Indio River and Gobea (Map 1), the average grain size of the Chagres Formation markedly decreases and the macrofossil content increases. The Toro Member The base of the Chagres Formation at the stratotype, is distinguished by a distinctive echinoid-mollusk-bar- nacle coquina, about 60 m thick, which Woodring (1957) separated as the Toro Member (Text-figs. 4, 7). It is well exposed in the cliffs west of Toro Point, the headland on the northwest side of Colon Harbor (Map 1). The Toro Member has a middle portion character- ized by about 10 m of steeply cross-bedded, prograd- ing foreset beds, 2-50 cm thick, consisting of alter- nating coquina and shelly, coarse, volcaniclastic sand- stone. The Toro Member wedges out about 15 km to the southwest of Toro Point. Its restricted distribution at the northern end of the Panama Canal Basin, with high energy cross bedded coquina and very coarse volcan- iclastics associated with bathyal Pacific benthic fora- minifera, led Collins et al. (1996) to suggest deposition from a transisthmian strait in which strong currents flushed Pacific sediments and benthos into deep Ca- ribbean waters. THE BOCAS DEL TORO BASIN An extensive series of exposures of upper Neogene sediments can be observed in the coastal region of Bocas del Toro, Panama where they form an extensive archipelago (Text-fig. 2). Mapping has revealed a Mio- cene basement of widely distributed basalt lava, flow 26 BULLETIN 357 breccia, and coarse, pyroclastic and volcaniclastic sed- iments. The Bocas del Toro Group lies nonconforma- bly on the underlying volcanics. The stratigraphic re- lations are shown in Text-figs. 5, 8a,b). The contact is well exposed in the Plantain Cays and on the coast south and west of Tobabe Point (Map 5, and inset B), where the volcanics form prominent bluffs of colum- nar basalt. Extinction, cooling, and subsidence of the volcanic arc locally in the region of the Bocas del Toro archipelago engendered a marine transgression repre- sented by the Bocas del Toro Group. The basal member of the Bocas del Toro Group is the Tobabe Sandstone, named herein. Messinian (7.2— 5.3 Ma) in age, it represents a basal, transgressive, near-shore marine facies that gradually gives way to the upper bathyal facies (Collins, 1993) of the over- lying Nancy Point Formation (Text-figs. 5, 8b). Con- tinued regional elevation of the isthmus initiated, about 5 Ma, a shallowing upward sequence represented by the Shark Hole Point, Cayo Agua, and Escudo de Ver- aguas formations. This culminated in extensive, shal- low marine, mixed volcaniclastic and coral reef de- posits, about two Ma, many of which are exposed in unnamed units on Bastimentos and Colon islands (Text-fig. 8a). Also included is the early Pleistocene Swan Cay Formation (Text-fig. 5), a deep fore-reef deposit (Collins, Appendix 1, in Jackson et al., this volume) exposed only on Swan Cay, a small island immediately north of Colon Island (Map 9). The new Tobabe sandstone and Swan Cay formations are here added to the Nancy Point, Shark Hole Point, Cayo Agua, and Escudo de Veraguas formations (Coates et al., 1992) to form the expanded Bocas del Toro Group. The Tobabe Formation Origin of the name.—The formation is named after the Guaymi village of Tobabe, located on the north coast of the Valiente Peninsula near the Plantain Cays (Maps). Location of stratotype.—The stratotype is on Small Plantain Cay; a small, unnamed island immediately to the west; and for about one km along the coast, be- tween Tobabe Point and the village of Tobabe (Inset B of Map 5). Reference sections.—Section 14 describes a lateral variation of the Tobabe Formation exposed on the Toro Cays, south of the entrance of Bluefields Bay, at the western end of the Valiente Peninsula (Inset B of Map 5). Stratal relations.—The Tobabe Formation sits non- conformably upon the Miocene basement of basalt la- vas, flow breccias, and volcaniclastic sediments. It passes conformably up into the overlying Nancy Point Formation (Text-fig. 8b). Lithology.—The Tobabe sandstone is the oldest unit of the Bocas del Toro Group. At the stratotype (Sec- tion 12) the basal unit of the Tobabe sandstone is a pebble and cobble conglomerate, about 15 m thick, with a variety of sedimentary and volcanic subangular clasts. It is unfossiliferous in its basal portion, but con- tains scattered, thick-shelled mollusks and occasional echinoids in the upper part, which grades insensibly into the overlying quartz sandstone that forms the rest of the formation. The sandstone is relatively clean, in- durated, massive and pervasively bioturbated, al- though thin but persistent horizons of pebble conglom- erate occur and larger volcanic cobbles are scattered throughout. Well-preserved burrows are present with particularly good examples of Ophiomorpha and thal- assinoid burrows. The unit is distinguished by very abundant, large, thin-shelled Amusium, numerous large sand dollars and spatangoid echinoids. Other mollusks are present as low-diversity, poorly preserved molds. Occasional specimens of wood bored by Teredo, and worms, including serpulids and vermetids, are also present. The Tobabe Formation cropping out on the Toro Cays (Section 14), is about 30 m thick. The lower 20 m consists of extensively burrowed, coarse, quartz and lithic volcanic sandstone with abundant and elaborate thalassinoid burrow systems, alternating with beds of 1-m-thick basalt and sandstone conglomerate, and oc- casional thin siltstone and muddy sandstone units con- taining scattered turritellids and other mollusks. The upper 10 m of the Tobabe Formation on the Toro Cays is composed of massive, shelly, volcaniclastic, rela- tively clean bioclastic sandstone, with strongly cross- cutting, laminated channels. Spectacularly large, shell- filled burrows about 10-15 cm in diameter are very characteristic, as is pervasive burrow mottling, and in other horizons, vertical 1—3 cm burrow tubes. Thick- shelled mollusks, including pectens, erect bryozoans and spatangoid echinoids, are common. The Nancy Point Formation Origin of the name.—The Nancy Point Formation (Text-fig. 5; Sections 12,15) was named by Coates et al. (1992), for the promontory called Nancy Point which lies 2.5 km south of the village of Tobabe (Map =e Location of stratotype.—The stratotype of the Nan- cy Point Formation lies along the northern coast of the Valiente Peninsula, starting at Nancy Point and run- ning south to near Chong Point (Map 5, and insets D, E, F): Reference sections.—There are no exposures be- tween the stratotypes of the Nancy Point and the To- babe Sandstone along the north coast of the Valiente LITHOSTRATIGRAPHY: COATES 27 A WNW Wild Cane Pliocene Long fs Beach > A amaere * a 4A “me: I pb oe ge r eof AA OA A a Az Boa A AS ATAS ATA ATAm Acker] Beane AeA ar a 7 EN aS agree Rae LAA rerwarts Perilli meetelcs: 7 : : ee co Se 3: + ESE Pliocene Fish Hole Reef ea Old Point Basalt flows mudstone NW Reef patches Foraminiferal Volcanic Arc System ———— Volcanic Arc Sequence Avispa Point Valiente Peninsula fo Bocas del Toro Group Chong Shark Hole Point Nancy Point Point Text-figure 8.—a. Schematic geologic cross section along the north coast of Bastimentos Island showing younger unnamed formations of the Bocas del Toro Group unconformably overlying the volcanic arc deposits. b. Schematic geologic cross section from the western tip of the Valiente Peninsula to Shark Hole Point, showing the volcanic arc deposits unconformably overlain by the older units of the Bocas del Toro Group. Peninsula. Assuming a constant dip (the type Tobabe and Nancy Point Formations have the same strike and dip), 400 m of section are not exposed. Much of this missing interval is exposed on the southern coast of the Valiente Peninsula between Warrie Point and the southern headland of the small peninsula of Toro Point (Map 5), and is diagrammed in Section 15. Litholog- ically it appears to be more typical of the Nancy Point Formation. Stratal relations.—The Nancy Point Formation is conformable with the Tobabe Formation below and the Shark Hole Point Formation above (Text-fig. 8b). Lithology.—The Nancy Point Formation consists of massive, pervasively bioturbated, shelly, muddy and silty sandstone, muddy siltstone, with scattered mol- lusks and occasional leaves and plant fragments, and occasional coarse volcaniclastic and bioclastic sand- stone beds. There are several low-diversity shell beds (Section 15) near the base and several diverse mod- erately abundant molluscan assemblages throughout the section. The base of Section 15, on the south coast of the Valiente Peninsula has a faulted contact with the underlying volcanics so that no typical Tobabe Sand- stone crops out. The transition from Tobabe Formation to Nancy Point Formation deposits is best seen on Toro Cay 28 BULLETIN 357 (Map 5, inset A), where dark blue-gray, silty sand- stone, typical of the Nancy Point Formation, contains occasional, clearly defined 50-cm-thick thalassinoid burrow systems and extremely abundant and diverse mollusks. It overlies coarse, channeled Tobabe sand- stone with only a 10-m gap. The transition from the Tobabe sandstone to the Nancy Point Formation thus appears to be conformable and to involve relatively rapid deepening from nearshore to upper slope facies. The Shark Hole Point Formation Origin of the name.—The Shark Hole Point For- mation was named by Coates et al. (1992) for the promontory of the same name that lies 3 km east of Chong Point (Map 5). Location of stratotype.—The stratotype lies along the coast between Chong Point and Bruno Bluff (Map 5, and inset F). Reference sections.—Section 15, along the south coast of the Valiente Peninsula contains the Shark Hole Point Formation as is indicated on Text-figure 5. Stratal relations.—The Shark Hole Point Formation conformably overlies the Nancy Point Formation (Text-fig. 8b) and is overlain by an unnamed conglom- eratic, cross bedded, coarser grained sequence of vol- caniclastics containing large pieces of wood and plant fragments. This unnamed unit is exposed only along the southern coast of the Valiente Peninsula, east of Secretario (Map 5). Lithology.—The formation is about 200 m thick and consists of micaceous, clayey siltstone that is perva- sively bioturbated and rich in large scaphopods. The uppermost part of the formation also contains abun- dant, thin, shelly beds and intraformational slumps with pillow folds and rip-up clasts. The Escudo de Veraguas Formation The stratigraphic order of the formations described above has been determined by physical superposition. The three remaining formations of the Bocas del Toro Group are known only on islands and their position relative to the other units has been determined by bio- stratigraphic evidence discussed in Appendix 1, (this chapter). Origin of the name.—The Escudo de Veraguas For- mation (Text-fig. 5) was named by Coates ef al. (1992) for the island of the same name that lies about 27 km east of Nancy Point (Map 4). Location of the stratotype.—The original stratotype, defined by Coates et al. (1992), is located along the north coast (Map 4, inset A and B), from Long Bay Point one km eastward (lower part of the formation), and for two km south of Long Bay Point on the west coast (upper part of the formation). We have since car- ried out more detailed field work that indicates that the coastal section immediately east of Long Bay Point is essentially along strike and thus probably exposes the same sequence of beds several times. A continuous section for the lower part of the formation is best ob- tained along the coast on the east side of the V-shaped embayment situated in the center of the north coast about two km east of Long Bay Point (Map 4C). This locality is now defined as the stratotype for the lower part of the Escudo de Veraguas Formation. The stra- totype for the upper part of the formation remains that originally defined by Coates ef al. (1992) along the west coast for one km south of Long Bay Point. Be- tween these upper and lower stratotypes, both of which have clearly documented physical superposition of strata, there is a small but unknown amount of section missing. The exposures along the north coast of Es- cudo de Veraguas, immediately east of Long Bay Point, and west of the V-shaped embayment in the center of the north coast, which were part of the orig- inal stratotype defined by Coates et al. (1992), are es- timated to fall in this gap. However, because the coast is irregular and only approximately parallel to strike, the stratigraphic order of samples from these exposures can not be determined. Collectively these samples were dated as 1.8—1.9 Ma and because they underlie the upper stratotype, they constrain it to be younger than 1.8—1.9 Ma. Stratal relations.—The upper and lower contacts of the Formation are not exposed. Lithology.—The lowest 10 m of the formation at the stratotype is moderately indurated, fine, silty sandstone and clayey siltstone, pervasively bioturbated and con- taining frequent, cemented, irregular burrow-concre- tions and horizons with dense thalassinoid burrow sys- tems. The overlying 30 m of clayey siltstone, silty claystone and silty, fine sandstone is also pervasively bioturbated, with frequent concretions, thalassinoid burrows and scattered mollusks with a distinctive basal 2 m thick marker bed rich in corals and mollusks. Following 70 m of no exposure, the section contin- ues with 13 m of clayey bioclastic siltstone, with some angular basalt grains, scattered mollusks and cupulad- rian bryozoans. The section is massive and pervasively bioturbated with scattered fine shell hash. About 5 m from the top, a second marker horizon is. defined by a densely packed coral biostrome that is also rich in echinoids and mollusks. The lower part of the Escudo de Veraguas Formation 2.6—3.5 Ma. The upper part of the Escudo de Veraguas Forma- tion (Section 10) consists of about 8 m of blue-gray, clayey siltstone and silty claystone, sparsely shelly and intensely burrow-mottled. Thalassinoid-type burrows are common, as are echinoids; the latter are very frag- LITHOSTRATIGRAPHY: COATES 29 ile and almost impossible to collect. Two distinctive marker beds within this section consist of slightly more indurated burrow zones, suggesting minor dis- conformities or slower depositional rates. The Cayo Agua Formation Origin of the name.—The Cayo Agua Formation (Text-fig. 5) was named by Coates er al. (1992) for the island of the same name in the Bocas del Toro archipelago, that lies about six km to the west of Toro Point, Valiente Peninsula (Map 6, and insets of Map 6). Location of stratotype.—More detailed section mea- suring on Cayo Agua has revealed that the stratotype is slightly more complex structurally than indicated by Coates et al. (1992). The stratotype for the formation (Section 19) is now calculated to be slightly thinner because a small block immediately to the south of Nis- pero point is rotated to dip to the northeast and repeats a portion of the section (Section 20). The newly de- fined stratotype (Section 19 runs from just south of North Point along the east coast southward to Nispero Point, and then from the northernmost to the south- ernmost exposures on the coast surrounding Tiburon Point. Reference sections.—Additional reference sections are Section 16, immediately west of North Point; Sec- tion 18, north of Red Rock Point; and Section 17, on the south coast immediately west of Red Rock Point The stratigraphic relations of these sections are shown in Text-figure 5. Stratal relations —The Cayo Agua Formation is equivalent in age to the upper part of the Shark Hole Point and the lower part of the Escudo de Veraguas formations and represents a shallower water facies. No contacts are known. Lithology.—The Cayo Agua Formation is distin- guished lithogically as a pervasively bioturbated gray blue, muddy, silty lithic sandstone with common ho- rizons of abundant thick shelled mollusks and aher- matypic corals. Occasional horizons of pebble con- glomerate and very coarse-grained volcaniclastic sand- stone are common in the middle of the formation. Compared to the Shark Hole and Escudo de Veraguas formations, the Cayo Agua Formaton is consistently coarser-grained, with common basalt grains and gran- ules, phosphatic pebbles, and wood fragments. A dis- tinctive marker bed of packed ahermatypic corals oc- curs near the top of the formation and is well exposed at Tiburon Point (Map 6) and the unnamed point to the south. In addition the mollusks and corals in the Cayo Agua Formation are larger and more heavily cal- cified than those of the Shark Hole and Escudo de Veraguas formations. Evidence from benthic forami- nifera (Collins, 1993) confirms the inference from grain size and fauna that the Cayo Agua Formation represents a more shallow-water facies than either the Shark Hole or Escudo de Veraguas formations. The Swan Cay Formation Origin of the name.—The Swan Cay Formation is named for the small island of the same name that lies 1.7 km off the north coast of Colon Island (Map 9). The location of the stratotype.—The stratotype (Text-fig. 5; Section 25) is the section that runs from north (youngest) to south (oldest) across Swan Cay. Reference sections.—No other sections of the Swan Cay Formation have been observed. Stratal relations.—No contacts are observed but the stratigraphic relationship of the Pleistocene Swan Cay Formation, within the Bocas del Toro Group, based on biostratigraphic data, is shown in Text-figure 5. Lithology.—The formation has three components. The lower 15 m is exposed on the southerly low hill of the island and consists of silty sandstone and shelly calcarenitic siltstone, with coral rubble and red algal fragments and balls. The middle four m consists of calcarenitic clayey siltstone, with dense, fine shell- hash horizons, and abundant large coral colonies in the lower part. The upper 60 m of the formation consists of massively thick-bedded, pale tan-white calcarenite. The upper 30 m are vuggy, sparry, and clean and in- clude a 4-m-thick coral bed with large Montastraea colonies, other corals and mollusks. The lower 30 m consist of more silty calcarenite with common red al- gae and large foraminifera, shell hash, and micromol- lusks. Cave deposits, about five m above the base of the calcarenite, consist of silty, shelly, volcaniclastic sandstone, mixed with abundant volcanic cobbles and boulders, and calcareous reef rubble containing an abundant and diverse molluscan assemblage. THE LIMON BASIN The southern part of the Limon Basin is located on Text-figure 2. The tectonic sedimentary history of the southern Limon Basin was recently reviewed by Bot- tazzi et al. (1994). They analyzed Campanian to Pleis- tocene sequences in detail, utilizing surface and sub- surface data that were generated by petroleum explo- ration since 1957. They list four upper Neogene units; the older Uscari and Rio Banano formations and the younger Suretka (terrestrial) and Limon (marine) for- mations. In a previous publication, Coates ef al. (1992) grouped these sediments (excluding the Suretka For- mation) in the Limon Group. However, the sediments of the Limon Formation (for which Bottazzi er al. (1994) give no type locality) were previously included in the Moin Formation, recognized originally by Gabb 30 BULLETIN 357 WSWw < Quebrada Chocolate >< _ Formation Buenos Aires < Member Mollusk lagoonal Formation Empalme orator i Moin Lomas del Mar LIMON Member eu Quebrada : Chocolate Fm Fore reef facies stratotype (Cangrejos Creek) Holocene Reefs pes ee U scan Fm 2 ee Text-figure 9,—Schematic geologic cross section across the southern Limon Basin from Chocolate Creek to the Port of Limon, slightly modified from McNeill ef a/. (in press). (1881) and Taylor (1975), and given formation status by Cassell (1986), Cassell and Sen Gupta (1989a), Coates et al. (1992) and Collins et al. (1995). The name Moin thus has priority over Limon for this unit. A recent detailed analysis of the upper Neogene sed- iments of the region immediately west of Limon, that combined litho- and biostratigraphy with paleomag- netic analysis (McNeill ef al., in press), confirmed the Moin Formation as the youngest unit of the Limon Group. It also named a new unit, the Quebrada Choc- olate Formation, that conformably overlies the Rio Banano Formation and underlies the Moin Formation (Text-fig. 6). The four formations of the Limon Group represent a genetically coherent, shallowing upward, marine, sedimentary sequence that reflects the rise of the Central American isthmus. Elsewhere in the south- ern Limon Basin, the Quebrada Chocolate and Moin Formations are laterally replaced by the Suretka For- mation (Bottazzi et al., 1994), which consists of ter- restrial (alluvial) volcanic conglomerate and breccia. Some of the reef deposits now included in both the Quebrada Chocolate and Moin formations were erro- neously identified by Coates et al. (1992, pp. 822, Fig. 7) as overlying, younger Pleistocene Reef Limestone. In part this was because these reef deposits commonly contain Acropora palmata, which was widely accepted to be restricted to the Pleistocene. Subsequent facies analysis by McNeill et al. (in press), and identification of the coral fauna by Budd ef al. (this volume), have established that this Reef Limestone consists of several reef members interbedded with the Pliocene to low- ermost Pleistocene siliciclastic and bioclastic sedi- ments of the Quebrada Chocolate and Moin Forma- tions. The stratigraphic relations of the formations of the Limon Group are shown in Text-figures 6, 9. In the southern Limon Basin, during the Early and Middle Miocene, sedimentation was dominated by bathyal, fine-grained, siliciclastic deposits typified by the oldest unit of the Limon Group, the Uscari For- mation (Cassell and Sen Gupta, 1989b; Collins ef al., 1995). While these sediments contain a rich microfau- na, to date, they have not yielded abundant macrofos- sils. Throughout the southern Limon Basin, the Uscari lithofacies is replaced diachronously by coarser, more variable, shallow-water sediments, interpreted as a nearshore marine and deltaic sequence, whose prove- nance was the rising Talamanca Range to the south- west. This near-shore marine and deltaic facies, rep- resented by the Rio Banano Formation, replaced the bathyal facies, represented by the Uscari Formation, during the latest Miocene. By the Pliocene, these fa- cies were present throughout most of the southern Li- mon Basin (Bottazzi et al., 1994). The Rio Banano Formation is highly variable in lithology, consisting of interfingering marine delta-front and delta-plain de- posits, and locally containing well-preserved, abun- dant, diverse, shallow-marine faunal assemblages. By the late Pliocene and Pleistocene, much of the southern Limon Basin had become emergent so that the marine deposits are unconformably overlain by coarse alluvial conglomerate and breccia, comprising the Suretka Formation. However, in the vicinity of Li- mon, marine deposition continued until the early Pleis- tocene, forming shallow-water, brackish and normal- marine claystone, sandstone, and reef deposits that rep- resent lagoonal, mangrove, and seagrass habitats, in- terfingering with a variety of tabular and patch reefs (Text-fig. 9). These deposits comprise in part the Que- brada Chocolate and Moin formations. The Quebrada Chocolate Formation is, in part, coeval with the upper portion of the Rio Banano Formation in its type area near Bomba on the Banano River (Text-fig. 6). LITHOSTRATIGRAPHY: COATES 31 The development of active folds and faults associ- ated with the active North Panama Deformed Belt, in association with the insertion of the Cocos Ridge (Col- lins et al., 1995), has meant continued rapid uplift of the Limon Group deposits. Today, the youngest unit, the Pleistocene Moin Formation, is now 40 meters or more above sea level. Modern reef deposits around Limon rose a maximum of 1.7 m in the earthquake of 1991 to form emergent coastal terraces, indicating that uplift continues. The presence of A. palmata (McNeill et al., 1997) in the upper Pliocene Buenos Aires Member (about 35 Ma) of the Quebrada Chocolate Formation is now the oldest known record of the species. The Uscari Formation Origin of the name.—The Uscari Formation (Text- fig. 6) was first named informally by Berry (1921) and later formalized by Olsson (1922) for deposits along the Uscari Creek, a tributary of the Amoura River that runs into the Sixaola River. Location of the stratotype.—The original stratotype was not precisely located along the Uscari River and is probably not now exposed. Reference sections.—The lack of good exposures at the original type locality led Cassell and Sen Gupta (1989a) to designate a new reference section along the Terciopelo Creek, about 62 km due west of Limon. The Uscari Formation also crops out widely in the southern Limon Basin, where it has been described by several authors (Olsson, 1922; Redfield, 1923; Palmer, 1925). Stratal relations.—The Uscari Formation passes abruptly, although conformably for the most part, into the coarser grained strata of the overlying Rio Banano Formation. However, this transition is strongly diach- ronous across the southern Limon Basin. Lithology.—The early authors, noted above, de- scribed the Uscari Formation as a friable, gray clay- stone, rich in montmorillonite, with minor limestone and calcareous sandstone, and a thickness between 600 and 1500 m. Later, Olsson (1942) proposed a two-fold division into a lower black shale, typically occurring around the type area, and the ‘‘upper grey shales’’, separated by a ‘“‘Dentalium Zone’. He noted that the upper unit contains sandstone and conglomerate, and is typically developed between Puerto Viejo on the coast and the Panamanian border to the south. The reference section described by Cassell and Sen Gupta (1989a) consists of a basal biocalcarenite, 12 m thick, formed of larger foraminifera and red algal rho- doliths interbedded with sandstone rich in pyroxene and foraminifera, that is overlain by 550 m of well- bedded, soft, dark shale, rich in planktic foraminifera. Foraminifera of the Uscari Formation were described by Goudkoff and Porter (1942), Cassell (1986) and Cassell and Sen Gupta (1989a, b), and an extensive facies analysis was given by Bottazzi et al. (1994). In this study, in the extreme south of the Limon Basin, claystone sections were measured near Carbon Dos, (Map 10; Section 28), and at the Sandbox River, near Catarata (Map 10; Section 27), both representing bathyal conditions (Collins et al., 1995). In the Sand- box River section, the claystone is overlain discon- formably by a basal conglomerate of the deltaic Rio Banano Formation. In the area of the Banano River, southwest of Limon, Taylor (1975) described a con- formable transition of increasing grain size between the Uscari Formation and the overlying Rio Banano Formation. The Rio Banano Formation Origin of the name.—The name Rio Banano For- mation is taken from the Banano River which flows into the Caribbean about nine km south of Limon. Location of the stratotype-—The formation was named by Taylor (1975) but the stratotype (Text-fig. 6) was designated by Cassell (1986) as the bluffs on the Banano River, 500 m southwest of the railroad bridge at Bomba. Reference sections.—The stratotype was measured by Coates et al. (1992), who extended the section along the Banano River from just west of Quitaria to the railroad bridge at Bomba (Inset C of Map 11, Sec- tion 29). In addition to the stratotype, sections of the Rio Banano Formation (Map 11) have been measured on the Bananito (Section 31), Peje (Section 30) and Vizcaya (Section 39) rivers and at Santa Rita (Section 32). Stratal relations.—The Rio Banano Formation sits diachronously with abrupt lithological transition on the Uscari Formation, mostly conformably and sometimes disconformably. The Rio Banano Formation passes conformably into the Quebrada Chocolate Formation (Text-fig. 9) in the area immediately west of Limon but throughout most of the southern Limon Basin it is unconformably overlain by the breccia and coarse grained volcaniclastics of the Suretka Formation. Lithology.—Some of the rocks designated by Taylor as the Rio Banano Formation were first recognized as a lithological unit by Howe (1907) and subsequently correlated with the Gatun Formation of Panama by Olsson (1922). Taylor’s (1975) definition of the Rio Banano Formation included not only the dominant sandstone lithofacies that crops out at the type locality but also a conglomerate, reef and claystone facies. Subsequently, Cassell and Sen Gupta (1989a) and Coates et al. (1992) restricted the formation to the eS) NO sandstone unit and separated the reef and claystone lithologies as the overlying Quebrada Chocolate and Moin formations and their respective reef members. The sections at Quitaria (Section 29) and Bomba (Section 29) show two relatively thin, richly fossilif- erous marine units within a thick deltaic section of burrow-mottled, coarse, tuffaceous, concretionary sandstone that frequently contains basalt pebbles, leaves, seeds, and wood fragments. The lower marine section is exposed along the road and adjacent river bank, about 200 m east of the banana loading station at Quitaria (inset C, Map 11). About 15 m of burrow- mottled, shelly, clayey siltstone and silty, tuffaceous sandstone contain frequent shelly stringers and lenses that contain an abundant and diverse marine mollusk and bryozoan assemblage. Thalassinoid burrow sys- tems are common, often packed with shell hash, as are slabby and irregular concretion zones. The upper ma- rine section is well exposed on both the north and south banks of the Banano River approximately 500— 700 m southwest of the railway bridge at Bomba (Inset C, Map 11). The lower part of the section crops out on the north bank and is about 17 m thick. It consists mostly of blue-gray, burrow-mottled, clayey siltstone with scattered, fine, volcanic pebbles. The unit is rich in diverse mollusks, including scaphopods, and in bryozoans. The remaining part of the marine section is exposed on the south bank of the Banano River immediately to the east of the lower unit. The section contains 10 m of massive, sporadically fossiliferous and finer grained, tuffaceous siltstone at the base. Per- vasive bioturbation is common, with slabby and irreg- ularly rounded concretions and occasional richly shelly zones. This is overlain by 6 m of dominantly blue- gray, tuffaceous, silty sandstone with scattered volca- nic pebbles and abundant burrow systems, packed with shell hash and volcanic pebbles. Some horizons are rich in spatangoid echinoids and all are very rich in diverse and well-preserved mollusks. The marine units described at the type locality ap- pear to persist along strike and were measured in the Vizcaya and Bananito rivers (Map 11). The section in the Vizcaya River extends upwards into a distinctive shoreface facies. It is dominated by coarse, laminated sandstone, with low angled, prograding foreset beds, discrete, complex, thalassinoid burrow systems as well as large, circular, vertical burrows. The molluscan as- semblages of these units are less diverse than in the stratotype, and are often dominated by Docinia; cu- puladrian bryozoans are also common. Fine pebble- to granule-sized volcanic clasts are common and l-cm diameter phosphatic pebbles are also present. The section in the Peje River (Map 11) is the highest in the Rio Banano Formation and is correspondingly BULLETIN 357 more proximally deltaic (German Gonzalez, oral com- mun., 1992). The section is relatively unfossiliferous, dominated by volcanic conglomerate and sandstone. Rip-up clasts, dispersed, low-diversity mollusks, wood fragments, and large seeds are common. The Rio Banano Formation crops out extensively in other parts of the southern Limon Basin (Bottazzi et al., 1994), where sequences have been interpreted as representing estuarine-bay, fan-delta, delta-plain, del- ta-front, and shoreface facies. The Quebrada Chocolate Formation Origin of the name.—The formation is named for the Chocolate Creek that flows north into the Carib- bean, crossing Route 32 at Buenos Aires, about six km west of Limon. Location of the stratotype-—McNeill et al. (in press) have recently revised the stratigraphy and pa- leoenvironmental interpretation of the Limon Group in the region of Limon. They established the stratotype of their new Quebrada Chocolate Formation (Text-figs. 6, 9) along Chocolate Creek for about two km south of Route 32 and along Route 32 from Buenos Aires to 1.2 km east of the junction of Route 32 and Route 240 (Old Moin road). Reference sections.—There are no other measured sections in the Quebrada Chocolate Formation, which only crops out in a restricted area west of Limon (McNeill et al., Text-fig. 2, in press) Stratal relations.—The Quebrada Chocolate For- mation conformably overlies the Rio Banano Forma- tion and is conformably overlain by the Moin For- mation. Lithology.—The Quebrada Chocolate Formation (Section 33) consists of coarse, volcaniclastic, clayey siltstone and sandstone, conglomerate, calcarenite, and reef-rubble limestone. In the lower part there are thin, low-diversity, recrystallized reef lenses with corals in growth position. The base of the Quebrada Chocolate Formation lies at the waterfall on Chocolate Creek, 1.5 km south of Route 32, where the first reef member is composed dominantly of Porites in a clayey siltstone matrix. This unit is overlain by shelly, volcaniclastic sandstone with extensive thalassinoid burrow systems, coarsely laminated, cross-bedded sandstone, fine con- glomerate, reef rubble, and calcarenite. Corals from these units are abundant and diverse (Budd ef al., this volume) but associated flank rubble and sandstone also contain a large, thick-shelled Spondylus and many small bivalves and oysters. The Buenos Aires Member The top of the Quebrada Chocolate Formation con- sists of a series of extensive tabular reefs that is sep- LITHOSTRATIGRAPHY: COATES 33 arated as the Buenos Aires Reef Member (Text-figs. 6, 9). It is named for the village of Buenos Aires, on Route 32 six km west of Limon, where it forms a low but distinct topographic feature. The stratotype is lo- cated from the intersection with Chocolate Creek east along Route 32 for 1.2 km. The Buenos Aires Reef Member (Section 33) is about 140 m thick and consists of a series of coral thickets with a silty claystone ma- trix, dominated by Porites, Acropora, Stylophora, Caulastrea, and agariciids. Interbedded with the reefs are coral- and mollusk-rich carbonate sandstone and siliciclastics. The top of the Buenos Aires Reef Mem- ber has an interfingering contact with the grey-blue claystone, siltstone, and channeled sandstone of the base of the Moin Formation. The Moin Formation Origin of the name.—The formation is named for the small port of Moin, at the mouth of the Moin River, about six km west of Limon. Location of the stratotype.-—The name Moin was first used by Gabb (1881) for the dark claystone and muddy sandstone along the coast between Moin and Limon. He named these deposits the “‘Moin Member” without specifying a stratotype. Taylor (1975) included the Moin Member in his broadly defined Rio Banano Formation. He specified the stratotype as the small, unnamed creek that flows northwestward from near the Barracuda road, about two km west of Limon, through the Cangrejos housing complex (Inset B, Map 11), which lies south of the main coastal road at Playa Bon- ita. Reference sections.—Sections 35, 36, and 38, in the Lomas del Mar suburb of western Limon. These in- clude the Empalme and Lomas del Mar reef members described below. Stratal relations.—The Moin Formation sits con- formably on the Quebrada Chocolate Formation and has no upper contact. Lithology.—The Moin Member was established as a formation by Cassell (1986) and Cassell and Sen Gupta (1989a), and was expanded in concept by Coates et al. (1992) to include its biostromal reef members as well as other lithologies now separated as the Quebrada Chocolate Formation. The Moin Formation (Text-figs. 6, 9) is here re- tained with Taylor’s originally designated stratotype along the creek that runs through the Cangrejos sub- division (Section 37). It consists of blue-gray, silty claystone and blocky, muddy, shelly, volcaniclastic sandstone, with common basalt granules. Also present are scattered mollusks and callianassid burrows, and the lower part of the section contains laminated, py- ritic, organic-rich claystone and erect, branching bryo- zoans. In general, siliciclastic sediments in the Moin Formation are finer grained than in the Quebrada Chocolate Formation. The reef members of the formation cap the high ground that forms a plateau at about 40 m of elevation immediately to the northwest of Limon. To the west of the plateau, the lower Moin Formation is evident in sporadic exposures to the north of Route 32, until about four km west of Limon. Alternating thin units of shelly, blue-grey siltstone and claystone, extremely rich in diverse mollusks, bryozoans, crabs, and oyster beds, alternate with sandstone, often containing dense, callianassid burrow systems, thin, small, coral-rich patch reefs, and tight, dark claystone with mangrove root systems, logs, and wood fragments. McNeill et al. (in press) separated two major reef build-ups as mem- bers within the upper Moin Formation, as described below. The Empalme Member Interbedded with the mollusk-rich claystone and siltstone of the lower Moin Formation is the first of the two major reef trends within the formation (Section 34). McNeill et al. (in press) named the Empalme Reef Member (Text-fig. 6) for the small settlement of Em- palme Moin near the intersection of Route 240 and the Empalme Moin to Limon Road. This unit is usually deeply weathered and poorly exposed, but temporary, fresh exposures near Route 240 (Insets A, B, Map 11), Empalme Moin, and west of Pueblo Nuevo Cemetery, reveal calcarenite, carbonate sandstone, and coral-rich sandstone, dominated by the hermatypic corals Pori- tes, Montastraea, Stephanocoenia, Caulastrea, Diplo- ria, and agariciids. The Lomas del Mar Member The youngest and most fossiliferous of the reef units, the Lomas del Mar Member (Text-fig. 6), is about 30 m thick and caps the plateau immediately to the west and northwest of Limon from 45-60 m in elevation (Text-fig. 9). Extensive fresh sections in the Lomas del Mar Reef Member were exposed at two sites that were bulldozed down to bedrock for housing projects. The eastern of these (Section 36) is now cov- ered by houses; as of 1998, the western section (38) was exposed south of Barracuda Avenue 300-800 m. west of the intersection with King Fish road. There are also scattered exposures underlying and lapping against the Lomas del Mar Member of a deeply weath- ered, unfossiliferous, coarse sandstone, named by Tay- lor (1975) the Pueblo Nuevo Sandstone. The Lomas del Mar Member consists of three litho- facies (Sections 36—38): 1) Bioclastic, rubbly, reef limestone forming irreg- 34 BULLETIN 357 ular lensing patches, each 20-50 m in diameter, and about 1—3 m in maximum thickness. The patches yield abundant large coral heads, particularly Montastraea cavernosa and Dichocoenia, and a diverse and abun- dant array of small colonies, including several species of Montastraea, Agaricia, Mycetophyllia, and Dicho- coenia, as well as numerous, slender, branching colo- nies of Madracis asperula (Budd et al., this volume) 2) Blue-grey, clayey siltstone and calcarenite that form flank beds immediately adjacent to the reef patches. Sediments are packed with small, diverse mollusks, including vermetids, small solitary and ocu- linid ahermatypic corals, bryozoans, especially cupu- ladrians, serpulids, and large echinoid spines. The reef facies laterally interfingers with these flank beds at the Cangrejos creek stratotype (Section 37). 3) Massive, blocky, grey-blue claystone between the reef patches with scattered mollusks and fine shell hash. CONCLUSIONS The Neogene stratigraphy of three depositional ba- sins along the Caribbean coast, from Limon, Costa Rica to the Panama Canal, is revised and several new formations created. Eleven maps plus detailed insets and 39 sections are presented in Appendices A and B, respectively, giving the location of all PPP samples studied for this volume and their stratigraphic position and relations. The Middle and Upper Miocene Gatun and Chagres formations of the Panama Canal Basin form an apron of marine sediments flanking the isthmian volcanic arc. In the Bocas del Toro Basin, in western Panama, an Upper Miocene to Pleistocene sequence, the Bocas del Toro Group, forms an apron of inner to outer ne- ritic marine sediments, comprising six formations, that overly a Middle Miocene basement of basalt and coarse volcaniclastics. The Miocene to Pleistocene ma- rine deposits of the southern Limon Basin consist of a shallowing upward (bathyal to inner neritic) series of four Neogene formations that form the Limon Group. THE PANAMA CANAL BASIN The Gatun Formation is described from four refer- ence sections. The lower and middle part of the section is well exposed between Sabanita and Payardi (Sec- tions 1, 2) and the upper part near Toro Point (Section 3) and Pina (Section 4). The Gatun Formation lies un- conformably on the Cretaceous volcanics in the east- ern part of the basin and on the Caimito Formation in the west. It consists of about 500 m of volcaniclastic bioturbated gray-green claystone, siltstone and sand- stone, with large concretions and shell beds packed with mollusks in the lower part and coarser sandstone and fine conglomerate in the upper part. The Chagres Formation lies disconformably on the Gatun Formation. Locally, near Colon, the Chagres is distiguished by a basal member, the Toro Point, which is an echinoid-barnacle-mollusk coquina, in part strongly cross-bedded with coarse sandy to pebbly vol- caniclastics, deposited in bathyal water depths and containing dominantly Pacific benthic foraminifera. Four small sections were measured along the North Coast of Panama (Sections 6, 7, 8, and 9) but are not currently assigned to any formation. In age and litho- logical affinity they most resemble the Gatun Forma- tion. THE BOCAS DEL TORO BASIN The Neogene sediments of the Bocas del Toro Basin are represented by the Bocas del Toro Group. The most continuous Neogene section lies along the north and south coasts of the Valiente Peninsula (Sections 12, 15). Here, the Bocas del Toro Group sits noncon- formably on the underlying columnar basalt, basalt flow breccia, coarse volcaniclastics, and coral reef lenses of the Middle Miocene ‘“‘basement’’. The sec- tion ranges from Upper Miocene (Messinian, 7.1—5.3 Ma) to upper Pliocene (3.5 Ma). Younger deposits are found on Escudo de Veraguas island (Sections 10, 11; 3.5—1.8 Ma) and Bastimentos, Colon, and Swan Cay islands (Sections 21—26; ~2.0—1.5 Ma) and a more shallow water facies of the lower Pliocene is found on Cayo Agua island (Sections 16—20; 5.0—3.5 Ma). The basal unit of the Bocas del Toro Group is the newly defined Tobabe Formation, the stratoytpe for which is on Little Plantain Cay, an unnamed small island to the west, and along the south coast of Tobabe Point. A basal pebble and cobble conglomerate passes up into an indurated clean quartz and lithic sandstone containing thin volcanic cobble horizons, large sand dollar echinoids, and abundant large thin shelled Amu- sium. Thalassinoid burrow systems, Ophiomorpha, and large (10-15 cm in diameter) unidentified shell-filled burrows are very distinctive of this unit. Laterally, the formation contains other mollusks, erect bryozoa, and cross-cutting? tidal channels with coarse volcaniclastic laminated sandstone infilling. By rapid increase in silt and clay content the Tobabe Sandstone grades con- formably into the Nancy Point Formation. The stratotype of the formation runs from Nancy Point to Chong Point but is also well exposed along the south coast of the Valiente Peninsula (Section 15 and in the Toro Cays (Section 14), where it is partic- ularly fossiliferous. The Nancy Point Formation is per- vasively bioturbated, shelly, muddy, and silty sand- stone with occasional plant fragments and, near the LITHOSTRATIGRAPHY: COATES top, a series of channels of coarse volcaniclastic sand- stone and fine conglomerate. It ranges in age from 8.2— 5.6 Ma and represents bathyal deposition shallowing toward the top. The Shark Hole Point Formation conformably over- lies the Nancy Point Formation (Sections 12, 15) and has its stratotype at the promontory of the same name. The formation is about 200 m thick and consists of micaceous, clayey siltstone that is pervasively biotur- bated and contains abundant large scaphopods. The upper part of the formation contains abundant thin shell beds and intraformational slumps with pillow folds and rip-up clasts. The Shark Hole Point is 5.6— 3.5 Ma and shows the continuation of the regression begun in the upper Nancy Point Formation. The Escudo de Veraguas Formation is known only from the island of the same name that lies 27 km east of Nancy Point. It is younger than the Shark Hole Point, ranging in age from 3.5—1.8 Ma, and represents an outer neritic to upper bathyal deposit. Its stratotype, for the lower part of the formation, is along the coast on the east side of the the V-shaped embayment situ- ated in the central part of the north coast, about one kilometer east of Long Bay Point. The stratotype for the upper part of the formation lies along the west coast for about one km. South of Long Bay Point. Lithologically, it consists of pervasively bioturbated clayey siltstone and silty claystone, with frequent con- cretions, and scattered shelly hash, often with scattered whole and diverse mollusks and ahermatypic corals. The lower part of the formation is more indurated with very common and densely packed cemented burrow concretions and thalassinoid galleries, as well as a dis- tinctive marker bed of corals and mollusks two meters from the base. The upper part of this sequence is capped by a coral biostrome. The Cayo Agua Formation has its stratotype (Sec- tion 19) along the east coast of the island of the same name that lies six km to the west of Toro Point, Val- iente Peninsula. Other good exposures of the formation are described in Sections 16, 18, and 20. The Cayo Agua Formation (5.0—3.5 Ma) was deposited in shal- lower water (inner neritic, 20-40 m) than the coeval Shark Hole Point Formation. Lithologically, the Cayo Agua Formation is distinguished as a pervasively bio- turbated, muddy, silty, lithic sandstone with common basalt grains and granules, phosphatic pebbles, and wood fragments. It has many horizons of abundant, diverse, and thick-shelled mollusk asemblages and there is a distinctive marker bed of densely packed ahermatypic corals near the top of the formation, ex- posed on the coast of Tiburon Point. The Swan Cay Formation is known only from a small island of the same name, 1.7 km off the north Ww Nn coast of Colon Island. The section is 79 m thick and youngs from south to north across the island. It con- sists, in the lower part, of shelly calcarenitic siltstone and silty sandstone with coral rubble and red algal fragments. The upper part is formed of massively thick bedded tan-white calcarenite with large Montastraea colonies and other corals and mollusks. About 5 m above the base of the calcarenite in large cavities in the calcarenite, deposits of silty, shelly, volcaniclastic sandstone, mixed with volcanic cobbles and boulders, have yielded abundant diverse Pleistocene mollusks and otoliths. THE SOUTHERN LIMON BASIN The Neogene of the southern Limon Basin is rep- resented by the Limon Group, which contains the Us- cari, Rio Banano, Quebrada Chocolate, and Moin for- mations. The name Moin Formations is shown to have priority over the Limon Formation of Bottazzi et al. (1994). These formation form a genetically coherent shallowing upward marine sequence that reflects the rise of the Central American isthmus. From the Uscari Formation (oldest; 8.3—5.6 Ma) to the Moin Formation (youngest; ~1.9-1.5 Ma) the depositional environ- ment shallows from bathyal to lagoonal, although one biofacies of the Moin Formation suggests either local upwelling or adjacent deep water. The basal formation of the Limon group is the Us- cari Formation. It consists of a basal 12-m biocalcar- enite formed of larger foraminifera and red algal rho- doliths interbedded with foraminiferal, pyroxene-bear- ing sandstone. The main portion of the formation, about 550 m thick at the stratotype but probably highly variable across the basin, is soft dark gray shale rich in planktic foraminifera. Sections 27 and 28, measured at Sandbox River and Carbon Dos respectively, indi- cate deposition at bathyal depths, and an age range of 8.3-5.6 Ma, although other parts of the formation may be younger because the transition to the shallow water conditions of the overlying Rio Banano is highly diachronous. The Rio Banano Formation has its stratotype from Quitaria to Bomba along the banks of the Banano Riv- er. Other sections were measured in the Bananito (Sec- tion 31), Peje (Section 30), and Vizcaya (Section 39) rivers and the village of Santa Rita (Section 32). Lith- ologically, the formation at the type locality is domi- nantly burrow mottled, coarse, tuffaceous, concretion- ary sandstone with frequent basalt pebbles, leaves, seeds, and wood fragments. Two richly fossiliferous units occur near Quitaria and about 500 m south of Bomba. These are shelly, clayey siltstone and silty sandstone with stringers and lenses of abundant di- verse marine mollusks with bryozoa and echinoids at 36 BULLETIN 357 some horizons. Thalassinoid burrow systems are com- mon as are slabby concretionary zones. The Rio Ban- ano Formation is about 750 m thick in its type locality but because of the highly diachronous boundaries with the Uscari and Quebrada Chocolate formations, its thickness varies considerably across the southern Li- mon Basin. The depositional environment of the for- mation is inner neritic, close to reef buildups, and it ranges in age from ~3.6—2.8 Ma. The Quebrada Chocolate Formation, about 500 m. thick at its stratotype (Section 33), has its stratotype along the creek of the same name, immediately west of Buenos Aires, six km west of Limon. The lithology consists of coarse, volcaniclastic, clayey siltstone and sandstone, conglomerate, calcarenite, and reef rubble limestone. Thin low-diversity reef lenses occur in the lower part, and thalassinoid burrow systems and coarse- ly laminated and cross bedded sandstone characterize the middle part of the formation. The Quebrada Choc- olate Formation represents shallow water lagoonal and patch reef depositional environments and ranges in age from 3.5—2.8 Ma. The top of the formation is formed by a distinctive coral reef member, about 140 m thick, whose stratotype is along Route 32 for 1.2 km east of the intersection with Chocolate creek. It consists of a series of coral thickets of Porites, Acropora, Stylopho- ra, Caulastrea, and agariciids embedded in a silty clay matrix. The contact with the overlying Moin Formation is either interfingering, with the gray-blue claystone and siltstone, or disconformable with the chanelled Pueblo Nuevo sandstone facies (Taylor, 1975) of the overlying Moin Formation. The Moin Formation is a mosaic of three facies. At the stratotype (Section 37) it consists of blue gray, silty claystone and blocky, clayey, shelly, volcanic sandstone. Basalt granules, scattered mollusks, arthropod burrows are common. Locally, laminated, py- ritic and organic rich claystone is present (see upwelling below). In general, the siliciclastic sediments of the Moin Formation are finer than those of the Quebrada Chocolate Formation. The sediments of the stratotype appear to be flank deposits close to associated reef buildups separated below as members of the formation. They may have formed close to local upwelling or down an adjacent, deeper water, shelf slope. The second facies is displayed by the Empalme and Lomas del Mar reef members, which cap the higher ground (about 40 m and above) that lies immediately to the northwest of Limon. The Empalme Member (older) is usually deeply weathered and poorly exposed but contains Montas- traea, Stephanocoenia, Caulastrea, Diploria, and agar- iciids in a calcarenite matrix (Section 38). The Lomas del Mar Member is about 30 m thick (Sections 36, 38) and consists of bioclastic rubbly reef patches, each about 20-50 m in diameter and 1-3 m thick. The unit yields large colonies of Montastraea cavernosa and Dichocoenia, small colonies of Mon- tastraea, Agaricia, Mycetophyllia, and Dichocoenia as well as numerous slender branching colonies of Mad- racis asperula. These reef patches grade into clayey siltstone and calcarenite at their margins, which are packed with small diverse mollusks, including ver- metids, as well as oculinid corals, bryozoa, especially cupuladrians, and serpulids. Between the coral reef patches is foraminiferal blue-gray claystone. The third facies consists of blue-gray siltstone and claystone, extremely rich diverse mollusks, bryozoa, crabs, and oyster beds alternating with tight dark clay- stone with abundant mangrove root systems, logs, and wood fragments. These sediments are interpreted to rep- resent shallow mangrove and lagoonal seagrass depos- its. REFERENCES CITED Astorga, A., Fernandez, J.A., Barboza, G., Campos, L., Obando, J.A., Aguilar, A., and Obando, L.G. 1991. Cuencas sedimentarias de Costa Rica: Evoluci6n geodi- namica y potencial de hydrocarburos. Revista Geologica América Central, vol. 13, pp. 25—59. Aubry, M-P. 1995. From chronology to stratigraphy: interpreting the lower and middle Eocene stratigraphic record in the Atlantic Ocean. in W.A. Berggren, D.V. Kent, M-P. Aubry, and J. Hardenbol, eds., Geochronology Time Scales and Global Stratigraphic Correlation. 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D. thesis, Columbia University, New York, 179 pp. Keigwin, L.D. 1978. Pliocene closing of the Isthmus of Panama, based on bio- stratigraphic evidence from nearby Pacific Ocean and Ca- ribbean cores. Geology, vol. 6, pp. 630—634. 1982. Isotopic paleoceanography of the Caribbean and East Pa- cific: role of Panama uplift in late Neogene time. Science, vol. 217, pp. 350-352. Kellogg, J.N., and Vega, V. 1995. Tectonic development of Panama, Costa Rica, and the Colombian Andes: Constraints from global positioning system geodetic studies and gravity. in Geologic and tec- tonic development of the Caribbean plate boundary in southern Central America. P. Mann, ed., Geological So- ciety of America Special Paper, no. 295, pp. 75-87. Kolarsky, R.A., Mann, P., and Montero, W. 1995. Island are response to shallow subduction of the Cocos Ridge, Costa Rica. in Geologic and Tectonic Develop- ment of the Caribbean Plate boundary in southern Central America. P. 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P- Mann, ed., Geological Society of America Special Paper, no. 295, pp. 111-130. McNeill, D.F., Budd, A.F., and Borne, P.F. 1997. Earlier (late Pliocene) first appearance of the Caribbean reef-building coral Acropora palmata. Stratigraphic, evo- lutionary implications. Geology, vol. 25, pp. 891-894. McNeill, D.F., Coates, A.G., Budd, A.F., and Borne, P.F. 1999. Integrated biological and paleomagnetic stratigraphy of the Late Neogene deposits around Limon, Costa Rica: A coastal emergence record of the Central American Isth- mus. Geological Society of America Bulletin (in press) Olsson, A.A. 1922. The Miocene of northern Costa Rica with notes on its stratigraphic relations. Bulletins of American Paleontolo- gy, vol. 9, no. 39, pp. 181-192. 1942. Tertiary and quaternary fossils from the Burica Peninsula of Panama and Costa Rica. Bulletins of American Pale- ontology, vol. 9, no. 39, 309 pp. Palmer, K.V. 1925. Foraminifera and a small molluscan fauna from Costa Rica. Bulletins of American Paleontology, vol. 10, no. 40, pp. 1-20. Redfield, A.H. 1923. Petroleum possibilities in Costa Rica. Economic Geology, vol. 18, pp. 354-381. Seyfried, H., and Hellmann, W. 1994. Geology of an evolving island arc. Institut fur Geologie und Palaontologie, Universitat Stuttgart, Profil, vol. 7, 433 Pp- Silver, E.A., Reed, D.L., Tagudin, J.E., and Heil, D.J. 1990. Implications of the north and south Panama deformed belts for the origin of the Panama orocline. Tectonics, vol. 9, pp. 261-282. Taylor, G.D. 1975. The geology of the Limon area of Costa Rica. unpub- lished Ph. D. thesis, Louisiana State University, Baton Rouge, 114 pp. Woodring, W.P. 1957. Geology and paleontology of the Canal Zone and adjoin- ing parts of Panama, description of Tertiary mollusks (Gastropoda: Trochidae to Turritellidae). U. S. Geological Survey Professional Paper, no. 306-A, pp. 1-145. 1970. Geology and palaeontology of the Canal Zone and ad- joining parts of Panama, description of Tertiary mollusks (Gastropoda: Eulimidae, Marginellidae to Helminthoglyp- tidae). U. S. Geological Survey Professional Paper, no. 306-D, pp. 299-326. 1977. Distribution of Tertiary marine molluscan faunas in south- ern Central America and northern South America. Insti- tuto de Geologia, Universidad Nacional Aut6noma de Mexico, vol. 101, pp. 153-165. 1982. Geology of the Canal Zone and adjacent parts of Panama, description of Tertiary mollusks (Pelecypods: Propeamus- siidae to Cuspidariidae; additions to families covered in P306-E; additions to gastropods, cephalopods). U. S. Geo- logical Survey Professional Paper, no. 306-F pp. 542— 845. APPENDIX 1 Newest Biostratigraphy MARIE-PIERRE AUBRY AND WILLIAM A. BERGGREN Woods Hole Oceanographic Institution Woods Hole, Massachusetts 02543 INTRODUCTION The Limon and Bocas del Toro groups were first defined and analyzed biostratigraphically by Coates et al. (1992). Biostratigraphic determinations for this chapter were made principally by L. Bybell (calcare- ous nannofossils) and H. Dowsett and M. Cotton (planktic foraminifera). This biostratigraphy is de- scribed in detail by Bybell (this volume) and Cotton (this volume). In addition, the Gatun and Chagres for- mations of the Panama Canal Basin have been exten- sively revised recently by Collins, Coates, Berggren, Aubry, and Zhang (1996). Subsequent fieldwork has added formations to the Limon and Bocas del Toro groups, and has in some cases revised and expanded the description of the other formations (see Coates, this chapter). We present here all the biostratigraphic research conducted since Coates et al. (1992). The calcareous nannofossil analyses have been done by Aubry and those for planktic foraminifera by Berggren. Calcare- ous nannofossil and planktic foraminiferal assemblag- es at most levels are generally scarce, of low diversity, and relatively poorly preserved. This is probably be- cause many of the sections represent relatively near LITHOSTRATIGRAPHY: COATES 39 shore, shallow marine deposition. Nevertheless, these collections have allowed us to develop a reliable bio- zonal assignment for all formations. It has not been possible to recognize precisely any sequence of bio- stratigraphic datums within zones, thus hampering a precise temporal interpretation of sections (see Aubry, 1995). We use the time scale of Berggren et al. (1995). NEW FORMATIONS AND SECTIONS OF THE BOCAS DEL TORO GRouP, PANAMA The Tobabe Formation The only samples dated from the Tobabe Sandstone are from Section 14, on the Toro Cays where the for- mation conformably underlies the Nancy Point For- mation but does not expose the unconformable contact with the underlying Miocene volcanics. Planktic foraminifera occur throughout Section 14 (29 samples) and include Globigerinoides extremus, G. obliquus, G. trilobus, and, less frequently, G. siegliei, together with the Globorotalia conomiozea/miotumida group and the menardine globorotaliids (G. menardii ““B”’, G. pseudomiocenica, and G. plesiotumida). Glo- bigerinoides conglobatus occurs in the lower part of the section and has its FAD near the Miocene/Pliocene boundary. G. siegliei is diagnostic of the Late Miocene (Messinian) and Early Pliocene (Zanclian), with a LAD at ~4.7 Ma, and occurs throughout the Tobabe Formation. Globorotalia plesiotumida (LAD near the Miocene/Pliocene boundary) occurs throughout the section and Globorotalia cibaoensis (LAD at ~ 4.7 Ma) also occurs in one sample. This association strongly suggests that the Tobabe Formation is Late Miocene and largely Messinian (about 7.2—5.3 Ma). The Chagres Formation, in the Panama Canal Basin, is of comparable age (late Tortonian to Messinian; 8.6— 5.3 Ma (Collins er al., 1996) and represents a deep- ening event from ~25 to 200 meters depositional depth, from the Gatun Formation to the bathyal (200— 500 m) basal Toro Member of the Chagres Formation. The Tobabe Formation is the basal, trangressive unit of the Bocas del Toro Group and grades upwards into the Nancy Point Formation, which is also bathyal in origin (Collins, 1993). Correlating the sea level rise represented by the Toro Member and the Tobabe For- mation would imply that the base of the Tobabe For- mation may be as old as late Tortonian (about 7—8 Ma). The Nancy Point Formation; Toro Cays and Toro Point Sections Samples from two new sections of the Nancy Point Formation have been analyzed for both nannofossils and planktic foraminifera. The section on Toro Cays yields common Discoaster quinqueramus, indicating Zone NN11 (5.6—8.2 Ma). Planktic foraminifera also suggest Late Miocene (Messinian) and the presence of Globigerinoides conglobatus, which normally first ap- pears near the Miocene/Pliocene boundary, and Glo- bigerinoides siegliei, which characterizes the Messi- nian and Zanclian (Early Pliocene), suggests correla- tion with the upper part of Zone NN11. The younger assignment also agrees with the original biostrati- graphic analysis of the Nancy Point Formation (Coates et al., 1992). Section 15, along the westernmost south coast of the Valiente Peninsula, south of Toro Point and west of Playa Verde, includes the Nancy Point Point Formation and contains a fauna that is also as- signed to Zone NN11. The Cayo Agua Formation The type section of the Cayo Agua Formation is along the east coast of the island (Section 19) and the oldest units are just south of Norte Point. Planktic fo- raminifera we have observed include Globigerinoides obliquus, G. extremus, G. trilobus, Globoquadrina al- tispira (LAD at 2.9 Ma), Globorotalia pseudomiocen- ica (LAD at 3.5 Ma), and members of the N. hume- rosa/dutertrei group. This assemblage suggests an ear- ly Pliocene age; there are no taxa definitive of the Messinian and there are no specimens of G. miocenica whose FAD is at ~3.5 Ma. Our data thus confirm an age of ~5.0—-3.5 Ma for the lower Cayo Agua For- mation. The top of the Cayo Agua Formation was poorly constrained by Coates et al. (1992) as = or > than 2.9 Ma based on the presence of Globoquadrina altispira (LAD at ~2.9 Ma). Work on the nannofossils from PPP 293-301 (Sec. 19) by Bybell (this volume) placed the top of the formation in the upper part of Zone NN15 (3.7-3.8 Ma). From different samples (PPP 348, Sec. 17), we have identified Discoaster brouweri, D. asymetricus, D. surculus, and S. neoabies, and also note the absence of Reticulofenestra pseudoumbilicus. This assemblage would suggest the lowermost Zone NN16 (3.4-3.7 Ma). An age of about 3.4—3.7 Ma seems much more secure than the 2.9 Ma postulated by Coates et al. (1992). Section 16 lies on the coast west of Norte Point and contains a lowermost Zone NN16 nannoflora (3.4—3.7 Ma). The Swan Cay Formation A series of samples was examined from the strato- type (Section 25) for both nannofossils and planktic foraminifera. The foraminifera include large, robust specimens of Globigerinoides ruber, G. conglobatus, G. trilobus and Globorotalia truncatulinoides, and in- dicate Zone N22 (Lower Pleistocene). One specimen of Globoturborotalita nepenthes (LAD at 4.0 Ma) was 40 BULLETIN 357 identified in a sample in the middle of the section. However, no specimens of Globorotalia pseudomio- cenica or any other diagnostic Pliocene species was found. Furthermore, the nannoflora is extensively re- worked, yielding specimens, in various samples, of the Eocene (e.g., Reticulofenestra reticulata) as well as Reticulofenestra pseudoumbilicus, Sphenolithus neoa- bies, Discoaster cf. D. surculus, Ceratolithus cristatus, and? Amaurolithus primus, the last two together char- acterizing Zones NN13 and NN14 (4.5—3.7 Ma). The large and abundant planktic foraminifera associated with the mollusk faunas in samples PPP 1180 and 1181 seem unequivocally Pleistocene', and strongly suggest that the one specimen of Globoturborotalita nepenthes is reworked. LIMON Group, COSTA RICA The Rio Banano Formation The Rio Banano Formation is characterized by the presence of the planktic foraminiferal taxa Sphaeroi- dinellopsis subdehiscens s. str., Globigerinoides con- globatus and Pulleniatina obliquiloculata. The occur- rence of these taxa constrains the age of the Rio Ban- ano Formation to Early Pliocene, ~ 3.5—5.3 Ma. Cal- careous nannoplankton taxa include: Sphenolithus abies (LAD at ~ 3.6 MA), and Pseudoemiliania la- cunosa (FAD at ~ 3.7 Ma) in the middle part of the Rio Banano Formation at Quitaria (Coates et al., 1992), and constrains this section to approximately 3.7 to 3.6 Ma in age. The upper part of the Rio Banano Formation, in the river section at Bomba, contains Discoaster pentara- diatus (LAD at 2.46 Ma) and small (<4 wpm) Gephy- rocapsa spp. (FAD at ~3.7 Ma), which constrain the upper part of the formation to Upper Pliocene (Coates et al., 1992). In the immediately overlying and re- cently exposed landslide section the occurrence of Globorotalia miocenica (FAD at 3.5 Ma) and Dento- globigerina altispira (LAD at ~3.1 Ma) constrains the age of the upper part of the Rio Banano Formation more precisely to the early Late Pliocene. The Quebrada Chocolate Formation Microfossils are rare in the newly defined Quebrada Chocolate Formation. A silty clay in Chocolate Creek ' Editors’ note: New paleomagnetic data (D. McNeill, written commun.) indicate pre-Brunhes Chron deposition for collections from Swan Cay, giving an age of Early Pleistocene, 0.78—1.77 Ma. has yielded small (<0.4 pm) gephyrocapsids, sug- gesting an age of ~< 3.7 Ma for this unit, consistent with biochronologic estimates of the underlying Rio Banano Formation. Age-diagnostic microfossils are absent in other coral-reef-bearing units. The Moin Formation The presence of large gephyrocapsids (FAD at ~ 1.5 Ma) near the top of Section 34, in Cangrejos Creek, indicate an Early Pleistocene (or younger) for this part of the section. More recent examination of the Cangrejos Creek samples by one of us (MPA) has revealed the presence of small (<0.4 wm) gephyro- capsids (LAD 0.96, FAD 2.5—3.7 Ma) which supports the Lower Pleistocene NN1I9 assignment by Bybell (this volume), based on G. truncatulinoides and C. ma- cintyrel. Large gephyrocapsids (FAD at ~ 1.5 Ma) occur in the lagoonal facies of the Moin Formation (Section 34), flanking the Empalme Member, as does Globo- rotalia ungulata, which is considered a Pleistocene marker. The Pueblo Nuevo Sandstone of the Limon For- mation (McNeill ef al., in press), has not yielded un- equivocal age-diagnostic microfossils. A latest Plio- cene age is estimated on the basis of its stratigraphic position between the underlying Quebrada Chocolate Formation and the overlying reef members of the Moin Formation. While the reefal Empalme Member has not yielded any age-diagnostic microfossils, Section 34 (west), newly described here, and Section 36 (east), dated by Bybell (this volume) and Cotton (this volume), rep- resenting the reefal Lomas del Mar Member, contain biostratigraphically useful microfossils. Section 38 is probably of latest Pliocene to Early Pleistocene age. The lower part of the section contains P. lacunosa (LAD at ~0.46 Ma), C. macintyrei (LAD at ~1.59 Ma) and Globigerinoides extremus (LAD at ~1.8 Ma) which provides a minimum age estimate of 1.8 Ma (or younger). The upper part of the section of the Lomas del Mar Reef Member contains G. truncatulinoides (FAD at ~1.9 Ma) and large gephyrocapsids (FAD at ~ 1.5 Ma), suggesting an early Pleistocene (or youn- ger) age. Gephyrocapsa oceanica, a form restricted to the Pleistocene, occurs in other samples. CHAPTER 2 NEOGENE CALCAREOUS NANNOFOSSIL BIOSTRATIGRAPHY OF THE CARIBBEAN COAST OF PANAMA AND COSTA RICA LAUREL M. BYBELL U.S. Geological Survey 926 National Center Reston, Virginia 20192, U.S.A. INTRODUCTION The data presented in this paper are part of the Pan- ama Paleontology Project (PPP) that has as its goal the documentation of the evolutionary and ecological con- sequences to marine species and environments of the final closure of the Isthmus of Panama (Coates et al., 1992; Collins et al., 1995). This event, which separat- ed the oceanic regimes of the Pacific and the Atlantic, contributed to important climatic, oceanographic, and biologic changes. Calcareous nannofossils were stud- ied from marine deposits in Costa Rica and Panama in order to provide precise biostratigraphic ages for sediments that were deposited before, during, and after the final closure of the Isthmus of Panama. The data presented in this paper were collected by the PPP from 1986 to 1991 and represent the initial results of cal- careous nannofossil studies on the Caribbean sections. Other calcareous nannofossil studies in the Caribbean and Gulf of Mexico include Aubry (1993a, 1993b), Gartner ef al. (1983, 1987), Lang and Watkins (1984), and Watkins and Verbeek (1988). Aubry and Berggren (this volume) review nannofossil work that was un- dertaken on PPP samples collected from 1991 to 1996. This later work has added new formations to the Li- mon and Bocan del Toro groups and thus extends and refines the biostratigraphic zonation developed in this first phase of the project. The numbered sections re- ferred to in this chapter are fully described by Coates (Chapter 1 and Appendix B, this volume). Sample numbers refer to the PPP site numbers cataloged in the PPP Database, all of which are also located stratigraph- ically and geographically by Coates (App. A, B, this volume). The nannofossil occurrence data are available at the internet site http://www.fiu.edu/~collins|/. ACKNOWLEDGMENTS I wish to thank Anthony G. Coates, Laurel S. Col- lins, and members of the Panama Paleontology Project for providing all the microfossil samples and for help- ful discussions concerning the study area. Xenia Guer- ra provided detailed computer drawings of the Costa Rica and Panama stratigraphic sections, and these were used to construct the simplified sections figured in this paper. I thank Jean M. Self-Trail for preparing the cal- careous nannofossil samples and drafting the text-fig- ures. I thank Eric de Kaenel, Jose-Abel Flores, and Jean M. Self-Trail for their thoughtful reviews of this paper. Field work was supported by two grants from the National Geographic Society to A. G. Coates and J. B. C. Jackson, and by grants from Sigma Xi and the Roger Tory Peterson Institute to L. S. Collins, and by the Smithsonian Tropical Research Institute (STRI). METHODS The calcareous nannofossil samples were dried in a convection oven to remove residual water, and the dry sediment was placed in vials for long-term storage in the calcareous nannofossil laboratory at the U. S. Geo- logical Survey in Reston, Virginia. A timed settling procedure was used to obtain the optimum sediment- size fraction. For this procedure, a small amount of sample was placed in a beaker, stirred, and settled through 2 cm of water. An initial settling time of one minute was used to remove the coarse fraction, and a second settling time of 10 minutes was used to remove the fine fraction. Smear slides then were prepared from the remaining suspended material. Cover slips were attached to the slides using Norland Optical Adhesive (NOA-65), a clear adhesive that bonds glass to glass and cures when exposed to ultraviolet light. Initially, all samples were examined using a Zeiss Photomicroscope III. A few samples, which were thought to have the best preservation and the highest abundances of calcareous nannofossils, were scanned later using a JEOL 35 scanning electron microscope (SEM). BIOSTRATIGRAPHIC ZONATION In this study, the biostratigraphic zonation of the Neogene strata came from 14 composite outcrop sec- tions in Panama and Costa Rica (Text-figs. 1 and 2). The sequence is based primarily upon the calcareous 42 BULLETIN 357 Pacific Ocean Text-figure 1—Map of Panama and Costa Rica showing general location of investigation. A, region in Bocas del Toro Basin that was examined in this study and is enlarged in Text-figure 2A; B, region in Limon Basin that was examined in this study and is enlarged in Text-figure 2B. nannofossil zonation of Martini (1971), and second- arily upon that of Bukry (1973, 1975, 1978) and Oka- da and Bukry (1980). There was considerable variation in the abundance and preservation of the calcareous nannofossil assemblages, but they generally were sparse with fair preservation. Most sections contained at least a few barren samples, and some sections had more than half of the samples barren of calcareous nannofossils. However, there were sufficient numbers of specimens, diversity of taxa, and preservational state to allow dating of almost all samples that did contain calcareous nannofossils. Because of the poor preservation in some of the samples, specimens could be identified only to the proper genus (i.e., Discoaster sp., Sphenolithus sp.). Table 1 is a list of calcareous nannofossil species that can be used to date sediments of Miocene, Plio- cene, and Pleistocene age throughout the world. Not all of these species are present in the study area. Zonal markers for the standard Martini zonation are indicated with an *, and a # indicates a zonal marker for the Bukry zonation. The remaining species have been found to be biostratigraphically useful by various au- thors. Placement of these additional first appearance datums (FAD’s) and last appearance datums (LAD’s) within a particular calcareous nannofossil zone is gen- erally accurate, but the relative positions of FAD’s and LAD’s to each other within an individual zone are much less accurate. Subdivision of Zone NN 19 is from Gartner (1977). Zone NN 19a is used to denote Gartner’s Cyclococcolithina macintyrei Zone, Zone NN 19b is his Helicopontosphaera sellii Zone, Zone NN 19c is his small Gephyrocapsa Zone, and Zone NN 19d is his Pseudoemiliania lacunosa Zone. Most species in Table 1 are illustrated in Perch-Nielsen (1985). The ages for the FAD’s and LAD’s are from Berggren et al. (1985). Text-figure 3 is a correlation chart from Berggren et al. (1985) that has been mod- ified to show only relative placement of calcareous nannofossil zones, planktic foraminiferal zones, ep- ochs, and the geochronometric scale. SOUTHERN LIMON BASIN The southern Limon sedimentary basin is located on the Caribbean (north) coast of Costa Rica near the bor- der with Panama (Text-fig. 1). Samples from seven locations were examined for calcareous nannofossils (Text-fig. 2B). They are Rio Sandbox (Section 27), Carbon Dos (Section 28), Rio Banano (Section 29), Santa Rita (Section 32), Pueblo Nuevo Cemetery (Sec- tion 35), Lomas del Mar, Eastern Sequence (Section 36), and Lomas del Mar, Western Reef Flank (Section 37). All the sediments that are exposed at these local- ities are included in the Limon Group (see Coates et al., 1992; Coates, Chapter 1, this volume). LIMON GROUP This group consists of the Uscari, Rio Banano, Que- brada Chocolate, and Moin formations, which range from Late Miocene to Late Pleistocene. The litholog- ical descriptions of the formations are given in Coates (this volume). Uscari Formation—Zone NN 11 (Upper Miocene)— 8.2—5.6 Ma This formation, the oldest in the Limon Group, is at least 565 m thick and consists of biocalcarenite and shale, which in places contains a basaltic sill in its upper part. Calcareous nannofossils were examined from two localities that expose the Uscari Formation: Rio Sandbox (Section 27) and Carbon Dos (Section 28; Text-fig. 2B). Both localities contain calcareous nannofossils that place this formation in the Upper Miocene Zone NN 11 of Martini (1971). See Cotton (this volume) for a discussion of the age of this for- mation at these two localities based on planktic fora- minifera. Rio Sandbox locality, Section 27 (Zone NN 11) Three samples were examined from the upper 20 m of the 40-m-thick section of the Uscari Formation at the Rio Sandbox locality. All three samples are placed in the Upper Miocene Zone NN 11 based on the pres- ence of Discoaster berggrenii or Discoaster quinquer- amus. Both species only occur in Zone NN 11. The following is a list of the species present in each of the CALCAREOUS NANNOFOSSIL BIOSTRATIGRAPHY: BYBELL 43 Valiente Peninsula Canbbean Sea Escudo de Veraguas Nancy Point Cayo Agua Chon Pont) Bruno Bluff Caribbean Sea Text-figure 2.—Locations of outcrops in the Bocas del Toro Basin, Panama (A), and in the Limon Basin, Costa Rica (B), that were examined for this study. Numbers refer to assigned section numbers, and more precise geographic positions and composite stratigraphic sections can be found in Appendices A and B, this volume. Map A—10, Escudo de Veraguas, Northern Coast; 11, Escudo de Veraguas, Southeastern Coast; 12, Valiente Peninsula, Bruno Bluff to Plantain Cays; 16, Cayo Agua, North Point, Western Side, 18, Cayo Agua, Piedra Roja Point, Eastern Sequence; 19, Cayo Agua, North Point to Tiburon Point. Map B—27, Rio Sandbox; 28, Carbon Dos; 29, Rio Banano; 32, Santa Rita; 35, Pueblo Nuevo Cemetery; 36, Lomas del Mar, Eastern Sequence; and 37, Lomas del Mar, Western Reef Flank Sequence. 44 BULLETIN 357 Table 1—Calcareous nannofossil species useful for dating Miocene, Pliocene, and Pleistocene sediments. * = zonal marker, Martini (1971) zonation. # = zonal marker, Bukry (1973, 1975, 1978) zonation. Ages (Ma) are from Berggren er al., 1985. Species Martini zones Bukry zones Age (Ma) PLEISTOCENE FAD Emiliania huxleyi acme in NN 21 in CN 15 0.085 *#FAD Emiliania huxleyi base NN 21 base CN 15 0.272 *#LAD Pseudoemiliania lacunosa top NN 19d top CN I4a 0.474 FAD dominant Gephyrocapsa larger base NN 19d in CN l4a LAD dominant Gephyrocapsa smaller top NN 19¢ in CN I4a LAD Helicosphaera sellii top NN 19b in CN I4a 1.37 LAD Calcidiscus macintyrei top NN 19a in CN l4a 1.45 LAD Ceratolithus rugosus in NN 19a in CN |4a #FAD Gephyrocapsa oceanica in NN 19a base CN |4a 1.68 #FAD Gephyrocapsa caribbeanica in NN 19a base CN 13b 1.74 PLIOCENE FAD Gephyrocapsa \arger species in NN 19a in CN 13a *#LAD Discoaster brouweri top NN 18 top CN 12d 1.9 LAD Discoaster triradiatus mid NN 18 in CN 12d 129, FAD Gephyrocapsa aperta in NN 18 in CN 12d DD. LAD Discoaster asymmetricus top NN 17 top CN 12c DD. *#LAD Discoaster pentaradiatus top NN 17 top CN 12c 2.4 *#LAD Discoaster surculus top NN 16 top CN 12b 2.4 #LAD Discoaster tamalis upper NN 16 top CN 12a 2.6 LAD Discoaster challengeri mid NN 16 in CN 12a LAD Discoaster decorus mid NN 16 in CN 12a 2.9 LAD Discoaster variabilis mid NN 16 in CN 12a 2.9 #LAD Sphenolithus neoabies top NN 15 top CN 11b 3.5 LAD Sphenolithus abies top NN 15 top CN 11b 35 *#LAD Reticulofenestra pseudoumbilicus top NN 15 top CN 11b Shs) FAD Pseudoemiliania lacunosa upper NN 15 in CN Ila/b 3.6 modified herein FAD Discoaster tamalis mid NN 15 in CN 1la/b 3.8? #LAD Amaurolithus primus top NN 14 top CN 10d 3h7/ *#LAD Amaurolithus tricorniculatus top NN 14 top CN 10d 37) *FAD Discoaster asymmetricus base NN 14 base CN 10d 4.1 *#FAD Ceratolithus rugosus base NN 13 base CN 10c 4.5 #LAD Ceratolithus acutus top NN 12 top CN 10b 4.6 #FAD Ceratolithus acutus mid NN 12 base CN 10b 5.0 MIOCENE #LAD Triquetrorhabdulus rugosus mid NN 12 top CN 10a 5.0 FAD Helicosphaera sellii base NN 12 base CN 10a 5.6 *#LAD Discoaster quinqueramus top NN 11 top CN 9d? 5.6 LAD Amaurolithus amplificus upper NN 11 top CN 9c? 5.6 FAD Amaurolithus amplificus upper NN 11 base CN 9c? S2) LAD Discoaster berggrenii upper NN 11 in CN 9b 5.6 FAD Amaurolithus tricorniculatus upper NN 11 in CN 9b 6.0 LAD Discoaster neohamatus upper NN 11 in CN 9b FAD Amaurolithus delicatus upper NN 11 in CN 9b 6.5 #FAD Amaurolithus primus upper NN 11 base CN 9b 6.5 #FAD Discoaster berggrenii base NN 11 base CN 9a 8.2 *FAD Discoaster quinqueramus base NN 11 base CN 9a 8.2 #FAD Discoaster surculus base NN 11 base CN 9a 8.2 LAD Discoaster bollii upper NN 10 in CN 8b 8.3 #FAD Discoaster neorectus mid NN 10 base CN 8b 8.5 #FAD Discoaster loeblichii mid NN 10 base CN 8b 8.5 LAD Catinaster coalitus lower NN 10 in CN 8a 9.0 LAD Catinaster calyculus lower NN 10 in CN 8a 8.8 LAD Discoaster exilis lower NN 10 in CN 8a 8.8 *#LAD Discoaster hamatus top NN 9 top CN 7b 8.9 FAD Discoaster neohamatus upper NN 9 in CN 7b #FAD Catinaster calyculus in NN 9 base CN 7b 10.0 FAD Discoaster pentaradiatus lower NN 9 in CN 7a *#FAD Discoaster hamatus base NN 9 base CN 7a 10.0 CALCAREOUS NANNOFOSSIL BIOSTRATIGRAPHY: BYBELL 45 Table 1.—Continued. Species *#FAD Catinaster coalitus FAD Discoaster challengeri *#FAD Discoaster kugleri #LAD Reticulofenestra floridana *#LAD Sphenolithus heteromorphus FAD Discoaster variabilis #FAD Calcidiscus macintyrei *LAD Helicosphaera ampliaperta #FAD Sphenolithus heteromorphus *LAD Sphenolithus belemnos *LAD Triquetrorhabdulus carinatus #FAD Sphenolithus belemnos *#FAD Discoaster druggii FAD Helicosphaera ampliaperta #LAD Dicytococcites bisectus *LAD Helicosphaera recta #LAD Sphenolithus ciperoensis Martini zones base NN 8 mid NN 7 base NN 7 top NN 6 top NN 5 base NN 5 base NN 5 top NN 4 in NN 4 top NN 3 top NN 2 in NN 2 base NN 2 base NN 2 top NP 25 top NP 25 top NP 25 samples. Species that are in bold are the most useful for dating the samples. Sample 737 (36.0 m above base of section) Discoaster quinqueramus Reticulofenestra pseudoumbilicus Sphenolithus abies Abundance: common Preservation: fair Age: Zone NN 11 Sample 735 (25.0 m above base of section) Calcidiscus macintyrei Helicosphaera carteri Reticulofenestra pseudoumbilicus Sphenolithus abies Abundance: common Preservation: fair Age: Zone NN 11 by superposition Sample 736 (21.0 m above base of section) Discoaster berggrenii Discoaster brouweri Helicosphaera carteri Reticulofenestra pseudoumbilicus Sphenolithus abies Abundance: frequent Preservation: fair Age: Zone NN 11 Bukry zones Age (Ma) base CN 6 10.8 mid CN 5b base CN 5b 13.1 top CN Sa 11.6 top CN 4 14.4 base CN 4 16.2 base CN 4 16.2 top CN 3 16.2 base CN 3 Nea in CN 2 17.4 in CN 2 19.0 base CN 2 2S base CN lc 232: base CN Ic PUEN PDN top CP 19b 2307. top CP 19b top CP 19b Zone NN 11. The following is a list of the species present in each of the two samples. Species that are in bold are the most useful for dating the samples. Sample 726 (1.0 m above base of section) Calcidiscus leptoporus ?Catinaster Coccolithus pelagicus Discoaster berggrenti Discoaster sp. aff. D. exilis Discoaster pentaradiatus Discoaster quinqueramus Helicosphaera carteri Reticulofenestra pseudoumbilicus Sphenolithus abies Abundance: frequent Preservation: fair Age: Zone NN 11 Sample 727 (3.0 m above base of section) Carbon Dos locality, Section 28 (Zone NN 11) Two samples were examined from the 15-m-thick section of the Uscari Formation at Carbon Dos, both of which are placed in the Upper Miocene Zone NN 11 based on the presence of Discoaster berggrenii and/ or Discoaster quinqueramus, which only occur within Calcidiscus macintyrei Coccolithus pelagicus Discoaster berggrenii Discoaster brouweri Discoaster pentaradiatus Helicosphaera carteri Reticulofenestra pseudoumbilicus Reticulofenestra small species Sphenolithus abies Abundance: frequent Preservation: fair Age: Zone NN 11 Rio Banano Formation—Zones NN 15-17 (upper Lower to Upper Pliocene )—3.6—2.2 Ma This formation, which overlies the Uscari Forma- tion, can be 750 m thick and consists primarily of silt- 46 ) ® x 3 Be} 2 @ ao [_d_INN 18} |b | 4 ® 8 is) a LN 18 J Planktonic Foraminifera Zone Calcareous Nannofossil Zone Bukry (1975) Calcareous Nannofossil Zone seme CN 11F-3-JNN 15 N19 ae sy = BULLETIN 357 stone and sandstone. Calcareous nannofossils were ex- amined from two localities: Rio Banano (Section 29) and Santa Rita (Section 32). These sediments can be placed in the upper Lower to Upper Pliocene at Rio Banano and in the Upper Pliocene at Santa Rita, where only the upper part of the formation is exposed. Rio Banano locality, Section 29 (upper Zone NN 15 to Zone NN 17) Twenty-one samples were examined from the 893- m-thick section of the Rio Banano Formation at the Rio Banano locality. Four of these samples were bar- ren of calcareous nannofossils (Text-fig. 4). The lowest 15 samples are placed in the upper part of Zone NN 15 because of the presence in this interval of Sphen- olithus abies, Sphenolithus neoabies, and Reticulofe- nestra pseudoumbilicus (LAD’s at the top of Zone NN 15) and Pseudoemiliania lacunosa (FAD near the top of Zone NN 15). The overlying samples 672 (789.0 m) and 671 (790.0 m) are probably in Zone NN 16; they do not contain representatives of the genus Sphen- olithus or the species R. pseudoumbilicus (which plac- es them above Zone NN 15) or any smaller represen- tatives of the genus Gephyrocapsa, which first appears in Zone NN 17 in the study area. The upper four sam- ples in this section, 670 (791.0 m), 669 (792.5 m), 668 (794.0 m), and 678 (832.5 m), tentatively are placed in Zone NN 17 based on the presence of smaller spec- imens of the genus Gephyrocapsa and specimens of Discoaster pentaradiatus (LAD defines the top of Zone NN 17). Santa Rita locality, Section 32 (Zone NN 16) Five samples were examined from the 60-m-thick section of the upper part of the Rio Banano Formation at Santa Rita. The samples from this locality tenta- tively were placed in Zone NN 16 (Upper Pliocene) (Text-fig. 5). There appears to be some reworking in this section, as evidenced by single specimens of Re- ticulofenestra pseudoumbilicus in samples 721 (38 m) and 720 (58 m) and a single specimen of the genus Sphenolithus in sample 723 (13 m). Single occur- rences of smaller specimens of the genus Gephyro- capsa also occur in samples 723 and 720. If these occurrences are considered to be due to contamina- tion or reworking, these samples may be placed in Zone NN 16 (younger than Zone NN 15 because of the absence of Sphenolithus and Reticulofenestra os Text-figure 3.—Correlation of calcareous nannofossil zones, planktic foraminiferal zones, geochronometric scale, and epochs (from Berggren er al., 1985) for the Neogene period. CALCAREOUS NANNOFOSSIL BIOSTRATIGRAPHY: BYBELL 47 Rio Banano Formation Formation @| Calcidiscus macintyrei Coccolithus pelagicus Cribrocentrum reticulatum @| Cyclococcolithusspp. Dictyococcites bisectus Discoaster asymmetricus @} Discoaster brouweri Discoaster challengeri Discoaster deflandrei ®) Discoaster pentaradiatus Discoaster tamalis @| Discoasterspp. @| Gephyrocapsa smaller species @| Helicosphaera carter Helicosphaera euphratis Helicosphaera sellii Lithostromation operosum Lithostromation perdurum @| Pontosphaera discopora Pontosphaeraspp. ®) Pseudoemiliania lacunosa Reticulofenestra pseudoumbilicus Rhomboaster sp. aff. R. orthostylus| Scyphospaera amphora Scyphosphaera apsteinii Sphenolithus abies Sphenolithus neoabies Sphenolithus sp. @| Thoracosphaera spp. F B F R F F R C_ ClAbundance P F Paar FF) Preservation Text-figure 4.—Calcareous nannofossil occurrences in the Rio Banano Section 29, Costa Rica. See Text-figure 2 for location of exposure. For Text-figures 4—10, the following symbols are used. Abundance: A, abundant or >10 specimens per field of view at X500; C, common or 1-10 specimens per field of view at X500; EF frequent or 1 specimen per 1-10 fields of view at 500: R, rare or 1 specimen for >10 fields of view; B, barren of calcareous nannofossils. Preservation: G, good; F fair; P, poor. Other symbols: R, specimens likely reworked;? , possible occurrence; numbers, number of specimens observed in the entire sample. 48 BULLETIN 357 Rio Banano Formation Formation Pliocene p Series Braarudosphaera bigelowii Calcidiscus leptoporus Calcidiscus macintyrei Coccolithus pelagicus Discoaster brouwen Discoaster pentaradiatus Gephyrocapsa smaller species Helicosphaera carteri Pontosphaera discopora Pseudoemiliania lacunosa Reticulofenestra pseudoumbilicus Reticulofenestra spp. Rhabdosphaera clavigera Sphenolithus spp. Thoracosphaera spp. Abundance Preservation Text-figure 5 —Calcareous nannofossil occurrences in the Santa Rita Section 32, Costa Rica. See Text-figure 2 for location of ex- posure and Text-figure 4 for explanation of abundance, preservation, and other symbols. pseudoumbilicus, and older than Zone NN 17 by the absence of smaller specimens of the genus Gephy- rocapsa). Even if the presence or absence of these three species were discounted, the presence of both Pseudoemiliania lacunosa (FAD very near the top of Zone NN 15) and Discoaster pentaradiatus (LAD de- fines the top of Zone NN 17) in these samples con- fines them to Zone NN 16 or Zone NN 17. See Cotton (this volume) for a discussion of the planktic fora- minifera from the Rio Banano locality. Moin Formation—Zone NN 17-19a or b (Upper Pliocene )—2.4—1.7 Ma possibly includes Zone NN 21 (Upper Pleistocene)—2.4—0.275 Ma This formation, which overlies the Rio Banano For- mation, can be 200 m thick and consists of alternating claystone and sandstone. Calcareous nannofossils were examined from three localities: Pueblo Nuevo Ceme- tery (Section 35), Lomas del Mar, Eastern Sequence (Section 36), and Lomas del Mar, Western Reef Flank (Section 37) (Text-fig. 2B). Most of the Moin For- mation is Upper Pliocene in age, but some Moin sed- iments may be as young as Upper Pleistocene. Pueblo Nuevo Cemetery locality, Section 35 (Zones NN 17-19) Six samples were examined from the 94-m-thick section of the Moin Formation at the Pueblo Nuevo Cemetery. From bottom to top, the samples and their meters above section base include 632 (85.5 m), 631 (89.0 m), 630 (89.5 m), 629 (90.5 m), 628 (90.5 m), and 633 (93.0 m). Five of these samples were barren. Only sample 631 (89.0 m) contained calcareous nan- nofossils, which indicate either an Upper Pliocene or a Lower Pleistocene age. Gephyrocapsa small species first appear in Zone NN 17 in the study area, and Pseu- doemiliania lacunosa last appears in the upper part of Zone NN 19. The following is a list of the species present in this sample. Species in bold are the most useful for dating the sample. Sample 631 Calcidiscus leptoporus Gephyrocapsa smaller species Helicosphaera carteri Pontosphaera spp. Pseudoemiliania lacunosa—1| specimen Reticulofenestra pseudoumbilicus—1 specimen (re- worked?) Thoracosphaera spp. Abundance: common Preservation: fair Age: upper Pliocene or lower Pleistocene, Zone NN 17-19 Lomas del Mar, Eastern Sequence, Section 36 (Zones NN 17/18 or NN 19 and NN 21) Thirteen samples were examined from the 70-m- thick composite section of the Moin Formation at Lo- mas del Mar, Eastern Sequence. This was a difficult section to date because of the large amount of mixing present (Text-fig. 6). There are Late Pliocene to Pleis- tocene specimens, some obvious Miocene specimens, probably some Oligocene specimens, and a large num- ber of Eocene specimens all occurring in the same samples. If one assumes that reworking is the most likely source for this mixing, then the youngest age present (Late Pliocene to Pleistocene) should reflect the age of deposition for the sediments. However, there are very few of these younger specimens. Only sam- ples 710 (7 m) and 738 (13.5 m) contain specimens of Gephyrocapsa larger species (FAD in Zone NN 19). They are absent from the other 11 samples in this sec- CALCAREOUS NANNOFOSSIL BIOSTRATIGRAPHY: BYBELL 49 Moin Formation Pliocene Pleistocene Seri Late Tate eres NN 17/1 alcareous Nannofossi wo Zz Zz nN Zone (Martini, 1971 ‘ ey if Meters Above Base of Section Sample Number Barren Braarudosphaera bigelowii Calcidiscus leptoporus 8 wo on rar) D o- a Olly) OL veg] SZL seg] SEL sez] OL LEQ] Sz Sy9] SO Zyg} O CE eg] S Org] Ss Leg} S 9EQ] S2z 9e ge s9 ello] Se Calcidiscus macintyrei Catinaster coalitus Ceratolithus spp Coccolithus pelagicus Cribrocentrum reticulatum Cyclococcolithus formosus Cyclococcolithus neogammation Dictyococcites bisectus Discoaster barbadiensis Discoaster berggrenii Discoaster brouweri Discoaster challengeri Discoaster deflandrei Discoaster pentaradiatus Discoaster trradiatus? Discoaster woodringii Discoaster spp Emiliania huxleyi Gephyrocapsa protohuxleyi Gephyrocapsa larger species Gephyrocapsa smaller species Helicosphaera bramlettei Helicosphaera carter Helicosphaera compacta Helicosphaera euphratis Helicosphaera sellii Markalius inversus Pentaster lisbonensis Pontosphaera discopora Pontosphaera multipora Pontosphaera spp Pseudoemiliania lacunosa Reticulofenestra floridana Reticulofenestra pseudolockeri Reticulofenestra pseudoumbilicus Reticulofenestra spp Rhabdosphaera clavigera Rhabdosphaera spp Scyphosphaera amphora Sphenolithus abies Sphenolithus tribulosus/predistentus Sphenolithus spp Syracosphaera pulchra Thoracosphaera spp. F | Abundance Preservation Text-figure 6.—Calcareous nannofossil occurrences in the Lomas del Mar, Eastern Sequence Section 36, Costa Rica. See Text-figure 2 for location of exposure and Text-figure 4 for explanation of abundance, preservation, and other symbols. 50 BULLETIN 357 tion. The presence of the planktic foraminifer Globor- otalia truncatulinoides (Cotton, this volume) demon- strates an age of <1.9 Ma, which supports placement in calcareous nannofossil Zone NN 19. If this is true, and the Gephyrocapsa specimens are in place, then specimens of Discoaster brouweri (LAD at top of Zone NN 18), which are found in six of the samples in this section, must be reworked. An alternative choice is to consider D. brouweri to be in place and position these samples in either Zone NN 17 or NN 18 based on the presence of smaller specimens of the genus Gephyrocapsa (FAD probably in Zone NN 17) and D. brouweri. The highest sample from this section, 627 (65.5 m), was collected from a small construction site just north of the Pueblo Nuevo Cemetery. Akers (1972) exam- ined a sample of the Moin Formation from an outcrop near here, and he placed his sample in the Pleistocene, primarily on the basis of the foraminifera. He found no discoasters in his sample, which he stated could indicate a post-Pliocene age. Sample 627 from the cur- rent study, which has been placed tentatively in the Moin Formation, contains Emiliania huxleyi (FAD at base of Zone NN 21), and this indicates placement in the uppermost Pleistocene Zone NN 21. The presence of E. huxleyi was confirmed with a scanning electron microscope. Lomas del Mar, Western Reef Flank Sequence, Section 37 (Zone NN 19) Eleven samples were examined from the 74-m-thick section of the Moin Formation at its type locality at Cangrejos Creek. All eleven samples contained fre- quent to common calcareous nannofossils with fair to good preservation (Text-fig. 7). There is a minor amount of reworking in this section. The nine lower samples all could be placed in Zone NN 19a because they do not contain Discoaster brouweri (LAD defines the top of Zone NN 18) and do contain Calcidiscus macintyrei (LAD at the top of Zone NN 19a). The presence of a noticeable amount of larger specimens of the genus Gephyrocapsa (FAD in Zone NN 19) in these samples also indicates that they are younger than Zone NN 18. Samples 654 (64 m) and 657 (71 m), the uppermost samples at this locality, each contain only a single specimen of C. macintyrei and no Dis- coaster species. If these occurrences are valid, then these samples should be placed in Zone NN 19a. How- ever, if these are reworked specimens, then samples 654 (64 m) and 657 (71 m) should be placed in Zone NN 19b. For the purposes of this paper, these samples are considered to be in either Zone NN 19a or Zone NN 19b. Planktic foraminifera from this locality (Cot- ton, this volume) indicate an age of 1.8—1.9 Ma, which is consistent with placement of these sediments within Zone NN 19a. BOCAS DEL TORO BASIN—PANAMA The Bocas del Toro sedimentary basin is located on the Caribbean coast of Panama near the border with Costa Rica in the Bocas del Toro Province (Text-fig. 1A). See Coates et al. (1992) and Coates (this volume) for a discussion of the geology of this region. Samples from seven locations were examined for calcareous nannofossils (Text-fig. 2B). They are Valiente Penin- sula, Bruno Bluff to Plantain Cays (Section 12), Cayo Agua, Punta Norte, Western Side (Section 16), Cayo Agua, Punta Norte to Punta Tiburon (Section 19), Cayo Agua, South of Punta Nispero (Section 20), Cayo Agua, Punta Piedra Roja, Eastern Sequence (Section 18), Escudo de Veraguas Northern Coast (Section 10), and Escudo de Veraguas, Southeastern Coast (Section 11). All of the sediments that were col- lected from these sections are in the Bocas del Toro Group, first described by Coates er al. (1992) and up- dated by Coates (this volume). BOcAS DEL TORO GROUP This group consists of five formations: the Tobobe Sandstone, the Nancy Point, Shark Hole Point, Cayo Agua, and Escudo de Veraguas formations, Upper Miocene to Upper Pliocene. See Coates, this volume, for lithologic descriptions. Valiente Peninsula, Bruno Bluff to Plantain Cays locality, Section 12 (Zones NN 11-15) Eleven samples were examined from the approxi- mately 1,960-m-thick section of the Bocas del Toro Group on the north coast of the Valiente Peninsula. The lower four samples in this section are in the Nancy Point Formation, and the upper seven samples are in the Shark Hole Point Formation. Text-figure 8 is a cal- careous nannofossil occurrence chart for this section. Nancy Point Formation—Zone NN 11 (Upper Miocene)—8.2—5.6 Ma The Nancy Point Formation, the oldest in the Bocas del Toro Group, can be 378 m thick and consists of volcanic sandstone with abundant clay and silt in the matrix. Between the lower three samples of the Nancy Point Formation and the uppermost sample from this formation, there is a thick interval that was not ex- amined for calcareous nannofossils. Sample 409 (1,232 m), the oldest sample examined from this for- mation, was barren of calcareous nannofossils. Sam- ples 408 (1,236 m), 407 (1,244 m), and 390 (1,606 m) were placed in the Upper Miocene Zone NN 11 be- cause they contain Discoaster berggrenii and/or Dis- CALCAREOUS NANNOFOSSIL BIOSTRATIGRAPHY: BYBELL 51 Series Calcareous Nannofossil Zone (Martini, 1971) Braarudosphaera bigelowii Calcidiscus leptoporus Calcidiscus macintyre! Ceratolithus sp. Coccolithus pelagicus Cyclococcolithus neogammation Cyclococcolithus spp Discoaster berggrenii Discoaster deflandrei Discoaster spp. Gephyrocapsa larger species Gephyrocapsa smaller species Helicosphaera bramlettei Helicosphaera carteri Helicosphaera sellii Lithostromation perdurum Pontosphaera discopora Pontosphaera millepuncta Pontosphaera multipora Pontosphaera spp. Pseudoemiliania lacunosa Rhabdosphaera clavigera Rhabdosphaera spp. Scyphosphaera amphora Scyphosphaera apsteinii Scyphosphaera spp. Sphenolithus abies Sphenolithus spp. Syracosphaera pulchra Thoracosphaera spp. Abundance Preservation Text-figure 7—Calcareous nannofossil occurrences in the Lomas del Mar, Western Reef Flank Sequence Section 37, Costa Rica. See Text- figure 2 for location of exposure and Text-figure 4 for explanation of abundance, preservation, and other symbols. BULLETIN 357 Nn NO Nancy Point Formation Shark Hole Point Formation Miocene 1 Pliocene Formation r o = oO Le | cs z z C4 v a3 az za zc i. 3 kel fe | Calcareous Nan Zo nofossil artini, 1971 Meters Above Base of Section Sample Number Gees | 60P |CECL 80 |9ECL LOv |vvcl O6E |9091 88E |1Lc9 68E |ZZ9L L8€ |6E91 98E |6E9L OLE |ZV6L LZLE |8V6L BLE |CSEL e e Calcidiscus leptoporus OOr O © one Calcidiscus macintyrei O O Ceratolithus acutus e Ceratolithus spp. ee ee Coccolithus pelagicus Cyclococcolithus spp. Discoaster berggrenii Discoaster brouweri Discoaster challengeri Discoaster sp. aff. exilis Discoaster pentaradiatus Discoaster quinqueramus Discoaster surculus Discoaster variabilis Discoaster spp. Helicosphaera carter Helicosphaera euphratis Helicosphaera lophota Helicosphaera sellii Markalius inversus Pontosphaera discopora Pontosphaera millepuncta Pontosphaera multipora Pseudoemiliania lacunosa Reticulofenestra pseudoumbilicus Reticulofenestra spp. Rhomboaster sp. aff. orthostylus Sphenolithus abies Thoracosphaera spp. Abundance Preservation Text-figure 8. figure 2 for location of exposure and Text-figure 4 for explanation of abundance, preservation, and other symbols. Calcareous nannofossil occurrences in the Valiente Peninsula, Bruno Bluff to Plantain Cays Section 12, Panama. See Text- CALCAREOUS NANNOFOSSIL BIOSTRATIGRAPHY: BYBELL 53 Table 2.—The approximate duration in millions of years for the Neogene NN Zones as presented in Berggren er al. (1985). Numbers in parentheses are revised ages from Berggren er al. (1995). Zone NN 21—0.275 (0.26) Zone NN 10—0.65 (0.8) Zone NN 20—0.199 (0.2) Zone NN 9—1.15 (1.8) Zone NN 19—1.4 (1.49) Zone NN 8—0.8 (0.6) Zone NN 18—0.3 (0.5) Zone NN 7—2.3 Zone NN 17—0.2 (0.15) Zone NN 6—1.3 Zone NN 16—1.1 (1.15) Zone NN 5—1.8 (2.0) Zone NN 15—0.2 (0.23) Zone NN 4—1.2 (2.7) Zone NN 14—0.4 (0.19) Zone NN 3—1.5 (0.7) Zone NN 13—0.4 (0.83) Zone NN 2—4.3 (4.2) Zone NN 12—1.1 (0.6) Zone NN 1—0.5 (0.7) Zone NN 11—2.6 (3.0) coaster quinqueramus. Both species only occur in Zone NN 11 (Table 1). This age is consistent with the planktic foraminiferal ages for this formation (Cotton, this volume). Shark Hole Point Formation—Zones NN 12-15 (upper Miocene to lower Pliocene )—5.6—3.6 Ma Seven samples of the Shark Hole Point Formation were sampled from the north coast of the Valiente Pen- insula. The lower part of this section exposes the Nan- cy Point Formation, while the upper part exposes the Shark Hole Point Formation. Here, the Shark Hole Point Formation can be 341 m thick and consists pre- dominantly of siltstone. Text-figure 8 is an occurrence chart for Valiente Peninsula section. Samples 388 (1,621 m) and 389 (1,627 m), from the lower part of the Shark Hole Point Formation, are placed in Zone NN 12 (Upper Miocene to Lower Pliocene) because they do not contain either Discoaster berggrenii or Discoaster quinqueramus (the marker species for Zone NN 11), which do occur in the underlying Nancy Point Formation. These samples occur below samples 387 and 386 (both from 1,639 m), which definitely are in Zone NN 12, based on the presence of Ceratolithus acutus. Berggren et al. (1985) placed the Miocene- Pliocene boundary approximately one-third of the way up into Zone NN 12. Samples 388 and 389 could therefore have been deposited either in the Upper Mio- cene or the Lower Pliocene. Sample 387 contains Cer- atolithus acutus, a species that only occurs in the upper part of Zone NN 12 (Lower Pliocene) between 5.0 and 4.6 Ma (Berggren et al., 1985). Sample 386 occurs at the same stratigraphic position as sample 387. Sample 376 (1,947 m) is placed in the upper part of Zone NN 15 (Lower Pliocene) because it contains Reticulofe- nestra pseudoumbilicus, Sphenolithus abies (both have their LAD’s at the top of Zone NN 15), and Pseudoem- iliania lacunosa (FAD near the top of Zone NN 15). Small, poorly preserved, rare, and difficult-to-identify specimens of P. lacunosa were described by Rio et al. (1990) from farther down in Zone NN 15. They stated that “‘it becomes abundant and more easily recogniz- able close to the extinction level of R. pseudoumbili- cus.” It is this upper horizon with more common and easily identified specimens that is considered signifi- cant for this paper. Sample 377 (1,948 m) was collected from slump material. This is corroborated by calcareous nannofos- sils because sample 377 contains an almost identical flora to sample 387. Both samples contain the very short ranging species Ceratolithus acutus (upper Zone NN 12), and these two samples are presumed to be from the same sedimentary deposit. Sample 378 (1,952 m), the highest sample examined in the section, is barren of calcareous nannofossils. Planktic forami- nifera from this location (Cotton, this volume) are able to restrict further the age of this formation to 5.3—3.4 Ma, or the Lower Pliocene. Cayo Agua Formation—upper Zone NN 15 (upper lower Pliocene )—3.6—3.5 Ma The Cayo Agua Formation either overlies or is a facies equivalent to the upper part of the Shark Hole Point Formation and is in general much coarser. Cal- careous nannofossils were examined from this forma- tion at four localities: North Point, Western Side (Sec- tion 16), North Point to Tiburon Point (Section 19), South of Nispero Point (Section 20), and Piedra Roja Point, Eastern Sequence (Section 18) (Text-fig. 2B). Cayo Agua, North Point, Western Side locality, Section 16 (upper Zone NN 15) Calcareous nannofossils were examined from one sample of the Cayo Agua Formation at the North Point locality. At this site, sample 57, which was collected 49 m above the base of the 50-m-thick section, can be placed in the upper Lower Pliocene in the upper part of Zone NN 15 based on the presence of Sphenolithus abies (LAD near the top of Zone NN 15) and Pseu- doemiliania lacunosa (FAD in the upper part of Zone NN 15). The following is a list of the species present in this sample. Species that are in bold are the most useful for dating this sample. Sample 57 (49 m) Calcidiscus macintyrei Discoaster brouweri Discoaster pentaradiatus Helicosphaera carteri Helicosphaera sellii Pontosphaera discopora Pseudoemiliania lacunosa Reticulofenestra small species 54 BULLETIN 357 Cayo Agua Formation Early Pliocene Formation Series Rib? comer Calcareous Nannofossil Zone (Martini, 1971 Meters Above Base of Section Sample Number | Braarudosphaera bigelowii Calcidiscus leptoporus Calcidiscus macintyrei Ceratolithus rugosus Coccolithus pelagicus Cyclococcolithus spp. Discoaster brouweri Discoaster pentaradiatus Discoaster spp. Helicosphaera carteri Helicosphaera euphratis Helicosphaera sellii Pontosphaera millepuncta Pontosphaera multipora Pseudoemiliania lacunosa Reticulofenestra pseudoumbilicus Reticulofenestra spp. Rhabdosphaera spp. Rhomboaster sp. aff. R. orthostylus Sphenolithus abies Thoracosphaera spp. AGC CAF RB BB FB BB R RB |Abundance FFFFFE F i> |p Preservation Text-figure 9.—Calcareous nannofossil occurrences in the Cayo Agua, North Point to Tiburon Point, Section 19, Panama. See Text-figure 2 for location of exposure and Text-figure 4 for explanation of abundance, preservation, and other symbols. Sphenolithus abies Thoracosphaera spp. Abundance: common Preservation: fair Age: upper Zone NN 15 Cayo Agua, North Point to Tiburon Point locality, Section 19 (upper Zone NN 15) Sixteen samples from the Cayo Agua Formation were examined for their calcareous nannofossil content from this 293-m-thick section. No samples were ex- amined for calcareous nannofossils from the lowest 45 m of the exposure. Of the 16 samples examined, 7 were barren, and the remaining nine samples contained rare to abundant calcareous nannofossils (Text-fig. 9). Most of the sediments from this section can be placed in calcareous nannofossil Zone NN 15. The co-occur- rence of Reticulofenestra pseudoumbilicus (LAD at top of Zone NN 15), Sphenolithus abies (LAD at top of Zone NN 15), and Pseudoemiliania lacunosa (FAD CALCAREOUS NANNOFOSSIL BIOSTRATIGRAPHY: BYBELL 55) in upper part of Zone NN 15) in these samples indi- cates that these sediments were deposited near the very top of Zone NN 15. The upper 25 m of this section, however, either was barren of calcareous nannofossils or contained insufficient assemblages for dating. Cayo Agua, South Nispero Point locality, Section 20 Three samples were examined from this 44-m-thick section: 307 (35.0 m), 305 (36.0 m), and 303 (43.5 m). All three samples were barren of calcareous nan- nofossils. Cayo Agua, Piedra Roja Point, Eastern Sequence locality, Section 18 (upper Zone NN 15) One sample was examined from the 22-m-thick sec- tion of the Cayo Agua Formation at the Piedra Roja Point locality. Sample 356, which was collected 5 m above the base of the section, could be placed in the upper part of Zone NN 15 based on the presence of Sphenolithus abies, Sphenolithus neoabies, and Pseu- doemiliania lacunosa. Planktic foraminifera (Cotton, this volume) only could restrict the age of this for- mation to between 5.3 and 3.5 Ma. The following is a list of species present in this sample. Species that are in bold are the most useful for dating this sample. Sample 356 (5 m) Helicosphaera carteri Pseudoemiliania lacunosa Reticulofenestra small species Sphenolithus abies Sphenolithus neoabies Abundance: rare Preservation: fair Age: upper Zone NN 15 Escudo de Veraguas Formation—Zones NN 15 to mid NN 18 (upper Lower to Upper Pliocene )—3.6— 2.1 Ma Escudo de Veraguas, Northern Coast locality, Section 10 (Zones NN 15-18) This formation, which probably overlies the Cayo Agua Formation, can be 60 m thick and consists dom- inantly of claystone and siltstone. Fifteen samples were examined from the 60-m-thick section exposed on the north coast of the island of Escudo de Veraguas. All fifteen samples contained rare to common calcar- eous nannofossils (Text-fig. 10) and could be placed in a specific calcareous nannofossil zone. Samples 369 (11.0 m) and 368 (16.5 m), the lowest samples studied from this section, are placed in the upper part of the Lower Pliocene Zone NN 15 by the presence of Sphenolithus abies (LAD at the top of Zone NN 15) and Pseudoemiliania lacunosa (FAD near the top of Zone NN 15). Overlying samples 367 (21.0 m), 366 (25.5 m), 365 (27.5 m), and 364 (34.5 m) are placed in Zone NN 16 because they contain neither S. abies (LAD at top of Zone NN 15) nor smaller species of the genus Gephyrocapsa (FAD oc- curs in Zone NN 17 in the study area). Sample 361 (38.5 m) is placed in Zone NN 17 because it contains Gephyrocapsa smaller species and Discoaster pentar- adiatus (LAD defines the top of Zone NN 17). Sam- ples 363 (40.0 m), 362 (41.5 m), 360 (45.5 m), and 358 (50.5 m) are placed in lower to middle Zone NN 18 based on the absence of D. pentaradiatus (LAD defines the top of Zone NN 17) and the presence in sample 358 of Discoaster brouweri (LAD at the top of Zone NN 18) and Discoaster triradiatus (LAD in middle part of Zone NN 18). Samples 174 (57.7 m), 173 (58 m), 172 (58.6 m), and 171 (59.7 m) also are placed in Zone NN 18 because they overlie samples placed in Zone NN 18 and do not contain larger spec- imens of Gephyrocapsa (FAD in Zone NN 19). Dis- coasters are absent from these last four samples, except for a few reworked specimens, and, therefore, this ge- nus was not used for age determination in these sam- ples. A minor amount of reworking is present through- out this interval. For example, there is a small amount of material from the Middle to Upper Eocene that is reworked into sample 368. Planktic foraminiferal ages (Cotton, this volume) are consistent with the calcare- ous nannofossil ages for this formation. Escudo de Veraguas, Southeastern Coast locality, Section 11 (Zones 17-18) Three samples were examined from an approxi- mately 20-m-thick section of the Escudo de Veraguas Formation on the south coast of the island of Escudo de Veraguas (Appendix B, this volume). All three sam- ples were collected approximately 11.5 m above the base of the section. This location is away from the type section, and these three samples cannot be located more precisely than being equivalent to some part of the type section. They yield frequent calcareous nan- nofossils with fair to poor preservation, and they can be placed no more accurately than in uppermost Zone NN 15, Zones NN 16, NN 17, or NN 18 due to the presence of Discoaster triradiatus (has its LAD in Zone NN 18) in sample 169 and the presence in all three samples of Pseudoemiliania lacunosa (FAD in uppermost part of Zone NN 15). There are few other diagnostic species, although the absence of larger specimens of the genus Gephyrocapsa could indicate an age no younger than Zone NN 18. All three samples have some specimens that probably are reworked from older material. If these specimens are discounted, then 56 BULLETIN 357 Escudo de Veraguas Formation Series Calcareous Nannofossil Zone (Martini, 1971) Meters Above Base of Section Braarudosphaera bigelowii Calcidiscus leptoporus Calcidiscus macintyrei Ceratolithus sp Coccolithus pelagicus Cribrocentrum reticulatum Cyclococcolithus neogammation Dictyococcites bisectus Discoaster brouweri Discoaster challengeri Discoaster deflandrei Discoaster pentaradiatus Discoaster triradiatus Discoaster spp Gephyrocapsa smaller species Helicosphaera carteri Helicosphaera sellii Pontosphaera discopora Pontosphaera multipora Pontosphaera scutellum Pontosphaera sp. aff. P wechesensis Pontosphaera spp Pseudoemiliania lacunosa Reticulofenestra flondana Reticulofenestra pseudoumbilicus Reticulofenestra spp Rhabdosphaera clavigera Rhabdosphaera spp Rhomboaster orthostylus Scyphosphaera amphora Scyphosphaera apsteinii Scyphosphaera gladstonensis Scyphosphaera pacifica Scyphosphaera spp. Sphenolithus abies Sphenolithus spp. Syracosphaera pulchra Thoracosphaera spp Abundance Preservation Text-figure 10.—Calcareous nannofossil occurrences in the Escudo de Veraguas, Northern Coast Section 10, Panama. See Text-figure 2 for location of exposure and Text-figure 4 for explanation of abundance, preservation, and other symbols. CALCAREOUS NANNOFOSSIL BIOSTRATIGRAPHY: BYBELL 57 Table 3—Summary of calcareous nannofossil ages for each formation. Formation Epoch Zone Age, Ma Limon Group Moin upper Pleistocene to upper Pliocene NN 17-21 2.4—-0.275 or upper Pliocene NN 17-19a or b 2.4-1.7 Rio Banano upper Pliocene to upper lower Pliocene NN 15-17 3.6—2.2 Uscari upper Miocene NN 11 8.2-5.6 Bocas del Toro Group Escudo de Veraguas upper lower—upper Pliocene NN 15-mid 18 3.6—2.1 Cayo Agua upper lower Pliocene upper NN 15 3.6-3.5 Shark Hole Point lower Pliocene to upper Miocene NN 12-15 5.6-3.6 Nancy Point upper Miocene NN 11 8.2-5.6 the samples can be placed in either Zone NN 17 or NN 18. Sample 168 Calcidiscus leptoporus Calcidiscus macintyrei Discoaster spp.—2 specimens (possibly reworked?) Helicosphaera carteri Pseudoemiliania lacunosa Reticulofenestra pseudoumbilicus—1 specimen (re- worked?) Rhabdosphaera clavigera Rhomboaster orthostylus—1 specimen (reworked?) Sphenolithus abies—1 specimen (reworked?) Syracosphaera pulchra Thoracosphaera spp. Abundance: frequent Preservation: fair Age: possibly Zone NN 17 or NN 18 Sample 169 Calcidiscus leptoporus Calcidiscus macintyrei Coccolithus pelagicus Hayaster perplexus Discoaster triradiatus—1 specimen (possibly re- worked?) Gephyrocapsa smaller species Helicosphaera carteri Pontosphaera sp. Pseudoemiliania lacunosa Rhabdosphaera clavigera Scyphosphaera amphora Sphenolithus abies—3 specimens (reworked?) Sphenolithus sp. aff. S. radians—1 specimen (re- worked?) Syracosphaera pulchra Thoracosphaera spp. Abundance: frequent Preservation: poor Age: possibly Zone NN 17 or NN 18 Sample 170 Calcidiscus leptoporus Calcidiscus macintyrei Coccolithus pelagicus Discoaster sp.—1 specimen (possibly reworked?) Gephyrocapsa smaller species Helicosphaera carteri Pontosphaera multipora Pseudoemiliania lacunosa Reticulofenestra pseudoumbilicus—1 specimen (re- worked?) Reticulofenestra sp. Rhabdosphaera clavigera Sphenolithus abies—3 specimens (reworked?) Syracosphaera pulchra Thoracosphaera spp. Abundance: frequent Preservation: fair Age: possibly Zone NN 17 or NN 18 CONCLUSIONS Calcareous nannofossils are present, although not abundant, in many exposures along the Caribbean coast of Panama and Costa Rica. The sediments from these outcrops were first described and named by Coates et al. (1992), and calcareous nannofossils pro- vided the primary means of determining ages for these marine sediments. Coates et al. (1992) divided these formations into two groups: the Bocas del Toro Group for sediments from the northwestern coast of Panama and the Limon Group for sediments from the northern coast of Costa Rica. Table 3 is a summary of the ages for these formations based on calcareous nannofossils. 58 BULLETIN 357 REFERENCES CITED Akers, W.H. 1972. Planktonic Foraminifera and biostratigraphy of some Neo- gene formations, northern Florida and Atlantic Coastal Plain. Tulane Studies in Geology and Paleontology, vol. 9, pp. 1-139. Aubry, M.-P. 1993a, Neogene allostratigraphy and depositional history of the De Soto Canyon area, northern Gulf of Mexico. Micro- paleontology, vol. 39, no. 4, pp. 327-366. 1993b. Calcareous nannofossil stratigraphy of the Neogene for- mation of eastern Jamaica. Geological Society of America Memoir 182, pp. 131-178. Berggren, W.A., Kent, D.V., Swisher, C.C., and Aubry, M.-P. 1995. A revised Cenozoic geochronology and chronostratigra- phy. in Geochronology, Time Scales and Global Strati- graphic Correlation. W.A. Berggren, D.V. Kent, M.-P. Au- bry, and J. Hardenbol, eds., Society of Economic Pale- ontologists and Mineralogists Special Volume no. 54, pp. 129-212. Berggren, W.A., Kent, D.V., and Van Couvering, J.A. 1985. The Neogene, pt. 2—Neogene geochronology and chro- nostratigraphy. in The Chronology of the Geological Re- cord, N.J. Snelling, ed., London, The Geological Society, Memoir no. 10: pp. 211—260. Bukry, D. 1973. Low-latitude coccolith biostratigraphic zonation. in Initial reports of the Deep Sea Drilling Project. N.T. Edgar, J.B. Saunders, et al., vol. 15, Washington, DC, U. S. Govern- ment Printing Office, pp. 685-703. 1975. Coccolith and silicoflagellate stratigraphy, northwestern Pacific Ocean, Deep Sea Drilling Project Leg 32. in Initial reports of the Deep Sea Drilling Project. R.L. Larson, R. Moberly, er al., vol. 32, Washington, DC, U. S. Govern- ment Printing Office, pp. 677—701. 1978. Biostratigraphy of Cenozoic marine sediments by calcar- eous nannofossils. Micropaleontology, vol. 24, pp. 44—60. Coates, A.G., Jackson, J.B.C., Collins, L.S., Cronin, T.M., Dow- sett, H.J., Bybell, L.M., Jung, P., and Obando, J.A. 1992. Closure of the Isthmus of Panama: The near-shore marine record of Costa Rica and western Panama. Geological So- ciety of America Builetin, vol. 104, pp. 814-828. Collins, L.S., Coates, A.G., Jackson, J.B.C., and Obando, J.A. 1995. Timing and rates of emergence of the Limon and Bocas del Toro basins: Caribbean effects of Cocos Ridge sub- duction? Geological Society of America Special Paper 295, pp. 263-289. Deflandre, G., and Fert, C. 1954. Observations sur les Coccolithophoridés actuels et fossi- les en microscopie ordinaire et électronique. Annales de Paléontologie, vol. 40, pp. 115-176. Gartner, S., Jr. 1977. Calcareous nannofossil biostratigraphy and revised zona- tion of the Pleistocene. Marine Micropaleontology, vol. 2, pp. 1-25. Gartner, S., Jr., Chen, M.P., and Stanton, R.J. 1983. Late Neogene nannofossil biostratigraphy and paleocean- ography of the northeastern Gulf of Mexico and adjacent areas. Marine Micropaleontology, vol. 8, pp. 17—S0. Gartner, S., Jr., Chow, J., and Stanton, R.J., Jr. 1987. Late Neogene paleoceanography of the eastern Caribbean, the Gulf of Mexico and the eastern equatorial Pacific. Ma- rine Micropaleontology, vol. 12, pp. 255-304. Grassé, P.P. 1952. Traité de zoologie. Paris, Masson, 1071 pp. Hay, W.W., Mohler, H.P., Roth, P.H., Schmidt, R.R., and Boud- reaux, J.E. 1967. Calcareous nannoplankton zonation of the Cenozoic of the Gulf Coast and Caribbean-Antillean area and trans- oceanic correlation. Gulf Coast Association of Geological Societies Transactions, vol. 17, p. 428-480. Lang, T.H., and Watkins, D.K. 1984. Cenozoic calcareous nannofossils from Deep Sea Drilling Project Leg 77: biostratigraphy and delineation of hiatus- es. in Initial Reports of the Deep Sea Drilling Project. R.T. Buffler, W. Schlager, et al., vol. 77, Washington, DC, U. S. Government Printing Office, pp. 629-648. Martini, EF. 1971. Standard Tertiary and Quaternary calcareous nannoplank- ton zonation. in Proceedings of the 2nd International Con- ference on Planktonic Microfossils. A. Farinacci, ed., Roma, Rome, Edizioni Tecnoscienza, vol. 2, pp. 739-785. Okada, H., and Bukry, D. 1980. Supplementary modification and introduction of code numbers to the low-latitude coccolith biostratigraphic zo- nation (Bukry, 1973; 1975). Marine Micropaleontology, vol. 5, no. 3, pp. 321-325. Perch-Nielsen, K. 1985. Cenozoic calcareous nannofossils. in Plankton stratigra- phy. H.M. Bolli, J.B. Saunders, and K. Perch-Nielsen, eds,, Cambridge, Cambridge University Press, pp. 427— 554. Rio, D., Raffi, I., and Villa, G. 1990. Pliocene-Pleistocene calcareous nannofossil distribution patterns in the western Mediterranean. in Initial Reports of the Deep Sea Drilling Project, K.A. Kastens, J. Mascle, et al., vol. 107, Washington, DC, U. S. Government Print- ing Office, pp. 513-533. Watkins, D.K., and Verbeek, J.W. 1988. Calcareous nannofossil biostratigraphy from Leg 101, northern Bahamas. in Initial reports of the Deep Sea Dril- ling Project. J.A. Austin, Jr., W. Schlager, er al., vol. 101, Washington, DC, U. S. Government Printing Office, pp. 63-85. APPENDIX 1 CALCAREOUS NANNOFOSSIL SPECIES CITED HEREIN * indicates the presence of the species in Panama or Costa Rica samples Amaurolithus amplificus (Bukry & Percival, 1971) Gartner & Bukry, 1975 Amaurolithus delicatus Gartner & Bukry, 1975 Amaurolithus primus (Bukry & Percival, 1971) Gartner & Bukry, 1975 Amaurolithus tricorniculatus (Gartner, 1967) Gartner & Bukry, 1975 *Braarudosphaera bigelowii (Gran & Braarud, 1935) Deflandre, 1947 *Calcidiscus leptoporus (Murray & Blackman, 1898) Loeblich & Tappan, 1978 *Calcidiscus macintyrei (Bukry & Bramlette, 1969) Loeblich & Tappan, 1978 Catinaster calyculus Martini & Bramlette, 1963 *Catinaster coalitus Martini & Bramlette, 1963 *Ceratolithus acutus Gartner & Bukry, 1974 CALCAREOUS NANNOFOSSIL BIOSTRATIGRAPHY: BYBELL 59 *Ceratolithus rugosus Bukry & Bramlette, 1968 *Coccolithus pelagicus (Wallich, 1877) Schiller, 1930 *Cribrocentrum reticulatum (Gartner & Smith, 1967) Perch-Nielsen, 1971 *Cyclococcolithus formosus Kamptner, 1963 *Cyclococcolithus neogammation Bramlette & Wilcoxon, 1967 *Dictyococcites bisectus (Hay, Mohler, & Wade, 1966) Bukry & Percival, 1971 *Discoaster asymmetricus Gartner, 1969 *Discoaster barbadiensis Tan Sin Hok, 1927 *Discoaster berggrenii Bukry, 1971 Discoaster bollii Martini & Bramlette, 1963 *Discoaster brouweri Tan Sin Hok, 1927 *Discoaster challengeri Bramlette & Riedel, 1954 Discoaster decorus Bukry, 1973 *Discoaster deflandrei Bramlette & Riedel, 1954 Discoaster druggii Bramlette & Wilcoxon, 1967 *Discoaster exilis Martini & Bramlette, 1963 Discoaster hamatus Martini & Bramlette, 1963 Discoaster kugleri Martini & Bramlette, 1963 Discoaster loeblichii Bukry, 1971 Discoaster neohamatus Bukry & Bramlette, 1969 Discoaster neorectus Bukry, 1971 *Discoaster pentaradiatus Tan Sin Hok, 1927 *Discoaster quinqueramus Gartner, 1969 *Discoaster surculus Martini & Bramlette, 1963 *Discoaster tamalis Kamptner, 1967 *Discoaster triradiatus Tan Sin Hok, 1927 *Discoaster variabilis Martini & Bramlette, 1963 *Discoaster woodringii Bramlette & Riedel, 1954 *Emiliania huxleyi (Lohmann, 1902) Hay & Mohler in Hay and others, 1967 Gephyrocapsa aperta Kamptner, 1963 Gephyrocapsa caribbeanica Boudreaux & Hay, 1969 Gephyrocapsa oceanica Kamptner, 1943 *Gephyrocapsa protohuxleyi McIntyre, 1970 *Hayaster perplexus (Bramlette & Riedel, 1954) Bukry, 1973 Helicosphaera ampliaperta Bramlette & Wilcoxon, 1967 *Helicosphaera bramlettei (Miiller, 1970) Jafar & Martini, 1975 *Helicosphaera carteri (Wallich, 1877) Kamptner, 1954 *Helicosphaera compacta Bramlette & Wilcoxon, 1967 *Helicosphaera euphratis Haq, 1966 *Helicosphaera lophota (Bramlette & Sullivan, 1961) Locker, 1973 Helicosphaera recta (Haq, 1966) Jafar & Martini, 1975 *Helicosphaera sellii (Bukry & Bramlette, 1969) Jafar & Martini, 1975 *Lithostromation operosum (Deflandre in Deflandre and Fert, 1954) Bybell, 1975 *Lithostromation perdurum Deflandre, 1942 *Markalius inversus Bramlette & Martini, 1964 *Pentaster lisbonensis Bybell & Gartner, 1972 *Pontosphaera discopora Schiller, 1925 *Pontosphaera japonica (Takayama, 1967) Nishida, 1971 *Pontosphaera multipora (Kamptner ex Deflandre, 1959) Roth, 1970 *Pontosphaera scutellum Kamptner, 1952 *Pontosphaera wechesensis (Bukry & Percival, 1971) Aubry, 1986 *Pseudoemiliania lacunosa (Kamptner, 1963) Gartner, 1969 *Reticulofenestra floridana (Roth & Hay in Hay et al., 1967) Theo- doridis, 1984 *Reticulofenestra pseudolockeri Jurasova, 1974 *Reticulofenestra pseudoumbilicus (Gartner, 1967) Gartner, 1969 *Rhabdosphaera clavigera Murray & Blackman, 1898 *Rhomboaster orthostylus (Shamrai, 1963) Bybell & Self-Trail, 1995 *Scyphosphaera amphora Deflandre, 1942 *Scyphosphaera apsteinii Lohmann, 1902 *Scyphosphaera gladstonensis Rade, 1975 *Scyphosphaera pacifica Rade, 1975 *Sphenolithus abies Deflandre in Deflandre and Fert, 1954 Sphenolithus belemnos Bramlette & Wilcoxon, 1967 Sphenolithus ciperoensis Bramlette & Wilcoxon, 1967 Sphenolithus heteromorphus Deflandre, 1953 *Sphenolithus neoabies Bukry & Bramlette, 1969 *Sphenolithus predistentus Bramlette & Wilcoxon, 1967 *Sphenolithus radians Deflandre in Grassé, 1952 *Sphenolithus tribulosus Roth, 1970 *Syracosphaera pulchra Lohmann, 1902 Triquetrorhabdulus carinatus Martini, 1965 Triquetrorhabdulus rugosus Bramlette & Wilcoxon, 1967 j ke 7 a\y | Mi i j Wy pe : ‘ sob i a me tO SB wmva — 1 hey fll ey ' rh, = ‘ i _ iJ <_ | i ie ; a ij i g t a ' i j poet , j > 2| mu" i lay? : weer & teal aay ; : m at pommel oe ¢ . iwi ' . sv'sQ “' an po tay Racdiyaer ai, it ee er - ‘ ‘ wi es Titi of _ ay Lo? Cael = , reer «| coil Whi WA ie au wild: wi bart (Va) tigen)? 'r _ h 7 (Pmtiyal! “ae > ~ me ‘ : Y we ow ie : =! eS 4 ae et saab =" ahi ee cade hpi iiqrave” SR TE caren Py os > eee ery ey oat” fae. Dent oom Perv age a rp ait? Mii: acre odes ee > Gia -—- Poe yey an oo) ee Te eset colainshemrestannielel* année? oh) ow at ad yam eg ao » hy PS ie —r S “% CHAPTER 3 NEOGENE PLANKTIC FORAMINIFERAL BIOCHRONOLOGY OF THE SOUTHERN CENTRAL AMERICAN ISTHMUS MATHEW A. COTTON Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605, U.S.A. INTRODUCTION It has been sixty years since Thalman (1934) pio- neered the use of planktic foraminifera for interregion- al correlation, illustrating their biostratigraphic utility. Early applications of planktic foraminifera in biostra- tigraphy occurred in three primary regions: the Alpine- Mediterranean region, Russia, and Trinidad, in the Ca- ribbean Basin (Bolli, 1974). Since then, planktic fo- raminifera have proven valuable in providing age con- straints on marine deposits around the world, both in land-based sections and in deep-sea cores. This study is part of the Panama Paleontology Proj- ect and the first major study of the planktic foramini- fera on the southern Central American isthmus (Cot- ton, 1990, 1991). The study area includes the Carib- bean coast of Costa Rica and Panama, specifically the Limon and Bocas del Toro basins (Appendix A, Maps 4—11). The purpose of the study is to provide a precise biochronologic framework for the stratigraphic units which encompass the Pliocene emergence of the isth- mus. Detailed biostratigraphic data offer age con- straints needed to resolve the tectonic history of the isthmus, in particular, the timing of emergence. Another product of this study is a Neogene strati- graphic correlation scheme for the Caribbean Basin based on data from planktic foraminiferal studies in Mexico (both Pacific and Caribbean coasts), Venezue- la, Colombia, Equador, St. Croix, Dominican Repub- lic, Jamaica, Puerto Rico, Florida, Virginia, North and South Carolina, and the Pacific coasts of Panama and Costa Rica. In some instances re-evaluation of earlier studies was necessary prior to correlation. ACKNOWLEDGMENTS Thanks are due to Laurel Collins and William Berg- gren for their constructive reviews of this manuscript. Special thanks go to Harry Dowsett (U. S. Geological Survey, Reston) who unselfishly contributed his advice and expertise as well as a thoughtful and prompt re- view of this manuscript. METHODS Seventy-five samples collected from ten locales were utilized in this study. Approximately fifty grams from each sample were disaggregated by soaking in water, and heated on a hot plate for several hours. If the sample proved difficult to dissaggregate, one of two methods was employed: (1) soaking the sample overnight in a solvent (such as paint thinner, kerosene or turpentine), followed by soaking it in water and heating; (2) adding Quaternary-O (a petroleum by- product) and heating in water (Zangula, 1968). Fol- lowing disaggregation, the samples were washed through <850 wm and >63 pm nested sieves to con- centrate the fraction larger than silt. The > 850 ym and < 63 wm portions were discarded. The remaining fraction was transferred to filter paper and oven dried at ~50°C. From a representative split of each sample, all planktic foraminifera (usually >200 specimens) were picked and identified to the species level follow- ing the taxonomy of Kennett and Srinivasan (1983) and Bolli and Saunders (1985). Additional compari- sons were made with specimens housed in the Cush- man Collection, which resides in the U. S. National Museum of Natural History, Smithsonian Institution, Washington, D.C. In samples with abundant speci- mens, planktic foraminifera were picked from random squares on the picking tray until 300 specimens were obtained. All identifications were made by the author. BIOSTRATIGRAPHY Thirty-nine species of planktic foraminifera were identified in the Neogene deposits along the Caribbean coast of western Panama and eastern Costa Rica (Ta- bles 1, 2). In general, the planktic foraminifera were both abundant and well preserved with the notable ex- ception of the Rio Banano Formation, in which they were sparse and poorly preserved. Total faunal diver- sity (number of planktic foraminifer species per sam- ple) varied from 2 to 20 and mean species diversity varied from 7 to 16 in the formations studied. Strati- graphic ranges of last appearance datums (““‘LADs’’) 62 BULLETIN 357 Table 1.—Faunal list. Candeina nitida d’ Orbigny, 1939 Dentogloboquadrina altispira (Cushman and Jarvis), 1936 Globigerina apertura Cushman, 1918 Globigerina bulloides (d’Orbigny), 1826 Globigerina decoraperta Takayanagi and Saito, 1962 Globigerina falconensis Blow, 1959 Globigerina nepenthes Todd, 1957 Globigerina woodi Jenkins, 1960 Globigerinella aequilateralis (Brady), 1879 Globigerinella calida (Parker), 1962 Globigerinita glutinata (Egger), 1893 Globigerinoides conglobatus (Brady), 1879 Globigerinoides obliquus Bolli, 1957 Globigerinoides obliquus Bolli, var. extremus Bolli and Bermudez, 1965 Globigerinoides ruber (d’Orbigny), 1839 Globigerinoides sacculifer Brady, 1877 Globigerinoides seigliei Bermudez and Bolli, 1969 Globorotalia crassaformis (Galloway and Wissler), 1927 Globorotalia exilis Blow, 1969 Globorotalia juanai Bermudez and Bolli, 1969 Globorotalia margaritae Bolli and Bermudez, 1965 Globorotalia menardii (Parker, Jones and Brady), 1865 Globorotalia miocenica Palmer, 1945 Globorotalia plesiotumida Blow and Banner, 1965 Globorotalia pseudomiocenica Bolli and Bermudez, 1965 Globorotalia puncticulata (Deshayes), 1832 Globorotalia scitula (Brady), 1882 Globorotalia tosaensis Takayanagi and Saito, 1962 Globorotalia truncatulinoides (d’ Orbigny), 1839 Globorotalia tumida (Brady), 1877 Orbulina universa d’Orbigny, 1839 Neogloboquadrina acostaensis (Blow), 1959 Neogloboquadrina dutertrei (d’ Orbigny), 1839 Neogloboquadrina humerosa (Takayanagi and Saito), 1962 Neogloboquadrina pachyderma (Ehrenberg), 1861 Pulleniatina obliquiloculata (Parker and Jones), 1865 Pulleniatina primalis Banner and Blow, 1967 Sphaeroidinella dehiscens (Parker and Jones), 1865 Sphaeroidinellopsis seminulina (Schwager), 1866 Turborotalita quinqueloba Natland, 1938 and first appearance datums (“‘FADs’’) of planktic for- aminifer species identified in this study (Table 3) are taken from Bolli and Saunders (1985) and Dowsett (1989). Bolli and Saunders (1985) document Globigerinoi- des ruber originating in the Early Miocene and dis- appearing from the fossil record between the late Mid- dle Miocene to within the Early Pliocene (approxi- mately 11.3—5.1 Ma), the so-called ‘“ruber-gap’’. However, based on data from Keller et al. (1989) on DSDP site 503A (Pacific side of the Central American isthmus, Text-fig. 1), and DSDP site 502A (Caribbean side of the Central American isthmus), G. ruber is missing from the fossil record from the base of both cores (Late Miocene) to approximately 3.5 Ma (Text- fig. 2). Globigerinoides ruber increases in number and NORTH oo AMERICA sf orktown Fm. Waccamaw Fm. s Cl TirabuzonFm. = / te: Si i. Bluff Fm. i Azua asin Cibao deposits~~ \ e i) / Jamaica \ Hoon @. ehuantepec ~—>— Maria Madre ora deposits — = | deposits® nF e SG ~~ DSDP 502. 0 nwe __f Pozon Fm. ‘@Munguido Fm. . Camuy Fm. Pal Bocas del Toro Gp Limon Gp.= Charco Azul Gp: vy @Esmeralda Fm. Lb ( SOUTH \AMERICA Text-figure 1.—Map of the Americas showing locations of land- based Neogene sediments and Deep Sea Drilling Project (DSDP) sites containing planktic foraminifera and discussed in text. percentage relative to G. obliquus to approximately 2.0 Ma, when G. ruber outnumbers G. obliquus about 9 to 1 (G. ruber ratio = 0.9) . The relatively rapid turn- over from a G. obliquus-dominated to a G. ruber-dom- inated fauna simultaneously in both cores is remark- able and led to the use of a value called “‘percent ruber” ( = G. ruber/G. ruber + G. obliquus X 100). It also suggests an oceanic connection between the Caribbean and the Pacific at that time (3.5 Ma). Globigerinoides obliquus in this study includes G. extremus, which is considered a morphologic variant of G. obliquus, al- though many planktic foraminiferal workers recognize a unique FAD for G. extremus in the Late Miocene. Previous studies of calcareous microfossils from the southern Central American isthmus include Cushman, 1918; Coryell and Mossman, 1942; Jenkins, 1964; Bold, 1967a, 1967b, 1972; Blacut and Kleinpell, 1969; Bandy, 1970; Akers, 1972; Taylor, 1975; Cassell, 1986; Pizarro, 1987; Berrangé et al., 1989; Cassell and Sen Gupta, 1989a, 1989b; Corrigan ef al., 1990; and Duque-Caro, 1990. Generally, these studies were lim- ited to a single formation and, for the most part, in- volved age determinations based on correlation to con- ventional calcareous microfossil zonations. There has been no comprehensive biostratigraphic work done for the Neogene deposits from the southern Central Amer- ican isthmus until this study (see Cotton, 1991, for original biostratigraphic data). All calcareous nannofossil data used in this chapter PLANKTIC FORAMINIFERAL BIOSTRATIGRAPHY: COTTON 63 DSDP site 503A 0 10 20 30 ao 4 50 oo 4 70 Aso 90 100 110 120 130 140 150 160 170 180 190 200 210 220 230 240 2.35 Ma 3.0Ma 3.5Ma CORE DEPTH in meters DSDP site 502A % ruber Pulleniatina coiling change =3.7Ma 40 A, 1.9 Ma 3.0Ma 3.4Ma CORE DEPTH in meters Text-figure 2.—Plot of percent Globigerinoides ruber relative to G.obliquus for DSDP sites 502A (Caribbean) and 503A (Pacific) versus core depth (m), based on data from Keller et al. (1989). 64 BULLETIN 357 Table 2.—Occurrences of planktic foraminifera at PPP sites (numbered across top). Numbers are in stratigraphic order from top = left, to bottom = right. Samples from the same section (App. B) are joined with a Uscari Fm. St. Rita “-” Data are available at internet site http://www. fiu.edu/ collinsl/. Rio Banano Fm. 726 737-735-736 720-721 700 678 -668 -670 -672 -690 -689 - 688 - 687 - 686 -685 - 684 - 683 - 682 -679 Candeina nitida Dentogloboquadrina altispira Globigerina apertura Globigerina bulloides Globigerina decoraperta Globigerina falconensis xX Globigerina nepenthes Xx Globigerina woodi x Globigerinella aequilateralis x x x x 8 Globigerinella calida Globigerinita glutinata x Globigerinoides conglobatus x Globigerinoides obliquus x Xx Xx Xx Xx Globigerinoides ruber X Globigerinoides sacculifer x x x x x ~ 6 KK Globigerinoides seigliei Globorotalia crassaformis x Globorotalia exilis Globorotalia juanai x Globorotalia margaritae Globorotalia menardti x x x x Globorotalia miocenica Globorotalia plesiotumida Xx Xx Xx x x Globorotalia pseudomiocenica Globorotalia puncticulata Globorotalia scitula x Globorotalia tosaensis Globorotalia truncatulinoides Globorotalia tumida Neogloboquadrina acostaensis X: % ire Neogloboquadrina dutertrei Neogloboquadrina humerosa Xx x Neogloboquadrina pachyderma Orbulina universa x x x X Pulleniatina obliquiloculata Pulleniatina primalis Sphaeroidinella dehiscens Sphaeroidinellopsis sp. x x Turborotalia quinqueloba are by the courtesy of L. Bybell, U. S. Geological Survey, who provided the identifications of the nan- nofossils for the initial part of the Panama Paleontol- ogy Project (Chapter 2, this volume). Nannofossil data from the isthmus are used to support, or show conflict with, the planktic foraminifera-based age constraints. LIMON GrRouP The Limon Group deposits occur in the Limon Ba- sin along the southeastern Caribbean coast of Costa Rica (Appendix A, Maps 10—11) with exposure south and west of the town of Limon. The Limon Basin con- tains approximately 10,000 meters of marine sedi- ments ranging from Paleocene to Pleistocene in age (Weyl, 1980). The Limon Group represents the Neo- x x x x xX x xX xX xX xX xX x xX x X x x xX xX ~* ” ~ x xX x x x x xX x xX xX x x x x xX Xx x xX x x x x xX x xX x x >< xX xX x x x x x X x xX x Xx x x x xX xX x gene portion of these deposits and is comprised of the Uscari, the Rio Banano and the Moin formations, to- taling over 1500 m of sediment. These Neogene sed- iments were deposited in a back-arc setting behind the uplifted Cordillera de Talamanca (Galli-Olivier, 1979; Escalante, 1990). The earliest published reference to these deposits was by Gabb (1895), who coined the term “‘Moin Clay Member” and described the molluscan fauna in the Limon region. Hill (1898) also made reference to fos- sils collected in the Limon area during his reconnais- sance of the isthmus. Nearly a quarter of a century later, Olsson (1922) collected and described mollusks from the Limon area and named the Uscari Formation after the Uscari creek about 50 km south of Limon. PLANKTIC FORAMINIFERAL BIOSTRATIGRAPHY: COTTON 65 Table 2.—Extended. Moin Fm. 631 638 - 643 - 645 - 637 - 636-635-710 658 - 657 - 654 - 656 - 653 - 655 - 652 - 650-648-647 705 712 Candeina nitida Dentogloboquadrina altispira Globigerina apertura x Globigerina bulloides Globigerina decoraperta X Globigerina falconensis Globigerina nepenthes Globigerina woodi Globigerinella aequilateralis Xx x x X Globigerinella calida Globigerinita glutinata x x x Globigerinoides conglobatus x x Globigerinoides obliquus Globigerinoides ruber x x Xs x x X Globigerinoides sacculifer x X x x x x Globigerinoides seigliei Globorotalia crassaformis Globorotalia exilis Globorotalia juanai Globorotalia margaritae Globorotalia menardti Globorotalia miocenica Globorotalia plesiotumida Globorotalia pseudomiocenica Globorotalia puncticulata Globorotalia scitula Globorotalia tosaensis Globorotalia truncatulinoides x x x Globorotalia tumida x Neogloboquadrina acostaensis x x Xx Neogloboquadrina dutertrei xi x x Neogloboquadrina humerosa Xx x x Xx Neogloboquadrina pachyderma Orbulina universa Xx x x x X Xx Pulleniatina obliquiloculata Pulleniatina primalis Sphaeroidinella dehiscens Xi x Sphaeroidinellopsis sp. Turborotalia quinqueloba He assigned the overlying post-Uscari fossiliferous beds to the Gatun Formation. For a detailed lithostra- tigraphy of the Limon Group see Coates et al. (1992) and Coates (Chapter 1, this volume). Uscari Formation Previous foraminiferal studies of the Uscari For- mation include Goudkoff and Porter (1942), Taylor (1975), Cassell (1986), Pizarro (1987), and Cassell and Sen Gupta (1989a). The following species of planktic foraminifera were identified in this formation in a study by Pizarro (1987): Globorotalia siakensis ( = Globorotalia mayeri in Bolli and Saunders, 1982; LAD = 10.2 Ma), Globigerina nepenthes (LAD = 4.0 Ma), Sphaeroidinellopsis seminulina and S. paenede- xX xX xX xX x x x Xx x x x x xX xX x x xX xX xX xX x xX xX xX x x x xX x xX x x xX xX xX xX x x x x xX xX x xX x x x x x X x x x xX xX x x x xX x xX xX xX xX x Xx x x x 2 ) hiscens (LAD = 3.0 Ma), Dentogloboquadrina alti- spira (LAD = 2.9 Ma), Globigerina venezuelana (LAD = 3.4 Ma) and Globigerinoides extremus (FAD = 7.2 Ma). Bolli and Saunders (1985) considered S. paenedehiscens to be a synonym of S. seminulina, which they gave a FAD of 7.2 Ma. Therefore, Pizarro’s extension of the Uscari Formation into the Late Mio- cene is confirmed by the presence of S. seminulina and G. extremus. Pizarro (1987) also identified Catapsydrax dissimilis and C. cf. C. unicavus, which he believes were re- worked and therefore not representative of the sedi- ments in which they were found. Upon viewing the scanning electron micrographs of these specimens, they appear to be Dentogloboquadrina venezuelana 66 Table 2.—Continued. BULLETIN 357 Shark Hole Point Fm. Nancy Point Fm. Cayo Agua Fm. 379 -376 -384 - 389 -388 -390- 393 - 396-397-401 -407-410- 411 Candeina nitida Dentogloboquadrina altispira Xx Globigerina apertura Globigerina bulloides x Globigerina decoraperta Pek eet ES ~ KK Globigerina falconensis Globigerina nepenche* Xx mx XK KKK ~*~ ~ Globigerina woodi at ins ioeetat otal tal ~~ Globigerinella aequilateralis Xx Globigerinella calida Globigerinita glutinata x x Globigerinoides conglobatus Globigerinoides obliquus X x Globigerinoides ruber ~ KKK MH Pad Pad ~*~ Globigerinoides sacculifer x x Globigerinoides seigliei Globorotalia crassaformis ~ Ke KK MK Globorotalia exilis Globorotalia juanai Globorotalia margaritae x x Globorotalia menardti x X X x Globorotalia miocenica Globorotalia plesiotumida x Xx Xx x Xx x Globorotalia pseudomiocenica x Globorotalia puncticulata Globorotalia scitula Globorotalia tosaensis Globorotalia truncatulinoides Globorotalia tumida Neogloboquadrina acostaensis x x x x x x x Neogloboquadrina dutertrei Neogloboquadrina humerosa x x x Neogloboquadrina pachyderma x Orbulina universa x Xx x Pulleniatina obliquiloculata Pulleniatina primalis x x Sphaeroidinella dehiscens Sphaeroidinellopsis sp. x x x x x Turborotalia quinqueloba mK KK 338 - 337 -336 -335 -334 - 300 - 298 - 293- x x xX xX x x x xX xX x x xX x xX xX x x xX x xX x xX x xX xX x x xX xX x xX xX xX x x xX xX x xX xX x Xx xX x x x x x x xX x x x x x x xX x x x x xX xX x xX xX xX x x x xX xX xX x x Xx x x x x x xX x x x xX xX x x xX x xX Xx x x xX xX x x x x xX xX x xX xX Xx x xX x x x Xx xX x x x x x xX x x xX xX x x x x x xX x x with an aberrant last chamber which sometimes forms in this species and leads to confusion in identification (Bolli and Saunders, 1985; p. 186). The specimens which Pizarro labelled as Catapsydrax dissimilis and C. cf. C. unicavus appear to have an aberrant last chamber with similar wall texture to the chambers of the test. True bullae usually exhibit a wall texture which differs from that of the chambers. Cassell and Sen Gupta (1989a) sampled three sep- arate sections from three different river valleys and analyzed the foraminifera. Based on their planktic fo- raminiferal identifications, they concluded that the Us- cari Formation spans Zones N3 to N10 (or Upper Oli- gocene to Middle Miocene). The Uscari Formation may possibly include sediments younger than Zone N10, but they did not have access to the upper portion of the type section along Quebrada Uscari. Globoro- talia kugleri was found at the base of the formation and is diagnostic of the earliest Miocene (approxi- mately 23.7—21.8 Ma; Zhang et al., 1993). The FAD of this species marks the Oligocene—Miocene bound- ary in the newly proposed Oligocene—Miocene bound- ary stratotype (Steinenger ef al., 1996). Less than 10 m above the base occur Catapsydrax stainforthi, C. dissimilis and Globorotalia fohsi peripheroronda, all indicative of the Early Miocene. In this study only the uppermost Uscari Formation was sampled. The four samples analyzed (PPP 726, 735-737; Table 2) appear to be approximately coeval with the lower portion of the Nancy Point Formation PLANKTIC FORAMINIFERAL BIOSTRATIGRAPHY: COTTON 67 Table 2.—Extended. Cayo Agua Fm. -61 - 374- 373-372-371 307-306 Candeina nitida Dentogloboquadrina altispira Xx Globigerina apertura x Globigerina bulloides Xx Globigerina decoraperta Globigerina falconensis x par palate ua bs ome Dae od Sasa ibd isd * Sap tint Den oe md Globigerina nepenthes Globigerina woodi x Globigerinella aequilateralis x Globigerinella calida Globigerinita glutinata Xx x x ~ ~ KKK KK MK Globigerinoides conglobatus Globigerinoides obliquus x x Xx x x Globigerinoides ruber Xx x x x Globigerinoides sacculifer x x x x Xx rat faltat sta! Globigerinoides seigliei Globorotalia crassaformis Globorotalia exilis x Globorotalia juanai Globorotalia margaritae x Xx x Globorotalia menarditi Xx x X Xx Globorotalia miocenica Globorotalia plesiotumida Xx x Xx X Globorotalia pseudomiocenica Globorotalia puncticulata x Globorotalia scitula Globorotalia tosaensis Globorotalia truncatulinoides Globorotalia tumida Neogloboquadrina acostaensis x x x x Neogloboquadrina dutertrei Neogloboquadrina humerosa x x Xx x Xx Neogloboquadrina pachyderma x Orbulina universa x x x x X Pulleniatina obliquiloculata Pulleniatina primalis x Sphaeroidinella dehiscens Sphaeroidinellopsis sp. x x X Turborotalia quinqueloba Escudo de Veraguas Fm. 168-169-170 358 - 360 - 361 - 362 - 364 - 365 - 366 - 367 - 368 -369 xX xX Xx xX xX x x x xX x xX x xX xX x xX x xX x xX xX x x xX xX x x x xX xX x xX xX Xx xX xX 4 xX xX xX x xX xX xX x xX xX xX x x xX x x x x xX x x x xX x x xX xX xX x xX xX x xX x x xX xX xX x x xX xX x xX xX xX x x xX Xx xX xX xX xX xX x x x x xX x xX x xX x x x x x xX x x x x xX xX xX x xX x x xX X x x x x x x x xX x xX Xx x x x x x x x (Late Miocene) of Panama and therefore are the youn- gest dated samples collected from the Uscari Forma- tion. These samples were collected from two locales (Appendix A, Map 10): (1) Carbon Dos Road, south of Limon and west of Punta Cahuita, and (2) Rio Sandbox, south of Limon and southwest of Punta Ca- huita. Planktic foraminifera identified from one sam- ple, PPP 726, of the Carbon Dos section (Section 28, Appendix B) include Globorotalia juanai (FAD = 10.4, LAD = 7.2 Ma) and left-coiled G. menardii and Globorotalia plesiotumida (FAD = 7.7; LAD = 4.0 Ma). Planktic foraminifera identified from the Rio Sandbox section (PPP 735-737, Section 27, Appendix B) include Globigerina nepenthes (LAD = 4.0 Ma) and Globorotalia plesiotumida, which undergoes an abrupt coiling change from sinistral to dextral between the middle and uppermost samples (PPP 735, 737) in the section. The total species diversity ranged from 6 to 13 per sample with a mean of 10 for the Uscari Formation, which is the highest for the Limon Group. The nannofossil results place 2 of 3 samples from Car- bon Dos and both Rio Sandbox samples in Zone NN 11 (approximate age = 8.2—5.6 Ma) based on the oc- currences of Discoaster berggrenii and/or D. quin- queramus. In summary, Pizarro (1987) apparently had samples of the Uscari Formation ranging in age from Middle to Late Miocene. Cassell and Sen Gupta (1989a) sam- pled the base of the formation, which has a maximum age of Early Miocene based on the occurrence of Glo- 68 BULLETIN 357 Table 3.—Neogene events of planktic foraminifera (F) and calcareous nannoplankton (N) used in this study (modified from Coates er al., 1992). Datums with an asterisk are held with lower confidence. Reapp. datum, LAD = last appearance datum. = reappearance, disapp. = disappearance, FAD = first appearance Type Taxon Pseudoemiliania lacunosa Calcidiscus macintyret Gephyrocapsa spp. (large) Globigerinoides obliquus/extremus Globorotalia truncatulinoides Discoaster brouweri Globorotalia miocenica Globorotalia menardii, lett-coiled Globorotalia menardii, right-coiled Pulleniatina spp. Discoaster pentaradiatus Gephyrocapsa spp. (small) Dentogloboquadrina altispira Sphaeroidinella dehiscens Sphaeroidinellopsis spp. Globorotalia tosaensis Pulleniatina spp. Globorotalia miocenica Globorotalia margaritae Globigerinoides ruber Sphenolithus abies Reticulofenestra pseudoumbilicus Pseudoemiliania lacunosa Pulleniatina, right-coiled Pulleniatina, left-coiled Globorotalia puncticulata Globorotalia plesiotumida Globigerina nepenthes Globorotalia crassaformis Globigerinoides seigliei Ceratolithus acutus Ceratolithus acutus Globorotalia tumida Globigerinoides conglobatus Globorotalia margaritae Discoaster quinqueramus Discoaster berggrenii Globorotalia plesiotumida Globorotalia juanai Neogloboquadrina humerosa Discoaster quinqueramus Globorotalia juanai Globigerina nepenthes lay! aa] YA Sag] ‘gl ‘asl Fa 2) lal Cool lol 4 A, teal Lag] lash Leal lea| Joshites| 74 74, 7409) lo Meal onl lool dasheoalilzs|iraur4 ps |elna| hes |alo ola Ae sl uln 9) Apa aA Event Age (Ma) Notes LAD 0.5 LAD 1.5 FAD 1.7 LAD 1.8 FAD 1.9 LAD 1.9 LAD Ded. Atlantic only FAD DD coiling event LAD Pip coiling event reapp. 22. Atlantic only LAD 2.4 FAD DSi LAD 2.9 FAD 3.0 LAD 3.0 FAD 3.1 disapp. 33 Atlantic only FAD 3.4 Atlantic only LAD 3.4 reapp. Shs LAD 3.5 LAD 35 FAD 3.6 FAD Sey coiling event LAD S60 coiling event FAD 4.0 LAD 4.0* LAD 4.0 FAD 4.3 LAD 4.3 LAD 4.6 FAD 5.0 FAD ae FAD Shs) FAD 5.6 LAD 5.6 LAD 5.6 FAD (3) Oe Wari LAD 7.2 FAD Tes FAD 8.2 FAD 10.4 FAD eS, borotalia kugleri. The stratigraphically highest sam- ples they dated belong to Zone N10 (estimated age = 14.8 to 13.9 Ma) or Middle Miocene based on the co- occurrence of Globorotalia fohsi peripheroronda and Orbulina universa. The limited samples from the up- permost Uscari Formation used in this study indicate a Late Miocene age with the combined planktic fora- miniferal-nannofossil estimate ranging from 7.7 to 5.6 Ma for the Rio Sandbox section, and 7.7 to 7.2 Ma for the Carbon Dos section. Rio Banano Formation The Rio Banano Formation was named by Taylor (1975), who assigned the strata to Zones N17—N21 (Upper Miocene to Upper Pliocene). Cassell (1986) and Cassell and Sen Gupta (1989b) restricted the mid- dle portion of the Rio Banano to Lower Pliocene (low- er Zone N18, ~5.3 Ma) using the stratigraphic overlap of planktic foraminifera, particularly G. extremus and G. ruber, and an ostracode species, Radimella ovata. Since both the upper and lower portions of the Rio PLANKTIC FORAMINIFERAL BIOSTRATIGRAPHY: COTTON 69 Banano Formation produced no age-diagnostic fora- miniferal species, they extrapolated the age of the Rio Banano Formation from the Late Miocene to the Late Pliocene. Sixteen samples of Rio Banano deposits used in this study were collected from two localities in southeast- ern Costa Rica. Map 11 and Inset C of Map 11, Ap- pendix A, show the localities, and Sections 29 and 32, Appendix B, show the stratigraphic order of samples in each of the sections. Fourteen samples (PPP 668, 670, 672, 678, 679, 682-690; Table 2) were collected from the type section (Section 29, Appendix B) along the Banano River, west of Bomba. Two samples (PPP 720 and 721) were collected from the Santa Rita lo- cality (Map 11, Appendix A; Section 32, Appendix B), south of Moin. In general, the planktic foraminifera were sparse, poorly preserved, and of low diversity (total species diversity ranged from 2 to 12 per sample with a mean of 7) in this formation. Species found in the Bomba section (Inset C of Map 11, Appendix A; Section 29, Appendix B) include Dentoglobigerina altispira (LAD = 2.9 Ma), Globi- gerinoides ruber, G. sacculifer, G. obliquus, and right- coiled Globorotalia plesiotumida (FAD = 7.7 Ma; LAD = 4.0 Ma). Dentoglobigerina altispira has a last occurrence in the upper fifth of the section. Globiger- inoides ruber and G. obliquus occur in equal numbers (% ruber ~ 0.5). The top of the section lacks D. al- tispira, but contains G. conglobatus (FAD = 5.3 Ma) and Globorotalia plesiotumida, which has a LAD in the Early Pliocene in mid-Zone N19 (or approximately 4.0 Ma based on Kennett and Srinivasan, 1983, and Bolli and Saunders, 1985. Neogloboquadrinids occur throughout the section with Neogloboquadrina duter- trei occurring in the top of the section and Neoglo- boquadrina acostaensis scattered throughout the sec- tion. Nannofossils were also rare or lacking in many of the samples processed from this formation. However, key taxa that were identified include Sphenolithus abi- es (LAD = 3.5 Ma) and Pseudoemiliania lacunosa (3.6—0.5 Ma) occurring together from the base through nine tenths of the section, where S. abies last occurs (at PPP 676). Uppermost samples contain small Ge- phyrocapsa sp. (FAD = 2.5) in association with Dis- coaster pentaradiatus (LAD = 2.4 Ma). The Santa Rita section (at PPP 720 and 721; Section 32, Appendix B) contains an equivalent planktic fauna to the Bomba section. Planktic foraminifera include Dentoglobigerina altispira, Globigerinoides conglo- batus, Globorotalia plesiotumida and right-coiled G. menardii. Calcareous nannofossils include P. lacuno- sa, D. pentaradiatus and a few isolated occurrences of small Gephyrocapsa spp. In summary, planktic foraminifera of the Rio Ban- ano Formation offer weak age information primarily relying on the presence of G. plesiotumida. This spe- cies puts the base of the formation (at PPP 679) be- tween 7.7 and 4.0 Ma. The top (at PPP 668) has a maximum age of 5.3 Ma based on the presence of Globigerinoides conglobatus. However, negative evi- dence suggests the top to be = 2.9 Ma based on the absence of Dentoglobigerina altispira, which last oc- curs near the top of the formation (at PPP 690). The presence of G. ruber suggests an age of = 3.5 Ma (Table 2). The presence of Globorotalia plesiotumida at the top of the formation offers conflicting evidence as it has a LAD of 4.0 Ma. However, in ranking the confidence of identifications, G. plesiotumida would rank low because of its similarity to other menardii- form globorotaliids. On the other hand, nannofossil data suggest a younger base at 3.6 to 3.5 Ma, while they suggest a more refined younger age for the top at 2.5 to 2.4 Ma. The conflict between the nannofossil- and the foraminifera-based age estimates lies in the identification and geologic range of G. plesiotumida. Since this author has less confidence in the identifi- cation and the geologic range for this species, the nan- nofossil-based age estimates are considered more re- liable for this formation. However, it is interesting to note that Cassell and Sen Gupta (1989b) showed the Rio Banano extending to the base of the Pliocene, which supports the older foraminiferal-based date from this study. Moin Formation Akers (1972) provided the first age determination for the Moin Formation (Early Pleistocene) using planktic foraminifera based on one locality—TU 954 of Vokes, a hill cut behind the Standard Fruit Com- pany, west of the Pueblo Nuevo Cemetery, approxi- mately 2 km west of Limon. His key taxa were Neo- globoquadrina dutertrei, Sphaeroidinella dehiscens, Globigerinoides obliquus, Globigerinoides congloba- tus and Pulleniatina obliquiloculata. Using the LADs and FADs of this association gives an age span of 3.0— 1.8 Ma or Late Pliocene. Globigerinoides obliquus be- came extinct 1.8 Ma and S. dehiscens evolved 3.0 Ma. Neogloboquadrina dutertrei evolved in the Pliocene and does not necessarily indicate a Pleistocene age (Bolli and Saunders, 1985, p. 211). If one uses the Atlantic reoccurrence date of Pulleniatina at 2.2 Ma (Saito, 1976; Keigwin, 1982) it would narrow the age restriction to 2.2-1.8 Ma. The Atlantic reoccurrence date of Pulleniatina refers to its disappearance from the Atlantic depositional record at 3.3 Ma and reap- pearance at 2.2 Ma, while its Pacific fossil record is continuous through this period. 70 BULLETIN 357 Cassell (1986) dated the Moin Formation as Pleis- tocene based on the occurrence of Globorotalia trun- catulinoides excelsa (FAD = 1.9 Ma) and Sphaeroi- dinella dehiscens-excavata, which Cassell considered to be restricted to the Pleistocene. However, Bolli and Saunders (1985) and Dowsett (oral commun., 1991) stated that this form (Sphaeroidinella dehiscens-exca- vata) originated in the Pliocene. Planktic foraminifera identified in this study came from twenty samples (Table 2) taken from six Costa Rica locales: 1) near Pueblo Nuevo Cemetery, | km west of Limon (PPP 631, Section 35, Appendix B; 635-637, Section 36, Appendix B); 2) Lomas del Mar, west of Limon (PPP 638, 643, 645; Section 36, Ap- pendix B); 3) an unnamed creek near Cangrejos (to be called ‘‘Cangrejos Creek” herein), west of Limon (PPP 647, 648, 650, 652-658, Section 37, Appendix B); 4) west of Rio Blanco, south of Limon (PPP 705, no drawn section); 5) southwest of Liverpool (PPP 710, in Section 36, Appendix B); 6) Route 32, west of Pueblo Nuevo Cemetery (PPP 712, in Section 34, Appendix B). All PPP numbers are shown in Appen- dix A on Inset B of Map 11! except PPP 705, which is shown on Map 11. Planktic foraminifera identified from near Pueblo Nuevo Cemetery (Section 36, Appendix B) include left-coiled Globorotalia truncatulinoides (FAD = 1.9 Ma), Sphaeroidinella dehiscens (FAD = 3.0 Ma), Glo- bigerinoides conglobatus (FAD = 5.3 Ma), and right- coiled Pulleniatina obliquiloculata, which reappeared in the Caribbean at 2.2 Ma. Since Globorotalia trun- catulinoides occurs near the base, the entire section is probably <1.9 Ma. The total species diversity of planktic foraminifera for the Moin Formation ranges from 2 to 15 per sample with a mean of 7, equal to the underlying Rio Banano despite the greater abun- dance and better preservation. At Lomas del Mar, G. truncatulinoides (FAD = 1.9 Ma) was found throughout this part of the section (middle of Section 36, Appendix B), confirming the age to be < 1.9 Ma. Also noted was a coiling change from right to left between samples PPP 643 and 638. However, due to the complex coiling history of G. truncatulinoides in the Caribbean, correlating such an event does not carry much confidence. The Cangrejos Creek locale (Section 37, Appendix B) produced abundant and diverse, well-preserved planktic foraminifera. Key taxa found consistently throughout the section are right-coiled Pulleniatina (P. obliquiloculata and P. primalis) and Sphaeroidinella dehiscens (FAD = 3.0 Ma). Globorotalia truncatulinoides (FAD = 1.9 Ma) was also identified throughout this section. Globigerinoides obliquus (LAD = 1.8 Ma) was found in one sample (PPP 658) with G. truncatulinoides, restricting the age of that sample to 1.9—1.8 Ma. Two specimens of sphaeroidi- nellids lacking secondary apertures as in Sphaeroidi- nellopsis sp. (LAD = 3.0 Ma) were identified in two samples (PPP 655, 658). These specimens are small relative to the larger, co-occurring Sphaeroidinella de- hiscens, and are considered by the author to be juve- nile forms of S. dehiscens. Bolli and Saunders (1985) discussed Sphaeroidinella and Sphaeroidinellopsis in detail and stated that juvenile forms of the former do not always exhibit the secondary aperture(s) which de- fine the genus Sphaeroidinella, and appear to be the ancestral Sphaeroidinellopsis, (also refer to Berggren, 1993, for discussion on Sphaeroidinella and Sphae- roidinellopsis). The remaining three locales of the Moin Formation did not produce any age-indicative planktic foraminiferal taxa. In summary, planktic foraminifera indicate that the Moin Formation is restricted to 1.9 Ma and younger based on the occurrence of G. truncatulinoides (FAD = 1.9 Ma). The consistent occurrence of right-coiled Pulleniatina obliquiloculata, suggesting an age no greater than 2.2 Ma (Saito, 1976; Keigwin, 1982), sup- ports this. Left-coiled Globorotalia menardii also sug- gests an age of 2.2 Ma or younger (Bolli and Saunders, 1985). Coates et al. (1992) date the base of the Moin at 3.0 Ma based on the co-occurrence of Sphaeroidinella and Sphaeroidinellopsis; however, the author believes the Sphaeroidinellopsis-like specimens lacking secondary apertures may be an example of ontogeny recapitulat- ing phylogeny (Gould, 1977). Further investigation, including ontogenetic studies of living Sphaeroidinella sp., is necessary to determine what characters are use- ful in distinguishing members in this genus from mem- bers of the ancestral genus, Sphaeroidinellopsis. Nannofossils were identified from 27 Moin Forma- tion samples taken from the Pueblo Nuevo Cemetery (4 samples), Lomas del Mar (6 samples), and Cangre- jos Creek (11 samples) locations. Key nannofossils identified from the Pueblo Nuevo (lower) part of the Lomas del Mar Eastern Sequence (Section 36, App. B) include Sphenolithus abies (LAD = 3.5 Ma), Pseu- doemiliania lacunosa (3.6—0.5 Ma), Calcidiscus ma- cintyrei (LAD = 1.5 Ma), Discoaster pentaradiatus (LAD = 2.4 Ma), D. brouweri (LAD = 1.9 Ma), and D. berggrenii (LAD = 5.6 Ma). Additional older nan- nofossils were found which are clearly reworked. In fact, D. berggrenii and some of the S. abies are ap- parently reworked as the planktic foraminiferal faunas discussed above are clearly younger than these two nannofossil species would indicate. Based on the spe- PLANKTIC FORAMINIFERAL BIOSTRATIGRAPHY: COTTON 71 cies consistently occurring in these samples, the lower section (12—24 m) is >1.9 Ma and <3.6 Ma. Stratigraphically above this, at Lomas del Mar (Sec- tion 36, App. B, 30-38 m), Pseudoemiliania lacunosa (3.6—0.5 Ma) occurs throughout this part of the section with Discoaster brouweri (LAD = 1.9 Ma), placing this part also at 3.6—1.9 Ma. The specimens of S. abies (LAD = 3.5 Ma) at the top of this part (at PPP 640) are probably reworked, based on the presence of Glo- borotalia truncatulinoides (FAD = 1.9 Ma) in a sam- ple (PPP 638) 1.5 m below. At Cangrejos Creek (Section 37, App. B), nanno- fossils were identified in eleven samples. The species occurring most frequently throughout the section are large Gephyrocapsa (FAD = 1.7 Ma), P. lacunosa (3.6—0.5 Ma) and C. macintyrei (LAD = 1.5 Ma), thus placing the entire section between 1.7 and 1.5 Ma. These dates concur with those based on the planktic foraminifera. Older nannofossils, including D. berg- grenii and S. abies, were identified in one sample (PPP 647) but are most likely reworked, based on the youn- ger fauna found in the same sample. Integrated nannofossil and planktic foraminifera data suggest that the top of the Moin Formation is restricted to 1.7—1.5 Ma (nannofossils) and the base is as old as 1.9 Ma based on the combined evidence of D. brouweri (LAD = 1.9 Ma) and G. truncatulinoides (FAD = 1.9 Ma). Bocas DEL ToRO GROUP The sediments that comprise this group in north- western Panama were only recently described in detail by Coates et al. (1992) for the Panama Paleontology Project. Prior to that study, the only published refer- ences were brief mentions by Terry (1956) and Olsson (1922, 1942). In general, these sediments consist of approximately 600 meters of re-worked, nearshore volcaniclastic sediments that were deposited in a back- arc setting in the Bocas del Toro Basin. The calcareous microfossils found in these units are abundant and well-preserved, and indicate an approximate age range of Late Miocene to Early Pleistocene. Nancy Point Formation The Nancy Point Formation is composed of approx- imately 500 m of sediments which are Late Miocene in age. These sediments crop out along the northeast- ern coast of Valiente Peninsula from the western mar- gin of Shark Hole Point to Nancy Point (Map 5, Ap- pendix A). Planktic foraminifera were identified from seven samples (PPP 393, 396, 397, 401, 407, 410, 411; Table 2) of the Nancy Point Formation. Their locations are shown on Insets D-F of Map 5, Appendix A, and rel- ative stratigraphic positions are plotted on Section 12, Appendix B. Key taxa identified near the base of the formation (at PPP 410—411) include Dentoglobigerina altispira (LAD = 2.9 Ma), Globigerina nepenthes (LAD = 4.0 Ma) and Globorotalia juanai (10.4—7.2 Ma). Coates et al. (1992) used the FAD of Globoro- talia exilis found (one specimen) at PPP 410 to place a maximum date of 6.5 Ma. However, Kennett and Srinivasan (1983) placed the FAD of G. exilis in mid- N18, which they equated with Late Miocene, while Bolli and Saunders (1985) placed the FAD in the Early Pliocene Zone N18 (Berggren ef al., 1985). The un- certainty of the FAD and the possibility that the spec- imen was a contaminant suggest that G. exilis is a less reliable age indicator. Planktic foraminifera that were identified near the top of the Nancy Point Formation (PPP 393) include Globigerinoides conglobatus, Den- toglobigerina altispira, Sphaeroidinellopsis sp. and G. seigliei (LAD = 4.3 Ma) to give an age range of 5.3— 4.3 Ma for the top of Nancy Point Formation. The total species diversity (planktic foraminifera) of the Nancy Point Formation ranged from 10 to 19 per sample with a mean of 16, the highest among the formations ex- amined in this study. Nannofossils from near the base and top of the Nan- cy Point Formation include Discoaster quinqueramus and D. berggrenii (both with a FAD of 8.2 Ma and a LAD of 5.6 Ma), which restrict the formation to NN 11 or 8.2—5.6 Ma. In summary, the Nancy Point For- mation is constrained to 5.6—5.3 Ma (Globigerinoides conglobatus and D. berggrenii) at the top and 8.2—7.2 Ma (Globorotalia juanai and D. quinqueramus) at the base. Shark Hole Point Formation Overlying the Nancy Point Formation is the ~340- m-thick Shark Hole Point Formation, which is Early Pliocene in age. The Shark Hole Point Formation crops out east of the Nancy Point Formation along the coast of the Valiente Peninsula from Bruno Bluff to the eastern margin of Shark Hole Point (Map 5 and Inset EK Appendix A). Planktic foraminifera were iden- tified from five samples (PPP 376, 379, 384, 388-390; Table 2) of the Shark Hole Point Formation. Their rel- ative stratigraphic positions are plotted on Section 12, Appendix B. Globigerinoides spp. are abundant, while Globorotalia plesiotumida (right-coiled), Globigerina bulloides and G. falconensis consistently occur throughout the formation. Neogloboquadrina spp. oc- cur in low abundance. The total species diversity for the Shark Hole Point Formation ranged from 8 to 18 per sample with a mean of 13, less than the underlying Nancy Point Formation but greater than the other for- mations in this study. WZ BULLETIN 357 The key planktic foraminifera identified from the base of the Shark Hole Point Formation (at PPP 388) are Dentoglobigerina altispira (LAD = 2.9 Ma), Sphaeroidinellopsis noitals (LAD = 3.0 Ma) and Glo- bigerina nepenthes (LAD = 4.0 Ma). The latter form occurs through this formation, which suggests a pre- 4.0-Ma age for most of the formation. Dentoglobiger- ina altispira, Globorotalia plesiotumida (right-coiled) Sphaeroidinellopsis sp., Pulleniatina primalis (left- coiled) and Globorotalia margaritae (at PPP 376) are present near the top of the section. Globorotalia mar- garitae restricts the maximum date to 5.6 Ma (Table 2); however, the presence of Globigerinoides conglo- batus in a lower sample, midway in the section (at PPP 384), restricts the age of the overlying samples to a post-5.3-Ma age. Pulleniatina primalis undergoes a coiling change from left to right at ~ 3.7 Ma. There- fore, the minimum age would be 3.7 Ma. Due to the lower confidence in G. plesiotumida, it is not used here. The nannofossils Sphenolithus abies (LAD = 3.5 Ma) and Reticulofenestra pseudoumbilicus (LAD = 3.5 Ma) occur consistently in the Shark Hole Point Formation, which suggests a minimum age of 3.5 Ma. The uppermost sample is barren of calcareous nanno- fossils; however, Pseudoemiliania lacunosa (3.6—0.5 Ma) occurs in the top quarter of the section at PPP 376, which limits the maximum age of the top of the formation to 3.6—-3.5 Ma. No maximum age for the base is suggested by the calcareous nannofossils. In summary, the combined data provide an age range of 5.3 to 3.7 Ma or Early Pliocene for the Shark Hole Point Formation. In a study by Dowsett and Cot- ton (1996) in which graphic correlation analysis was applied to both the nannofossil and planktic forami- nifera data, the age estimates for the Shark Hole Point Formation were 5.68 to 3.28 Ma (note: samples now considered the top of Nancy Point Formation were originally considered the base of the Shark Hole Point Formation when the graphic correlation was applied. Thus, the maximum date of 5.68 Ma would represent an age for the top of Nancy Point Formation and the base of Shark Hole Point Formation would be younger than 5.68 Ma by necessity). Cayo Agua Formation Sediments of the Cayo Agua Formation crop out along the northeastern coast of the island of Cayo Agua which lies along the northern limit of the Chi- riqui Lagoon, west of the Valiente Peninsula (Map 6 and Insets, Appendix A). These sediments are approx- imately Early to middle Pliocene in age. Planktic fo- raminifera were identified from fifteen samples of the Cayo Agua Formation (PPP 371-374, 61, 306, 307, 293, 298, 300, 334-338; Table 2). Their relative strati- graphic positions are plotted on Sections 19—20, Ap- pendix B. The faunas in this formation typically have few Neogloboquadrina spp. and abundant Globigerinoides spp. with G. obliquus consistently dominant relative to G. ruber. Dentoglobigerina altispira occurs consis- tently throughout the section and in greater relative abundance than in any other formation in this study. Globigerina bulloides and G. falconensis are subdom- inant in parts of the section. The total species diversity for the Cayo Agua Formation ranged from 3 to 20 per sample with a mean of 12, lower than the Nancy Point and Shark Hole Point formations, but higher than the Limon Group formations. The total species diversity range, however, was the greatest in this study. The lowermost samples examined in Section 19 (Appendix B, PPP 61, 371-374) contain Sphaeroidi- nellopsis sp., Globorotalia margaritae, Globigerina nepenthes and a rare occurrence of Globorotalia punc- ticulata. The age of this lower part of the Cayo Agua Formation appears to be ~4.0 Ma using the extinction of Globigerina nepenthes at 4.0 Ma and the fact that Globorotalia puncticulata did not migrate into the Ca- ribbean earlier than 4.0 Ma (Dowsett, 1989; note: based on this, G. puncticulata does not appear at DSDP site 502 until approximately 3.4 Ma, at the boundary between the Gauss and the Gilbert chrons). In the uppermost samples of Section 19 (PPP 335- 338), key taxa include Dentoglobigerina altispira, Globigerinoides conglobatus, G. obliquus (significant- ly dominant relative to G. ruber), Sphaeroidinellopsis sp. and Globorotalia plesiotumida. The age range for the top of the formation is 5.3—3.5 Ma using the FAD of G. conglobatus and the date of 3.5 Ma when G. ruber becomes dominant relative to G. obliquus in the Caribbean (disregarding Globorotalia plesiotumida due to its lower confidence, which would suggest a minimum age of 4.0 Ma). Key calcareous nannofossil taxa identified include Sphenolithus abies, Reticulofenestra pseudoumbilicus, Pseudoemiliania lacunosa and Ceratolithus acutus. The co-occurrence of S. abies and P. lacunosa (PPP 371-374) indicates an age of 3.6—3.5 Ma; however, C. acutus (5.0 to 4.6 Ma) was found in a sample near the base of the Cayo Agua (at PPP 61) and provided an age estimate for the lowermost Cayo Agua Formation for Coates et al. (1992). On the other hand, the sample with C. acutus also contains a large number of re- worked planktic foraminifera, implying that C. acutus too, may be reworked. In summary, the planktic foraminifera constrain the top of Cayo Agua to 5.3—3.5 Ma based on the domi- nance of G. obliquus relative to G. ruber and the oc- PLANKTIC FORAMINIFERAL BIOSTRATIGRAPHY: COTTON 73 currence of G. conglobatus. The base is restricted to ~4.0 Ma based on the co-occurrence of Globorotalia puncticulata and G. nepenthes. Nannofossil data for the base indicate both older and younger dates relative to the planktic foraminiferal data; 3.6—-3.5 Ma by the co-occurrence of S. abies and P. lacunosa and 5.0—4.6 Ma by the presence of Ceratolithus acutus (if it is not a product of re-working). Based on the same micro- fossil occurrences (listed above), graphic correlation analysis (Dowsett and Cotton, 1996) concluded that the uppermost Cayo Agua is 3.51 Ma and the base 5.03 Ma. Escudo De Veraguas Formation The Escudo de Veraguas Formation crops out along the north and southeast coasts of the island of Escudo de Veraguas (Map 4, Appendix A), which lies due east of the Valiente Peninsula. These Late Pliocene sedi- ments are the youngest of the Bocas del Toro Group. Planktic foraminifera were identified from thirteen samples of the Escudo de Veraguas Formation (PPP 168-170, 358, 360-362, 364-369; Table 2). Their rel- ative stratigraphic positions are plotted in Sections 10— 11, Appendix B. The general character of the fauna in the Escudo de Veraguas Formation is an overwhelming dominance of Globigerinoides ruber and G. sacculifer, with Globorotalia spp. subdominant. The total species di- versity for the Escudo de Veraguas Formation ranged from 6 to 19 per sample with a mean of 11, which is the lowest value for the Bocas del Toro Group, but higher than the Limon Group formations. The globor- otaliids found in abundance in the main (North coast) section of this formation (Globorotalia miocenica and G. pseudomiocenica) were not found in the other sec- tions of this study (except one occurrence of the latter in the Nancy Point Formation), which suggests differ- ent oceanic conditions at this locale. Key planktic foraminifera for biochronology in the Escudo de Veraguas Formation include Globigerinella calida, which Bolli and Saunders (1985) considered indicative of the Pleistocene. However, Kennett and Srinivasan (1983) showed this species to range down into the Lower Pliocene. The only globorotaliid in the southeast section (PPP 168-170) is Globorotalia cras- saformis (FAD = 4.3 Ma), which occurs in low num- bers. Pulleniatina obliquiloculata was identified at PPP 168 and suggests a post-2.2-Ma age based on its reappearance in the Atlantic (Saito, 1976; Keigwin, 1982). Samples (PPP 360-369) in the middle and low- er north section are characterized by abundant G. mio- cenica (LAD = 2.2 Ma) and G. pseudomiocenica, which indicates a minimum date of 2.2 Ma. Other key taxa include Globigerinoides obliquus (LAD = 1.8 Ma) in all samples, Globorotalia tosaensis (FAD = 3.1 Ma) and Sphaeroidinella dehiscens (FAD = 3.0 Ma) at PPP 362 (middle section), and G. truncatuli- noides (FAD = 1.9 Ma) at PPP 360 (upper section). One notable difference between the faunas of the Escudo de Veraguas Formation and the other Bocas del Toro formations (Cayo Agua, Shark Hole Point and Nancy Point) is the increase in the numbers of Globigerinoides ruber relative to G. obliquus. The switch from G. obliquus-dominant to G. ruber-domi- nant faunas occurs at DSDP site 502A between 3.5 and 3.0 Ma (Text-fig. 2), which supports ages of = 3.5 Ma for the Escudo de Veraguas Formation. Nannofossils were abundant in this formation in general. The key taxa near the top were the small Ge- phyrocapsa spp. (FAD = 2.5 Ma) and Calcidiscus macintyrei (LAD = 1.5 Ma), which restrict the age to between 2.5—1.5 Ma. Near the base of the formation key taxa include: Sphenolithus abies (LAD = 3.5 Ma), Pseudoemiliania lacunosa (3.6—0.5 Ma), Discoaster brouweri (LAD = 1.9 Ma) and D. pentaradiatus (LAD = 2.5 Ma), which restrict the age to between 3.6—3.5 Ma. To summarize the biostratigraphic data for Escudo de Veraguas, the planktic foraminifera suggest 1.9—1.8 Ma near the top, and possibly as old as 3.5 Ma at the base. The nannofossil data indicate assignment to Zone NN18 (age estimate: 2.2—1.9 Ma) designation for the top and an upper Zone NN15 (age estimate: 3.6—3.5 Ma) assignment for the base. REGIONAL CORRELATION Correlations between formations described in this study and other Neogene marine deposits of the Ca- ribbean region were made in the following manner. Age designations based on a particular zonal scheme were converted to a numerical framework based on the Neogene time scale of Berggren et al. (1985). Dates based on LADs and FADs of taxa were altered to cur- rently accepted LADs and FADs based on Berggren et al. (1985) and Dowsett (1989). Studies which pro- vided taxonomic data were critically assessed as to stratigraphic importance of key taxa. When possible, taxonomic consistency was maintained by viewing fig- ured specimens and comparative material at the U. S. National Museum and the U. S. Geological Survey, Reston, Virginia. Text-figure 1 shows the localities of these eastern Pacific and Caribbean Neogene deposits, Text-figure 3 is an age chart of the formations analyzed herein, and Text-figure 4 is a correlation chart showing the relations among all of these land-based Neogene deposits. The Neogene sediments found on the Pacific side of the southern Central American isthmus were studied 74 Panama BOCAS DEL TORO GROUP BULLETIN 357 Costa Rica LIMON GROUP. PLEISTOCENE Moin 1.8 (F) = Formation MS) escudo ae =the ' 7 H ; Veraguas ! U Formation Rio Banano| Formation i) Zz <9) S) ys ~— = = i-© Point Formation Nancy Point Formation MIOCENE Usean Formation Text-figure 3—Age chart of Caribbean, southern Central Amer- ican formations, adapted from Coates et al. (1992). Dates with “N” or “F” next to them represent dates based on a nannofossil or fo- raminiferal datum, respectively. Dates in brackets represent age es- timates made by graphic correlation analysis (Dowsett and Cotton, 1996) by Coates et al. (1992), and a more detailed bio- chronology by Cotton (1991). The results are sum- marized as follows. The Charco Azul Group deposits occur on the Burica and Osa peninsulas of Panama and Costa Rica and were dated using planktic foraminifera and calcareous nannofossils. The Charco Azul Group consists of three formations: Penita, Burica and Ar- muelles. The Penita Formation is the oldest in the group and the planktic foraminifera were sparse and poorly preserved in general. The base of this formation could only be constrained to = 3.5 Ma by Sphaeroi- dinellopsis (LAD = 3.0 Ma) and the calcareous nan- nofossil Sphenolithus abies (LAD = 3.5 Ma). Low- ermost samples from the overlying Burica Formation provided a minimum age for the Penita Formation of 3.6-3.5 Ma based on the co-occurrence of S. abies and Pseudoemiliania lacunosa (3.6—0.5 Ma). This would make the Burica Formation partially coeval with the Rio Banano and Escudo de Veraguas formations. However, Cotton (1991) noted an absence of Globi- gerinoides obliquus (LAD = 1.8 Ma) throughout the Burica Formation, whereas G. ruber was consistently present. This evidence combined with left-coiled Glo- borotalia menardii (FAD = 2.2 Ma) suggest a Late Pliocene date of =1.8 Ma for a maximum age of the Burica Formation, which would make it coeval with the Moin and upper Escudo de Veraguas formations. The top of the Burica Formation and the base of the Armuelles Formation were dated at 1.7—1.5 Ma based on the presence of both large Gephyrocapsa spp. and Calcidiscus macintyrei. The minimum age for the Ar- muelles Formation was constrained to 0.5 Ma by the presence of P. lacunosa and absence of C. macintyrei. The Armuelles Formation is younger than the youn- gest Caribbean formation in this study, the Moin For- mation. Neogene marine sedimentary sequences have been studied in detail from two regions in the Dominican Republic: the Cibao Basin in the north (Saunders ef al., 1986) and the Azua Basin in the southwest (McLaughlin, 1989). In the Cibao Basin, the Mao For- mation was dated as Zones NN14—15 (approximate age: 4.0—3.4 Ma), which correlates with the lower Rio Banano, Cayo Agua, and upper Shark Hole Point for- mations. The Gurabo Formation was dated as Zones NN12-13 (approximately 5.6—4.0 Ma), which corre- lates with the lower Shark Hole Point Formation and uppermost Nancy Point Formation. In the Azua Basin, Dominican Republic, the Trinch- era Formation was dated by McLaughlin (1989) from the base of the Globorotalia menardii Zone to the top of the G. margaritae margaritae Subzone (approxi- mately 10.4—3.6 Ma). It is not apparent what species the base is defined by as the key species G. mayeri (LAD = 10.4) was not listed for this formation. How- ever, Globoquadrina dehiscens (LAD = 5.3 Ma) co- occurs with Globorotalia margaritae (5.6—3.4 Ma) in the upper part of the formation, which gives it an age of 5.6—5.3 Ma. Therefore, the top of the Trinchera For- mation correlates with the uppermost Nancy Point For- mation. The Quita Coraza Formation was dated by McLaughlin (1989) from the middle of the Globoro- talia margaritae margaritae Subzone to the top of the G. margaritae Zone, and (based on my interpretation) using the time scale of Berggren et al. (1985) this cor- responds to approximately 5.15 to 3.4 Ma. Globigerina nepenthes occurs throughout this formation and sug- gests a minimum age of 4.0 Ma (Dowsett, 1989). Glo- bigerina nepenthes co-occurs once with Globorotalia margaritae, which further restricts the minimum age to 5.6—4.0 Ma. The maximum age relies on the oc- PLANKTIC FORAMINIFERAL BIOSTRATIGRAPHY: COTTON TS) PANAMA-COSTA RICA MEXICO CARIBBEAN PACIFIC PACIFIC AGE Maria BURICA PLEISTOCENE | EPOCH AGUEGUE XQUITE fa ESMERALDAS TIRABUZON UPPER CONCEPTION a HOLE POINT LOWER CONCEPTION wi z i oO Q jj a _—--=-. PENITA CUBAGUA NANCY USCARI POINT JAMAICA DOMINICAN REPUBLIC Virgin Islands “ ARROYO JACKSON BLANCO BLUFF upper BUFF BAY NOT GURABO NAMED KINGSHILL lower,"type” BUFF BAY Text-figure 4—Regional correlation chart comparing the Neogene sequences of this study to other formations of the Central American isthmus and in the Caribbean and eastern Pacific basins, based on studies mentioned in text. currence of Globigerinoides ruber (McLaughlin, 1989, used a FAD of mid-Zone N18, approximately 5.15 Ma) in the underlying Trinchera Formation. This for- mation correlates with the lower Shark Hole Point For- mation. The youngest formation in the Azua Basin, the Ar- royo Blanco Formation, was dated (McLaughlin, 1989) from the upper Globorotalia humerosa Zone through the Globigerinoides trilobus-fistulosus Zone (approximately 5.6-3.0 Ma). Dentoglobigerina alti- spira and Sphaeroidinellopsis sp. are the limiting taxa for the upper age estimate. The maximum date relies on the FAD of Candeina nitida, due to the absence of other data. This formation correlates with the Rio Ban- ano Formation, and possibly the Cayo Agua, Shark Hole Point and Nancy Point formations, depending on the true base of the Arroyo Blanco Formation. Bermudez and Seiglie (1970) identified planktic fo- raminifera from the Neogene Camuy Formation of Puerto Rico. They recorded Sphaeroidinellopsis sp. (LAD = 3.0 Ma), Globorotalia margaritae (5.6—3.4 Ma), D. altispira (LAD = 2.9 Ma) and G. twmida (FAD = 5.2 Ma), which indicate the age of 5.2—3.5 Ma. The Camuy Formation is correlative with the bas- al Rio Banano, basal Escudo de Verguas, Cayo Agua and Shark Hole Point formations. Palmer (1945) was the first to identify numerous planktic foraminifera from the Bowden Formation of Jamaica. Included in her species list is G. truncatuli- noides (FAD = 1.9 Ma), which would limit part of the Bowden to =1.9 Ma. Globorotalia miocenica (listed as a variety of G. menardii), which became extinct in the Latest Pliocene, is also listed. Sphaeroidinella de- hiscens (FAD = 3.0 Ma) is also listed, which limits a portion of the Bowden to = 3.0 Ma. Lamb and Beard (1972), however, did not find G. truncatulinoides be- low the Manchioneal Formation, which overlies the Bowden. Berggren (1993), in a detailed biostratigraph- ic analysis, identified Globoquadrina dehiscens, Glo- bigerina nepenthes, D. venezuelana, and Globorotalia 76 BULLETIN 357 margaritae in the Bowden Formation, which is ap- proximately Late Miocene to Early Pliocene and cor- relative with the upper Uscari, lower Rio Banano, low- er Cayo Agua, Shark Hole Point and Nancy Point for- mations. In the Buff Bay Formation of Jamaica, Lamb and Beard (1972) recorded G. margaritae (LAD = 3.4 Ma), Sphaeroidinellopsis sp. (LAD = 3.0 Ma), Sphae- roidinella sp. and G. miocenica in the upper portion, and Globigerinoides conglobatus (FAD = 5.3 Ma), N. humerosa and N. acostaensis in the lower, “‘type” Buff Bay. This puts a boundary of 3.4—3.0 Ma on the upper section and <5.3 Ma on the lower section. Also found in the lower Buff Bay was Globoquadrina dehiscens, which last occurs in the Late Miocene (LAD = 5.3 Ma). The lower, ‘type’? Buff Bay correlates with the Nancy Point and the Uscari formations, while the up- per Buff Bay is correlative with the lower Rio Banano Formation and possibly with the lower Escudo de Ver- aguas Formation. Neogene sediments from the island of St. Croix, Lesser Antilles, yielded planktic foraminifera indica- tive of the Late Miocene and Early Pliocene (Lidz, 1982). Key taxa reported include Globigerina nepen- thes (LAD = 4.0 Ma) in association with Globigeri- noides conglobatus (FAD = 5.3 Ma), providing an age range of 5.3-4.0 Ma for some of the samples. Glo- boquadrina dehiscens (not found in Panama or Costa Rica) is recorded in numerous samples, indicating an age = 5.3 Ma. Notably absent is Sphaeroidinella sp., while Sphaeroidinellopsis sp. is recorded frequently, supporting an age = 3.0 Ma. Globorotalia margaritae is absent from these deposits despite the fact that its stratigraphic range overlaps with the age of these sed- iments, suggesting that environmental conditions were not conducive to G. margaritae. The deposits from St. Croix are correlative with the lower Shark Hole Point Formation and Nancy Point Formation. Neogene deposits from the Caribbean side of the Isthmus of Tehuantepec, Mexico, were dated by Akers (1979, 1984), who identified key nannofossils and planktic foraminifera. Akers assigned Zone N20 to the Agueguexquite Formation, which contains Sphaeroi- dinella sp. (FAD = 3.0 Ma) and nannofossils P. la- cunosa (FAD = 3.6 Ma) and D. pentaradiatus (LAD = 2.4 Ma), fixing the age between 3.0 and 2.4 Ma. The absence of S. abies (LAD = 3.5 Ma) supports this age. This formation is correlative with the Rio Banano Formation and possibly Escudo de Veraguas Forma- tion. The Filisola Formation (Akers, 1979) contains no age-indicative planktic foraminifera; however, the co- occurrence of P. lacunosa and S. abies limits the age to 3.6-3.5 Ma. Planktic foraminifera identified from the Concepcion Inferior Beds (Akers, 1979) include G. margaritae (5.6—3.4 Ma), Sphaeroidinellopsis sp. (LAD = 3.0 Ma), and Globigerinoides conglobatus (FAD = 5.3 Ma), which yield an age of 5.3—3.4 Ma, but the same key nannofossil taxa occur as in the Fil- isola, restricting the age to 3.6-3.5 Ma. The Concep- cion Superior Beds contain Globigerina nepenthes (LAD = 4.0 Ma) and Globigerinoides conglobatus (FAD = 5.3 Ma), constraining the age to 5.3—4.0 Ma. However, Akers also identified both nannofossil taxa P. lacunosa (FAD = 3.6 Ma) and S. abies (LAD = 3.5 Ma), which suggests a younger age. The Filisola, Concepcion Inferior and Concepcion Superior beds correlate with the Rio Banano, Cayo Agua, and Shark Hole Point formations and possibly the basal Escudo de Veraguas Formation. In the eastern Falcon region, Venezuela, Blow (1959) reported numerous planktic foraminifera from the Poz6n Formation, most of which are indicative of a Miocene age (e.g., Globoquadrina dehiscens). How- ever, he also recorded S. dehiscens and G. miocenica, both indicative of the Late Pliocene. If these identifi- cations are correct, the upper Pozon Formation corre- lates to the Rio Banano, Cayo Agua, and Shark Hole Point formations. The Esmeraldas Formation of Ecuador yields well preserved, abundant planktic foraminifera which pro- vided Hasson and Fischer (1986) with an age range of 3.5—3.2 Ma. However, in their table of species occur- rences are G. tosaensis and S. dehiscens, which sug- gest younger dates. Hasson and Fischer discussed the discrepancy of G. tosaensis in the samples as its FAD = 3.1 Ma, and S. dehiscens has a FAD = 3.0 Ma. Further, Globigerinoides ruber is found in most of the samples while G. obliquus is found in less than half of the samples (Text-fig. 2). This supports the mini- mum age indicated by the above mentioned FADs of 3.0 Ma, to give an age of 3.5—3.0 Ma. The Esmeraldas is correlative with the Rio Banano Formation and pos- sibly lower Escudo de Veraguas Formation. Duque-Caro (1990) dated the Munguido Formation, in northwestern Colombia, as 3.4 Ma to Late Miocene Zone N17 based on Globorotalia margaritae (5.6—3.4 Ma) found near the top and G. conomiozea subcon- omiozea found at the base. The Munguido is correla- tive with the Cayo Agua, Shark Hole Point, upper Nancy Point, and lower Rio Banano formations. The Neogene of Maria Madre island, off the Pacific coast of Mexico, was studied by Carreno (1985). She identified Globigerinoides fistulosus (FAD = 2.9 Ma), S. dehiscens (FAD = 3.0) and the nannofossils P. la- cunosa (FAD = 3.6 Ma) and D. pentaradiatus (LAD = 2.4 Ma). These taxa provide an age of 2.9—2.4 Ma and are correlative with the upper Rio Banano For- mation and possibly the lower Escudo de Veraguas PLANKTIC FORAMINIFERAL BIOSTRATIGRAPHY: COTTON igh Formation. Carrefio (1981) identified planktic forami- niferal faunas from the Neogene Tirabuzon Formation, Baja, California, and assigned a date of Early to mid- dle Pliocene. This formation approximately correlates to the Rio Banano, Cayo Agua, and Shark Hole Point formations. Dowsett and Wiggs (1992) showed that planktic fo- raminiferal faunas from the Yorktown Formation in southeastern Virginia support an age of 4.0—3.0 Ma based on the presence of Sphaeroidinellopsis seminu- lina (LAD = 3.0 Ma), Dentoglobigerina altispira (LAD = 2.9 Ma) and Globorotalia puncticulata (FAD = 4.0 Ma). These sediments correlate with the Rio Banano, Cayo Agua, and upper Shark Hole Point for- mations. Akers (1972) analyzed planktic foraminiferal faunas from the Waccamaw Formation of North and South Carolina, and the Jackson Bluff Formation of south- western Florida. The Waccamaw Formation yielded G. truncatulinoides (FAD = 1.9 Ma) and G. obliquus (LAD = 1.8 Ma) at one locale, which limits the age of at least part of the Waccamaw to 1.9—1.8 Ma. This portion of the Waccamaw, therefore, correlates to the Moin Formation of this study. Akers (1972) recorded the presence of Sphaeroidinella sp. (FAD = 3.0 Ma), Sphaeroidinellopsis sp. (LAD = 3.0 Ma), G. margar- itae (5.6—3.4 Ma), D. altispira (LAD = 2.9 Ma) and Globigerinoides conglobatus (FAD = 5.3 Ma) in the Jackson Bluff Formation. The co-occurrence of Sphaeroidinella and Sphaeroidinellopsis in the same horizon/locality indicates an age of 3.0 Ma. Globigerinoides conglobatus co-occurring with Glo- borotalia margaritae at another locality restricts the age of that sample to 5.3-3.4 Ma. A third locality which contains Sphaeroidinella sp. and D. altispira, but not Sphaeroidinellopsis sp., is restricted to 3.0—2.9 Ma. The overall age assigned by Akers is Early Plio- cene to early Late Pliocene. Thus the Jackson Bluff was deposited contemporaneously with the lower Rio Banano, Shark Hole Point and the Cayo Agua for- mations of this study. SUMMARY Using planktic foraminifera (this study) and calcar- eous nannofossil data (Bybell, Chapter 2, this volume), I have established a biochronologic framework for the Neogene strata along the Caribbean coast of the south- ern Central American isthmus (Text-fig. 1). In the Bocas del Toro Group, the oldest formation in the study, the Nancy Point Formation, is Late Mio- cene, between 8.2 to 5.3 Ma in age. The Shark Hole Point Formation overlies the Nancy Point Formation and is Early to mid-Pliocene, between approximately 5.3 and 3.7 Ma. The Cayo Agua Formation is contem- poraneous with the Shark Hole Point Formation and is Early to middle Pliocene (between 4.0, possibly 5.0, and 3.5 Ma). The Escudo de Veraguas Formation is the youngest of the Bocas del Toro Group and is dated as middle to Late Pliocene (between 2.2, possibly 3.5, and 1.8 Ma). In the Limon Group, the uppermost Uscari Forma- tion is dated as Late Miocene (between 7.7—5.6 Ma) from this study; previous studies indicate that the base is Early Miocene (23.7 Ma). The Rio Banano For- mation is dated as middle to early Late Pliocene (3.6 to 2.4 Ma). The Moin Formation is dated as uppermost Pliocene to Lower Pleistocene (1.9 to 1.5 Ma). Mean species diversity of planktic foraminifera per sample in the Bocas del Toro Group decreases from the oldest (16 species) to the youngest (11 species) formation, which coincides with an increasingly emer- gent isthmus. Evidence from benthic foraminifera (Collins, 1993; Collins et al., 1995) suggests a general shallowing from the Nancy Point Formation to the Es- cudo de Veraguas Formation. Faunal diversity in the Limon Group is lower than the Bocas del Toro Group with a mean species diversity of 7 species, compared to 13 for Bocas del Toro. The frequently found co-occurrence of Sphenolithus abies and Pseudoemiliania lacunosa in this study im- plies that during the 100,000 year period between 3.6 Ma and 3.5 Ma there was a widespread sedimentation event which was preserved extensively in both the Li- mon and Bocas del Toro basins, as well as other lo- cales in the Caribbean Basin (e.g., Akers, 1979, 1984). The 3.6—3.5 Ma period corresponds to a eustatic sea- level rise of approximately 60 m, the TB3.6 sea-level cycle of Haq er al. (1987), dated as 3.8—2.9 Ma. The Atlantic Coastal Plain Model of Krantz (1991) also documented a sea-level rise between 4.0 Ma and 3.3 Ma. An alternative explanation to this phenomenon is that one or more of the accepted LADs and FADs for these two species may not be applicable in these shal- low, nearshore deposits. Pseudoemiliania lacunosa may have originated earlier than 3.6 Ma or S. abies may not have become completely extinct at 3.5 Ma. Global diachrony in first and last appearances of Late Neogene planktic foraminifera and nannofossils was demonstrated by Dowsett (1988, 1989), among others, and diachrony may exist between deep-sea occurrenc- es and those on the continental shelf. The age restrictions placed on the Neogene sedi- ments examined in this study aid in geologic investi- gations of the Central American isthmus, especially those related to the history of its emergence, temporal shifts in its faunal assemblages, and evolutionary and ecological changes of marine organisms living adja- 78 BULLETIN 357 cent to the isthmus. The biochronology provided in this study also contributes to the regional correlation of Neogene deposits in the Caribbean and eastern Pa- cific basins. REFERENCES CITED Akers, W.H. 1972. Planktonic foraminifera and biostratigraphy of some Neo- gene formations, Northern Florida and Atlantic coastal plain. Tulane Studies in Geology and Paleontology, vol. 9, no. 1—4, pp. 1-139. 1979. Planktonic foraminifera and calcareous nannofossil bio- stratigraphy of the Neogene of Mexico, part I—middle Pliocene. Tulane Studies in Geology and Paleontology, vol. 15, no. 1, pp. 1=32. 1984. Planktonic foraminifera and calcareous nannofossil bio- stratigraphy of the Neogene of Mexico, part II—Lower Pliocene. Tulane Studies in Geology and Paleontology, vol. 18, no. 1, pp. 21-36. Bandy, O.L. 1970. Upper Cretaceous-Cenozoic paleobathymetric cycles, eastern Panama and northern Colombia. Gulf Coast As- sociation of Geological Societies, Transactions, vol. 20, pp. 181-193. Berggren, W.A. 1993. Neogene planktonic foraminiferal biostratigraphy of east- ern Jamaica. Geological Society of America, Memoir 182, pp. 179-217. Berggren, W.A., Kent, D.V., and Van Couvering, J.A. 1985. Neogene geochronology and chronostratigraphy. in The chronology of the geological record. N.J. Snelling, ed., Geological Society of London, Memoir 10, pp. 211-260. Bermudez, P.J., and Seiglie, G.A. 1970. Age, paleoecology, correlation and foraminifers of the Uppermost Tertiary formation of northern Puerto Rico. 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Occasional Pa- pers of the California Academy of Sciences, vol. 23, 91 PP- Thalmann, H.E. 1934. Die regional-stratigraphische verbreitung der oberkreta- zischen foraminiferen-gattung Globotruncana Cushman, 1972. Ecologae Geologiae Helvetiae, vol. 27, no. 2, pp. 413-428. Weyl, R. 1980. Geology of Central America. Gebruder Bornitraeger, Ber- lin, 312 pp. Zangula, R.P. 1968. A new breakthrough in sample washing. Journal of Pa- leontology, vol. 42, no. 4, p. 1092. Zhang, J., Miller, K.G., and Berggren, W.A. 1993. Neogene planktonic foraminiferal biostratigraphy of the northeastern Gulf of Mexico. Micropaleontology, vol. 39, no. 4, pp. 299-326. CHAPTER 4 A PALEOENVIRONMENTAL ANALYSIS OF THE NEOGENE OF CARIBBEAN PANAMA AND COSTA RICA USING SEVERAL PHYLA LAUREL S. COLLINS Department of Earth Sciences Florida International University Miami, Florida 33199, U.S.A. ORANGEL AGUILERA Centro de Investigaciones Arqueolégicas, Antropoldgicas y Paleontologicas Universidad Francisco de Miranda Coro, Estado Falcon, Venezuela PAMELA E BORNE Louisiana Sea Grant College Program Louisiana State University Baton Rouge, Louisiana 70803, U.S.A. STEPHEN D. CAIRNS Department of Invertebrate Zoology Smithsonian Institution Washington, D.C. 20560-0163, U.S.A. INTRODUCTION The Neogene sedimentary environments of the Pan- ama Canal, Bocas del Toro, and Limon basins were controlled by local and basin-wide Caribbean ocean- ographic conditions, regional tectonic movements, changes in sea level, and seafloor topography. Through the approximately 9 million years spanned by the sed- iment samples of the Panama Paleontology Project (PPP) examined in this study, only a few large-scale oceanographic and tectonic changes can be discerned above the “‘noise”’ of the more locally expressed con- trols. Local paleoenvironments, determined at the level cf individual sedimentary samples to formations, are interpreted by members of the PPP from four taxo- nomic groups: benthic foraminifera, corals, ostracodes and teleost fish. The data from this diverse array of taxa are used to summarize the depositional histories of the three Neogene basins. One of the most important contributions of paleoen- vironmental analyses to the PPP is that all of the pa- leobiologic data can be identified and compared within the framework of space and habitat, as well as time. Data on taxonomic occurrences of marine organisms used in studies of biodiversity, evolution (origination and extinction), biogeography and ecology begin by defining as precisely as possible their environment and their chronologic position. To make valid comparisons of taxa from different habitats, environment must be held constant to standardize the results and reduce sampling bias. For example, an analysis of Pliocene changes in the ecological structure of southern Central American mollusks would be meaningless if early Pliocene inner neritic and late Pliocene outer neritic taxa were compared. Inner and outer neritic faunas would be expected to differ without any change in time. Environmental bias is often not a problem with studies of change through time at large chronologic or geographic scales, such as for all Paleozoic shelf fau- nas or global biodiversity, but when faunal changes are examined at the relatively fine scale targeted by the PPP to unravel evolutionary histories and examine biogeographic changes, paleoenvironmental analyses become critical to standardizing faunal occurrence data. 82 BULLETIN 357 Paleoenvironmental studies are also valuable geo- logically for interpreting tectonic changes and the stra- tigraphy of paleobasins. For example, when paleoen- vironments are expressed as water depths, the paleo- bathymetric changes can be partitioned into tectonic and eustatic components, and rates of subsidence and uplift can be calculated (Collins et al., 1995). The large-scale structure of paleobasins can also be deter- mined from the paleobathymetry, from hydrographic data such as oxygen content infered from paleofaunas, and from the biogeographic affiliations of the faunas. Four taxonomic groups were used to interpret Neo- gene paleoenvironments of Caribbean Panama and Costa Rica: 1. Benthic foraminifera, which are part of the mei- ofauna, are the most commonly used fossils for ana- lyzing Cenozoic paleoenvironments because of their great abundance and diversity, their presence in vir- tually all marine-influenced environments, and the ex- tensive knowledge of their ecology. 2. Ostracodes, which are found in marine, brackish, hypersaline and freshwater habitats, are the next most frequently used taxon. This is probably a result of low- er abundances rather than individual taxa having larger environmental ranges. Furthermore, studies of envi- ronmental distributions of modern ostracodes of the Gulf of Mexico and Caribbean region have lagged be- hind studies of modern assemblages elsewhere, such as Europe. 3. Otoliths (fish earbones) are widely distributed and often abundant in PPP sediment samples. Because they are not as abundant as benthic foraminifera and ostracodes, have fewer taxonomic experts, and are commonly identifiable only to the genus level, they have traditionally been less useful for paleoenviron- mental analysis. However, although their originally pe- lagic habit results in wider geographic distributions of taxa, they are shown herein to be a powerful comple- mentary tool in determining paleobathymetry for most of the formations analyzed. 4. Scleractinian corals that lack symbiotic zooxan- thellae are usually not associated with reefs and live in a wide range of depths and temperatures. They are the least abundant and diverse taxon employed in this study (having only about 12 extant cosmopolitan spe- cies), but have also proved useful for paleoenviron- mental analysis. As with the other three taxa, their spe- cies distributions can be described by ranges in water depths. Ahermatypic corals occur down to 6300 m and geographic ranges are wider with increasing depth, so relatively shallow taxa, such as those reported herein, are commonly confined to one side of an ocean basin, e.g., the tropical Western Atlantic. Mollusks, including bivalves, gastropods and ceph- alopods, are quite useful for paleoenvironmental de- terminations. However, they are not included in this chapter because the necessary ecological studies of this vast tropical fauna have not been done. ACKNOWLEDGMENTS We thank Martin Buzas, Marie-Pierre Aubry and Barun Sen Gupta for their helpful reviews of this chap- ter. This is contribution number 11 to The Program in Tropical Biology at Florida International University. COMPARISON OF METHODS USED FOR DIFFERENT TAXA The methods employed in specimen preparation and paleoenvironmental determinations are described in more detail by Collins (benthic foraminifera, this vol- ume), Aguilera and Aguilera (teleost otoliths, this vol- ume), Borne et al. (ostracodes, this volume), and Cairns (ahermatypic corals, this volume). Below, we summarize briefly the similarities and differences in their paleoenvironmental methods and uses. For the benthic foraminiferal analyses, ecological relationships of modern species of the Caribbean Sea and Gulf of Mexico are applied to their fossil coun- terparts mainly in terms of ranges in water depths (but see Collins, this volume, for ranges in terms of habi- tat). Depth itself is not an ecological control, but rather, a proxy for a combination of correlated environmental parameters that generally vary consistently within, and even between, oceanic basins. About 90% of the spe- cies that are common (i.e., occur at =1% frequency in at least one sediment sample) in the Bocas del Toro and Limon groups are extant. For extinct taxa, depth ranges are determined through their association with extant species in fossil sediments of the tropical Amer- ican region. The techniques used for interpreting paleoenviron- ments are most similar for ostracodes and benthic fo- raminifera. In this study, species-level data for these fossils are compared to modern distributions of the same species found within the same Caribbean ocean basin, and separate depth determinations are made for each sample from a collecting site, which has a PPP number. As with benthic foraminifera, most ostracode species are still living; approximately 76% of identi- fied ostracode species of the Limon Group, Caribbean Costa Rica, are extant. The techniques differ in two ways. First, benthic foraminiferal interpretations are based on the deepest dwelling taxa (to allow for pos- sible downslope redeposition of sediments) using the overlap of the species’ paleobathymetric ranges, whereas ostracodes use the overall similarities between Recent and fossil faunas. Second, relative abundances PALEOENVIRONMENTS: COLLINS ET AL. 83 FORMATIONS PANAMA BOCAS DEL TORO BASIN LIMON BASIN CANAL BASIN Moin. Chagres,} Cayo Swan Escudo de Nancy|Rio Lomas Moin, Gatun Rio Indio 500 400 300 200 s 100 PALEOBATHYMETRY (M) KW Overlap (m): 15-40 50-80 37-46 Agua Cay Veraguas SG gypsy gIH44s4g4s9]9 II IIIIIO Point |Banano del Mar type TAXA benthic foraminifera ahermaty pic corals ostracodes teleost fish - g Y Z CONE Te 80-100 100-150 300-500 20-40 50-75 150-250 Text-figure 1.—Paleobathymetric comparison of members or formations (columns) of the Panama Canal, Bocas del Toro, and Limon basins using four different taxa. Shading denotes the overlap in depth ranges shown across the bottom. were used in the foraminiferal analyses, whereas the presence vs. absence of taxa was used for ostracodes. The use of paleoenvironmental methods with aher- matypic corals is similar to those with microfossils in that they are at the species level and use the overlap of bathymetric ranges. However, specimens are too large and sparsely distributed to make determinations for single samples, so bathymetric ranges of taxa from multiple samples of each formation or section are com- bined into one paleoenvironmental analysis. This method carries the assumption that the depositional en- vironment did not change within the formation/section, which was generally supported in this case by previ- ously published results for separate samples (Collins et al., 1995). Extinct taxa in the analyses (only about 20% of the ahermatypic coral species are extant) were compared to closely related living species for about 30% of the taxa. Paleoenvironmental analysis of teleost otoliths was standardized by Nolf and Brzobohaty (1992) and dif- fers from the methods used for the other taxa. Taxo- nomic identifications of fish from otoliths are generally at the generic level. Approximately 98% of the fossil genera are extant. Relative abundances of genera are calculated per 100-m-depth interval and the interval in which the most genera peak is considered the most likely depositional environment. If several peaks occur, reworking or some other process is implied and the results have less confidence. Otoliths, like ahermatypic corals, are too sparsely distributed in sediments to cal- culate paleobathymetries for single samples, so genera found within a formation or other stratigraphic unit are combined into one analysis. PALEOENVIRONMENTAL DATA TRENDS IN PALEOENVIRONMENTAL DETERMINATIONS FROM DIFFERENT TAXA A side-by-side comparison of paleoenvironmental determinations from benthic foraminifera, ostracodes, ahermatypic corals and otoliths indicates the extent to which they agree. Text-figure 1 plots paleobathyme- tries determined from the different taxa at the finest stratigraphic level at which they can all be compared: either biofacies, members or formations. Results from different taxa support each other quite well. For all stratigraphic units, the paleodepths from different taxa overlap each other. Except for possible prior know]l- edge of the previously published benthic foraminiferal results, each paleontologist worked independently. Even if the benthic foraminiferal results are excluded, the remaining water depth ranges overlap. To some extent, the differences in results (Text-fig. 1) mirror the differences in the methods and the tax- 84 BULLETIN 357 onomic level of identification. The benthic foraminif- eral paleobathymetric ranges always overlap those of another taxon and are commonly the most constrained. In % of the cases, the benthic foraminiferal range de- fines the complete range overlap. Benthic foraminifera tend to fall at the deeper end of the ranges of the other taxa, as happens with the Gatun Formation, Rio Indio biofacies of the Chagres Formation, Nancy Point For- mation, Swan Cay Formation, and Lomas del Mar Member of the Moin Formation. This is at least partly because paleodepths based on benthic foraminifera emphasize the shallowest ranges of the deepest dwell- ing taxa to compensate for possible downslope trans- port, which would tend to bias paleodepths toward deeper values. Depth ranges from ostracodes and tel- eost fish always overlap and generally extend beyond those of the benthic foraminifera at both the shallow and deep ends. In the case of ostracodes, this may be because estimates are based on presence/absence of species rather than relative abundance, their depth preferences are less well studied, the planktic species are more mobile, and/or proportionately fewer species are still extant. In the case of teleost fish, it may be because depth ranges of genera are wider than indi- vidual species and/or because planktic taxa are gen- erally more widespread than benthic ones. Ahermatyp- ic coral ranges overlap the other ranges in all three instances. Applying the modern depth ranges of four different taxa (each with its own analytic method) to paleo- depths of stratigraphic units results in paleoenviron- mental interpretations with a high degree of confi- dence. How constrained the depths are depends on the approach used to arrive at consensuses. Estimating the paleodepth of each formation based on the union of all taxonomic ranges gives the highest degree of con- fidence but the lowest resolution. Paleodepth estimates based on the intersection (overlap) of the ranges (Text- fig. 1, bottom) give the highest resolution and a lower degree of confidence, but one that is higher than any estimated from a single taxon. These comparisons are based at the levels of formations, members or biofa- cies, but Table 1 presents all paleoenvironmental de- terminations at a finer scale, for PPP sections on the Caribbean side of the Isthmus of Panama (App. B, this volume). COMPARISON OF PALEOENVIRONMENTS FROM THE CARIBBEAN COASTAL BASINS The main Neogene sedimentary basins of the Carib- bean side of southern Central America are, from east to west, the Panama Canal, Bocas del Toro, and Limon basins (Coates, this volume). They are dissimilar in their geographic spread, local to regional tectonic in- fluences, and thickness of sediments. The Panama Ca- nal Basin of central Panama (App. A, Maps 1-2, this volume) contains Eocene (Stewart and Stewart, 1980) to Upper Miocene, marine and terrestrial sediments, and as late as 6 Ma the basin connected Caribbean and tropical Pacific surface waters (Collins ef al., 1996a). The Panama Paleontology Project has examined the upper Middle to Upper Miocene Gatun Formation and the Upper Miocene Chagres Formation. Benthic fora- minifera and otoliths (Text-fig. 1) show the siliciclastic sediments of the Gatun Formation to be nearshore to middle neritic (Table 1, Sections 1—2, 6). Benthic fo- raminifera of the Chagres Formation indicate that the sediments were deposited in deep water (Table 1, Sec- tions 3—4). However, its Rio Indio biofacies reflects a much shallower depth (Text-fig. 1; Table 1, Section 5; Appendix A, Map 1) according to the benthic fora- minifera (SO—80 m) and otoliths (<100 m). The Panama Canal Basin was episodically an inter- oceanic strait. In the Early Miocene, Pacific and Ca- ribbean waters mixed to bathyal depths, as shown by depth preferences of benthic foraminifera from the La Boca Formation (Blacut and Kleinpell, 1969). With the late Middle Miocene deposition of the Gatun Forma- tion, waters had shallowed to the point that the strait was closed, as indicated by the 84-m elevation of the lowest-lying Pacific-Caribbean passageway in the re- gion, and by the biogeography of the benthic forami- nifera, which are Caribbean to cosmopolitan in oceanic affiliation. By 6 Ma, basinal waters had deepened to upper bathyal depths to allow Pacific inflow, as indi- cated by the paleobathymetry and oceanic affiliation of benthic foraminifera from the Chagres Formation (Collins et al., 1996a). Late Miocene sea level rise cannot account for the large (approximately 200-m) increase in water depth, so some tectonic event must have been the cause. The central Panama region was apparently tectonically quiescent in Late Miocene time (de Boer et al., 1988), but fault blocks in the Panama Canal Basin (Mann and Corrigan, 1990) may have dropped to produce the Late Miocene reopening of the strait. The shallower Rio Indio biofacies of the Cha- gres Formation (Table 1, Section 5) delimits the west- ern edge of the deep-water strait. The paleoenvironmental history of the back-are Bo- cas del Toro Basin of western Panama (Appendix A, Maps 4-9) can be traced from Early Miocene, deep- water sediments (Appendix B, Section 12) that formed the earliest Isthmus of Panama. Coastal sections of both the Bocas del Toro Basin and Limon Basin (see below) typically expose a sequence comprised of Up- per Miocene bathyal sediments, Pliocene neritic sedi- ments and Lower Pleistocene coral reefs (Appendix B, Section 25). Although most composite sections (Table PALEOENVIRONMENTS: COLLINS ET AL. 85 Table 1.—Paleoenvironmental information for geologic sections (Appendix B) of the Panama Canal, Bocas del Toro, and Limon basins. Superscripts refer to the following references: | Collins, 1993; * Collins er al., 1995; * Collins er al., 1996a; * Collins in Jackson et al. (this volume); * Cairns (this volume); © Borne ef al. (this volume); ’ Aguilera and Aguilera (this volume); * Coates (Appendix B, this volume). References 5, 6 (in part), and 7 determined paleoenvironments for members or formations rather than sections, and are cited where samples from those sections were included. Paleoenvironments pertain to whole geologic sections except where intervals of vertical thickness are specified. The most constrained paleodepth ranges, equal to the overlap of individual determinations, are cited. Sec. No. Section name (interval, m) Paleodepth (m) Additional descriptors PANAMA CANAL BASIN AND NORTH COAST OF PANAMA 1347 Sabanita to Payardi 15—40* Pas: Margarita to Gatun 10—40+ sh) Toro Point 200—S00* Pacific influence* 43 Pina 200-500 Pacific influence* s Indio River 50-80 637 Miguel de la Borda, | km to the East DIS BOCAS DEL TORO BASIN Olea Escudo de Veraguas, Northern Coast 100-150! muddy! Li}'5.6 Escudo de Veraguas, Southeastern Coast 100-150! carbonate shoal/reef influence! 12147 Valiente Peninsula, Bruno Bluff to Plantain Cays (1957-1937) 150-200! Bruno Bluff Mbr. (1780-1615) 100—200%” Shark Hole Point Fm., possible coastal upwelling (1607-1428) 200-5007 upper Nancy Point Fm. (1245-1168) 300-500!” lower Nancy Point Fm. 144 Valiente Peninsula, Toro Cays 60—100* ilSy Valiente Peninsula, Southern Coast (421) 150—300# (O) 100—200* 16Gb Cayo Agua, Norte Point, Western Side 20-40! proximal carbonate shoals/reef! gp Cayo Agua, Piedra Roja Point, Western Sequence 10—75+ proximal carbonate shoals/reef?* 1914-7 Cayo Agua, Norte Point to Tiburon Point (293-264) 20—70* proximal carbonate shoals/reef* (54-0) 40-80! proximal carbonate shoals/reef! PAYS Cayo Agua, South of Nispero Point inner-middle shelf® Pape Bastimentos Island, Fish Hole, Eastern Sequence 40—100* proximal carbonate shoals/reef* 2B es Bastimentos Island, Fish Hole, Western Sequence (1 1—2.6) 75-1007 (2.6-0) 40-1007 proximal coral reef** 25en8 Swan Cay, North of Colon Island 80—120+7 coral reef** 268 Colon Island, Hill Point coral reef® LIMON BASIN Dif Sandbox River ~200° continental shelf edge? 282 Carbon Dos (Dindiri) 300-5002 29281, Banano River 20-402 proximal carbonate shoals/reef* Sie Bananito River <100* 522° Santa Rita 20-407 proximal carbonate shoals/reef* 389% Chocolate to Buenos Aires (665-557) 1-96 carbonate®, coral reef® 34408 Empalme (65-108) 10-301 coral reef* & restricted nearshore® S67555 Lomas del Mar, Eastern Sequence 50-73" coral reef?* e728 Lomas del Mar, Western Reef Sequence 150-250? possible coastal upwelling® Bens Lomas del Mar, Western Reef Track Sequence 50-100" coral reef** 39% Vizcaya River 0-25+ 1, Sections 10-11, 14-15, 17, 20, 22-23, 25, 26) in- clude no biochronological changes, the few that do transit geologic time shallow upwards. The thick, northern Valiente Peninsula coastal section (Appendix A, Map 5) extends from the Upper Miocene Nancy Point Formation to the middle Pliocene Bruno Bluff Member of the Shark Hole Point Formation. Ostra- codes of the Nancy Point Formation indicate outer ne- ritic to upper bathyal deposition, whereas benthic fo- raminifera give the more constrained, upper bathyal depth (Text-fig. 1). Benthic foraminifera of the Bruno Bluff Member indicate an outer neritic paleodepth (Ta- ble 1, Section 12). All taxa (Text-fig. 1) indicate an inner to middle neritic depositional setting for the Cayo Agua Formation, on the small island of Cayo Agua (Appendix A, Map 6). Ahermatypic corals show the most constrained, middle neritic paleodepth. Ben- thic foraminifera show a carbonate shoal or reef influ- ence and some shallowing from the early Early Plio- cene (Table 1, Section 19) to late Early Pliocene (Table 86 BULLETIN 357 1, Section 16). The Escudo de Veraguas Formation (Appendix A, Map 4) does not reflect any measurable change in water depth through time, but exposures along the southern coast (Table 1, Section 11) show a stronger carbonate shoal to reef influence than the northern coastal section (Table 1, Section 10). All pa- leobathymetric estimates, except for the otolith deter- mination, which spans 0—300 m, support outer neritic depths (Text-fig. 1). Coral reefs and reef-associated sediments of Bocas del Toro are primarily Upper Pliocene (Table 1, Sec- tions 22, 26) to Pleistocene (Table 1, Section 25). Reefal deposits of the Pleistocene Swan Cay Forma- tion (Text-fig. 1) were relatively deep, as indicated by the benthic foraminifera and supported by the otoliths, with paleodepth ranges overlapping at 80—100 m. AI- though the Middle Miocene Valiente volcanics con- tains both hermatypic reef patches and ahermatypic corals, they are patchy and small in scale and diameter, and there appears to be an increasing abundance of larger scale reefs through time in the region. The pat- tern of an increasing occurrence of reefs is consistent with the hypothesis of an increasing carbonate regime in the Caribbean during the Late Miocene to Pliocene, associated with the progressive constriction of the Pa- cific-Caribbean seaway (Collins ef al., 1996b). Sediments of the Limon Basin, Costa Rica (Appen- dix A, Maps 10—11) were also deposited in a back-arc volcanic setting. The basin contains Upper Oligocene, carbonate-bank sediments that underlie bathyal silici- clastics of the thick, uppermost Oligocene to Upper Miocene Uscari Formation (Cassell and Sen Gupta, 1989a; Collins et al., 1995; Appendix A, Map 10; Ta- ble 1, Sections 27, 28). Upper Pliocene to lowermost Pleistocene sediments are exposed in relatively thin sections (Coates, this volume) and have a strong car- bonate shoal to reef influence (Table 1, Sections 29, 32-34, 36-38). The Rio Banano and Moin formations span measurable geologic time and display varying fa- cies in different exposures, but paleowater depths within their composite sections did not change mea- surably. The Rio Banano Formation (Appendix A, In- set C of Map 11; Table 1, Section 29) exposes a pre- dominantly siliciclastic facies at Quitaria, whereas the type section at Bomba (Cassell and Sen Gupta, 1989b) shows a strong carbonate shoal to reef influence. Ben- thic foraminifera, ostracodes and otoliths all indicate an inner to middle neritic setting, although the benthic foraminifera suggest the more constrained range of 20—40 m. The Moin Formation includes extremely dif- ferent paleoenvironments. Sediments of the type Moin section (Table 1, Section 37), exposed along an un- named creek in the E] Cangrejo community (Appendix A, Inset B of Map 11), were deposited in relatively deep water near the shelf edge, as indicated by benthic foraminifera and ostracodes (Text-fig. 1), possibly within an area of coastal upwelling (Borne et al., this volume). Up the steep hill from this exposure and forming one in a series of coral-reef ridges (Taylor, 1973, 1975) are the in-place coral reefs (Table 1, Sec- tions 36, 38) of the Lomas del Mar area, also included within the Moin Formation. Paleoenvironmental de- terminations using benthic foraminifera, ahermatypic corals and ostracodes, which have an overlapping pa- leobathymetry of 50—75 m (Text-fig. 1), indicate that the coral reefs were relatively deep. The Bocas del Toro and Limon basins have the same general back-arc setting and history of uplift, but sed- imentation patterns and some microfaunas indicate dis- similar paleoenvironments for the same ages and water depths. Similar Pliocene to Recent patterns of tectonic uplift between the two basins are indicated by covary- ing rates of coastal emergence and increased rates of Pleistocene to Recent uplift caused by the arrival of the subducted Pacific Cocos Ridge at the Caribbean coast (Collins et al., 1995). Sediment sources differed. Sediments of the Rio Banano Formation, derived from erosion of the Cordillera de Talamanca, formed a thick deltaic wedge that was part of the Miocene to Recent, broad coastal fan system of Costa Rica to southern Nicaragua (Sheehan et al., 1990). Coeval deposits of the Bocas del Toro Basin have no such facies, and faunas also differed. Cluster analyses of benthic fora- minifera (Collins et al., 1995) and ostracodes (Borne et al., this volume) generally group assemblages with similar ages and environments, but within these large clusters, benthic foraminiferal assemblages separate completely the uppermost Miocene, upper bathyal Nancy Point Formation of Bocas del Toro and Uscari Formation of Limon. They also distinguish the middle Pliocene, inner to middle neritic Cayo Agua Formation of Bocas del Toro and Rio Banano Formation of Li- mon. Additionally, oxygen isotopes of mollusks from the same ages and paleodepths, but from the two dif- ferent basins, exhibit different ranges in 6!°O, sug- gesting that marine conditions in the two basins were different (Terranes et al., 1996). SUMMARY 1. Paleoenvironmental methods among the four tax- onomic groups differ, mainly in the level of taxonomic and stratigraphic units considered. Benthic foraminif- era and ostracode methods are based on species and sediment samples, the ahermatypic coral method uses species and members to formations, and the teleost otolith method uses genera and members to forma- tions. The differences in methods are reflected in the differences in results to some extent. PALEOENVIRONMENTS: COLLINS ET AL. 87 2. Nonetheless, results of the paleoenvironmental interpretations using four different taxa overlap each other in all formations studied, resulting in more con- strained and confident estimates of paleodepths than with a single taxon. 3. Neogene environmental histories among the three basins differed. The Panama Canal Basin was a shallow, Middle Miocene, Caribbean basin until deep- ening about 6 Ma caused an inflow of deep, Pacific water. The Bocas del Toro and Limon basins were alike in their back-arc tectonic setting, histories of up- lift, and sequence of sediments: bathyal Miocene, ne- ritic Pliocene and lower Pleistocene coral reefs. The two basins differed in sediment source and coeval, iso- bathyal microfaunas. REFERENCES CITED Blacut, G., and Kleinpell, R.M. 1969. A stratigraphic sequence of benthonic smaller foraminif- era from the La Boca Formation, Panama Canal Zone. Contributions from the Cushman Foundation for Fora- miniferal Research, vol. 20, pt. 1, pp. 1-22. Cassell, D.T., and Sen Gupta, B.K. 1989a. Foraminiferal stratigraphy and paleoenvironments of the Tertiary Uscari Formation, Limon Basin, Costa Rica. Journal of Foraminiferal Research, vol. 19, no. 1, pp. 52— Tk 1989b. Pliocene foraminifera and environments, Limon Basin of Costa Rica. Journal of Paleontology, vol. 63, pp. 146— 157. Collins, L.S. 1993. Neogene paleoenvironments of the Bocas del Toro Basin, Panama. Journal of Paleontology, vol. 67, no. 5, pp. 699— 710. Collins, L.S., Budd, A.F., and Coates, A.G. 1996b. Earliest evolution associated with closure of the Tropical American Seaway. Proceedings of the National Academy of Sciences, vol. 93, pp. 6069-6072. Collins, L.S., Coates, A.G., Berggren, W.A., Aubry, M.-P., and Zhang, J. 1996a. The late Miocene Panama isthmian strait. Geology, vol. 24, no. 8, pp. 687-690. Collins, L.S., Coates, A.G., Jackson, J.B.C., and Obando, J.A. 1995. Timing and rates of emergence of the Limon and Bocas del Toro Basins: Caribbean effects of Cocos Ridge sub- duction? in Geologic and tectonic development of the Ca- ribbean plate boundary in southern Central America. Geo- logical Society of America Special Paper, no. 295. P. Mann, ed., Geological Society of America, Boulder, Col- orado, pp. 263-289. de Boer, J.Z., Defant, M.J., Stewart, R.H., Restrepo, J.F., Clark, L.F., and Ramirez, A.H. 1988. Quaternary calc-alkaline volcanism and implications for the plate tectonic framework. Journal of South American Earth Sciences, vol. 1, pp. 275-293. Mann, P., and Corrigan, J. 1990. Model for late Neogene deformation in Panama. Geology, vol. 18, pp. 558-562. Nolf, D., and Brzobohaty, R. 1992. Fish otoliths as paleobathymetric indicators. Paleontologia i Evolucio, vol. 24—25 (1992), pp. 255-264. Sheehan, C.A., Penfield, G.T., and Morales, E. 1990. Costa Rica geologic basins lure wildcatters. Oil and Gas Journal, vol. 88, pp. 73-79. Stewart, R.H., and Stewart, J.L. 1980. Geologic map of the Panama Canal and vicinity, Republic of Panama. U. S. Geological Survey Miscellaneous In- vestigations Series, map 1—1232. Taylor, G.D. 1973. Preliminary report on the stratigraphy of Limon, Costa Rica. Publicaciones Geologicas del ICAITI, no. IV, pp. 161-165. 1975. The geology of the Limon area of Costa Rica. unpub- lished Ph. D. thesis, Louisiana State University, Baton Rouge, 114 pp. Terranes, J.L., Geary, D.H., and Bemis, B.E. 1996. The oxygen isotopic record of seasonality in Neogene bi- valves from the Central American isthmus. in Evolution and environment in tropical America. J.B.C. Jackson, A.F Budd, and A.G. Coates, eds., University of Chicago Press, Chicago, pp. 105-129. a = B x \¢) 2 Fy o s y te pS ‘Dia > @o= 7 ” = = &- : C. pemura) is Cn} we" a - J Ge = “ys Os ay = Aye is nie — Mis net © pony = me ve Mey ot elise | ot ante ne maa 2ar hk 7 < PP ap ai Lae porary Las & So4by hig$2 ~~ Senet inaty ; ‘iii q f ius oo naaldaad re dita = iit'§ As di aS io A ae _ a & 14 oo Coney yor 1 om. ier banks Vanrmyin@ rete a iat fst wh eee maites ieee aun ep ojo ork phe On y im. 8] PART 2 PALEOBIOTIC SURVEY er, 9 lS Oe“ eS) Le oy oi < CHAPTER 5 THE MIOCENE TO RECENT DIVERSITY OF CARIBBEAN BENTHIC FORAMINIFERA FROM THE CENTRAL AMERICAN ISTHMUS LAUREL S. COLLINS Department of Earth Sciences Florida International University Miami, Florida 33199, U.S.A. INTRODUCTION The diversity of modern benthic foraminifera in tropical America is lower in the Eastern Pacific than in the Caribbean. Culver and Buzas (1982, 1986, 1987) used all published literature on the coastal wa- ters of North America to enumerate synonymized spe- cies: 1189 in the Caribbean, 798 from the Pacific coast between California and Baja, and 377 from the Mex- ican to Central American Pacific coast. Whereas the latter compilation by itself may suffer from inadequate sampling (Culver and Buzas, 1987), when it is com- bined with the Pacific California to Baja data set it is robust, as is the Caribbean one (Buzas and Culver, 1991). Diversity indices that take into account the rel- ative abundance as well as the number of these same species further indicate that benthic foraminiferal di- versity in the modern Caribbean is almost twice as high as in the tropical Eastern Pacific from Central America to California (Buzas and Culver, 1991). This pattern contrasts with those of most other invertebrate species: gastropod diversity is not demonstrably dif- ferent in the Recent tropical Western Atlantic than in the Recent tropical Eastern Pacific (Allmon et al., 1993), crustacean diversity does not appear apprecia- bly different (Jones and Hasson, 1985), and echinoid diversity is approximately equal (Chesher, 1972). Only reef corals seem to have a similar Eastern Pacific-Ca- ribbean contrast in the diversity of species: about 20 living in the East Pacific (Glynn, 1972) versus about 50 in the Caribbean (Johnson et al., 1995), although species richness is substantially lower than for benthic foraminifera. The historical reasons for differences in tropical At- lantic vs. Eastern Pacific diversity hinge on the evo- lutionary and biogeographic divergence of taxa caused by the closure of the Caribbean-Pacific seaway. Sur- face waters of the Caribbean and Pacific mixed before the isthmus emerged to constrict and close the seaway, so that assemblages were more similar and diversity lower before seaway closure than they are today. This is because the contrast in benthic foraminiferal diver- sity between the Caribbean and tropical Eastern Pacific almost certainly reflects oceanic conditions rather than the influence of biotic interactions (Buzas and Culver, 1998). Constriction of the seaway was advanced in Late Miocene time (Mikolajewicz et al., 1993; Collins et al., 1996a), and complete closure occurred approx- imately 3.5 Ma (Keigwin, 1982). Caribbean effects of the cutoff of Pacific waters should have been most pronounced closest to the seaway. This study uses latest Middle Miocene to Recent benthic foraminifera from upper bathyal to inner ne- ritic deposits of Caribbean Panama and Costa Rica to investigate the pattern of Caribbean diversity through the time of seaway constriction to closure. “Diversity” as used in this chapter refers to both the number of species and the distribution of individuals among spe- cies, i.e., the relative abundances of the species (Peet, 1974). A fossil assemblage is comparable to a live + dead Recent assemblage. As the number of individuals in a community is counted, the number of species commonly increases. Three hundred ninety-five species (Appendix 1), in- cluding undescribed ones (in some cases, preservation was too poor to distinguish between the two), were recognized in the 130 Miocene to Recent sediment samples of this study. There are probably another 50 species that I have not recognized in these assemblages because of their rarity and/or lack of good preservation of diagnostic features. All of the fossil species, from deposits of the Panama Canal Basin, the Bocas del Toro Basin, Panama, and the Limon Basin, Costa Rica (Coates, this volume), have been enumerated and form the basis of previous paleoenvironmental interpreta- tions (Collins, 1993; Collins et al., 1995, 1996b; Col- lins in Jackson et al., this volume). Proportions of all common fossil species, defined herein as those repre- sented by >1 specimen in a split of a sample collected by the Panama Paleontology Project (PPP), are pre- sented in electronic form at the PPP internet site: http: //www.fiu.edu/~collinsl/. Abundances of the Recent taxa were listed by Havach and Collins (1997). 92 BULLETIN 357 9°30' acces : Je: CARIBBEAN SEA “ta 2000 m...” CARIBBEAN PACIFIC OCEAN 9°00' .., Escudo de wre Veraguas |... “Fig Golfo de los Mosquitos e Chiriqut + 22 Comat aoc hip station number Chiriqui eologic section Grande 8°55! xe 5, 82°15' 82°10' 82°00' 81°50' 81°40' 81°30' 81°20! Text-figure 1.—Ship stations (+) for Recent foraminiferal assemblages, and land-based geologic sections (MM) for fossil assemblages of the province of Bocas del Toro, northwestern Panama. The diversity of a taxon such as benthic foramini- fera or mollusks typically varies considerably across neritic depths, which effect or bias should be consid- ered when examining biodiversity at this study’s rel- atively small, regional scale. The ecological relation- ships of Recent foraminifera from 37 ship stations (Text-fig. 1), covering a wide range of habitats in the archipelago of Bocas del Toro, northwestern Panama (Havach and Collins, 1997), were applied to interpret- ing paleoenvironments of Costa Rica and Panama. In this study, the species proportions and diversity of the Recent and fossil assemblages are analyzed quantita- tively and compared for similar environments. Because of generally strong associations of species with partic- ular marine environments, the relatively long geologic ranges of species in this study (mean first occurrence of 21.4 Ma, similar to that in the U. S. Atlantic coastal plain calculated by Buzas and Culver, 1998), and the few Pliocene to Recent extinctions of Caribbean taxa common to these deposits (Collins, 1996), the expec- tation was that assemblages from the same environ- ment, rather than age, would be most similar. Previous, smaller scaled studies of Miocene to Pliocene taxa of the Bocas del Toro and Limon basins (Collins ef al., 1995), and of Miocene taxa within the Panama Canal Basin (Collins et al., 1996b), showed that paleoenvi- ronment had a stronger influence than age on assem- blage composition. This study combines all the basins and adds Pleistocene to Recent data in an evaluation of 130 late Middle Miocene to Recent Caribbean as- semblages. The main purpose of this chapter is to evaluate ben- thic foraminiferal diversity of the past 11 m.y. within similar marine environments of Caribbean Panama and Costa Rica. The chapter is divided into three parts: (1) a classification of the assemblages of fossil and Recent benthic foraminifera, using cluster analysis, to evaluate environmental and chronologic patterns. (2) an iden- tification of the differences in diversity between Re- cent habitats/environments, using analyses of variance (ANOVAs), to apply as a yardstick in the paleobiodi- versity study. (3) a study of diversity over time, taking into account the expected diversity differences be- tween environments, to evaluate trends in relation to the Neogene closure of the Caribbean-Pacific seaway. BENTHIC FORAMINIFERA: COLLINS 93 ACKNOWLEDGMENTS Many people contributed to the collection, age dat- ing, and preparation of these samples (Introduction, this volume). I especially thank Anthony Coates and Tim Collins, who first invited me to join them in the field, and Jeremy Jackson for sponsoring a Smithson- ian Tropical Research Institute (STRI) postdoctoral fellowship to begin research on PPP material. STRI has continued to support related research with field vehicles and research vessels, collecting logistics, visas and permits. I am grateful to Martin Buzas, C. Wylie Poag and Barun Sen Gupta for insightful comments on an earlier manuscript. This study was supported primarily by National Science Foundation grants BSR90-06523, RII-9002977, and DEB-9300905 to Collins and others. The main repository for slides con- taining these foraminiferal faunas is the U. S. National Museum of Natural History. This chapter is contribu- tion number 12 to The Program in Tropical Biology at Florida International University. METHODS FIELD AND LABORATORY METHODS Members of the Panama Paleontology Project (PPP) collected the geologic samples between 1986 and 1994. The PPP assigned each sampling site a PPP number (Appendix A, this volume), which is the main reference for all sediment samples and foraminiferal assemblages. I selected for preparation 50-g portions of ninety-three sediment samples (Table 1) from geo- logic sections (Appendix B, this volume) with biochro- nologic ages restricted to the shortest intervals of time. For all cases except the Gatun Formation, at least one sample within each geologic section was biochrono- logically dated. Gatun samples are from many small exposures measured over a wide area at the Caribbean end of the Panama Canal (Appendix A, Maps 1-2). For lower, middle and upper parts of the Gatun For- mation, Panama Canal Basin, biochronologic indica- tors were either poorly preserved (calcareous nanno- plankton) or sparse (planktic foraminifera) due to the shallowness of the paleoenvironment. Therefore, I transferred ages from dated portions to undated por- tions of the sections by correlating physical strati- graphic relationships among the sections. For comparison with fossil material, I collected Re- cent marine sediment samples of 40 ml each from 3.7 to 240 m water depth in the Bocas del Toro archipel- ago (Text-fig. 1) using STRI’s R/V Benjamin. Ship sta- tions were chosen to maximize the estimated range of habitats in the area and to include habitats similar to those interpreted for the geologic samples. The cate- gories of habitats covered were: lagoon, the main channel (Tiger Channel) between the lagoon and open ocean, nearshore, coral reef, open-ocean middle nerit- ic, Outer neritic, and upper bathyal. Havach and Col- lins (1997) described sampling and habitats in detail. Fossil and Recent sediments were washed through a 63-y.m sieve, which retains all adults and identifiable juveniles of benthic foraminifera. For samples in which foraminifera were exceedingly sparse (e.g., the Bruno Bluff section, Shark Hole Point Formation), heavy-liquid separations of heavier grains from lighter foraminiferal tests were performed with sodium poly- tungstate. Each sample was split to yield > 400 ben- thic foraminifera, except for the few cases in which all washed residue yielded < 400 specimens. Four hun- dred is a sufficient number of individuals to accurately represent the proportional abundances of the species in the samples with a margin of error (confidence lim- its) of no more than + 0.05 (Buzas, 1990). Specimens were sorted into species on cardboard micropaleonto- logical slides, and species with >1 individual per sample were identified using comparative collections of specimens and the literature on Caribbean, Gulf of Mexico, and East Pacific Neogene to Recent benthic foraminifera. These methods for sample preparation are described in more detail by Collins (1993) and Havach and Collins (1997). METHODS OF NUMERICAL ANALYSES The relationships among the assemblages of 395 recognized species were summarized by a Q-mode cluster analysis of relative abundances, which joins as- semblages by their similarity of species proportions. I present results from Ward’s method (Systat, 1998), in which the sum of the squares of the Euclidean dis- tances was calculated between two clusters of assem- blages added up over all the variables (taxa), and at each generation, the within-cluster sum of squares was minimized over all partitions obtainable by merging two clusters from the previous generation. Relative abundance data were transformed using the relation- ship 2 arcsin VP, where P = the proportion of a spe- cies in an assemblage. Data were then standardized by the calculation of z values to approximate a multivar- iate normal distribution of the data matrix, which gives equal weight to all species in the calculation of the distances between assemblages. Cluster analysis, an exploratory, classificatory meth- od, was employed instead of more sophisticated, hy- pothesis-testing methods, such as discriminant analy- sis, because cluster analysis is one of the few methods in which the number of variables (species) can exceed the number of observations (assemblages). In tropical waters, the diversity of benthic foraminifera in a single cubic centimeter of washed, normal marine sediment 94 BULLETIN 357 Table 1.—Paleobathymetry (range midpoints), ages (range midpoints) and diversity (a, Fisher’s alpha) of benthic foraminiferal assemblages. The mean (1), standard deviation (6) and standard error of the mean (6,,), which is the average deviation of sample means from the expected value, are calculated for a in each stratigraphic unit or Recent habitat. Fossil assemblages are from the Panama Paleontology Project (PPP) and Recent assemblages are from Bocas del Toro (BDT) sites of Text-figure 1. * = environmental settings which are larger scaled than Recent habitats and used in Tables 2—6 and Text-figures 3—6. Stratigraphic unit Age Depth PPP or or Recent habitat (Ma) (m) BDT No. a pe rc) 5, Fossil, PPP Collections Lower Gatun Fm. 11.6 25 6 11 13.0 T2, 2:5 *Open Inner Neritic 14 5 15 9 16 13 35 13 1037 29 1038 15 1040 9 Middle Gatun Fm. 9.0 25 18 27 20.3 5.8 Sh} *Open Inner Neritic 19 17 34 17 Upper Gatun Fm. 9.0 25 17 12 19.7 Teall 2S) *Open Inner Neritic 20 23 Di 15 28 18 160 32 1660 18 Chagres Fm., Rio Indio facies Ts 65 24 21 25.7 4.5 2.6 *Middle Neritic 26 30 1645 26 Chagres Fm. 6.1 350 1088 22 19.5 Sb! 9) *Upper Bathyal 1097 15 1173 1 1174 23 Nancy Point Fm. 6.0 400 407 21 23.4 43 1.9 *Upper Bathyal 408 19 410 30 411 25 412 22 Shark Hole Point Fm., 3.6 175 376 25 22.5 1.9 1,0 Bruno Bluff section 377 21 *Outer Neritic 378 21 379 23 Cayo Agua Fm., NE section 4.8 60 59 19 2ileS) 2.9 1.4 *Middle Neritic 60 22 61 19 62 25 Cayo Agua Fm., NW section 3.6 30 57 23 17.8 3)-7/ 1.5 *Open Inner Neritic 63 17 195 21 196 13 197 18 198 15 Cayo Solarte section 3.6 50 68 21 *Middle Neritic Escudo de Veraguas Fm., SE 3.6 125 168 41 43.3 4.0 Pins *Mixed M. Neritic & O. Neritic/U. 169 48 Bathyal 170 41 BENTHIC FORAMINIFERA: COLLINS Table 1.—Continued. Stratigraphic unit Age Depth PPP or or Recent habitat (Ma) (m) BDT No. a pL 3 5 Escudo de Veraguas Fm., NE 3.6 125 365 34 39.2 5.4 2.4 *Mixed M. Neritic & O. Neritic/U. 366 38 Bathyal 367 48 368 40 369 36 Escudo de Veraguas Fm., NW 1.8 125 358 48 39.2 7.6 3.4 *Mixed M. Neritic & O. Neritic/U. 360 39 Bathyal 36] 45 362 35 363 29 Swan Cay section 1.4 100 1181 37 41.0 Sai/ 4.0 *Coral Reef 1789 45 Uscari Fm., Carbon Dos section SS) 300 726 24 28.6 5.3 2.4 *Upper Bathyal 727 27 728 32 729 36 730 24 Uscari Fm., Rio Sandbox section S)s) 200 735 30 2531. 4.5 2.6 *Upper Bathyal 736 26 WSy/ 21 Lower Rio Banano Fm. 3.6 30 679 43 32.4 8.4 3.8 *Open Inner Neritic 680 37 681 21 682 28 683 33 Upper Rio Banano Fm. Pr) 20 668 31 29.3 4.3 1.8 *Open Inner Neritic 669 28 670 29 671 26 677 37 678 25 Moin Fm., type section 1.8 200 647 41 32.4 10.3 3.0 *Mixed M. Neritic & O. Neritic/U. 648 21 Bathyal 649 27 650 38 651 25 652 57 653 25 654 40 655 32 656 32 657 22 658 29 Moin Fm., Lomas del Mar section 1.8 75 638 60 Ses) 18.8 9.4 *Coral Reef 640 28 641 17 642 44 Recent, BDT Collections Recent, shallow lagoon 0 18 21A 23 19.8 8.3 4.2 *Lagoon 12 22A 14 14 24A 12 14 25A 30 Recent, deep lagoon 0 21 4C 24 27.8 3.9 1.9 *Lagoon 34 SA 28 35 23A 26 25 26A 33 96 BULLETIN 357 Table 1.—Continued. Stratigraphic unit Age Depth or Recent habitat (Ma) (m) Recent, channel 0 34 *Middle Neritic 33 27 70 45 Recent, nearshore 0 24 *Open Inner Neritic 6 15 11 Recent, inner middle neritic 0 39 *Middle Neritic 56 Recent, outer middle neritic 0 73 *Middle Neritic 80 Recent, coral reef 0) 18 *Coral Reef 35 64 4 49 82 Recent, outer neritic 0 120 *Outer Neritic 168 120 180 Recent, upper bathyal 0 230 *Upper Bathyal 235 Recent, outer neritic/upper bathyal 0 113 mixed with middle neritic 164 *Mixed M. Neritic & O. Neritic/U. 203 Bathyal 240 is commonly > 100. Reducing the number of species to less than the number of assemblages in a data set that covers such a large range of time and space would be too great a loss of information to assess relation- ships among the assemblages. Conversely, increasing the number of assemblages to more than the number of species, even if few additional ones were recovered, is impracticable. The diversity of foraminiferal assemblages was measured for Recent habitats and for sections of geo- logic formations with Fisher’s alpha (a), an index which assumes that the proportions of species within assemblages are distributed in a log series. This dis- tribution has successfully predicted the frequency of benthic foraminiferal species occurrences from the PPP or BDT No. a vy ro) 5, 1B 37 37.0 er 5.0 2A 33 3B 44 27A 50 28A 21 8A 29 36.5 5.4 207. 15B 36 16A 5 17A 40 6C 22 24.5 BFS) DES) 7B Dali 29A 50 48.0 2.8 2.0 33A 46 9B 41 36.5 11.2 4.6 10B 30 ie 39 12B 46 13A 46 14C 17 30A 40 37.0 DES 1.2 Se 35 34C 38 35A 35 32A 32 BIS 0.7 0.5 36A 33 37A 55 67.3 IPI 6.1 38A 65 39A 65 40B 84 North American Atlantic coast, Gulf of Mexico and the Caribbean (Buzas et al., 1982). Calculations for a use the number of taxa and the number of specimens; iterative solutions for a for combinations of these pa- rameters were obtained from Hayek and Buzas (1997, Appendix 4). Analyses of variance (ANOVAs) were performed on diversity data and relative abundances and propor- tions of carbonate-associated taxa to test relationships among mean values for different environments or time intervals, entities that were defined a priori. Analysis of variance tests whether the variances of the means among the groups are greater than expected on the basis of the variances within the groups. An F-test quantifies the significance of the overall differences = Text-figure 2.—Cluster analysis (Wards algorithm) of assemblages (rows) grouped by similarity of standardized species abundances. Fossil sites are referenced by PPP numbers (‘P’ prefix). Recent sites are referenced by BDT numbers (‘B’ prefix). Section numbers in left column refer to the stratigraphic sections of Appendix B (this volume). Assemblages cluster primarily by formation sections (shaded), which cluster by epoch (Miocene, Pliocene to Pleistocene, or Recent) before depositional environment. BENTHIC FORAMINIFERA: COLLINS 97 FORMATIONS & SITE NO. & EUCLIDEAN DISTANCES SECTIONNUMBERS ~~ SECTION ,0 1 2 3 4 5 6 Pp Chagres Fm., type (3, 4) 61{98§&hae B40 82tt Panama ~ ae (Le Canal Basin main sections P P Pp P. Gatun Fm., Be a Pp (1, 2, 3, 4, 6) Z ARENA) ge Chagres Fm., Rio Indio (5) Uscari Fm., Carbon Dos section (28) Uscari Fm., : Rio Sandbox section (27) Nancy Point Fm. (12) Ri RI . MIOCENE 6 Nane FOSSIL VUVVUVUUVDUVUUVUDUUDUD Pr Rio Banano Fm. (29) poet Rant ie PI Moin Fee ti PLIOCENE - E. ee (37) Pees Mon PLEISTOCENE + Swan Cay (25) pea! on Shark Hole Point Fm., p37/ Brun Bruno Bluff section (12) Pol CAN + Cayo Agua Fm., P63 NE section (19) bare Bun ple? 365 EscN P369 EscN Escudo de Veraguas Fm., BI E all sections (10, 11) Paso Bey Paes ESN P358 EscN P366 N P196 C. Agua Fm., ae AN NW section (16) ae Pep CSol Moin Fm., peao toma Lomas del Mar section (36) 5824 Wor eS B28A ged Recent an on deep lagoon + channel ee ead B21A lags Recent shallow lagoon an ie shallow coastal Recent nearshore BIZA nrsh Bette Recent coral reef BiaB feet RECENT 14C reef 11C reef B36A upba open ocean Recent outer neritic B33 Oub. P & upper bathyal Bic out : yc BEC Gl Recent inner middle neritic B78 mich Recent si B39A tran outer middle neritic BSA role) & transported sediments 3234 mido 98 BULLETIN 357 among the groups to test whether the group means are different because of differences in the underlying pop- ulation means. In this study, if the probability that the means of the groups were drawn from the same pop- ulation is low (P < 0.05), the groups of environments or time intervals were considered significantly differ- ent. SIMILARITY OF LATE MIOCENE TO RECENT ASSEMBLAGES Age (epoch), rather than environment, has a stron- ger influence on the similarity of the 130 Miocene to Recent benthic foraminiferal assemblages. A cluster analysis (Text-fig. 2) shows the principal division on the right between fossil and Recent assemblages, and secondarily within fossil assemblages between the Miocene and Pliocene to Early Pleistocene (~1.4 Ma). These results differ from those of several previous, smaller scale cluster analyses in which isthmian ben- thic foraminiferal assemblages from different ages but similar environments were the most similar. Cluster analyses were previously performed on assemblages from the Late Miocene to Late Pliocene of the Bocas del Toro Basin (32 samples; Collins, 1993), the Late Miocene to Late Pliocene of the Bocas del Toro and Limon basins (67 samples; Collins et al., 1995), and the late Middle to Late Miocene of the Panama Canal Basin (9 samples; Collins et al., 1996b). In these ear- lier analyses, clusters were joined by paleobathymetry rather than age. However, in the present study, ages span a longer time interval (late Middle Miocene to Recent) but not a greater bathymetric range (although the Recent data include a new, distinctive lagoonal en- vironment). Extending the ages analyzed downward to the late Middle Miocene (Lower Gatun Formation) and upward to the Recent (of Bocas del Toro) clearly increases the disparity among assemblages more than adding additional depositional environments. Assemblages cluster by age despite an overlap in comparable modern and fossil environments, based on ecologic relationships of environment-diagnostic, ex- tant taxa (Collins ef al., 1995, 1996b). For example, environments of the Late Miocene Gatun Formation (~25 m deep), the Early Pliocene Cayo Agua For- mation (~20—40 m deep), the Late Pliocene Rio Ban- ano Formation (upper part ~20 m and lower part ~20—40 m deep), and the modern nearshore, middle neritic, and lagoon habitats are all characterized by similar depth ranges with many of the same species, but their assemblages cluster according to the three ages. Similarly, outer neritic to uppermost bathyal sed- iments that are mixed with shallower, reefal and sili- ciclastic-associated species, such as those of the Late Pliocene type Moin and Escudo de Veraguas forma- tions and Recent ship stations 38 to 40, are separated on the basis of age. Coral-reef-associated species of the latest Pliocene Lomas del Mar (paleodepth ~75 m) and Early Pleistocene Swan Cay (paleodepth ~ 100 m) sections are within the same age cluster but not grouped most closely. The four Early Pleistocene as- semblages are more similar to those of the Late Plio- cene than Recent, despite the wide range of water depths sampled off living coral reefs of the island of Escudo de Veraguas (Text-fig. 1). In the Panama Canal Basin, most Gatun Formation species are inner neritic and of Caribbean affiliation, whereas the Chagres For- mation species are of Pacific affiliation and upper bathyal (Collins et al., 1996b), two very different pa- leoenvironments. However, they cluster together when compared with Pliocene to Recent assemblages. Assemblages from the same formation sections or Recent habitats form coherent groups that represent discrete subenvironments. This cluster analysis sup- ports previous results that show the closest relation- ships (Text-fig. 2, on the left) to be within formation sections for fossil material (Collins, 1993; Collins et al., 1995, 1996b) and within habitats or groups of sim- ilar habitats for Recent material (Havach and Collins, 1997). Only 12% of the assemblages were classified outside of their sections or habitats. The cluster dendrogram (Text-fig. 2) was produced with the Wards algorithm, only one of several methods available. Other explored methods (average, single, centroid, median and complete linkage) yielded few clusters and “‘stringy’’ dendrograms, mostly joining assemblages one at a time to preceding ones. Similar- ity is more interpretable from clusters rather than strings, so Wards method, which is among the best available (Milligan, 1980), was preferred. In general, the few groupings of some Recent and fossil assem- blages produced by the other methods, such as a few lagoon to middle neritic Recent and Rio Banano For- mation assemblages (complete linkage algorithm), of- fer little insight into paleoenvironmental or other in- terpretations. How do Miocene, Pliocene to Early Pleistocene, and Recent assemblages differ in composition? Most of the Miocene assemblages, from the Uscari, Nancy Point and Chagres formations, are from deeper (bathyal) bio- facies than the Recent samples (~240 m deep), and most of the common Chagres species are Pacific. (Col- lins et al., 1996b, explain this anomalous Caribbean occurrence with a breached isthmus.) The Miocene type Gatun Formation (inner neritic) and the Rio Indio facies (middle neritic) of the Chagres Formation most- ly contain species associated with siliciclastic sedi- ments with low abundances of carbonate-associated species, unlike the other neritic assemblages. BENTHIC FORAMINIFERA: COLLINS 99 MIDDLE NERITIC LAGOON OPEN INNER NERITIC ZPD —_——— OS oa Oo A Relative Abundance of Carbonate-Associated Taxa oO = Nh wo CORAL OUTER MIXED M. NERITIC & REEF NERITIC O. NERITIC/U. BATHYAL UPPER BATHYAL oo Recent Pleistocene -Pliocene Miocene See: @ Text-figure 3.—Relative abundances of taxa primarily associated with carbonate shoals and reefs, plotted for the 130 assemblages. Assem- blages are grouped by sections of formations (fossil) or marine habitat (Recent), as in the cluster analysis, then ordered by the marine environments across the top. Within comparable environments, Recent assemblages include the most carbonate-associated taxa, Pliocene to Early Pleistocene the next highest, and Miocene the least (see Table 2 for means.) Table 2.—Mean relative abundances of taxa that are generally associated with carbonate facies (Miliolina and larger foraminifera) for the groupings by age and environment of Text-figure 3. Within environments, values increase over time. Pliocene to Recent Pleistocene Miocene Lagoon 3.0 Inner Neritic 47 Dal 0.3 Middle Neritic 3.0 1.1 0.1 Coral Reef SiS} 1.6 Outer Neritic 2 0.2 Mixed M. Neritic & O. Neritic/U. Bathyal 3.0 0.9 Upper Bathyal 0.6 0.06 An important age-related trend in the composition of assemblages is an increase in individuals and spe- cies associated with carbonate shoals and coral reefs. These carbonate-associated species are primarily of the suborder Miliolina (those with calcareous, imperforate tests) and, to a much lesser extent, larger foraminifera (those having morphologically complex, calcareous, perforate chambers and sizes larger than most benthic foraminifera). Their mean relative abundance (Text- fig. 3, Table 2) and proportions of total taxa at the neritic to bathyal depths show significant increases within different environments over the three age inter- vals (Table 3, ANOVAs). Of 98 Recent species that are not represented as fossils, about one-fourth are Miliolina and larger foraminifera, mostly the former. If 39 rare species (they occur in only one sample) are excluded to reduce potential sampling biases, and 3 100 BULLETIN 357 Table 3.—ANOVA tables, mean relative abundances (left column) and proportions (right column) of Miliolina and larger foraminifera (Text- fig. 3). All ANOVAs indicate overall significant differences among the three age categories (Miocene, Pliocene to Early Pleistocene, and Recent) within different environments, except for proportions of taxa in the reef environment (P = 0.0563). RELATIVE ABUNDANCE PROPORTION OF SPECIES Source of Sum of Mean Source of Sum of Mean variability df squares square F p(F) variability df squares square Ie) p(F) OPEN INNER NERITIC Age 2 66.61 33.30 49.53 0.0000 Age 2 7264.23 3632.11 47.23 0.0000 Error 34 22.86 0.67 Error 35 2691.58 76.90 MIDDLE NERITIC Age 2 23.63 11.81 25.89 0.0000 Age 2 4167.91 2083.96 48.95 0.0000 Error 14 6.39 0.46 Error 14 596.09 42.58 CORAL REEF Age 1 9.01 9.01 31.26 0.0002 Age 1 533.33 533.33 4.66 0.0563 Error 10 2.88 0.29 Error 10 1145.33 114.53 OUTER NERITIC Age ] 2.00 2.00 13.41 0.0106 Age 1 840.50 840.50 160.1 0.0000 Error 6 0.90 0.15 Error 6 31.50 515) Mrxep MippLe Neritic & OUTER NERITIC/UPPER BATHYAL Age 1 15.79 15.79 70.13 0.0000 Age 1 1365.55 1365.55 737) 0.0003 Error 27 6.08 0.23 Error 27 2123.00 78.63 UpprER BATHYAL Age ] 0.43 0.43 50.21 0.0000 Age I 215.37 2N5'37, 75.13 0.0000 Error 17 0.15 0.01 Error 17 48.74 2.87 species are excluded because they are fragile (delicate agglutinated or thin-walled calcareous), to reduce po- tential preservational biases, about half (30) of the re- maining 56 species are in the Miliolina or larger fo- raminifera. This is an approximation, as not all of the Miliolina are primarily carbonate-associated (e.g., Sig- moilina tenuis is in most assemblages) and some hya- line foraminifera (e.g., Neoeponides repandus) or ag- glutinated species (e.g., Bigenerina irregularis) that are carbonate-associated are not included in this count. However, the relatively large proportion of Miliolina plus larger foraminifera that appears in the Recent as- semblages suggests that carbonate-associated taxa have increased substantially in Caribbean isthmian waters during Pleistocene to Recent time. DIVERSITY IN RECENT HABITATS OF BOCAS DEL TORO Differences in diversity due to differences in envi- ronments should be taken into account when analyzing changes in diversity through time if environments have also changed. In this study, paleoenvironments of the Caribbean side of the Central American isthmus have varied in paleobathymetry from upper bathyal to inner neritic. They also vary from predominantly siliciclastic to carbonate sediments produced in situ by coral reefs (Collins et al., 1995, 1996b). Based on general bathy- metric subdivisions, as well as the results of a previous study (Havach and Collins, 1997), the Recent assem- blages are assigned a priori to the following seven environments: lagoon, open-ocean inner neritic, mid- dle neritic, coral reef, outer neritic, mixed middle to outer neritic/uppermost bathyal, and upper bathyal. These divisions were also distinguished by the pa- leoenvironmental studies based on benthic foramini- fera. The ‘‘mixed ”’ assemblages, i.e., those that con- tain many taxa transported from middle neritic depths, have artificially high values of diversity because they combine shallower, middle neritic and deeper, outer neritic or uppermost bathyal (~240 m deep) species. The interpretation of mixed assemblages is also sup- ported by clearly middle neritic values of stable iso- topes of oxygen from the benthic foraminiferal tests (Havach and Collins, 1997). The isotopic signals do not reflect a ‘“‘mixed” signal because the species ana- lyzed, Cibicides pachyderma, lives predominantly at middle neritic depths. Analyses of variance were performed to identify sig- nificant differences in diversity among the seven bathy- metric divisions. The first ANOVA (Table 4, top) in- dicates an overall, significant difference in diversity among environments (P < 0.00005), but dropping out the anomalously high mixed middle to outer neritic/ uppermost bathyal group (with an almost doubled ap- parent biodiversity) reduces differences among environ- ment means to insignificant levels (Table 4, bottom). BENTHIC FORAMINIFERA: COLLINS 101 Table 4—ANOVA tables, mean diversity (a) of Recent environ- ments of Text-figure 4. The top table indicates significant differences in diversity among environments (lagoon, open-ocean inner neritic, middle neritic, reef, outer neritic, Outer neritic/uppermost bathyal with transported sediments, and uppermost bathyal). However, this is due to the artificially high values for the assemblages that include transported middle neritic sediments. If these are excluded (bottom table), average diversity among the other environments is not sig- nificantly different. All 7 Environments Source of Sum of Mean variability df squares square F p(F) Environment 6 5108.64 851.44 9.74 0.0000 Error 30 2623.25 87.44 Excluding Mixed M. Neritic & O. Neritic/U. Bathyal Source of Sum of Mean variability df squares square In p(F) Environment 5 990.05 198.01 2.45 0.0592 Error 27 2182.50 80.83 Excluding the ‘“‘mixed” group, the main differences are for the lagoonal assemblages, which have the lowest diversity, and the upper bathyal assemblages, which have a lower diversity than open-ocean neritic assem- blages (Text-fig. 4). Not only are the differences among the open neritic groups insignificant, including open in- ner neritic, middle neritic, reef, and outer neritic, but the mean as for these environments are also quite sim- ilar, all falling between 36 and 37. The similarity of Open-ocean neritic diversities is surprising because pre- vious studies, based in areas of cooler waters, show that benthic foraminiferal species diversity increases from the shoreline to shelf edge (summarized by Sen Gupta, 1982), so that the pattern in this study may be a Carib- bean or warm-water phenomenon. In applying the re- sults to the fossil data below, the following should be noted: (1) A lagoonal facies is not represented in the paleoenvironments analyzed for this study. (2) The Open-ocean inner neritic, middle neritic, coral reef, and outer neritic environments are analyzed together as well as separately. (3) The upper bathyal assemblages are analyzed separately. LATE MIOCENE TO RECENT BIODIVERSITY On the basis of the similar diversity of benthic fo- raminifera among subdivisions of Recent, open-ocean neritic depths (open inner neritic, middle neritic, coral reef, and outer neritic), the diversities of fossil and Recent assemblages were analyzed together in an AN- OVA with age (Miocene, Pliocene to Early Pleisto- cene, or Recent) as the classification criterion. Twenty- five assemblages from the type section of the Moin and Escudo de Veraguas formations were excluded be- cause they contain greatly varying amounts of down- (Ee ALL RECENT ASSEMBLAGES iversity () ayeue D Text-figure 4.—For Recent environments, least squares means of diversity (a) and their standard deviation (error bars) predicted by the ANOVA of Table 4 (top). The data suggest that diversity does not differ significantly among environments of Bocas del Toro, and the most similar are open-ocean inner neritic, middle neritic, coral reef and outer neritic depths. The high value is an artefact of mixing assemblages from different environments and does not reflect true biodiversity. shelf to downslope transport of shallower taxa, so di- versity values are influenced primarily by sedimentary processes rather than true biodiversity. The environ- ments represented by the type Moin and Escudo de Veraguas formations are similar to the Recent “‘mixed assemblages”’ containing many shallow-water, coral- reef-associated species that mixed with typical outer neritic to upper bathyal species. In fact, the sections of the Moin Formation form a relict Pliocene relief. The type Moin section is located at the base of a steep hill composed largely of in-place hermatypic corals (Lomas del Mar section, Moin Formation). Both sec- tions are latest Pliocene, and many of the benthic fo- 102 BULLETIN 357 Table 5.—ANOVA table, mean diversity (a) of open-ocean, inner to outer neritic assemblages for Miocene, Pliocene to Early Pleis- tocene, and Recent ages (plotted in Text-fig. 5). Outer neritic assem- blages of the type Moin and Escudo de Veraguas formations have been excluded because the varying proportions of downshelf-trans- ported taxa result in unreliable estimates of biodiversity. Open Inner to Outer Neritic Source of Sum of Mean variability df squares square EB p(F) Age 2 3731.45 1865.73 21.52 0.0000 Error Ue? 6241.22 86.68 raminiferal species which I collected (1989) between fronds of Pliocene corals are also found at the base of the slope in the type Moin deposits. Overall, average neritic diversity for Miocene, Pli- ocene to Early Pleistocene, and Recent ages are sig- nificantly different (P < 0.00005; Table 5). Text-figure 5 shows a large increase from a mean of 18 in the Late Miocene (N = 20, which includes the open inner ne- ritic to outer middle neritic Gatun Formation and Rio Indio biofacies of the Chagres Formation), to a mean of 27 in the Pliocene to Early Pleistocene (N = 32, which includes the open inner neritic, middle neritic, coral reef and outer neritic environments of the Swan Cay Formation, Lomas del Mar section of the Moin Formation, Rio Banano Formation, Cayo Agua For- mation, and Cayo Solarte section), to a mean of 35 in the Recent (N = 23, which includes open inner neritic, middle neritic, coral reef, and outer neritic). The data indicate that the diversity of Caribbean benthic fora- minifera from neritic depths along the Central Amer- ican isthmus increased from Late Miocene to the Re- cent. Within-environment ANOVAs would be useful to examine patterns at a scale smaller than the entire open-ocean neritic interval (Table 5, Text-fig. 5). How- ever, data are too few for meaningful analysis of most of the environmental divisions, except perhaps for Open inner neritic and upper bathyal (Table 6, Text- fig. 6). Where statistically significant, within-environ- ment ANOVA results do support the pattern of in- creasing diversity (a) over time with no reversals, as follows: 1. Open Inner Neritic. For Ny =el/Npiccee= 17, Neecen = 4, age differences are significant and show increasing diversity with time (Table 6, top; Text-fig. 6, left). 2. Middle Neritic. Data are too few (Nyjiccene = 3; Npiiccene = 5+ Neecen. = 9) for convincing results (not shown), although overall they are significant (P = 0.0202). An ANOVA indicates that Miocene and Pli- ocene values of alpha are not significantly different (standard deviations from the least squares means Miocene OPEN OCEAN, INNER-OUTER NERITIC iversity (Q) D Text-figure 5—For Recent and fossil, open-ocean, inner to outer neritic environments (the four divisions are inner, middle, coral reef, outer), least squares means of diversity (a) and their standard error (bars) predicted by the ANOVA of Table 5. The data suggest that oOpen-ocean, neritic diversity in the isthmian region increased from the Late Miocene to Recent. Table 6—ANOVA tables, mean diversity (a) of open-ocean inner neritic and upper bathyal environments for Miocene, Pliocene to Early Pleistocene, and Recent ages (plotted in Text-fig. 6). These environments have sufficient data to test within-group differences per age. The data suggest that diversity has increased in these Ca- ribbean isthmian environments since the Miocene. Open Inner Neritic Source of Sum of Mean variability df squares square Ii p(P) Age P 1586.65 19332 13.33 0.0000 Error 35 2083.35 59.52 Upper Bathyal Source of Sum of Mean variability df squares square le) p(F) Age 1 117.07 117.07 4.32 0.0531 Error 17 460.62 27.10 BENTHIC FORAMINIFERA: COLLINS 103 44 40 open ocean, upper inner neritic bathyal — 33 33 = = 2 ® = QO 22 26 11 19 a a F F ca <0

, m ae - r eee 5 i ; : . rs é ae = Ja oun alee > 7 Pv" 9: nape to—ghor ; ~ é . ist Prves tiora Sa = we SPOR asa ter) NAA ‘ ' I oP 5 Shai eae ae @2senp © > bese” 1 een AS els crm ie An —- : 7 iy re ee rm - a a. i mc ureters] 7 Baan { pe. : a CaO wite=t. : en aay _ y CHAPTER 6 STRATIGRAPHIC DISTRIBUTION OF NEOGENE CARIBBEAN AZOOXANTHELLATE CORALS (SCLERACTINIA AND STYLASTERIDAE) STEPHEN D. CAIRNS Department of Invertebrate Zoology Smithsonian Institution Washington, D.C. 20560-0163, U.S.A. INTRODUCTION This paper presents occurrence data and analyses of the azooxanthellate Caribbean Scleractinia, and, to- gether with the paper of Budd ef al. (1994), provides a listing of all known Caribbean Scleractinia from the Miocene to Recent. Whereas the zooxanthellate coral compilation was based almost exclusively on previ- ously reported fossil faunas (Budd et al., 1994: Table 2), this paper is based not only on historical collections (Table 1), but also on extensive new material from the Pliocene of Panama and Costa Rica. Azooxanthellate corals are sometimes incorrectly re- ferred to as “‘deep-water’’, “‘solitary’’, or ‘‘ahermatyp- ic’ (non-reef) corals. Whereas a majority of azooxan- thellate corals do occur in water deeper than 200 m, the depth range of this ecological class of corals is intertidal to 6328 m. Because their distribution is not limited by the light requirement of algal symbionts (zooxanthellae), they not only occur below the eupho- tic zone, but also at temperatures of —1° to 29° and at latitudes ranging from the Arctic Circle to continental Antarctica (Cairns and Stanley, 1982). A majority of azooxanthellate coral genera are solitary in growth form, but one-third of the Recent genera are colonial, some colonies even forming extensive deep-water (to 1300 m) banks, and one azooxanthellate species, Tu- bastraea micranthus, forming shallow-water herma- typic reefs (Zibrowius, 1989). From the point of view of biodiversity, there is currently an equal number of Recent azooxanthellate and zooxanthellate scleractin- ian genera (i.e., 114 of each) and approximately 620 valid Recent azooxanthellate species, compared to a range of 640-833 valid Recent zooxanthellate scler- actinian species (Veron, 1995). Numbers of fossil spe- cies have not been tabulated. Therefore, although azooxanthellate corals are usually small and incon- spicuous, they are widespread in the marine environ- ment and have a biodiversity equal to their zooxan- thellate ecological counterparts. ACKNOWLEDGMENTS The new material on which this paper was based was collected by the Panama Paleontology Project, and was prepared by Yira Ventocilla (Smithsonian Tropical Research Institute (STRI), Panama) and Rene Pan- chaud (Naturhistorisches Museum, Basel, Switzer- land). Laurel Bybell and Harry Dowsett (United States Geological Survey) provided the biostratigraphic dat- ing of the PPP samples. I thank Harry Filkorn, Helmut Zibrowius, and Nancy Budd for thoughtful review of the manuscript. MATERIAL AND METHODS The new specimens mentioned above were collected during the first seven years of the Panama Paleontol- ogy Project (PPP) 1986—1992. Descriptions of the lith- ostratigraphy and biostratigraphic correlations of the PPP region were given by Coates et al. (1992), Collins (1993) and Collins et al. (1995), as well as being sum- marized in this volume (Coates). The PPP corals are listed in Table 2, the new species and stratigraphic range extensions having been reported by Cairns (1995). Absolute dates for many of the PPP collecting sites were derived from biostratigraphic dating using planktic foraminifera, calcareous nannoplankton, and the Neogene time scale of Berggren et al. (1985). Ages for previously collected Neogene corals were derived from the original publications (Table 1), Budd ef al. (1994: Table 3), and Cairns and Wells (1987). In the analyses of taxonomic turnover rates, species and generic richnesses were calculated using the “range through method”, wherein a species is as- sumed to be present during all time intervals between its earliest and latest occurrences, even if it was not found in all intermediate intervals. In the paleoecolog- ical analysis, the method of inferring the depth range of a formation from which a coral assemblage was collected was done in the following manner. The re- ported bathymetric ranges of the species with both fos- sil and Recent occurrences (Table 3) were combined with those of living species believed to be closely re- lated as determined by morphological similarity (i.e., Recent counterparts) to others fossil species found in a formation, e.g., Cayo Agua Formation, which in this example results in a range of 1-653 m. 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Data are at the PPP internet site, http://www.fiu.edu/ ~collinsl/. Septastraea altispina Septastraea marylandica Antillocyathus gracilis x x x x Antillocyathus cristatus Trochocyathus chevalieri Paracyathus henekeni Paracyathus adetos x x x x x Oxysmilia pliocenica Asterosmilia profunda x Asterosmilia irregularis Asterosmilia exarata Sphenotrochus hancocki Gardineria minor Guynia annulata Schizocyathus fissilis Balanophyllia pittieri Xx Strylaster roseus x tors have somewhat broad bathymetric ranges, a re- stricted (or narrower) range was calculated based upon the overlap of the ranges of the species being consid- ered. For example, if a similar extant counterpart spe- cies is known to occur at 10—SO m and another species from 20—60 m, the hypothetical restricted range (over- lap) of the formation in which they both occur would be 20-50 m. The same method was used by Cairns (1979) to infer restricted bathymetric ranges of Recent deep-water corals that were based on collecting sta- tions some of which covered a broad range of depths. RESULTS BIODIVERSITY The most diverse fossil azooxanthellate coral fauna known from the Caribbean consists of 20 species from Table 2.—Continued. 168 170) 177 178: 179) 180" 189) 193" 1945 195 9G eaS 7 xX x x xX x x xX xX x x x x x xX x xX X x the Dominican Republic (Cairns and Wells, 1987). Slightly less diverse faunas are known from the Ca- ribbean Panamanian Neogene (15 stony coral species) and from the Caribbean Costa Rican Neogene (11 spe- cies). Taken together, a total of 18 fossil azooxanthel- late stony coral species are known from this southern Central American region (Table 1). All 15 species from Panama are new records for that country (Table 1), whereas four of the 11 species from Costa Rica were previously reported from Limon, primarily by Vaughan (1919): Asterosmilia exarata Duncan, 1867 (as A. hilli Vaughan, 1919); Balanophyllia_ pittieri Vaughan, 1919; Sphenotrochus sp. cf. S. hancocki Durham and Barnard, 1952 (as S. intermedius by Wells, 1983); and Archohelia limonensis Vaughan, 1919. In addition to the Dominican Republic and Pan- 357 358 359 368 379 422 423 451 458 465 466 468 475 478 479 481 627 634 635 Septastraea altispina x x Septastraea marylandica Antillocyathus gracilis x x Antillocyathus cristatus Trochocyathus chevalieri x Paracyathus henekeni Paracyathus adetos Oxysmilia pliocenica Asterosmilia profunda x x Asterosmilia irregularis Asterosmilia exarata x x Sphenotrochus hancocki Gardineria minor Guynia annulata Schizocyathus fissilis x Balanophyllia pittieri Stylaster roseus xX xX x x x x xX x xX Xx x »¢ x x x xX x x AZOOXANTHELLATE CORALS: CAIRNS 113 Table 2.—Continued. 198 201 205 208 212 294 295 298 306 307 308 310 311 x x x xX xX xX xX x x x x x xX x x xX xX x x x xX xX Xx x 313) 324 326 334 335 339 345 346 348 350) 352) 355 amanian-Costa Rican areas, other localities within the Caribbean region (Table 1) from which Neogene azooxanthellate Scleractinia have been reported in- clude: Colombia (Vaughan, 1919), Venezuela (Weis- bord, 1968), Trinidad (Vaughan and Hoffmeister, 1926), Carriacou (Wells, 1971), Martinique (Wells, 1945), and Jamaica (Duncan, 1864; Vaughan, 1919; Cairns and Wells, 1987). Budd et al. (1994) reported 175 species of zooxan- thellate corals from the Caribbean Neogene through Recent, and 142 species of azooxanthellates are known from the same time period and region, resulting in a total of 317 species. The 101 azooxanthellate species known from the Recent are listed in Table 4 and the 49 fossil azooxanthellate scleractinian species are list- ed in Table 1, eight species common to both lists. Table 2.—Continued. There are, as yet, no Caribbean stylasterid species known exclusively from the fossil record (Table 5). Of the 175 zooxanthellate species reported by Budd et al. (1994), 23 or 13% are as yet undescribed, and 18 (also 13%) of the 142 azooxanthellates are undescribed. TURNOVER RATES Text-figures 1—2 illustrate the stratigraphic ranges of the 50 azooxanthellate stony coral species known from the Caribbean Neogene and Table 6 summarizes the numbers of origins and extinctions together with spe- cies and generic richness for the Neogene to Recent. When taxa found at only one locality or time level are omitted in order to decrease signal “‘noise”’ (species 1, 3, 6, 11, 13-15, 18, 20, 22, 23, 27-33, 35, 38—40, 43, 49), and the apparent sudden diversity increase of Re- 639 640 642 646 670 708 710 720 738 757 767 962 1101 1102 1103 1104 1105 1107 1118 1119 x x xX x x x x x x xX x x x x xX x x x x x xX x xX x x x x xX x x 114 BULLETIN 357 Table 3.—Inferred bathymetric ranges, general gross morphological characters, and stratigraphic occurrences of 10 Neogene Panamanian- Costan Rican azooxanthellate stony corals. Neogene species Recent counterpart Astrangia conferta Astrangia conferta * Septastraea altispina Septastraea marylandica Stylaster roseus Balanophyllia pittieri Balanophyllia grandis Asterosmilia marchadi * Asterosmilia exarata Gardineria minor Sphenotrochus hancocki Oxysmilia rotundifolia * Spheno. ct. hancocki Oxysmilia pliocenica Guynia annulata Schizocyathus fissilis * Depth range (m) of Attached/ Colonial/ counterpart unattached solitary Formations 9-37 A (¢ CA, SHP 9-37 A (e CA 1-73 A Ec CA, Mn 40-96 U S CA, Mn 32-229 U S CA, EV, Mn 2-242 A S Mn 18-274 U S EV, Mn, RB 46-640 A S CA 28-653 U S CA, EV, Mn 88-1300 U S EV * Counterpart species same as Neogene species. CA, Cayo Agua Formation; SHP, Shark Hole Point Formation; Mn, Moin Formation; EV, Escudo de Veraguas Formation; RB, Rio Banano Formation. A, attached; U, unattached. C, colonial; S, solitary. cent species is ignored, it appears that the highest spe- cies origination rate occurs in the Middle to Late Mio- cene, and the highest extinction rate in the Late Plio- cene, resulting in the highest species and generic rich- ness in the early Late Pliocene. The sudden increase in Recent species and genera is discussed below. The high species turnover rate in the Late Miocene to early Late Pliocene is especially apparent in the Car- yophylliidae, whereas most species in the Rhizangi- idae, Guyniidae, and Dendrophylliidae appear to be longer-lived (Text-fig. 1). In the case of the Dendro- phylliidae, however, these results may be influenced by the difficulty of discriminating fossil species. No azooxanthellate species is known to have crossed the Oligocene—Miocene boundary. Twenty-four azooxan- thellate scleractinian genera occurred in the Caribbean Neogene (Table 1), only three of which do not also occur in the Recent: Septastraea, Antillocyathus, and Dominicotrochus. PALEOECOLOGY Four of the 18 Panamanian—Costa Rican Neogene stony corals are also known from the Recent and an additional six species have closely related, if not iden- tical, counterparts in the Recent. These 10 species are listed in Table 3 in order of their shallowest to deepest maximum depth ranges of the Recent or counterpart species. Table 3 also lists whether the species is un- attached or attached, colonial or solitary, and the for- mations in which it was found. The restricted depth range (see Material and Methods) for the Cayo Agua Formation (Appendix A, Map 6 and Insets; Appendix B, Sections 16—20) is 37—46 m; Moin Formation, Lo- mas del Mar (Appendix A, Inset B of Map 11; Ap- pendix B, Section 36), 40-73 m; and Escudo de Ver- aguas Formation (Appendix A, Map 4, Insets A-C; Appendix B, Sections 10-11), 88-229 m. It is ac- knowledged that the actual range of a fossil assem- blage is probably broader than this rather conserva- tively determined figure, and that the depth ranges of extant species is incomplete and not necessarily di- rectly applicable to the Pliocene epoch. Other assump- tions implicit in this method are that the fossil species actually co-occurred in the formation and that the wa- ter depth within the formation did not appreciably change over time. Nonetheless, in an analysis based on foraminiferal assemblages, Collins (1993) and Col- lins et al. (1995) found similar inferred bathymetric ranges for the Cayo Agua Formation (20—80 m), Moin Formation, Lomas del Mar (SO—100 m), and the Es- cudo de Veraguas Formation (100—150 m). Using the same method described above, the re- stricted depth range of fossil species having attached coralla is 37-82 m and unattached coralla, 88-96 m, suggesting that greater depth (e.g., a higher probability of a soft substrate) favors unattached coralla. The re- stricted range for colonial species is 37—40 m and for solitary species, 82-88 m, suggesting that greater depth favors solitary coralla. Among the 114 extant azooxanthellate scleractinian genera, two-thirds have solitary coralla and one-third have colonial coralla. All of the colonial genera consist of species that are at- tached, whereas only 37% of the solitary genera con- sist of attached species, 51% of unattached species, 3% of a mixture, and 9% of transversely dividing species. To date there has been no study correlating generic depth ranges to growth form: both solitary and colo- nial as well as attached and unattached species and AZOOXANTHELLATE CORALS: CAIRNS nS Table 4.—List of the 101 species of Recent Caribbean azooxan- thellate Scleractinia, taken primarily from Cairns (1979) and Hub- bard and Wells (1986). An asterisk (*) indicates that the species is known to occur in water depths of less than 183 m, and a cross (+) signifies that the species has a fossil record and thus is also listed in Table 1. *Agaricia cailleti *Anomocora fecunda Anthemiphyllia patera *A. rathbuni *Astrangia solitaria *Asterosmilia marchadi *A. prolifera *Balanophyllia bayeri *B. caribbeana *B. cyathoides *B. dineta *B. floridana *B. goes *B. grandis B. hadros *B. palifera B. wellsi Caryophyllia ambrosia caribbeana *C. antillarum C. barbadensis *C. berteriana *C. cornuformis C. corrugata *C. parvula C. paucipalata C. polygona *C. zopyros *Cladocora debilis *Coenosmilia arbuscula *Colangia immersa Concentrotheca laevigata Crispatotrochus cornu *Dasmosmilia variegata Deltocyathus agassizi *D. calcar D. eccentricus +D. italicus D. moseleyi D. pourtalesi Dendrophyllia alternata +*D. cornucopia *D. gaditana D. alternata *Desmophyllum cristagalli *D. striatum Enallopsammia profunda E. rostrata Flabellum atlanticum F. moseleyi Fungiacyathus crispus F. marenzelleri F. pusillus F. symmetricus +*Gardineria minor *G. paradoxa *G. simplex +*Guynia annulata *Javania cailleti J. pseudoalabastra Labyrinthocyathus langae Leptopenus discus *Leptopsammia trinitatis *Lophelia prolifera *Madracis myriaster *M. asperula *M. brueggemanni *M. pharensis pharensis *Madrepora carolina *M. oculata *Oxysmilia rotundifolia +*Paracyathus pulchellus *Phacelocyathus flos +*Phyllangia americana Peponocyathus folliculus *P_ stimpsonit Placotrochides frustum *Polycyathus mullerae *P. senegalensis *Polymyces fragilis + Pourtalocyathus hispidus *Pourtalosmilia conferta *Rhizopsammia manuelensis *Rhizosmilia gerdae *R. maculata +*Schizocyathus fissilis Solenosmilia variabilis *Sphenotrochus auritus *Stenocyathus vermiformis Stephanocyathus coronatus S. diadema S. laevifundus S. paliferus *Tethocyathus cylindraceus T. recurvatus T. variabilis *Thalamophyllia riiset Trematotrochus corbicula *Trochocyathus rawsonii T. fossulus T. fasciatus Trochopsammia infundibulum *Tubastraea coccinea Table 5—Numbers of species of Neogene to Recent Caribbean Scleractinia and Stylasteridae reported as zooxanthellate and azoo- xanthellate components. Total Zooxan- Azooxan- Sclerac- — Stylas- thellate thellate tinia teridae Neogene to Recent 175 142 317 42 Recent 68 101 169 42 Exclusively Recent 18 93 111 41 genera occur at a broad range of depths (i.e., O—2000 m). Nonetheless, the data presented in Table 3 suggest that attached colonial coralla are more common in shallow water, whereas solitary and unattached species occur in deeper water. Consideration of inferred depth, coloniality, and attachment shows a consistent trend within the three studied Caribbean formations. The as- semblage from the Cayo Agua Formation is inferred to be the shallowest (37—46 m), its stony coral fauna consisting of two colonial attached, three solitary at- tached, and six solitary unattached species. The Moin Formation (Lomas del Mar section) is inferred to be of intermediate depth (40—73 m), its assemblage con- sisting of one colonial attached and seven solitary un- attached species. The Escudo de Veraguas Formation is inferred to be the deepest assemblage among the three (88-229 m), containing exclusively solitary un- attached species. Nonetheless, in the broader context of all azooxanthellate corals, these three formations are typical of a relatively shallow-water fauna; in the Re- cent fauna, azooxanthellates are more common at slope depths (especially 200-800 m), one species (Leptopenus discus Moseley, 1881), occurring as deep as 3475 m in the Caribbean (Cairns, 1979). DISCUSSION The uneven numbers of fossil azooxanthellate spe- cies from various regions of the Caribbean (Table 1) reflect the uneven collecting efforts made in those re- gions, the Dominican Republic and Panama being the most intensively sampled and thus having the highest species richness. The taxonomic composition of the list of 50 fossil species also shows a decided prepon- derance of shallow-water (<183 m) species, except for those species reported from the Early to Middle Mio- cene of Carriacou (Wells, 1971), which include pri- marily deeper water (bathyal) genera and species. Azooxanthellate species constitute 45% (142/317) of the Caribbean Neogene to Recent scleractinian fau- na (Table 5), which is probably a low estimate, since the deeper water fossil species are poorly known. By comparison, in the Recent fauna, for which the deep- 116 BULLETIN 357 ep) 4 PLIOCENE] © oO o 2 ® = = oO 20 & © a) > = w Ww ag © =| 2 D Ae] = > = o Ww (e) be G) | ATO | |i I | | | | 123 4 6 67 8 9 101112131415 1617 18 192021222324 25 2627 2829303 132333435 3637 3839 4041 42 43 44 45 46 47 484950 co) o o oO ~ oO oO = £18 = o = sn. o|/ 5 |® = a) 2 © = |=s > = oO x= — is oO a oe 6 a |o a °o 2 D °o —N A) Cc Cc fo) c = = = oO @}| o |= > Te) 2 Cc Ae) BH c N =| + = a > c >> 3 fe o 9 =] i) = ® =F el ce IO oO = ir 0) Q As Text-figure 1.—Stratigraphic ranges for all Caribbean azooxanthellate stony corals that have a fossil record in the Neogene. Taxa arranged by families. Numbers for species correspond with those in Table 1. and shallow-water species are well known, Cairns (1979: Table 5) found that azooxanthellates constituted 66% (116/177 species) of the tropical western Atlantic scleractinian fauna. If a similar ratio were applied to the Caribbean Neogene—Recent coral fauna, one might expect to find 340 (instead of 142) azooxanthellate species in addition to the 175 reef species. The almost order of magnitude increase in species originations and species and generic richness for Re- cent azooxanthellates (Table 6) is not interpreted as an evolutionary explosion, but rather as the common pa- leontological collecting artifact known as the “‘pull of the Recent’. Most of the fossil azooxanthellates known from the Caribbean were inferred to have lived at shelf depths (=183 m) (see Paleoecology section), a facies more likely to be preserved on land than a deeper water facies. However, 42% of the Recent Ca- ribbean azooxanthellates (Table 4) occur deeper than 183 m exclusively, many species are restricted to the lower slope, and one species, Leptopenus discus, oc- curs as deep as 3475 m. This large segment of the azooxanthellate fauna is relatively well known in the Recent (Cairns, 1979) but virtually unknown in the fossil record. Consistent with this explanation, Che- valier (1961) reported 112 azooxanthellate species from the Mediterranean Miocene, a fauna character- AZOOXANTHELLATE CORALS: CAIRNS ILI E7/ > an < Za (ae LU HK < =) Se) PLIOCENE OD | | th | SieZllOm2* 16" 16°20" 26 29* 25 37 41842442 46 © = x xt-figure 2—Age ranges of coral species in PPP samples. Species numbered as in Table |. Line = maximum range. Bar = minimum range, except for starred species, where bar = interval within which the species occurs. ized by many deep-water genera. Also in this context, only one of the 42 Recent species of western Atlantic stylasterids occurs in shallow water (Cairns, 1983), i.e., Stylaster roseus, which is also the only species known from the fossil record. Another reason for the small number of fossil azooxanthellates may be due to the small size of their corallum, some solitary species having an adult calicular diameter of only 1 mm. Table 6.—Estimates of species richness and numbers of origins and extinctions of Caribbean azooxanthellate Scleractinia. Richness estimated by using range-through method. Numbers in parentheses represent number of species known from only one locality. Total Total number Extinc- number of — of Origins tions species genera Recent 93 101 51 Pleistocene 0 1 (0) 9 9 Late Pliocene 3 (1) 9 (1) 18 15 early Late Pliocene 7 (4) 6 (4) 21 14 Early Pliocene 4(1) i) 19 1] Late Miocene 13 (2) 2 (2) 17 11 Middle Miocene 11 (7) 9 (7) 13 10 Early Miocene 11 (9) 9 (9) 11 8 CONCLUSIONS A total of 142 azooxanthellate scleractinian species are known from the Caribbean Neogene and Recent based upon new collections from the Pliocene of Pan- ama and Costa Rica and previous literature. Forty-one of those 142 species are known exclusively from fos- sils, and of the remaining 101 extant species, only eight have a fossil record. One stylasterid is also known from the Caribbean Neogene. Adding the 175 species of reef (zooxanthellate) species reported by Budd et al. (1994) results in 317 known species of Scleractinia from the Caribbean Neogene to Recent. The number of fossil azooxanthellates is considered to be relatively low because lithologies consisting of shallow-water (continental shelf) facies have been pre- dominantly studied thus far; most Recent azooxan- thellates occur in deeper (continental slope) water. The highest known species diversities of fossil azooxan- thellate stony corals within the Caribbean region are in the Dominican Republic (20 species) and Panama (15 species). Examination of the stratigraphic ranges of the azooxanthellate coral species suggests that the highest origination rate occurred in the Middle to Late 118 BULLETIN 357 Miocene, and that the highest extinction rate occurred in the Late Pliocene, following a maximum of both generic and specific taxonomic diversity in the early Late Pliocene. Using depth ranges of fossil taxa in- ferred from those of the same or closely related extant species, it is suggested that the Panamanian Cayo Agua Formation supported corals living at depths of 37—46 m, the Costa Rican Moin Formation at depths of 40—73 m, and Panamanian Escudo de Veraguas For- mation at depths of 88-229 m, the three formations listed in order from shallowest to deepest. REFERENCES CITED Berggren, W.A., Kent, D.V., Flynn, J.J., and Van Couvering, J.A. 1985. Cenozoic geochronology. Bulletin of the Geological So- ciety of America, vol. 96, pp. 1407-1418. Budd, A.F., Stemann, T.A., and Johnson, K.G. 1994. Stratigraphic distributions of genera and species of Neo- gene to Recent Caribbean reef corals. Journal of Paleon- tology, vol. 68, no. 5, pp. 951-977. Cairns, S.D. 1979. The deep-water Scleractinia of the Caribbean Sea and ad- jacent waters. Studies on the fauna of Curagao, vol. 57, no. 180, 341 pp., 40 pl. 1983. A revision of the Northwest Atlantic Stylasteridae (Coe- lenterata: Hydrozoa). Smithsonian Contributions to Zo- ology, no. 418, 131 pp., 53 pl. 1995. New records of azooxanthellate stony corals (Cnidaria: Scleractinia and Stylasteridae) from the Neogene of Pan- ama and Costa Rica. Proceedings of the Biological So- ciety of Washington, vol. 108, no. 3, pp. 533-550, 36 fig. Cairns, S.D., and Stanley, G.D. 1982. Ahermatypic coral banks: living and fossil counterparts. Proceedings of the Fourth International Coral Reef Sym- posium, Manila, vol. 1, pp. 611-618. Cairns, S.D., and Wells, J.W. 1987. Neogene Paleontology in the northern Dominican Repub- lic. Part 5. The suborders Caryophylliina and Dendro- phylliina (Anthozoa: Scleractinia). Bulletins of American Paleontology, vol. 93, no, 328, pp. 23—43, 52-55, 68-74, pl. 8-11. Chevalier, J.-P. 1961. Recherches sur les Madréporaires et les formations réci- fales Miocénes de la Méditerranée occidentale. Mémoires de la Société Géologique de France, vol. 40, 562 pp., 35 pl. Coates, A.G., Jackson, J.B., Collins, L.S., Cronin, T.M., Dowsett, H.J., Bybell, L.M., Jung, P., and Obando, J.A. 1992. Closure of the Isthmus of Panama: the near-shore marine record of Costa Rica and western Panama. Geological So- ciety of America Bulletin, vol. 104, pp. 814-828. Collins, L.S. 1993. Neogene paleoenvironments of the Bocas del Toro Basin, Panama. Journal of Paleontology, vol. 67, no. 5, pp. 699— 710, Collins, L.S., Coates, A.G., Jackson, J.B.C., and Obando, J.A. 1995. Timing and rates of emergence of the Limon and Bocas del Toro basins: Caribbean effects of the Cocos Ridge subduction? Geological Society of America Special Paper 295, pp. 263—287. Duncan, P.M. 1863. On the fossil corals of the West Indies Islands. Part 1. Quarterly Journal of the Geological Society of London, vol. 19, pp. 406—458. 1864. On the fossil corals of the West Indies Islands. Part 2. Quarterly Journal of the Geological Society of London, vol. 20, pp. 20—45, pl. 2-5. 1867. On the genera Heterophyllia, Palaeocyclus, and Asteros- milia. Philosophical Transactions of the Royal Society of London, vol. 157, pp. 643-656. Hubbard, R.H., and Wells, J.W. 1986. Ahermatypic shallow-water scleractinian corals of Trini- dad. Studies on the fauna of Curagao, vol. 68, no. 211, pp. 121-147, 40 fig. Vaughan, T.W. 1919. Fossil corals from Central America, Cuba, and Porto Rico, with an account of the American Tertiary, Pleisto- cene, and Recent coral reefs. Bulletin of the United States National Museum, vol. 103, pp. 189-524, pl. 68-152. Vaughan, T.W., and Hoffmeister, J.E. 1925. New species of fossil corals from the Dominican Repub- lic. Bulletin of the Museum of Comparative Zoology, vol. 67, pp. 315-326, 4 pl. 1926. Miocene corals from Trinidad. Publications of the Car- negie Institute of Washington, vol. 344, pp. 105-134, 7 pl. Vaughan, T.W., and Woodring, W.P. 1921. A geological reconnaissance of the Dominican Republic. Memoirs of the United States Geological Survey, vol. 1, 268 pp. Veron, J.E.N. 1995. Corals in space and time: the biogeography and evolution of the Scleractinia. Cornell University Press, Ithaca, NY, 321 pp. Weisbord, W.E. 1968. Some late Cenozoic stony corals from northern Venezue- la. Bulletins of American Paleontology, vol. 55, no. 246, pp. 288, 12 pl. Wells, J.W. 1945. West Indian Eocene and Miocene corals. Memoirs of the Geological Society of America, vol. 9: 23 pp., 3 pls. 1971. [Fossil corals from Carriacou] in Fossil Mollusks from Carriacou, West Indies. by P. Jung, Bulletins of American Paleontology, vol. 61, no. 269, pp. 158-160. 1983. Annotated list of the scleractinian corals of the Galapagos. in Corals and Coral Reefs of the Galapagos Islands. P. W. Glynn and G. M. Wellington, eds., University of Califor- nia Press, Berkeley, pp. 212-291, pl. 1-20. Zibrowius, H. 1989. Mise au point sur les Scléractiniares comme indicateurs de profondeur (Cnidaria: Anthozoa), Géologie Méditer- ranéenne, vol. 15, n. 1, pp. 27—47. CHAPTER 7 PLIOCENE TO PLEISTOCENE REEF CORAL ASSEMBLAGES IN THE LIMON GROUP OF COSTA RICA ANN FE Bubb DEPARTMENT OF GEOSCIENCE UNIVERSITY OF IOWA Iowa City, IOwA 52242, U.S.A. KENNETH G. JOHNSON Scripps Institution University of California at San Diego La Jolla, California 92093, U.S.A. THOMAS A. STEMANN Department of Geography and Geology University of the West Indies Mona, Kingston 7, Jamaica BRIDGET H. TOMPKINS Department of Geoscience University of Iowa Iowa City, lowa 52242, U.S.A. INTRODUCTION The Pliocene through Pleistocene sequence in the Limon Group of Costa Rica provides some of the rich- est and best preserved fossil material documenting an episode of accelerated evolution that transformed the Caribbean reef coral fauna between 4 and 1 Ma (Budd et al., 1994a; Johnson et al., 1995; Budd et al., 1996; Jackson et al., 1996; Budd and Johnson, 1997). During faunal turnover, approximately 80% of the >100 Mio- Pliocene reef coral species (32% of 38 genera) living in the Caribbean became extinct, and >60% of the species now living in the region originated. The pat- tern of turnover was unusual in that increased speci- ation preceded increased extinction by 1—2 million years. As a consequence, reef assemblages consisted locally and regionally of a mix of extinct and living species. To better understand the cause of turnover and the complex pattern of evolutionary events involved, we have been making large, bed-by-bed collections of reef corals through well-preserved Plio-Pleistocene reef sequences at scattered Caribbean locations and comparing patterns of replacement across the region. The purpose of this paper is to describe collections of reef corals made through reefal portions of the Limon sequence of Costa Rica during four initial field expe- ditions, and to discuss their potential for future anal- ysis of faunal turnover. In the Limon sequence, reef corals occur in three formations (the Rio Banano, Quebrada Chocolate, and Moin formations) that crop out in small, isolated ex- posures near the town of Limon. Each of these three formations contains one or more reef ‘trends’, which consist of a continuous series of coral buildups aligned parallel to paleoshoreline (Text-fig. 1). As detailed in McNeill er al. (in press) and Coates (this volume), the reef trends become progressively younger from south- west to northeast, and appear to correspond in timing with eustatic sea level highstands. The oldest known buildup in the sequence, the Brazo Seco ‘patch’, occurs within the Rio Banano Formation, and preliminary strontium-isotope analyses suggest that it has an Early 120 BULLETIN 357 Caribbean Sea Text-figure 1—Map of the Limon area showing four of the five reef trends (shaded) and the locations of 33 of the 34 analyzed localities. Locality BS (Brazo Seco trend) is located along Quebrada Brazo Seco approximately 13 km due west of C4. Locality abbreviations are given in Table 1. QC, Quebrada Chocolate trends; BA, Buenos Aires trend; EM, Empalme trend (including Santa Rosa patch); LM, Lomas del Mar trend. Pliocene age between 5.2—4.3 Ma (McNeill er al., in press). Two more extensive trends, the Quebrada Choc- olate and Buenos Aires trends, occur within the Que- brada Chocolate Formation. A combination of age-di- agnostic biostratigraphic markers (planktic foraminifera, nannofossils), strontium-isotope age ranges, and mag- netic polarity data indicate an early Late Pliocene age between 3.5—3.3 Ma for the Quebrada Chocolate trend, and 3.2—2.9 Ma for the Buenos Aires trend (McNeill er al., 1997; McNeill et al., in press; Coates, this volume). The two youngest trends, the Empalme and Lomas del Mar trends, occur within the Moin Formation. A com- bination of age-diagnostic biostratigraphic markers (planktic foraminifera, nannofossils), strontium-isotope age ranges, and magnetic polarity data indicate a Plio- Pleistocene age between 2.9—1.9 Ma for the Empalme trend (including the Santa Rosa patch), and 1.9—1.5 Ma for the Lomas del Mar trend (McNeill et al., in press; Coates, this volume). Despite apparent gaps in the se- quence at 4.3—3.3 Ma as well as between the respective reef trends, the trends completely bracket the period of Caribbean faunal change, and thus provide valuable in- formation for understanding patterns of species evolu- tion within the basin during faunal turnover. Collections of ‘hermatypic’ or ‘reef-building’ corals (zooxanthellate members of the Order Scleractinia, Class Anthozoa, Phylum Cnidaria) were made through the Limon reefal units as part of four field expeditions associated with the Panama Paleontology Project (PPP): (1) April 1989 (233 specimens, 49 species), (2) January 1992 (302 specimens, 37 species), (3) March 1992 (647 specimens, 55 species), and (4) July 1993 (1209 specimens, 76 species). During these different expeditions and on different days within the same ex- pedition, samples were taken both from newly discov- ered sites and from previously sampled sites. The col- lected specimens were shipped to the University of Iowa, prepared, identified to species, and entered into a specimen database that is available on the World- Wide Web at http://nmita.geology.uiowa.edu. Al- though currently still at the University of Iowa, most of the material will be deposited at the U. S. National Museum of Natural History, Department of Inverte- brate Zoology (NMNH). Selected voucher specimens REEF CORALS: BUDD ET AL. 121 of each species will be deposited at the Paleontology Repository of the Department of Geology, University of Iowa (SUI), and at the Escuela de Biologia of the University of Costa Rica. Altogether, the collections comprise 2392 specimens (2356 identifiable to spe- cies) and 82 species. Prior to 1989, the only published faunal list of fossil reef corals from the Limon region was that of Vaughan (1919), who recorded only four species and one variety. Since 1993, additional collec- tions of reef corals have been made in the region on two subsequent PPP-associated expeditions (February 1995, July 1996); these newer collections are still in the process of being prepared and identified, and are therefore not included herein. As part of our description of the Limon collections, we provide details on sampling methods and consider potential biases that these methods pose for quantita- tively analyzing species evolution and community change through geologic time. In contrast to other fos- sil groups treated in this volume, reef coral specimens are typically large in size (sometimes over a 0.5 m in diameter) and therefore cannot be sampled using con- ventional bulk sampling procedures. Furthermore, be- cause specimens are often worn and fragmented, spe- cies are difficult to identify in the field, especially in pre-Quaternary deposits such as those treated herein. In addition, reefal units tend to be relatively rapidly deposited and patchily distributed through space and time; therefore, long, continuous sequences are rare. Because of their symbiotic algae, diverse accumula- tions of reef corals are generally restricted to depths of <40—50 m, where microfossils that are most useful in biostratigraphy are poorly preserved (see Budd and Kievman, in press). As a consequence, estimated age dates are low in resolution. Given these difficulties, we provide preliminary an- alyses of data derived from the collections to assess the diversity and abundance of reef coral taxa within the Limon reef sequence and to provide a general overview of the fauna. In this assessment, we use num- bers of species (i.e., species richness) to estimate coral diversity, and numbers of specimens per species to es- timate relative abundance. Other than species richness, we do not attempt to calculate diversity indices be- cause of sampling inconsistencies. We examine fre- quencies of different colony shapes to provide a rough interpretation of the paleoenvironments of different reef trends, and we compare global first and last oc- currences of species among trends to evaluate the evo- lutionary significance of the fauna. We analyze the as- semblages using multivariate statistical procedures (cluster analysis, detrended correspondence analysis) to describe patterns of replacement and faunal change within the Limon region during Plio-Pleistocene turn- over. Lastly, to present a broader picture of Neogene and earliest Quaternary reef faunas along the north- eastern Costa Rican and Panamanian coast, we provide descriptions of small initial PPP collections from the Bocas del Toro region of Panama. Our conclusions focus on the significance of the Limon reef fauna in understanding Plio-Pleistocene faunal turnover across the entire Caribbean region. We emphasize that the analyses presented herein are preliminary and exploratory in nature, and were per- formed mainly to assess the potential for more rigor- ous faunal analysis in the future. In future analyses, Borne and Budd plan to compare coral and ostracode assemblages to interpret reef environments more pre- cisely. Quadrat or line transect sampling methods (see Budd er al., 1989; Stemann and Johnson, 1992; Pan- dolfi, 1996) also need to be applied at selected well- preserved exposures to evaluate more accurately di- versity, relative abundance, and species associations. ACKNOWLEDGMENTS We are especially grateful to Jeremy Jackson and Tony Coates for bringing our attention to these excep- tional exposures of Plio-Pleistocene reef corals; and to Tony Coates, Don McNeill, Pam Borne, and Jorge Ob- ando for providing the stratigraphic and sedimentolog- ic framework for our work. We thank Harry Filkhorn, Terry Hughes, John Pandolfi, and Brian Rosen for re- viewing the manuscript; Alan Cheetham for statistical advice; J. Cortés, P. Denyer, J. Wineberg-Swedberg, S. Schellenberg for help with field work; J. Golden, K. Ketcher, K. Saville, T. Coffer, and S. Cairns for help with specimen preparation and curation; and Y. Ven- tocillo, H. Fortunato, and X. Guerra for help with ship- ping and integrating our collections into the PPP Da- tabase, maps and stratigraphic sections. This research was supported by a grant from the U.S. National Science Foundation (EAR-9219138, to Budd) and a U.K. Natural Environment Research Council Advanced Postdoctoral Fellowship in Taxon- omy (to Johnson). Johnson’s 1992 trip was supported by a Smithsonian Walcott Fund grant to J. Jackson and a MUCIA grant from the University of Iowa (to Budd). Stemann’s 1989 trip was supported by a Na- tional Geographic Society Grant to A. G. Coates. Tompkins’ participation was supported by an REU supplement to EAR-9219138. DEFINITIONS OF TERMS Because our collecting methods differ from those employed for other fossil groups, our usage of collec- tion-related terms is unique and requires explanation. In our work, an ‘individual specimen’ is defined as a single colony (or fragments of what was presumed to 122 BULLETIN 357 Table 1.—List of 1989-1993 Costa Rica and Panama zooxanthellate coral collection sites arranged in stratigraphic order from oldest to youngest. Strat. PPP. section Field number — number Site name number Costa Rica: AB93-05 1381 Brazo Seco none AB93-67 1386 Q. Chocolate 33 AB93-68 1384 Q. Chocolate—road 33 AB93-37 1347 Rt. 32—Dole 33 AB93-49 1359 Rt. 32-CTA fence 33 AB93-50 1360 Rt. 32—CTA fence 33 AB93-70-1 1387 Rt. 32-—CTA fence 33 AB93-06 1316 Rt. 32-CTA fence 33 AB93-52 1362 Rt. 32-—CTA fence 33 AB93-70-2 1388 Rt. 32-—CTA fence 33 AB93-36 1346 Old Moin Road-south 33 AB93-53 1363 Old Moin Road-south 33) KJ-C-1 1125 Old Moin Road-south 33 AB93-35 1345 Old Moin Road-south 33 AB93-54 1364 Old Moin Road-south 33 AB93-55 1365 Old Moin Road-south 33 AB93-60 1370 Rt. 32—Chiquita 33 AB93-38 1348 Rt, 32—La Colina 33 KJ-32-1 1124 Rt. 32 33 AB93-32 1342 Moin flat field—sw 33 AB93-33 1343 Moin flat field—mid 33 AB93-34 1344 Moin flat field—mid 33 AB93-56 1366 Moin flat field—north 83) AB93-31 1341 Old Moin Road-north 33) TS-CR-8 719 Pueblo Nuevo 34 AB93-63 1373 Rt. 32—swimming pool 34 JW93-16 1499 Rt. 32—swimming pool 34 JW93-17 1500 Rt. 32—swimming pool 34 TS-CR-7 715 Pueblo Nuevo 34 AB93-41 1351 Rt. 32—stadium 34 AB93-62 S72) Santa Rosa Road 34 AB93-57 1367 Rt. 32—Santa Marta Soda 34 AB93-84 1428 Pueblo Nuevo 34 AB93-30 1340 Old Moin Road-north 34 AB93-64 1374 Corales 1—Tajo 36 AB93-39 1349 St. Eduviges 36 AB93-40 1350 ocean view 36 TS-CR-5 771 St. Eduviges 36 TS-CR-6 a2. St. Eduviges 36 TS-CR-9 646 Lomas del Mar 36 TS-CR-1 639 Lomas del Mar 36 CJ-92-06-21 962 Lomas del Mar 36 CJ-92-06-22 963 Lomas del Mar 36 KJ-LM-16 1115 Lomas del Mar 36 KJ-LM-17 1116 Lomas del Mar 36 KJ-LM-18 1117 Lomas del Mar 36 KJ-LM-19 1118 Lomas del Mar 36 KJ-LM-20 1119 Lomas del Mar 36 KJ-LM-21 1120 Lomas del Mar 36 KJ-LM-22 1121 Lomas del Mar 36 KJ-LM-23 1122 Lomas del Mar 36 KJ-LM-24 23 Lomas del Mar 36 KJ-LM-25 1971 Lomas del Mar 36 KJ-LM-26 1972 Lomas del Mar 36 CJ-92-06-07 948 Lomas del Mar 36 CJ-92-06-08 949 Lomas del Mar 36 Num- ber of speci- mens species S7 10 65 NNDWOON OO 2 94 Lo = aA Num- ber of 19 6 tN wWwn = npr WONneNNNA Ore Nv & ENF Wee NFFOOOMAAANAIYK OONK ONKH DOO — —_ Formation Rio Banano . Chocolate . Chocolate . Chocolate . Chocolate . Chocolate Chocolate Chocolate Chocolate Chocolate Chocolate Chocolate Chocolate Chocolate Chocolate Chocolate Chocolate Chocolate Chocolate . Chocolate . Chocolate . Chocolate . Chocolate . Chocolate Moin PPPLPLLLLLLLADLLOLLLLLLOLLO Locality Reef trend code Brazo Seco BS Q. Chocolate (CP) Q. Chocolate C4 Buenos Aires BAI Buenos Aires BAI Buenos Aires BAI Buenos Aires BAI Buenos Aires BA2 Buenos Aires BA2 Buenos Aires BA2 Buenos Aires BA3 Buenos Aires BA3 Buenos Aires BA3 Buenos Aires BA4 Buenos Aires BA4 Buenos Aires BA4 Buenos Aires BAS Buenos Aires BA6 Buenos Aires BA6 Buenos Aires BA7 Buenos Aires BA7 Buenos Aires BA7 Buenos Aires BA7 Buenos Aires BA8 Santa Rosa SRI Santa Rosa SR2 Santa Rosa SR2 Santa Rosa SR2 Santa Rosa SR2 Santa Rosa SR3 Santa Rosa SR3 Santa Rosa SR4 Empalme El Empalme E2 Lomas del Mar LEO Lomas del Mar LEl Lomas del Mar LE1 Lomas del Mar LE1 Lomas del Mar REI Lomas del Mar LE2 Lomas del Mar LE3 Lomas del Mar LE4 Lomas del Mar LE4 Lomas del Mar LES Lomas del Mar LES Lomas del Mar LES Lomas del Mar ES. Lomas del Mar LES Lomas del Mar LES Lomas del Mar EES Lomas del Mar LES Lomas del Mar LES Lomas del Mar LES Lomas del Mar LES Lomas del Mar LE6 Lomas del Mar LE6 REEF CORALS: BUDD E7 AL. 123 Table 1.—Continued. Num- ber Num- Strat. of ber REE section — speci- of Locality Field number number Site name number mens_ species Formation Reef trend code KJ-LM-09 1108 Lomas del Mar 36 15 11 Moin Lomas del Mar LE7 KJ-LM-10 1109 Lomas del Mar 36 21 10 Moin Lomas del Mar LE] KJ-LM-11 1110 Lomas del Mar 36 8 i Moin Lomas del Mar LE7 KJ-LM-12 1111 Lomas del Mar 36 14 6 Moin Lomas del Mar LEZ KJ-LM-13 1112 Lomas del Mar 36 11 7 Moin Lomas del Mar LE7 KJ-LM-14 1113 Lomas del Mar 36 19 8 Moin Lomas del Mar LE7 KJ-LM-15 1114 Lomas del Mar 36 38 13 Moin Lomas del Mar LE7 CJ-92-06-01 942 Lomas del Mar 36 30 17 Moin Lomas del Mar LE8& CJ-92-06-02 943 Lomas del Mar 36 28 13 Moin Lomas del Mar LE8& KJ-LM-O1 1100 Lomas del Mar 36 55 16 Moin Lomas del Mar LEO KJ-LM-02 1101 Lomas del Mar 36 7 5) Moin Lomas del Mar LE9 KJ-LM-03 1102 Lomas del Mar 36 21 13 Moin Lomas del Mar LE9 KJ-LM-04 1103 Lomas del Mar 36 11 5 Moin Lomas del Mar LE9 KJ-LM-05 1104 Lomas del Mar 36 22 12 Moin Lomas del Mar LE9 KJ-LM-06 1105 Lomas del Mar 36 21 9 Moin Lomas del Mar LE9 KJ-LM-07 1106 Lomas del Mar 36 22 10 Moin Lomas del Mar LE9 KJ-LM-08 1107 Lomas del Mar 36 17 9 Moin Lomas del Mar LE9 AB93-71 1389 Lomas del Mar 36 63 24 Moin Lomas del Mar LE10 JW93-01 1412 Lomas del Mar 36 15 10 Moin Lomas del Mar LE10 JW93-02 1413 Lomas del Mar 36 14 10 Moin Lomas del Mar LE10 JW93-03 1414 Lomas del Mar 36 14 9 Moin Lomas del Mar LE10 JW93-04 1415 Lomas del Mar 36 19 14 Moin Lomas del Mar LE10 JW93-05 1416 Lomas del Mar 36 24 15 Moin Lomas del Mar LE1O JW93-06 2005 Lomas del Mar 36 18 14 Moin Lomas del Mar LE10 JW93-07 2006 Lomas del Mar 36 24 15 Moin Lomas del Mar LEO JW93-08 2007 Lomas del Mar 36 6 6 Moin Lomas del Mar LE1O JW93-09 2008 Lomas del Mar 36 10 8 Moin Lomas del Mar LEI1O JW93-10 2009 Lomas del Mar 36 15 12 Moin Lomas del Mar LE10 JW93-11 2010 Lomas del Mar 36 9 i Moin Lomas del Mar LE10 JW93-12 2011 Lomas del Mar 36 21 11 Moin Lomas del Mar LE10 JIW93-13 1385 Lomas del Mar 36 12 10 Moin Lomas del Mar LE10 JW93-14 1410 Lomas del Mar 36 11 8 Moin Lomas del Mar LE10 JW93-15 1309 Lomas del Mar 36 45 25 Moin Lomas del Mar LE10 AB93-65 1375 Avy. Barracuda—dorms 38 i 5) Moin Lomas del Mar LW1 AB-93-21 1331 Avy. Barracuda 38 16 12 Moin Lomas del Mar LW2 AB-93-22 1332 Av. Barracuda 38 33 15 Moin Lomas del Mar LW2 AB93-47 1357 Ay. Barracuda—dirt 38 9 ih Moin Lomas del Mar LW3 AB93-48 1358 Av. Barracuda—dirt 38 13 10 Moin Lomas del Mar LW3 AB93-23 1333 apt. complex none 19 13 Moin Lomas del Mar Pl AB93-24 1334 apt. complex none 30 16 Moin Lomas del Mar Pl AB93-25 1335 apt. complex none 22 15 Moin Lomas del Mar Pl AB93-26 1336 apt. complex none 26 17 Moin Lomas del Mar Pl AB93-27 i233}7/ apt. complex none 14 11 Moin Lomas del Mar Pl AB93-28 1338 apt. complex none 13 9 Moin Lomas del Mar Pl AB93-29 1339 apt. complex none 5 4 Moin Lomas del Mar Pl AB93-72 1390 apt. complex none 11 9 Moin Lomas del Mar Pl AB93-42 1352 Portete none 6 6 Moin Lomas del Mar P2 AB93-43 1353 Portete none Ci 5 Moin Lomas del Mar P2 KJ-P1 1126 Portete Reef #1 none 5 4 Moin Lomas del Mar P2 KJ-P2 1127 Portete Reef #2 none 35 11 Moin Lomas del Mar P2 AB93-45 1355 Bahia Portete none 4 4 Moin Lomas del Mar P3 Costa Rica: total no. collections = 107 total no. specimens = 2392 median no. specimens per collection = 15 min—max no. specimens per collection 1-146 124 BULLETIN 357 Table 1.—Continued. Num- ber Num- Strat. of ber PPP: section — speci- of Locality Field number — number Site name number mens_ species Formation Reef trend code total no. species = 87 median no. species per collection = 10 min—max no. species per collection = 1—31 Panama: AB93-74 1423 Paunch none 27 13 — — PA AB93-75 1424 Paunch none 5 5 — — PA AB93-76 2002 Swan Cay 25) 6 6 — — SC AB93-77 1285 Ground Creek—west none 6 2; — — GC AB93-79 1260 Hill Point-south 26 12 6 a — HP AB93-80 1425 Hill Point—west 26 18 14 — — HP CJ-93-20-02 943 Isla Bastimentos 22 20 10 os — FH Panama: total no. collections = 7 total no. specimens = 95 median no. specimens per collection = 12 min—max no. specimens per collection = 5—27 total no. species = 36 median no. species per collection = 6 min—max no. species per collection = 2—14 have been a single colony) collected at a single site. A ‘site’ is defined as a coral-rich horizon exposed at an outcrop on a particular day. ‘Sites’ may vary in vertical and lateral dimensions. A ‘collection’ is made at a ‘site’. Different site numbers are assigned if the same horizon at a given outcrop is recollected on dif- ferent days. On the other hand, we use the term ‘lo- cality’ to mean a group of ‘collections’ made within an area having precisely defined vertical and lateral dimensions. In the present work, the vertical dimen- sion of a locality is defined as 3—5 m, and the lateral dimension as 300-500 m. Therefore, ‘sites’ are des- ignated in the field and assigned field numbers (and corresponding PPP numbers; see Kaufmann, this vol- ume); whereas ‘localities’ are defined subsequent to field work, with reference to specific maps and strati- graphic sections. Our usage of the term ‘locality’ therefore differs from that in other fossil groups (e.g., bryozoans) treated in this volume. Our ‘localities’ are roughly equivalent to correlated site codes (CSC num- bers) in the PPP Database (Kaufmann, this volume). Finally, we use the term ‘assemblage’ to refer to the taxa collected at a given locality. LIST OF ASSOCIATED MAPS AND SECTIONS The collection sites treated in this paper are shown on the following maps and columns in Appendices A and B. Unless otherwise indicated, citations to maps and sections in this paper refer only to those given on this list. Costa Rica: Appendix B, Section 33: Chocolate to Buenos Aires Appendix B, Section 34: Empalme Appendix B, Section 35: Pueblo Nuevo Cemetery Appendix B, Section 36: Lomas del Mar, Eastern Sequence Appendix B, Section 38: Lomas del Mar, Western Reef Track Sequence Panama: Appendix B, Section 22: Isla Bastimentos, Fish Hole, Eastern Sequence Appendix B, Section 25: Swan Cay, North of Isla Colon Appendix B, Section 26: Isla Colon, Hill Point COLLECTING SITES AND LOCALITIES Collections of hermatypic corals were made at a to- tal of 107 sites within a 25 X 15 km area near Limon (Table 1; Appendix 1; Appendix A): one in the Brazo Seco patch (Rio Banano Formation), two in the Que- brada Chocolate trends (Quebrada Chocolate Forma- tion), 21 in the Buenos Aires trend (Quebrada Choc- olate Formation), 10 in the Empalme trend and asso- ciated Santa Rosa patch (Moin Formation), and 73 in the Lomas del Mar trend (Moin Formation). Detailed REEF CORALS: BUDD ET AL 125 descriptions of the general geology of the Limon area and of particular reef trends are provided by Coates (this volume). Of importance to the present study, these descriptions indicate that the fossil reef corals that crop out in the five trends show evidence of only minor, local transport. At each site, some colonies were found upright and in place, and most show no signs of excessive breakage or wear. Furthermore, none of the collected horizons reveal strong internal stratification. Thus, throughout our assessment and an- alyses of the collections that follow, we assume that each of the collected coral assemblages consists large- ly of species that lived together in life. To make the collections, individual coral specimens were extracted from the face of the outcrop using a rock hammer. Specimens were selected so that the col- lections would be qualitatively representative of the species composition of each site and their relative abundance. The species composition of each site was assessed by visually examining the exposure and dis- tinguishing species without assigning names. Relative abundances of species were determined following a similar qualitative approach. Only specimens that ap- peared to be potentially identifiable to species were collected. In the Limon area, outcrops usually occur along recently bulldozed roads or construction sites and along creek banks, and are generally small in size (usually <5 m high, <15 m wide). Because of extreme variability in the amount, size, and preservation of cor- al specimens at different outcrops as well as in the objectives of different collectors; equivalent volumes of material were not collected at each site. Similarly, sizes of sites ranged from meter-square quadrats (e.g., the “KJ-LM’ and ‘JW’ field numbers at Lomas del Mar east, see Table 1) to outcrops that were 5 X 15 m in dimension. Because of these inconsistencies, data from the collections cannot be analyzed statistically without grouping the collections into more uniform and mean- ingful sampling units. Comparisons among reef trends (Text-fig. 2) show that many more specimens and species were collected overall in the Lomas del Mar trend, and fewer were collected in the Brazo Seco patch and Quebrada Choc- olate trends. Because of exceptional preservation and abundance, collecting was especially intense in six coral-rich siltstone horizons (0.5—3 m thick) exposed at the southeast side of the Lomas del Mar trend (Ap- pendices A; B, Section 36): two collections (PPP 639, 646; 163 specimens, 32 species) were made in 1989 in the two lowest horizons (Appendix B, Section 36: 32-34 m, 37-38 m); 32 collections (PPP 942-943, 948-949, 962-963, 1100-1123, 1971-1972; 772 spec- imens, 50 species) were made in 1992 in the middle three horizons (Appendix B, Section 36: 45-47 m, Total number of species Reef trend 1600 hy NO So oO Total number of specimens @ 3 400 Reef trend MM platy free-living (_] massive branching Text-figure 2.—Bar charts showing the total numbers of species and specimens collected in the Limon reef trends. Bars are shaded according to four colony shape categories. The trends are arranged in chronological order from oldest (left) to youngest (right). BS, Brazo Seco patch; QC, Quebrada Chocolate trends; BA, Buenos Aires trend; EM, Empalme trend (including Santa Rosa patch); LM, Lomas del Mar trend. 49.5—50 m, 52.5—53 m); and 16 collections (PPP 1309, 1385, 1389, 1410, 1412-1416, 2005-2011; 320 spec- imens, 39 species) were made in 1993 in the upper horizon (Appendix B, Section 36: 61—64 m). A similar reef coral fauna, also exceptionally rich and well-pre- served, was collected in 1995 on the south side near the middle of the Lomas del Mar trend (PPP 2037; Appendix B, Section 38), and is currently being washed and identified. Collecting was more limited in the Buenos Aires and Empalme trends primarily because of poorer pres- ervation. We surveyed numerous small exposures of 126 BULLETIN 357 these two trends, but the corals were often extensively recrystallized. Collecting efforts were most reduced in the Quebrada Chocolate trends and in the Brazo Seco patch, mainly because of lack of field time. Clearly, these older trends and other sites in the Santa Rita and Rio Blanco areas could prove essential to a more com- prehensive documentation of Pliocene events preced- ing and concurrent with Plio-Pleistocene turnover in Caribbean reef communities and therefore warrant more thorough study in the future. To ensure that localities were consistently defined in statistical analyses of the collections, all specimens collected within 300-500 m of one another laterally and 3—5 m of one another vertically were grouped to- gether into localities (34 total) (Text-fig. 1; Tables 1, 2). These groupings were made by careful study of the 1:10000 Ciudad de Limon (Edicion 2—IGNCR 1989) and the 1:50000 Rio Banano (3545-I, Edicion 2— IGNCR 1989) and 1:50000 Moin (3546-I, Edicion 3—IGNCR 1989) topographic sheets and Appendix B, Sections 33, 34, 36, 38. Based on these groupings, only one locality was represented within the Brazo Seco patch, two in the Quebrada Chocolate trend, eight in the Buenos Aires trend, seven in the Empalme trend, and 16 in the Lomas del Mar trend (Text fig. 1; Table 1). In general, numbers of specimens collected per lo- cality range from two to 320 (median = 45), and num- bers of species collected per locality range from one to 39 (median = 19.5). Study of the frequencies of numbers of species and specimens per locality sug- gests that a disproportionally high number of localities contain low numbers of species and specimens (Text- fig. 3). Ideally, if sampling were equal in different lo- calities and the localities were equal in species rich- ness, the two histograms in Text-figure 3 should be more or less bell-shaped; however, the distribution for number of species is platykurtic and the distribution for specimens is skewed to the right. Therefore, seven of the 34 localities with fewer than 10 specimens (those with double asterisks in Table 2) were dropped in subsequent statistical analyses, leaving a total of 27 localities in the analyzed data set. In addition, as shown in the scatterplot in Text-figure 3, the relation- ship between numbers of species per locality and num- ber of specimens per locality changes sharply near a locality having 30 specimens and 18 species (BA3). Six additional localities that have fewer than 30 spec- imens were thus treated with caution in subsequent Statistical analyses. Results of Kruskal-Wallis Non-parametric One-way Analysis of Variance (using the 27 localities with > 16 specimens) indicate that no significant difference exists among trends in numbers of specimens (Cor- rected Chi-Square = 5.372, D.E = 4, p-value = 0.251) or species (Corrected Chi-Square = 5.492, D.E = 4, p-value = 0.240) collected per locality (Text-fig. 4). These results suggest that sampling intensity was roughly equivalent within localities in the different reef trends. TAXA IDENTIFICATION AND GENERAL DESCRIPTION Budd and Stemann identified a total of 82 species belonging to 31 genera in the collections (Table 3). Five additional species were present, but could not be identified due to poor preservation. To guide us in making identifications, we used a consistent set of characters and character states developed on the basis of morphometric analyses of Neogene and Recent cor- al samples collected across the Caribbean region (Fos- ter 1986, 1987; Foster et al., 1988; Budd, 1991; Ste- mann, 1991, in press; Budd et al., 1994a, b; Swedberg, 1994; Johnson and Budd, 1996; Budd and Johnson, 1999). Lists and illustrated definitions of these char- acters along with information on species authorship and synonyms are currently available on the World- Wide Web at http://nmita.geology.uiowa.edu. A spec- imen database (Appendix) is also available at the same address. As part of the identification procedure, we assigned identification confidence codes to each spec- imen as follows: 1 = 100% confident, 2 = 75% con- fident, 3 = 50% confident, 4 = 25% confident. The total number of species (87) recognized in the Costa Rican collections is less than the total number of species (107) estimated to have lived in the entire Caribbean region between 5.5 and 1.5 Ma, as docu- mented in the 1996 Cenozoic Coral Database (CCD) compiled by Johnson and Budd in S-plus using STAT- POD (Johnson and McCormick, 1995; Budd and John- son, 1997). Nevertheless, cumulative number of spe- cies curves for the 27 better-sampled localities (Text- fig. 5) suggest that a major proportion of the species in the Limon reef trends has been sampled. These curves differ from more traditional species area curves used in ecology (Ricklefs, 1990) in that our localities do not represent repeated samples from a single statis- tical population. Instead, the localities are samples of a fauna that is undergoing change and therefore rep- resent different stages in a faunal transition. Because they are not replicate samples, curves were constructed using two methods: (1) the localities were added in temporal order beginning with the stratigraphically oldest reef trend and continuing to the youngest (Text- fig. SA), and (2) the data were randomly resampled to determine the average number of species as a function of numbers of localities sampled (PC-ORD, Version REEF CORALS: BUDD ET AL. 277; Table 2.—Numbers of identified specimens and species collected within localities and reef trends. Localities with double asterisks were not included in statistical analyses; localities with single asterisks were treated with caution. All Branching Freeliving Massive Platy No. Locality No. No. No. No. No. speci- No. No. No. No. Reef trend code species specimens species specimens species mens species specimens species specimens Brazo Seco BS 19 56 5 25 4 11 a 14 3) 6 All localities n=I1 19 56 5) 25 4 11 7 14 3 6 Q. Chocolate @zs 6 10 3 5 0 0 3 5) 0 0 Q. Chocolate C4 27 59 7 14 2 4 15 31 3 10 All localities n=2 29 69 7 19 2 4 NY 36 3} 10 Buenos Aires BAI* 7 20 3 14 0 0 3 4 i 2) Buenos Aires BA2 20 86 7 39 1 I 10 31 2 15 Buenos Aires BA3 18 30 9 18 i l 6 8 2; 3} Buenos Aires BA4* 12 16 5 8 0 0 7 8 0 0 Buenos Aires BAS** 1 2 1 2 0 0 0 0 0 0 Buenos Aires BA6 20 114 TV 61 2 13 9 23 D, ay Buenos Aires BA7 36 160, 9 66 4 1s 21 69 2 10 Buenos Aires BA8* 13 21 4 9 0 0 8 11 1 1 All localities n=8 42 449 iil PRG) 5! 30 24 154 2 48 Median n=7 18 30 Uf 18 1 1 8 11 1 2 Minimum n=7 i 16 3} 8 O 0 3 4 0 0 Maximum n=7 36 160 9 66 4 / i) 21 69 2 ey Empalme Bits 6 9 1 1 0 0 4 6 1 2 Empalme E2e* 4 4 1 1 (0) 0 3 3 0 0 Empalme E3 10 18 1 1 0 0 9 17 0 0 Santa Rosa SR1 24 30 7 9 22 2 14 18 1 1 Santa Rosa SR2 26 74 7 25 2 4 16 39 1 6 Santa Rosa SR3 25) 56 3 7 1 2 17 38 4 9 Santa Rosa SR4* 13 23 3 8 0 0 10 15 0 0 All localities n=7 46 214 8 52 5} 8 3] 136 4 18 Median n=5 24 30 5} 8 1 2 14 18 1 1 Minimum n=5 10 18 i 1 0 0 9 15; O 0 Maximum n=5 26 74 if. 25 7 4 17 39 4 9 Lomas del Mar LE1 22 41 5 10 1 2 11 16 5 13 Lomas del Mar LE2 31 135 4 10 3 7 16 4] 8 Wil Lomas del Mar LE3* 12 28 1 1 0 0 5 5 6 22 Lomas del Mar LE4 32 242 3 28 4 14 18 93 7 10 Lomas del Mar LES 28 168 2 8 1 1 19 97 6 62 Lomas del Mar LE6** 3 3 0 0 0 0 1 1 2 7 Lomas del Mar LE7 26 126 1 5 1 1 19 85 5 35 Lomas del Mar LE8 29 79 4 8 4 14 iI) 40 5 16 Lomas del Mar LE9 31 175 3 7 4 8 18 69 6 91 Lomas del Mar LE10 39 320 3 22. 3 27 25 159 8 112 Lomas del Mar LW1** 5) 7 1 1 0 0 PA 2 2, 4 Lomas del Mar LW2 24 49 7 11 2 2 11 21 4 15 Lomas del Mar LW3* 12 22 0 0 1 2 3) 8 6 12 Lomas del Mar Pl 34 139 3) 8 1 1 22 100 8 30 Lomas del Mar P2 19 52 4 6 1 1 10 23 4 Pap) Lomas del Mar B3cx 4 4 1 1 0 0 3 3 0 0 All localities n= 16 65 1590 12 123 Yi 67 3}5) 758 11 619 Median n = 13 29 126 3 8 1 2, 16 41 6 30 Minimum n= 13 12 19 0 0 0 0 5) 5 4 12 Maximum n = 13 OZ 320 7 28 4 27 PDs) 159 8 112 Panama PA 14 32 0 0 0 0 11 28 3 4 Panama SGFS 6 6 3} 3 0 0 2 2, i} I Panama GG , 6 2 6 0 0 0 0 0 0 Panama HP 18 29 5 10 1 1 10 14 2 4 Panama FH 11 21 2 3 4 13 4 4 1 1 All localities n=5 515) 94 10 22 5) 14 17 48 3 10 Minimum n=3 11 21 0 0 0 O 4 4 1 it Maximum n=3 18 32 3} 10 4 13 eh: 28 &) 4 128 BULLETIN 357 5 5 4 - 3 3 fe 58 ° zo , 0 0 4 8 12 16 20 24 28 32 3% 40 Number of species per locality 10 oS 8 5B 6 oy 5a z= 2 0 0 30 60 90 120 150 180 210 240 270 300 330 Number of specimens per locality 3 40 30 Ze Mes 5 8 20 oO a Ee 2 10 5 z 0 0 50 100 150 200 250 300 350 Number of specimens per locality Text-figure 3.—Histograms and scatterplot showing the numbers of species and specimens collected per locality. 2.0, McCune and Mefford, 1995; Text-fig. 5B). The first curve (Text-fig. 5A) levels off in a series of stepped plateaus. The steps appear to correspond with reef trends and are best developed for localities within the three younger trends. This result indicates that the species that lived within each reef trend are more or less adequately sampled. On the other hand, the second curve (Text-fig. 5B) levels off at between 10 to 15 localities, again indicating that the sampled localities adequately estimate species richness. Of the 82 identified species, 49 are living, and 33 are extinct (Table 4). The 49 living species represent 81.7% of the 60 hermatypic species in the Caribbean today (Budd er al., 1994a). Among the living species a ® o 40 ” oe 0 i 83 4 _ Ew = 10 0 BS @C BA EM LM 2 400 @ £ oO a a 200 Ue o= 33 sa] > Ex. 0 za BS @C BA EM LM Text-figure 4.—Maxima, medians, and minima of numbers of spe- cies and specimens collected per locality within each of five reef trends. The trends are arranged in chronological order from oldest (left) to youngest (right). BS, Brazo Seco patch; QC, Quebrada Chocolate trends; BA, Buenos Aires trend; EM, Empalme trend (in- cluding Santa Rosa patch); LM, Lomas del Mar trend. are all of the species that dominate modern Caribbean shallow and deep reef communities (Goreau, 1959; Goreau and Wells, 1967), including Acropora palma- ta, A. cervicornis, Undaria agaricites ( = tenuifolia), Agaricia lamarcki, Siderastrea siderea, Porites astreo- ides, P. furcata, Diploria strigosa, members of the Montastraea annularis complex, and Colpophyllia na- tans. Also included are exceptionally well-preserved specimens of the modern, typically deep forereef spe- cies described by Wells (1973) [e.g., species of Mad- racis, Agaricia, Mycetophyllia, Dichocoenia] and Por- ites colonensis. Eight of the species identified in the collections, represented by 76 specimens, are unde- scribed, as are species in at least two species com- plexes (Montastraea ‘limbata’, 88 specimens; M. ‘cav- ernosa’, 39 specimens). Among the extinct corals are several species of Stylophora that are common or abundant through Plio-Pleistocene intervals of the Ba- REEF CORALS: BUDD FT AL. 129 Table 3.—List of species identified in collections, arranged by country in taxonomic order. Phaceloid colony shapes are classified as branching; solitary coralla as massive. CCD No. of No. of No. of species Colony speci- collec- _local- Family Genus Species ID no. shape mens tions ities Costa Rica: Astrocoeniidae Stephanocoenia intersepta 2 massive 69 36 12 Astrocoeniidae Stephanocoenia duncani 3 massive 113 33 20 Astrocoeniidae Stephanocoenia spongiformis 4 massive 8 7 5 Pocilloporidae Stylophora affinis 5) branching 2 1 1 Pocilloporidae Stylophora granulata 7 branching 14 5) 4 Pocilloporidae Stylophora minor g) branching 7 3 3 Pocilloporidae Stylophora monticulosa 10 branching 8 7 6 Pocilloporidae Pocillopora crasssoramosa 15 branching 19 3 3 Pocilloporidae Madracis asperula 16.5 branching 26 18 5 Pocilloporidae Madracis decactis 17 massive 110 42 11 Pocilloporidae Madracis mirabilis 20 branching 55 24 11 Pocilloporidae Madracis pharensis 21 massive 1 1 1 Pocilloporidae Madracis sp. A 2iES branching 2 1 1 Acroporidae Acropora cervicornis 22 branching 74 22 15 Acroporidae Acropora palmata 23 branching 45 21 14 Agariciidae Agaricia grahamae 29 platy 137 39 13 Agariciidae Agaricia lamarcki 30 platy 97 36 13 Agariciidae Agaricia undata 32 platy 67 32 12 Agariciidae Undaria agaricites 33 platy 324 9) 25 Agariciidae Undaria crassa 34 massive 27 18 11 Agariciidae Undaria pusilla 35 platy 4 4 4 Agariciidae Helioseris cucullata 43 platy 27 19 14 Siderastreidae Siderastrea radians 56 free-living 3 2 2 Siderastreidae Siderastrea siderea 58 massive 42 28 20 Poritidae Porites astreoides 63 massive 52 30 7 Poritidae Porites portoricensis 65 branching 12 10 8 Poritidae Porites waylandi 68 massive 5 ) 5 Poritidae Porites baracoaensis 69 branching 26 15 11 Poritidae Porites branneri 70 massive 18 14 10 Poritidae Porites colonensis 73 massive 14 9 4 Poritidae Porites furcata 76 branching 63 24 11 Poritidae Porites porites Wi branching 3 3 2 Poritidae Goniopora imperatoris 80 massive 1 1 1 Faviidae Caulastraea portoricensis 83 branching 58 26 17 Faviidae Favia fragum 88 massive 7 5 Faviidae Diploria clivosa 94 massive 12) 7 7 Faviidae Diploria labyrinthiformis 95 massive 15S 9 5 Faviidae Diploria sarasotana 96 massive 2 2 2 Faviidae Diploria strigosa 97 massive 38 24 19 Faviidae Manicina areolata 100 free-living 37 19 10 Faviidae Manicina mayori 101 massive DD) 18 12 Faviidae Manicina puntagordensis 102 free-living 7 11 7 Faviidae Thysanus sp. A 109 free-living 19 11 d Faviidae Colpophyllia amaranthus 112 massive 3 2 2 Faviidae Colpophyllia natans 114 massive 39 28 20 Faviidae Colpophyllia sp. A 114.5 massive 14 12 7 Faviidae Montastraea faveolata 117 massive 50 33 14 Faviidae Montastraea franksi 118 massive 34 19 11 Faviidae Montastraea limbata-| fart massive 10 6 5 Faviidae Montastraea limbata-2 121.2 massive a) 38 21 Faviidae Montastraea sp. A 122 massive 29 16 11 Faviidae Montastraea canalis 124 massive 11 9 8 Faviidae Montastraea cavernosa-2 126 massive 32 27 15 Faviidae Montastraea cavernosa-3 127 massive 7 6 6 Faviidae Montastraea cylindrica 128 massive 113 44 14 Faviidae Solenastrea bournoni 131 massive 23 6 5 Table 3.—Continued. Family Trachyphyllidae Meandrinidae Meandrinidae Meandrinidae Meandrinidae Meandrinidae Meandrinidae Meandrinidae Meandrinidae Meandrinidae Meandrinidae Oculinidae Mussidae Mussidae Mussidae Mussidae Mussidae Mussidae Mussidae Mussidae Mussidae Mussidae Mussidae Mussidae Caryophyllidae Caryophylliidae Costa Rica: Panama: Astrocoeniidae Pocilloporidae Pocilloporidae Pocilloporidae Pocilloporidae Pocilloporidae Pocilloporidae Acroporidae Acroporidae Agariciidae Agariciidae Agariciidae Siderastreidae Poritidae Poritidae Poritidae Faviidae Faviidae Faviidae Faviidae Faviidae Faviidae Faviidae Faviidae Faviidae Faviidae Faviidae Faviidae BULLETIN 357 Genus Species Antillophyllia Sawkinst Meandrina braziliensis Meandrina meandrites Meandrina sp. A Placocyathus trinitatis Placocyathus variabilis Dichocoenia caloosahatcheensis Dichocoenia eminens Dichocoenia stokesi Dichocoenia stellaris Dichocoenia tuberosa Archohelia limonensis Antillia dentata Scolymia cubensis Scolymia lacera Mussa angulosa Mussismilia aff. M. hartti Tsophyllastrea sp. B Mycetophyllia aliciae Mycetophyllia danaana Mycetophyllia ferox Mycetophyllia lamarckiana Mycetophyllia reest Mycetophyllia sp. A Eusmilia fastigiata Eusmilia sp. A total no. genera = 31 total no. species = 82 total no. specimens 2356 Stephanocoenia duncant Stylophora affinis Stylophora granulata Stylophora monticulosa Pocillopora crassoramosa Madracis asperula Madracis decactis Acropora cervicornis Acropora palmata Undaria agaricites Undaria crassa Helioseris cucullata Siderastrea siderea Porites astreoides Porites baracoaensis Porites furcata Caulastraea portoricensis Diploria labyrinthiformis Diploria strigosa Manicina areolata Manicina puntagordensis Thysanus corbicula Colpophyllia natans Colpophyllia sp. A Montastraea faveolata Montastraea franksi Montastraea limbata-2 Montastraea cavernosa-2 CCD species ID no. 137 138 139 139:5 143 144 145 146 148 149 150 152.5 153 155 157 158 160.5 164.5 166 168 169 170 171 171.5 173 175 100 102 111 114 114.5 117 118 PAY 126 Colony shape free-living free-living massive massive free-living free-living massive massive massive massive massive branching free-living massive massive branching branching massive platy massive platy platy platy platy branching branching massive branching branching branching branching branching massive branching branching platy massive platy massive massive branching branching branching massive massive free-living free-living free-living massive massive massive massive massive massive No. of speci- mens to oS) onn oe nN = NAQDANK WON NN NY OOH to 455 WNn + I it) WNrRe NN eRe Ree env fe a io We io Wi — ii al me ote Sala) No. of collec- tions tN AWD BRE eee UNO bh S95) ee COG Th TF ON KON Oe 1 —_ Re ON Ree EP PNY KF RP EF WWNEK KE NWNK UN Kee ee eer No. of local- ities — ORR eB KB WOAH LP ee (>) =H BPNANAANKHUANWR Kf Ree EP WNNRF KF EF NNNK KEP NNN KK ENR RRP RP ee eb REEF CORALS: BUDD ET AL. 131 Table 3.—Continued. Family Genus Species CCD species ID no. No. of speci- mens No. of local- ities No. of collec- tions Colony shape Meandrinidae Meandrina braziliensis Meandrinidae Meandrinidae Meandrinidae Meandrinidae Mussidae Mussidae meandrites sp. A variabilis Meandrina Meandrina Placocyathus Dichocoenia eminens Mycetophyllia Mycetophyllia danaana ferox Panama: total no. genera = 19 total no. species = 35 total no. specimens 94 eS) 138 free-living 139 massive 139.5 144 free-living massive 146 massive 168 massive 169 platy — NN — ON — l 1 1 1 2 2 l ] 1 | 1 1 1 | Cumulative number of species 23> 0 10 20 30 Number of localities 40 20 Average number of species 00 0 10 20 30 Number of localities Text-figure 5—Cumulative number of species curves assessing sampling adequacy. (A) Curve constructed by adding localities in stratigraphic order beginning with the oldest and continuing to the youngest. Each point represents a locality, the abbreviations for which are given in Table 1. The curve levels off in a series of steps corresponding to the different reef trends. (B) Curve constructed by randomly resampling localities. See text for details. hamas Drilling Project cores (Budd and Kievman, in press). Only five of the identified species are known only from the Limon area of Costa Rica (Table 4); there- fore, most species (77 of 82) appear to have been widely distributed geographically. However, study of the numbers of localities and specimens per species shows that most species occur at relatively few local- ities and are represented by relatively few specimens (Text-fig. 6). A regression analysis of these two vari- ables (y-intercept = —13.595, regression coefficient = 5.509) yields an adjusted R-square of 0.5127 and a p- value <0.001. Thus, most species are less common or rare within localities, and species with lower abun- dances tend to occur at fewer localities. ENVIRONMENTAL SIGNIFICANCE OF THE COLLECTED TAXA Hermatypic coral assemblages are commonly used in sedimentology and stratigraphy to interpret ancient depositional environments (e.g., James, 1984; Scoffin, 1987). They are especially important in shallow car- bonate environments where environmentally diagnos- tic microfossils are rare. For example, comparisons be- tween corals and lithologic data in lithostratigraphic units in the Bahamas Drilling Project cores show that interpretations based on corals correspond well with those based on independent sedimentologic criteria (Budd and Kievman, in press). Three characteristics of coral assemblages are examined in making these in- terpretations: (1) frequencies of different colony shapes, (2) species richness, and (3) occurrences of indicator species. Following Geister (1983) and Graus and Macintyre (1989), an abundance of species with plate-shaped col- onies can be interpreted to indicate deep forereef or low light environments. Mound-shaped or encrusting colonies indicate shallow platform or backreef condi- tions with high wave action. Mound-shaped and BULLETIN 357 132 S ISLES ENSCé Ops (RW) asuvl ase [eqo1D qua00y jua09y lie eps| 1ua09y jud00y ]UZ9ayY jua.ay jUD00y jUD99y jus90y qua00y qua00y quay qua0ay quaday —¢T) evolewmes ‘spog kod PIO (RW L'I-8']) seweyeg ‘7-epup (RW 8 [-6 [) seureyeg ‘¢-epup (RI L'1-8']) seweyeg ‘7-epuy (RW 9 I-S'€) RAND “Wy zm eT JUD99yy (PI TEL -€'L1) dey ‘wog ‘wy voneg (PW €-S'€) PLOY ‘Sg Ise199UIg (RI OL —¢°g) doy ‘wog ‘uly oprs1a9 (RIN OL —¢°g) ‘doy wog ‘wy opesisD (RW 9°S -¢'1) day ‘wog ‘wy oqeiny (RW OL —¢'°g) ‘day ‘woqd ‘wy opersiaD (RIN TIT —¢]) pepluny ‘uy euRWiey (RI OL —¢'°g) ‘day “wog ‘wy oqriny (RIN Tel -€L1) day woq ‘uy eoueg (RW Tel -¢L1) day “wog ‘uly eourg (RW TI-L’€T) Bpuoly ‘uy edurey, (RW 9'LI 77) vureueg ‘wy 10peiodurg (RIN OLI -TZ) eueurg ‘wy 10peiodwiq (RIN 9S —¢'1) ‘day ‘wog ‘uy oqeiny (PW 79I —CT) Pyinsuy ‘wy eypinsuy (RW €-C'€) BPLIO[ *S§ JSIIOSIUT (OTD 20UaLINI90 IsET [RQOID (Od) 29ueLIND90 IsIy [RGOTD (Od) x x x x (OA) X x x x x x x x x x é x (OA) X x Mi (OH) x x x Xs x x x x x x x x x x (RWW 91-61) Ivy [ep seuoT (IN (@NE-9'¢) (CPN 91-61) SoIly €-9'¢) out souong aq] -jedwiq -o00yD To) é (OT ‘OD x (RW pes) Jequinu 099§ ozrig S lon CNTICIRGNIIQN Ca: =m NM Ol C di soroads (eee) Dasapis SUDIPDA pID]JNINI pypisnd DSSDAI SaIADSD pyppun 1YOADUD] ADUDYDAS pipujod SIUAODIALAD (JW “ds sisuaipyd SIJIQDAIU syop2ap pynsadsp DSOWDAOSSDAD psojnoyuou 4oulu pDID]NUuvAS siuyffo DAAISDAIPIS DAAISDAAPIS suasoyay pDiuppuy) piuppuy) piuppuy) DIIUDSY DIIUDSY DIDIUDSY piodosoy piodosoy SIOLpDW SIODAPDIV SIODApDW S1IODAPDW SIODAPDW piodopp1a0g pioydojAlg paoydojAlg psoydojg paoydojAlg smusofisuods piuaosouvydalg lupounp viuaoouvydalg vidasiajul_ viuaozouvydaig satoadg “saSueI UL papnyourl jou pur ‘syeUI uonsanb Kq paieorpur ore saouapyuos mor YIM poynuapr satsadg “vose uoWT] ay) WOIy A[UO UMOUY are Systajse YM satoadg *spusd joor UUM] a4} UT SsadudLMNIIO—p A1qeL 133 REEF CORALS: BUDD E7 AL. (RN TTI J I-SI —C[) pepluuly “Wy euelUey ( (OT) x x x Tei l¢cal [-DIpquiy vavalsvIUuop O-9'€ JUIIIY x x (OA) X é SII isyudaf DADAISDIUOPB 0-9'€ yU299y x x (OH) X LII pipjoaanf DavAISDIUOP 0-9'€ JUII9y x x (Od) x Crii v ds pyyjAydodjoy (RI Sb 0-9'¢ quasoy «-—9°¢) ‘day ‘wiog ‘wy oqeing x x x é rll supjpUu piyjdydodjo) 0-9’ JUDIIY x3 (OH) X TIT snyjuvapwud piyyjaydodjoy 9 1-9'¢ (OT) x x x (OH) x 601 v ds snuvsxy [ (RWW 9'I-8'T) 9I1-9'E Pplol “Wy soysyeyRsooye+) (OT) x (Ox) x th TOI sisuapsospjund pulnluvypy 0-9'¢ JUIIIY x x (OH) X é IOI LoxADUL DUuloIUuUD (RI O-9'€ JUIIIY €-C'¢) PPUO]Y “Sg ISa1D9UId x x (Od) x OoT DID]OAAD pUulolUuDpy (eI 0-9'€ JUus99y €—-G'¢€) BPPHO] “Sg ISaID9UI x x (OH) X L6 DSOS1AIS poi diq (RW (RW eSSis €—-G'¢) BPHOLY Sg ISa1D9UT €-°¢) PPUO]{ “Sg ISa1D9UTd é é 96 DUDIOSDADS piuojdiq 0-9'¢ qusd9y x x (OA) X 66 SIUMOfIYIUIKAAGD] pioj}diq eS JUIIOY x x (OH) X +6 DSOAI]I puojdiq (RW O-S't JUQIOY €—-G°¢) BPHO] “Sg ISa1O9UId x (OA) X 88 unsvif DIADY (RW I-81) (RI I=Z' TT poreuer ‘UL. [ROUOTYOUR|L €S-T 11) euopy ‘uy ory (OTD xX x Xe €8 sisuansoj4od DavAISDINDD (RW (eI ONSLSG 8' 1-6 1) seueyeg ‘e-epuy) = TZ—L'€T) PPUOLA ‘uly eduey, x 08 stiopsaduar paodouoy (eA Os9ie JUIIOY €—-¢°¢) BPHO]{ “Sg Isa1D9UT x (OH) X LL saqiiod SANsOd 0-9'¢ JUIIIY x x (OH) x OL pinpoint Sao 0-61 WUII9yY (OH) x €L SISUIUO]OI SaOd (RW O-S'€ JUIIOY €-C'¢) BPLIO][Y “sg 1so1D9UIg x é OL LaUuUudiqg SAVMOd (PIN (RW OLI 0-77 L1-8'|) seweyeg ‘7-epuy, —ZZ) eureuRg ‘wT 10priodwiq (OTD x x x x 69 SISUADOIDADG Salod (RW ONS ESe TC-LET) PPUOTY “uLy eduey, (OT) x é 89 IpunjAnm Sayod (eI CPI SAKE TI-L'€T) BpuUOTy ‘uy edurey, (OT) x x x 69 sisuansoj4od Saylod (RW 0-9'€ quss9y €-S'¢€) BPO] ‘sg Isa1s9uUTg x x (OH) X é €9 SAP1OIAISD Salo (PW) (OT) 299uU91IND90 JsPT [RGO[H (OA) 29U9aLIND50 ISI [RQO[H (eI 9 T-6' 1) (RW (RWW ¢€-9'¢) (eI (RW p-¢E'S) Jequinu sa1oadg asuri o3e Ie 9'I-6'1) Somly €-9'€) 099g OzRIg dl [egolp [ep seuoT] oul souong ae] sorsads -yedwiq me doo ‘ponunuog—"p 21qPL 0-9'S BULLETIN 357 (RW) asuri ase [PGOID 134 juD90y 1UD90y (RW b-S"b) ‘day ‘wog ‘uy oqeinn, 1Ud90y JUIDIy (RW 9 I-81) PPHOLY ‘ULy syoeyRsooyeD (RW 9'I-8'1) PPUOLy ‘UL IayieyRsoojeD (eI T-¢) PoreUIes ‘UL Uspmog quas0y Ua99y JUDIIY jua.90y JUDIIY 1Ua99y (RW 9S —¢'1) day ‘wog ‘uLj oqriny (RW OL —¢°g) day ‘wog ‘uly oprsia9 (RW 9° —¢'L) ‘day ‘wog ‘uj oqeiny PIN €-S'€) PPHOLY “Sg Is91d9UTg (eI PEe-L'¢) ‘doy ‘wog ‘uy orpy (RW I€l —¢'L1) ‘day ‘woqg ‘wy eroleg (RIN ES-TID pepluuy ‘wy eypluezueyy (IN Sb -9°¢) day ‘wog ‘uny oqeiny (EW €-$°¢) BPUO] ‘sg Isa1D9UIg (RIN SL —¢'g) ‘day wog ‘uy oproia5 (RW T9I TZ) Blinsuy ‘uy eypinsuy (PIN TT-L'€T) PPHOLY ‘wy eduey, (QW TIT —S]) pepluuy ‘wy vueuey (RIN Tel ELI) day ‘wog ‘uy vourg (eI TT-L'€T) PPHOTY “uy eduey (OW TTI S|) pepruny, “wy euRUEL, (QJ) 29uUd1IND90 IsE] [RGoTH (Od) 29ueLIND90 IsIY [eqOTD (Od) x x é (OT (Od) X (OT x x Xx (Od) x x x (OA) X (OT) x (OT x é é x é x x (OD x x ys (OD x x x (Ox) x x x (OD x x é (OT) x x x (OT x x x (RINDI-61) (RN (PIN €-9'¢) RIN 91-61) SOIL, Joep sewoT oul souong -jedwiq LSI é 8rl orl Stl bri é 8el é (Ox) x m cIcl (RI p-E'S) Joquinu ooag ozeig ss I sarsads (a(e)e) (RIN €—-9'¢) aI] -00049 x0) DAJIV] sisuaqno pivjuap (,,)sisuauoUutl] pso1aqni SLID] JIS 1sayo]s suaunua SISUJIYIIDYDSOO]VI SUIQDIADA sypyiUut4y v ‘ds Sajlipupaut SISUIT[IZDAG 1SULYMDS 1u0usnog DI1ApUuyArd €-DSOULAADI T-DSOUAIADI SIJDUDID Gow ds T-pIpquay DIWUA]OIS DIWUAJOIS HIRENY pYayoydIy DIUZOIOYIIG. DIUaOIOYIIG. DIU2OIOYIIG. DIU2OIOYIIGD DIU2OIOYIIG SNYIDKIOID] J SNYIDAIOID] g DULIpUubapy DULIpUuDapy DULIpUDa piyyAydo]yuy padjspuajog DIDAISDJUOpy DIDAISDJUOP DIDAISDIUOPy DAIDAISDJUO Py DADAISDJUO PY pavilsD]Uuop sa1sadg ‘ponunuodj—fp 2GRL 135 = SI9yIO ‘(8661) TON pue ppng = vorewres ‘(996]) 7P Ja siapunesg = o1[qnday uvotulM0g OMI Ve O-S't 91-61 0-6'1 0-61 0-61 0-61 0-61 CoE REEF CORALS: BUDD FT AL. OPIS 71L O-S'L (RI) asuvl ase reqolD (OTD) 29uaLINDd90 sey [RGo[D 0c I 0 II £ oI 8°89 OSL S19 t9 Ip 6c (RIN 9°¢ —¢L) ‘day ‘wog ‘why oqring (OT x (eI jua09yf €-C$'¢) BPLUO[Y ‘sg Isa1D9UIg x x (OT ‘Ox X Udy (OAD) X jud90y (OAD X é JUddNy X (OH) xX jUa99y (Od) X JUID9y (OH) X (OT OH) X (RW 9'¢ —¢L) ‘doy ‘wog ‘uy oqeiny (OTD x x x (RIN 9S quasaye — GL) ‘day “wog ‘wy oqeindy x x x (Od) 99UaLINIDO IsIY [RQOID (RIN DI-6'1) (PN (BRIN €-9'¢) Ie 91-61) SoITy jap seuloT oul souong -jeduiq 0 (eI €-9'¢) arr] -O5 OUD oO (L661) UOsuYyor puke ppng pur (9861) 7) J2 ppng (8661) 72 J2 ppng = ovSeinD ‘(ssoid ul) URWAary pue ppng = seweYyRg :sadInos I (RW b-¢E'S) Jequinu 092g ozeig SLI cLl SILI IZI OL! 691 891 991 S Pol S091 8ST di satoads GOD S9DU9LINIIO ISP] jo Joquinu [R10], $39U9.IN550 ISI jo Jaquinu [R10], soiseds SuIAT] % peynuapr satsads jo Joquinu [R10], vy ds DIDISYSD{ (x)W ‘ds 18aad DUDIYIADUID] xosaf puppuvp av19D («)a “ds uly “YW “ye psojnsuv piyynusngq DIMUsSNy pyyjAydojankp pypAydojaod py pyypAydojaodp pyyjdydojaokp piyyjAydojakp pypAydojaoAp vaslsp]jAydos] piynusissnpy DSSnW satoads ‘ponunuoy— Pf a1qeL 136 BULLETIN 357 12 10 Number of species 2 4 6 8 10 12 14 16 18 20 22 24 26 Number of localities per species Number of species 8 24 40 56 72 88 104 120 136 152 168 Number of specimens per species 160 140 ” ‘Oo 2D 12 - O° 100 SES w On Soo ZG8 “ 20 0 0 5 10 15 20 25 Number of localities per species Text-figure 6.—Histograms and scatterplot showing the numbers of localities and specimens collected per species. branching colonies indicate intermediate depths with moderate wave action on exposed forereefs. Exclu- sively branching species indicate muddy conditions with reduced wave action. Abundant free-living colo- nies indicate unstable substrates, often associated with seagrass flats. Following Done (1983) and Geister (1983), high diversities can be interpreted to indicate moderately exposed forereef environments at shallow to intermediate depths (5-20 m) on open marine lee- ward platforms; whereas low diversities indicate pro- tected environments, such as shallow (<5 m) platform or deepest (40-100 m) shelf areas, or highly exposed (<5 m) windward reefs. Recent species with narrow depth ranges (Goreau and Wells, 1967) that were identified in our collections include three shallow (<10 m) reef crest species (Ac- ropora palmata, Diploria strigosa, Colpophyllia amaranthus) and four deeper (>20 m) forereef species (Agaricia lamarcki, Mussa angulosa, Stephanocoenia intersepta, Madracis decactis). Of these, Diploria stri- gosa may also occur at intermediate (10—20 m) depths, and the four deeper forereef species may also occur at shallower depths under turbid conditions. Therefore, the presences of indicator species should be interpreted with caution. Of the shallow reef crest indicators, Ac- ropora palmata is perhaps the most definitive (see McNeill et al., 1997). To evaluate colony shapes of the collected corals, each species identified in the collections was assigned to one of four colony shape categories (Table 3), and percentages of species with different shapes at each lo- cality were compared among reef trends. Following Johnson et al. (1995), the four categories consist of: branching (19 species, 439 specimens), free-living (10 species, 133 specimens), massive and encrusting (43 species, 1100 specimens) [hereafter termed ‘massive’ ], and platy (11 species, 702 specimens). Species exhib- iting more than one colony shape were assigned to the colony shape category that they most frequently pos- sess. Comparisons among trends indicate that species and specimens of platy corals are more frequent in localities in the Lomas del Mar trend; branching corals are more frequent in the Buenos Aires trend; and massive corals are more frequent in the Empalme trend (Table 5; Text- fig. 7). In general, corals in the less-collected older trends (the Brazo Seco patch and Quebrada Chocolate trend) appear to have colony shapes most like the Buen- os Aires trend, although free-living corals are more common in the Brazo Seco patch (Text-fig. 7). The differences in colony shapes among trends sug- gest that environmental conditions may have differed among trends. However, the high numbers of species collected in most localities suggest that most assem- blages formed in exposed reef environments at shallow to intermediate depths (<30 m). The high percentages of platy corals in the Lomas del Mar trend suggest low light intensities, and thus either deep reef (30—40 m) = Text-figure 7.—Maxima, medians, and minima of percentages of species with branching, free-living, massive, and platy colony shapes within each of five reef trends. The trends are arranged in chronological order from oldest (left) to youngest (right). BS, Brazo Seco patch; QC, Quebrada Chocolate trends; BA, Buenos Aires trend; EM, Empalme trend (including Santa Rosa patch); LM, Lomas del Mar trend. 137 BUDD ET AL. REEF CORALS: ) ¢ - Ayyeoo) sed suewpeds bulyoueg % i oS — ' ¢ Sa) Se aeesS Ayye00| Jed seeds Bulyoueig % BS QC BA EM LM BS QC BA EM LM ® & 2 © Aye90) Jed suewpeds BulAl|-ee14 % be | _ ae ' e.c8 S55 Ayiye00| 16d sepeds BHulal|-ea4 % BS QC BA EM LM BS QC BA EM LM Se Sa See fap Ay)}290| 19d suewjoeds eAIsseW] % be | — — 8 8 8 F< & — Ay\je00] 16d saioeds anisseyy % BS QC BA EM LM BS QC BA EM LM — Ll bo] ' ¢ See) one. aan Aye00| Jad sueweds Ajeiq % I+] _ le ¢ ' Bi 3 las Ay\je00] 18d saeds Ayeid % BS QC BA EM LM BS QC BA EM LM 138 BULLETIN 357 Table 5.—Results of non-parametric tests comparing colony shape frequencies and % living species among the Limon reef trends. Kruskal-Wallis test Correc- ted Species Chi- Mann-Whitney subgroup Datatype Square df p-value U test results branching — species 16.987 4 0.002 BA >EM>LM specimens 19.360 4 0.001 BA >EM>LM free-living species 4.835 4 0.305 BA = EM = LM specimens 4.673 4 0322 BA =EM=LM massive species 9.687 4 0.046 EM > BA = LM specimens 11.696 4 0.020 EM > BA = LM platy species 20.526 4 0.000 LM > BA = EM specimens 20.475 4 0.000 LM > BA = EM % living species 11.625 4 0.024 BA = EM = LM; BA < LM or muddy environments at intermediate depths (10—30 m). The high percentages of branching corals in the Buenos Aires trend (and possibly the Quebrada Choc- olate trends) suggest moderate exposure and shallow to intermediate water depths (<20 m). The high per- centages of massive corals in the Empalme trend and associated Santa Rosa patch suggest exposed shallow conditions (<10 m) with high wave energies. The high percentage of free-living corals at the Brazo Seco patch suggests a shallow (<10 m) unstable substrate. Shallow-water indicators are common or abundant in localities within all five trends, and Acropora palmata is common or abundant at localities within the three younger trends. Nevertheless, deep-water indicators are common or abundant only in the Lomas del Mar trend, suggesting deep forereef environments. Thus, a mix of reef environments may be involved within each trend; and assignment of uniform depths to individual trends may be an over-simplification. In summary, hermatypic corals indicate that reef en- vironments changed from moderately exposed shallow and intermediate depth environments (Quebrada Choc- olate and Buenos Aires trends), to exposed shallow environments (Empalme trend), to deep forereef en- vironments (Lomas del Mar trend) within the Limon sequence through geologic time. Preliminary compar- isons with microfossil and ahermatypic coral data col- lected in nearby sites generally support the interpre- tations based on hermatypic corals. Assemblages of benthic foraminifera suggest water depths of 50—100 m for the Lomas del Mar trend (Collins et al., 1995) and 10—30 m for the Empalme trend (Collins in Jack- son et al., this volume). Ahermatypic corals suggest water depths of 40-73 m for the Lomas del Mar trend (Cairns, this volume). As mentioned above, more de- = So So te) o hi % Living species per locality 3 > So ——_- _ _». BS QC BA EM LM Text-figure 8.—Maxima, medians, and minima of percentages of living species within each of five reef trends. The trends are arranged in chronological order from oldest (left) to youngest (right). BS, Brazo Seco patch; QC, Quebrada Chocolate trends; BA, Buenos Aires trend; EM, Empalme trend (including Santa Rosa patch); LM, Lomas del Mar trend. tailed comparisons between hermatypic corals and os- tracodes are planned in future analyses. EVOLUTIONARY SIGNIFICANCE OF THE COLLECTED TAXA Survey of the 73 Neogene to Recent Caribbean stratigraphic units in the 1996 Cenozoic Coral Data- base (CCD) compiled by Johnson and Budd (Budd and Johnson, 1997) indicates that unusually high numbers of global first and last occurrences of species occur within the Limon sequence (Table 4). A total of 39 first occurrences and 22 last occurrences takes place in the sequence. Among the first occurrences are those for 32 of the 60 species that currently live in the Ca- ribbean. Several of these first occurrences are for im- portant modern reef dominants, including Acropora palmata (see McNeill et al., 1997), A. cervicornis, Porites astreoides, P. furcata, Diploria strigosa, Mon- tastraea faveolata, and M. franksi. Last occurrences take place in many species that are abundant or com- mon in the Mio-Pliocene of the Cibao Valley of the Dominican Republic (Budd et al., 1996), including two species of Stephanocoenia, three species of Por- ites, and four species of Montastraea. Closer examination of the first and last occurrence information (Table 4) shows that first occurrences take place in all five Limon reef trends, although the high- est numbers of first occurrences are in the Buenos Ai- res trend and, to a lesser extent, the Lomas del Mar trend (Table 4). In fact, six of the seven modern reef dominants listed in the paragraph above first occur in the Buenos Aires trend. Two of these six (Porites as- treoides, Diploria strigosa) also occur at approximate- ly the same time in the Pinecrest Sandstone of Florida. In contrast, almost all of the observed last occurrences are concentrated in the Lomas del Mar trend. REEF CORALS: BUDD E7 AL. 139 Table 6.—Occurrences of species of Stylophora, Acropora, and Caulastraea within collections. PPP Locality S. : S. monti- A. cervi- A. C. porto- number code Reef trend affinis granulata — S. minor culosa cornis palmata ricensis 1381 BR Brazo Seco x Xx 1386 C2 Q. Chocolate Xx 1384 C4 Q. Chocolate x X x 1316 BA2 Buenos Aires x Xx x Xx 1362 BA2 Buenos Aires X Xx x 1388 BA2 Buenos Aires Xx 1346 BA3 Buenos Aires X x 1125 BA3 Buenos Aires x x x 1345 BA4 Buenos Aires Xx 1364 BA4 Buenos Aires Xi 1348 BA6 Buenos Aires x x 1124 BA6 Buenos Aires x x x 1342 BA7 Buenos Aires x x x x 1343 BA7 Buenos Aires X x x x x 1344 BA7 Buenos Aires x x x 1366 BA7 Buenos Aires x x x 1341 BA8 Buenos Aires x x x 1428 El Empalme x 1340 E2 Empalme x 1374 E3 Empalme x 719 SRI Santa Rosa x X x 1373 SR2 Santa Rosa x x 1499 SR2 Santa Rosa x x 1500 SR2 Santa Rosa x x 1351 SR3 Santa Rosa x 1372 SR3 Santa Rosa x x 1367 SR4 Santa Rosa x x x 1349 LE Lomas del Mar X Te? LE1 Lomas del Mar x 646 LE2 Lomas del Mar X 1106 LE9 Lomas del Mar x 1375 LW1 Lomas del Mar x 1331 LW2 Lomas del Mar 1335 Pl Lomas del Mar 1336 Pl Lomas del Mar x 1337 Pl Lomas del Mar 1338 Pl Lomas del Mar xX x 1339 Pl Lomas del Mar 1353 jew) Lomas del Mar x 1126 |e) Lomas del Mar x 1127 Pe Lomas del Mar x x 1355 P3 Lomas del Mar x Of the 82 species that occur in the Limon reef trends, 61 species (74.4%) originated within the past 11 million years (Table 4). Of these 61 species, 12 originated during the Late Miocene time (11.2—5.3 Ma), and five originated during Earliest Pliocene time (5.3—4 Ma). Twenty-seven of the remaining 41 species (65.8%) originated between 4—3 Ma, and 14 originated over the past two million years. Thus, approximately one-third of the fauna appears to have originated dur- ing a one-million year peak of origination at 4—3 Ma. In contrast, 29 of the 33 extinct species in the fauna (87.9%) became extinct during a one-million year peak of extinction at 2-1 Ma. These calculations (see Budd and Johnson, 1997, for further discussion of evolu- tionary rates) suggest that accelerated origination pre- ceded accelerated extinction in these corals by 1—2 million years and, together with field observations, they indicate that members of the ‘pre-turnover’ (i.e., Mio-Pliocene) and ‘post-turnover’ (i.e., Recent) Carib- bean reef coral faunas co-existed within the Limon se- quence though the critical interval of faunal change on Caribbean reefs. Percentages of living species collected within each trend range from <40% in the two older reef trends to >70% in the two younger trends, and thus further sup- port the transitional interpretation for the fauna. Per- 140 BULLETIN 357 Table 7.—Occurrence matrix of species of 27 Costa Rica and 3 Panama localities. Species are in taxonomic order; localities are arranged by reef trend. Occurrences are coded relative to abundance: ‘R’ = rare, ‘C’ = common, ‘A’ = abundant, ‘F’ = super-abundant. CCD species ID Genus Species number BS C4 BAI BA2 BA3 BA4 BA6 BA7 BA8 E3_ SRI SR2_ SR3 Stephanocoenia _ intersepta 2 R Stephanocoenia — duncani 3 A R (e R Cc Cc (e Stephanocoenia — spongiformis 4 Stylophora affinis 5 Cc Stylophora granulata 1 (Cc iG Cc (C Stylophora minor 9 Cc (c R Stylophora monticulosa 10 R cS Cc (S ic R Pocillopora crassoramosa 15 F Madracis asperula 16.5 Madracis decactis 17 R Madracis mirabilis 20 R R Madracis pharensis 21 Madracis sp. A 2S) Cc Acropora cervicornis 22 € 18) A Ec A A Cc Cc Acropora palmata 23 A A (e A (e Cc (e Agaricia grahamae 29 Agaricia lamarcki 30 Agaricia undata 32 Undaria agaricites 33 A Cc A Ec A ec (e R A A Undaria crassa 34 Cc ec (E Cc (S Undaria pusilla 35 R R Helioseris cucullata 3 & R (S (e R (cc Siderastrea radians 56 Cc Siderastrea siderea 58 G! Cc € (cS A c Porites astreoides 63 R (e R A (e R (E (e Porites portoricensis 65 R (e Cc (S Porites waylandi 68 Porites baracoaensis 69 R (@ Cc R (€ Porites branneri 70 Porites colonensis 73 Porites furcata 76 Ie! (e (G cC A (© R A Porites porites Wil ‘ec Cc Caulastraea portoricensis 83 (c € A A (c A (c Favia fragum 88 R (S iS ic (S Diploria clivosa 94 R Cc R (e ( R Diploria labyrinthiformis 95 R (C A Diploria strigosa 97 (e Cc (E R (S A ¢€ R (e Manicina areolata 100 (e (e R (C (E Manicina mayort 101 Cc Manicina puntagordensis 102 cc Thysanus sp. A 109 Gc R Ee Gc GC Thysanus corbicula 111 Colpophyllia amaranthus A: R (C Colpophyllia natans 114 (o Cc (e A R R Colpophyllia sp. A 114.5 R Cc Montastraea faveolata MN9/ Cc Cc (c Montastraea franksi 118 A E Cc R Montastraea limbata-| 121.1 (e (e R Montastraea limbata-2 121.2 (e (S A ic E ¢ Cc Montastraea sp. A 122. (cc ie c G R Cc Montastraea canalis 124 R Montastraea cavernosa-2 126 (S Cc R (e ie Montastraea cavernosa-3 127 E R Montastraea cylindrica 128 (S Solenastrea bournoni 131 (S A A REEF CORALS: BUDD ET AL. 141 Table 7.—Continued. No. of Costa No. of Rica Panama local- — local- SR4 LE] LE2 LE3 LE4 LES LE? LE8 LES LEIO LW2 LW3 Pi P2 HP PA FH ities ities Cc Cc (ec (@ (S Cc SG A Ee R 11 0 (@ Cc A (G A A A A A (© Cc © (cS 19 1 @ R R 3 0 © 1 1 A 4 1 3 0 Cc 6 1 Cc 2 1 R Cc Cc ‘Ee A 5) 0 Cc A A (C Ec A (c Cc R (e 11 1 A A Cc A Cc Cc Cc Cc c 11 0 R 1 0 1 0 (C (ce R Cc R Cc 14 1 A R R ‘Ee iE Cc 12 1 (Cc In A A A Cc Cc A A Cc Cc 11 0 (c A A A (Cc R Cc A A (e (S Cc 12 0 (O A (C Cc Cc R A A A Cc 10 0 F F IE A A A A A A A A A FE A € (c 23 3} (C (e R R A (c 10 1 R R 4 0 Cc R (© R R R (e R (e (e 14 2 € 2 0 Cc R Cc Cc Cc R (cS R Cc Cc 15 I Cc Cc R (Cc ie A R (e Cc 15 2; Cc 5 0 R 1 0 Cc 6 0 Cc Cc Cc R R 5) 0 Cc Cc (e Cc 4 0 A cc 10 0 2 0 Cc € Cc R A 11 1 iS 0 (e 7 0 Cc (e Cc 5 1 A Cc R R R A A A (e 16 2 R ( R R Cc (S 10 1 (c (e R (E (cc Cc 7 0 R Cc R Cc € (G 6 1 5) 0 CG 0 1 2 0 (e R R (C (S (cc R Cc A 14 I R Cc R A (C 5 2 (e Cc (S R Cc (C A A ( A 11 2 R Cc R cS Cc Cc Cc 10 1 R 4 0 (c Cc (C (S A R Cc 14 0 (e R 8 0 (Cc (G (c R Cc R 7. 0 (© Cc Cc A R Cc € Cc 13 0 Cc R R R 6 0 (e (C (C A A A A A (c (ce R 12 0 Cc Cc 5 0 142 Table 7.—Continued. BULLETIN 357 BA2 BA3 BA4 BA6 BA7 BA8 £3 SRI SR2_ SR3 CCD species ID Genus Species number BS C4 Antillophyllia sawkinsi 137 Meandrina braziliensis 138 Meandrina meandrites 139 Meandrina sp. A 1395 Placocyathus trinitatis 143 (C Placocyathus variabilis 144 A R Dichocoenia caloosahatcheensis 145 Dichocoenia eminens 146 (e Dichocoenia stokesi 148 Dichocoenia stellaris 149 Dichocoenia tuberosa 150 R R Archohelia limonensis 152.5 Scolymia cubensis 155 Scolymia lacera 157 Mussa angulosa 158 Mussismilia aff. M. hartti 160.5 Isophyllastrea sp. B 164.5 Mycetophyllia aliciae 166 Mycetophyllia danaana 168 Mycetophyllia ferox 169 Mycetophyllia lamarckiana 170 Mycetophyllia reesi 171 Mycetophyllia sp. A ES Eusmilia fastigiata 173 Eusmilia sp. A 175 e Number of species 80 1S 18 E R (Cc (Se (S (C Cc R R R R E R R R R centages of living species collected within each locality are statistically lower in the Brazo Seco and Quebrada Chocolate trends than in the Buenos Aires, Empalme, and Lomas del Mar trends (Table 5, Text-fig. 8). Also supporting the transitional interpretation are the co-occurrences of species of Stylophora, Acropora, and Caulastraea at individual collection sites, and within localities (Table 6). Stylophora and Caulas- traea, two genera that are now extinct in the Carib- bean, dominated shallow and intermediate depth reef environments in pre-turnover faunas; whereas Acro- pora has dominated these same environments in post- turnover faunas (Budd and Kievman, in press). Much of the shift between these two distinctly different sets of community dominants takes place in the Late Pli- ocene and Early Pleistocene, between 4—1 Ma. How- ever, Acropora palmata, the species that sometimes dominates modern Caribbean reef crests, does not be- come extremely abundant in reef coral assemblages until the Late Pleistocene (Jackson, 1994; Jackson and Budd, 1996). Four species of Stylophora, two species of Acropora, and one species of Caulastraea occur at a total of 42 of the 107 collection sites and 24 of the 34 localities (Table 6). Species of Stylophora occur at 10 collection sites (7 localities) in the three older reef trends; species of Acropora occur at 30 collection sites (18 localities) in the four younger trends; and the one species of Cau- lastraea occurs at 25 collection sites (16 localities) in the three younger trends. Species of Stylophora and Ac- ropora co-occur at seven sites (PPP 1125, 1316, 1342, 1343, 1346, 1362, 1388) and five localities (BA2, BA3, BA4, BA6, BA7) in the Buenos Aires trend; Acropora palmata itself co-occurs with species of Stylophora at three Buenos Aires sites (PPP 1125, 1342, 1343). Spe- cies of Caulastraea and Acropora co-occur at 16 sites and 12 localities within the three younger trends. These co-occurrences support the notion that dominant mem- bers of pre- and post-turnover faunas lived side by side in the same environment. ASSEMBLAGE ANALYSES OCCURRENCE MATRIX We used the specimen database to assemble an oc- currence matrix (78 species X 27 localities) containing codes for relative abundances (Table 7). Counts of specimens were obtained for species within each lo- REEF CORALS: BUDD ET AL. 143 Table 7.—Continued. No. of Costa No. of Rica Panama local- local- SR4S UE SILE2) 1bE3 E44 EES LET) LES® LES EEO) EwWw2 Iswsis Pl P2 HP PA FH ities ities R 1 0 A 0 1 R Cc R € 4 1 E Cc (C A 2 2; 1 0 (e Cc R Cc (cc R Cc F 9 1 1 0 (e 1 i (G R R 8 0 (C R (C e R R R R 9 0 Cc (e 5 0 Cc 1 0 (c (e R 3 0 R 1 0 R R R Cc 6 0 Cc 3 0 1 0 R R 2 0 R (EC (eS (e (C (S c 6 1 R R (e R (Cc (e (S 7 1 R R Cc 3 0 Cc Cc (G: C Cc Cc Cc R 8 0 R R 2 0 R R Cc 4 0 R 2 0 12 20 29. 11 28 25 24 20 30 38 19 11 31 16 15 13 10 524 38 cality, and codes for rare, common, abundant, and su- per-abundant were assigned using a modified version of the ‘proportion of species’ method described by Gaston (1994). In this procedure, beginning with the lowest specimen counts for a given locality, approxi- mately 25% or less of the species were designated as rare, 50% or more were designated as common, and 25% or less were designated as abundant. Only in cas- es where the highest count exceeded the next highest count by two times were species designated as super- abundant. In counting specimens, specimens with identification confidence codes of <25% were counted as only one-half. If the total specimen count for a spe- cies within a locality was only one-half, the species was deleted from the data set. CLASSIFICATION (CLUSTER ANALYSIS) To determine if the assemblages could be separated into discrete groups, we first analyzed the occurrence matrix using average linkage cluster analysis (SPSS for Windows, version 6.1, 1994). We performed ana- lyses for both localities (Q-mode) and species (R- mode), and used both relative abundance (frequency count) codes and presence-absence (binary) data. When analyses were performed with relative abun- dance codes, the Phi-square coefficient was used. As explained by Shi (1993) and Hayek (1994), the Phi coefficient is a traditional measure of association, sim- ilar to a chi-square statistic but normalized relative to frequency so that it is less affected by sample size. When analyses were performed with binary data, the Lance and Williams coefficient was used. The Lance and Williams coefficient is similar to a Dice or Bray- Curtis similarity coefficient; both give more weight to joint presences and exclude joint absences. The results using relative abundance codes (Text- fig. 9) suggest four clusters of localities [two large clusters (I and II) and two small clusters (III and IV)] and eight clusters of species [(four large clusters (A to D) and four small clusters (E to H)]. Locality cluster I consists of 10 localities within the Lomas del Mar trend; locality cluster II consists of 12 localities within the Buenos Aires trend, the Lomas del Mar trend, and the Santa Rosa patch of the Empalme trend; locality cluster III consists of two localities within the Buenos Aires trend that have low sample sizes; and locality cluster IV consists of the two localities in the two strat- igraphically older trends. The one locality within the 144 BULLETIN 357 Localities Species LE10 aagn00},00|I00 Bowaal> >>r>oar>>>> >aOrO0F>F>|O0 >rQOQgNANFN>Y nm Q200F7F00 zany >rOrYyrrKrTNOO OFOF>9N00BBOSI 2N9F0OBBOHOVNANVANBWO F = ee >>oO} 0g900090200000 0200 REEF CORALS: BUDD E7 AL. 145 Empalme trend proper does not belong to any of the four clusters. Removal of locality cluster I (the Lomas del Mar trend) from the analysis results in exactly the same pattern of relationships among localities within locality cluster II; thus, the unexpected grouping of localities from three stratigraphically separate reef trends in locality cluster II appears to be robust. The results using relative abundance codes (Text- fig. 9) further indicate that the similarities among spe- cies are low; therefore, clusters can only be defined at very high levels. Several species are abundant or com- mon in more than one locality cluster, implying con- siderable overlap in species composition among local- ity clusters. Locality cluster I is characterized by spe- cies clusters B and E, which contain low numbers of extinct species (5 out of 19) that are predominantly platy and massive. Locality cluster II is characterized by species clusters D, E G, and H, which contain slightly higher numbers of extinct species (7 out of 19) that are predominantly branching and massive. Lo- cality cluster IV is characterized by species cluster A, which contains a high proportion of extinct species (13 out of 19). The single Empalme branch appears to be characterized by species cluster C; however, species within cluster C are common throughout all four lo- cality clusters. Locality cluster III is composed of two localities with low sample sizes and does not seem to correspond with any species clusters. Q-mode results using binary data reveal exactly the same locality clusters as those found using relative abundance codes; however, R-mode results using bi- nary data show important differences among species clusters. Most notably, species previously belonging to species cluster C no longer group together and are scattered across the dendrogram. In general, the large amount of overlap in species composition among locality clusters and the instability of the R-mode results indicate that distinct clusters and associations among species do not exist within these data, and that the assemblages are not discrete. Distinct clusters would be expected if species within commu- nities responded similarly either to short-term changes in the local environment or to long-term evolutionary changes. When localities in the deep forereef trend (Lomas del Mar) are removed and only shallow and intermediate environments are included in the analysis, the clusters continue to overlap, indicating that evo- lutionary changes in the fauna during this interval were not simultaneous. These results agree with the origination and extinction data of the previous section (Table 4) and the increasing percentages of living spe- cies from older to younger reef trends (Text-fig. 8), and they provide additional support for the interpre- tation that the fauna was transitional during the two million year interval between 3.6—1.6 Ma. ORDINATION (DETRENDED CORRESPONDENCE ANALYSES) In order to determine the major directions of vari- ation among the assemblages in the sequence and to search further for environmental and evolutionary gra- dients of faunal change in the data, we analyzed the occurrence matrix using a linear ordination technique known as ‘detrended correspondence analysis’ (PC- ORD, version 2.0, McCune and Mefford, 1995). The purpose of ordination is to produce a representation of the data in low-dimensional space, in which similar species and samples are close together and dissimilar entities are far apart. The resulting axes are interpreted using independent environmental and evolutionary data. We selected a linear ordination technique, de- trended correspondence analysis (DCA), over another commonly used nonlinear ordination technique, non- metric multidimensional scaling (NMDS), because DCA is more effective at revealing linear gradients in the data (Gauch, 1982). DCA has the added advantage of using chi-square distances (metric values) and not rank-order dissimilarities (as in NMDS). Unlike DCA, NMDS preserves relative and not absolute dissimilar- ities. Furthermore, DCA simultaneously ordinates lo- calities and species in 3-dimensional space, thus alle- viating problems associated with choice in numbers of axes and axis interpretation (Shi, 1993). DCA is a form of reciprocal averaging in which species ordination scores are averages of sample or- dination scores and, reciprocally, sample ordination scores are averages of species ordination scores. The procedure is iterative and begins with arbitrary species ordination scores, which in turn are used to calculate sample ordination scores. The sample ordination scores are then used to obtain species ordination scores. Iterations are continued until the scores stabi- lize. Detrending is applied to sample scores at each iteration to rescale the axes and correct for arch effects (Gauch, 1982). —_ Text-figure 9 —Q-mode (localities) and R-mode (species) cluster analysis of relative abundance code data. Clusters of localities (labeled ‘I through ‘IV’) and clusters of species (labeled ‘A’ through ‘H’) are defined on the basis of their descriptive utility and not on the basis of cutoff levels. Abbreviations for localities are given in Table 1. Occurrences are coded as ‘R’ for rare, ‘C’ for common, ‘A’ for abundant, and ‘F’ for super-abundant. 146 BULLETIN 357 0 100 200 300 AXIS 1 Text-figure 10.—Scatterplots of DCA scores for 27 Costa Rican localities determined using relative abundance code data. Each point represents a locality, the abbreviations for which are given in Table 1. Shaded areas encompass localities within the Buenos Aires (BA), Empalme (EM), and Lomas del Mar (LM) reef trends. The Lomas del Mar localities are divided into two subsets based on cluster anal- ysis results: (1) 10 tightly clustered localities (LE2, LE3, LE4, LES, LE7, LE8, LE9, LE10, LW2, LW3) that are scattered across the top of the topographic ridge formed by the Lomas del Mar trend, and (2) three localities (LE1, P1, P2) along the northern margin of that ridge. The results of DCA analyses using relative abun- dance codes (Text-fig. 10) suggest that, with the ex- ception of locality E3 and three northern localities within the Lomas del Mar trend (LEI, Pl, P2), the localities within the three better-sampled reef trends group together. The two clusters of localities within the Lomas del Mar trend are separated by a pro- nounced discontinuity; whereas localities within the Buenos Aires trend and Santa Rosa patch appear to overlap slightly. The Quebrada Chocolate locality is isolated in the plot of axis 1 vs. axis 2, but groups with the Buenos Aires locality in the plot of axis 1 vs. axis 3. These findings are consistent with those of the cluster analyses (Text-fig. 9). Visual examination of the two DCA plots (Text-fig. 10) suggests that axis 1 corresponds with relative stratigraphic position and geologic age. Possible ex- ceptions exist for locality E3 and the three northern localities within the Lomas del Mar trend. Neverthe- less, older localities generally have high values along axis 1; whereas younger localities tend to have low values. DCA scores for species (Text-fig. 11) are also generally higher for extinct species than for living spe- cies along axis 1. In contrast, axes 2 and 3 appear to bear no relationship to stratigraphic position or to numbers of extinct vs. living species. Thus, axis | ap- pears to be partially related to time; whereas axes 2 and 3 do not. The distributions of branching, massive, and platy species along the three DCA axes (Text-fig. 11) sug- gest that: (1) on axis 1, platy species tend to have low values, branching species have high values, and mas- sive species are more evenly distributed; (2) on axis 2, platy species tend to have high values, whereas branching and massive species have low values; and (3) on axis 3, platy, branching, and massive species are haphazardly scattered. These results suggest that environment may explain some of the variation along axes 1 and 2. The low values for platy species along axis 1 suggest that muddy or deep water conditions may be responsible for the tight cluster of Lomas del Mar localities on the extreme left sides of the plots in Text-figure 10. The low values for massive and branching species along axis 2 suggest that axis 2 may be more closely related to exposure and distance to shore, with more nearshore localities having low val- ues. Unlike the other localities in the analysis, the two localities with the lowest values along axis 2 (BAI, BA4) are both Porites thickets, which today are most common in shallow nearshore areas. None of the three axes appear to be related to the number of specimens or species collected within each locality. Thus, both axes 1 and 2 appear to be somewhat related to the environment, but axis 3 does not. The distribution of localities along axis 1 in the DCA analysis based on binary data (Text-fig. 12a) is generally similar to that for relative abundances (Text- fig. 10). The main exceptions are: (1) the gap between REEF CORALS: BUDD ET AL. 147 Number of species 0 -200 -100 0 100 200 300 400 500 AXIS 1 Number of species fa) a: We | ee Ti: -200 -100 0 100 200 300 400 AXIS 2 Number of species Number of species 0 -200 -100 0 100 200 300 400 500 AXIS 1 Number of species 0 -200 -100 0 100 200 300 400 500 AXIS 2 Number of species 0 -200 -100 0 100 200 300 400 500 -200 -100 0 AXIS 3 AXIS 3 Text-figure 11.—Histograms of DCA scores for species determined using relative abundance code data. The analysis is the same as Text- fig. 10. Species are grouped by colony shape (left side) and by survivorship (right side). “b’, branching; ‘m’, massive; ‘p’, platy, “E’, extinct; ‘L, living. the two clusters of Lomas del Mar localities is less distinct, and (2) there is less overlap among the Santa Rosa and Buenos Aires localities. Even with these ex- ceptions, axis 1 still appears to be related to time. The distribution of localities along axis 2, however, is somewhat different from the relative abundance re- sults; the two stratigraphically older localities (BS, C4) no longer have relatively high values, localities with more massive species are no longer distinct from lo- calities with more platy species, and the two localities in the Buenos Aires Porites thickets (BA1, BA4) are separated from the others by a large gap. Thus axis 2 appears to distinguish the two nearshore environments but is less clearly related to exposure. As in the relative abundance results, axis 3 is uninterpretable. In order to better understand how the 10 tightly clustering localities from the Lomas del Mar trend af- fected the results, a final analysis was performed after these 10 localities were removed. The analysis was run using relative abundance codes. The results (Text-fig. 12b) show even stronger overlap among the Buenos Aires, Santa Rosa, and three remaining Lomas del Mar localities along axis | as well as considerable overlap among the Buenos Aires and Santa Rosa localities along axis 2. Thus, as in the cluster analysis results, the assemblages appear to intergrade, and although variation among assemblages appears to be coarsely related to evolution and environment, more refined re- lationships are difficult to decipher. 148 BULLETIN 357 0 100 200 300 400 AXIS 1 Text-figure 12.—Scatterplots of DCA scores for Costa Rican lo- calities determined: (A) using binary data for 27 localities, (B) using relative abundance code data after the 10 tightly clustered localities in the Lomas del Mar trend were removed from the data set. Each point represents a locality, the abbreviations for which are given in Table 1. As in Text-figure 10, shaded areas encompass localities within the Buenos Aires (BA), Empalme (EM), and Lomas del Mar (LM) reef trends. PRELIMINARY COLLECTIONS FROM THE BOCAS DEL TORO REGION OF PANAMA During August 1993, small collections of corals were made at seven sites in the Bocas del Toro region of Panama (Table 1; Appendix 1; Appendix A): five on Isla Colon (68 specimens, 26 species), one on Isla Bas- timentos (21 specimens, 11 species), and one on Swan Cay (6 specimens, 6 species). Preliminary age estimates based on planktic foraminifera and nannofossils (Coates, this volume) are: 3.5—1.7 Ma for the sites on Isla Colon, 3.0—2.2 Ma for the site on Isla Bastimentos, and 1.6—1.2 Ma for the site on Swan Cay. Study of the 1:50000 Isla Colon (3744 III, Edicion 1-AMS, 1993) and the 1:50000 Bocas del Toro (3744 II, Edicion 1- AMS, 1993) topographic sheets and stratigraphic sec- tions HP, FH, SC (Coates, this volume) indicates that the five sites on Isla Colon could be grouped into three localities: GC (PPP 1285), PA (PPP 1423, 1424), and HP (PPP 1260, 1425). Based on this grouping, five lo- calities (HP, FH, SC, PA, GC) are represented in the Bocas del Toro collections (Table 1). Numbers of spec- imens collected per locality range from 6 to 32 (median = 21), and numbers of species collected per locality range from 2 to 18 (median = 11). A total of 35 species (12 extinct, 23 living) belong- ing to 19 genera were identified in the Bocas del Toro collections (Table 3). Only one species [Thysanus cor- bicula; first occurrence = Chipola Fm, Florida (18-15 Ma), last occurrence = Old Pera Beds, Jamaica (2.5— 1.8 Ma)], which occurred on Isla Bastimentos (FH), was recognized that was not found in the Limon col- lections. At the three Isla Colon localities, only two extinct branching species were found at Ground Creek (GC), 18 species (83.3% living) were found at Hill Point (HP), and 14 species (92.8% living) were found at Paunch (PA). In contrast, the fauna collected on Isla Bastimentos (FC) consisted of 11 species (27.2% liv- ing), and the collection made at Swan Cay (SC) con- tained 6 species (100% living). The fauna at Hill Point (HP) is distinctive among the Bocas del Toro collec- tions in that it contains both Acropora palmata and species of Stylophora. The Isla Bastimentos fauna con- tains species of Stylophora, but lacks Acropora. The Swan Cay fauna contains only Acropora palmata, but lacks Stylophora. Thus, collections from Isla Basti- mentos (FH) and Ground Creek (GC) consist primarily of the pre-turnover fauna, whereas those from Paunch (PA) and Swan Cay (SC) consist primarily of the post- turnover fauna. The Hill Point (HP) fauna consists of a mix of the pre- and post-turnover faunas. Most of the species collected at Hill Point (HP) and Paunch (PA) had massive colony shapes; whereas those collected at Isla Bastimentos (FH) possessed a variety of branching, free-living, massive, and platy shapes. The small collection at Swan Cay (SC) con- tained species with branching, massive and _ platy shapes. The apparently high diversity at all but the Ground Creek (GC) locality suggests that the assem- blages lived in open reef habitats, with the most ex- posed conditions occurring at Hill Point (HP) and Paunch (PA). To further compare the Bocas del Toro assemblages REEF CORALS: BUDD E£T AL. 149 300 200 0 100 200 300 200 AXIS 1 Text-figure 13.—Scatterplots of DCA scores for 27 Costa Rican and 3 Panamanian localities determined using relative abundance code data. Each point represents a locality, the abbreviations for which are given in Table 1. Panamanian localities are indicated by black dots. As in Text-figure 10, shaded areas encompass localities within the Buenos Aires (BA), Empalme (EM), and Lomas del Mar (LM) reef trends. with the Limon assemblages, DCA was performed on the three Bocas del Toro localities with more than 10 specimens (HP, PA, FH) and the 27 Limon localities using relative abundance code data. Codes for the three Bocas del Toro assemblages (Table 7) were determined by the same methods as described above for the Limon analyses. The results (Text-fig. 13) reveal a similar configuration of clusters of Limon localities as found earlier for just the 27 Limon localities using relative abundance codes (Text-fig. 10). The Isla Bastimentos (FH) assemblage lies in an isolated portion of the axis 1 vs. axis 2 plot, closest to assemblages from the two older Limon reef trends (Quebrada Chocolate, Brazo Seco). It therefore can be interpreted as composed of a pre-turnover fauna that differs from the Limon pre- turnover fauna probably because of the environment. The Hill Point (HP) assemblage lies near the line of overlap between clusters of Buenos Aires trend local- ities and Santa Rosa patch localities, and appears to have a similar mixed pre- and post-turnover compo- sition. The Paunch (PA) assemblage lies closest to the Empalme trend locality and the three northern locali- ties within the Lomas del Mar trend. It therefore can be interpreted as composed of a post-turnover fauna in a shallow reef environment similar to portions of the Lomas del Mar trend. CONCLUSIONS Our initial collecting efforts in the Limon and Bocas del Toro basins have clearly shown that coral reef communities were plentiful and diverse along the Ca- ribbean coast of Costa Rica and Panama during the 1— 3 million year interval of Plio-Pleistocene turnover in the reef coral fauna of the Caribbean region. In fact, the diversity of reef corals in the collected fossil as- semblages exceeds that reported in the region today, as well as at other individual locations in the Carib- bean during Plio-Pleistocene time. Although patchily distributed through geologic time, fossil assemblages in the two basins provide a wealth of data for explor- ing and comparing pathways of faunal replacement and community change. In the Limon area, evidence for faunal change be- gins to appear between 3.6—3 Ma (the Quebrada Choc- olate and Buenos Aires reef trends) and has almost ended by 1.9—1.6 Ma (the Empalme and Lomas del Mar trends). Similarly, in Curacao, faunal change be- gins between 5.6—3 Ma and has almost ended by 2.6— 2 Ma (Budd ef al., 1998), and in Jamaica, faunal change has ended in exposed shallow reef environ- ments by 2—1.8 Ma (Budd and McNeill, 1998). In con- trast, in Bocas del Toro, faunal change may not have begun until after 3—2.2 Ma in some places (e.g., Isla Bastimentos), but was complete by 1.6—1.2 Ma (Swan Cay) or even earlier (Paunch). This delay in turnover has also been observed in protected reef margin en- vironments in Jamaica (the Bowden-Old Pera se- quence), in which faunal change did not begin until after 3.3-1.8 Ma (Budd and McNeill, 1998). These results indicate that the timing and pace of faunal change may have varied from place to place across the Caribbean region as a whole, and along the Caribbean coast of Costa Rica and Panama in particular. Transi- 150 BULLETIN 357 tional faunas in which extinct species of Stylophora and living species of Acropora co-occur can be found in both the Limon (Buenos Aires reef trend, 3.6—3 Ma) and Bocas del Toro (Hill Point, 3.5—1.7 Ma) areas, suggesting that faunal change may have occurred ei- ther gradually or in a series of steps in both areas. To interpret the significance of these results, the ob- served patterns of faunal change need to be further refined, and to do this, clearly more samples need to be analyzed. In particular, large collections that are well-documented geographically, environmentally, and stratigraphically need to be collected at more sites throughout the Bocas del Toro area, and in Early Pli- ocene reefal deposits west of Limon. More precise en- vironmental interpretations are needed for individual localities within trends to effectively tease apart the effects of environment and evolution in faunal analys- es. Because examining co-occurrences of species has been found to be critical to understanding patterns of faunal change, the present analyses demonstrate the need for collections containing at least 50 specimens per site. Line transect or quadrat data documenting co- occurrences and relative abundances of species in the field, together with detailed taphonomic information, could further help to explain the variability in what preliminary results suggest to be a complex system. In addition, comparisons with patterns of change in other fossil groups could potentially assist in pinpointing and understanding the larger-scale external factors driving Plio-Pleistocene faunal change, as well as the apparently staggered and patchy response of reef com- munities to these factors. In addition to tracing patterns of faunal change, our initial collections are, for the most part, exceptionally well-preserved, and contain many global first occur- rences. They are therefore ideal for future morpho- metric studies of speciation in the fossil record, and for reconstructing phylogenies. For example, the two species of Acropora that dominate modern Caribbean reefs both have first occurrences in the Buenos Aires trend (McNeill er al., 1997), and so do the oldest mem- bers of the equally important Montastraea annularis sibling species complex (Knowlton et al., 1992). What was the pattern of speciation in these two clades, and what sort of morphologic innovations were involved? Did speciation within each clade involve an initial evo- lutionary burst or were speciation events staggered through time? What sorts of ecological conditions were associated with speciation? Collections such as those described herein provide an important starting point in searching for answers. REFERENCES CITED Budd, A.F. 1991. Neogene Paleontology in the northern Dominican Repub- lic. 11. The Family Faviidae (Anthozoa: Scleractinia). Part I. The Genera Montastraea and Solenastrea. Bulle- tins of American Paleontology, vol. 101, no. 338, pp. 5— 83, pls. 1-29. Budd, A.F., and Johnson, K.G. 1997. Coral reef community dynamics over 8 million years of evolutionary time: Stasis and turnover. Proceedings of the 8th International Coral Reef Symposium, vol. 1, pp. 423— 428. 1999. Neogene Paleontology in the Northern Dominican Re- public. 19. The Family Faviidae (Anthozoa: Scleractinia), Part II. Bulletins of American Paleontology, no. 356, pp. 5-83, pls. 1-21. Budd, A.F., Johnson, K.G., and Edwards, J.C. 1989. Miocene coral assemblages in Anguilla, BWI, and their implications for the interpretation of vertical succession on fossil reefs. Palaios, vol. 4, pp. 264—275. Budd, A.F., Johnson, K.G., and Potts, D.C. 1994b. Recognizing morphospecies in colonial reef corals: 1. Landmark-based methods. Paleobiology, vol. 20, pp. 484-505. Budd, A.F., Johnson, K.G, and Stemann, T.A. 1996. Plio-Pleistocene turnover and extinctions in the Caribbean reef coral fauna. in Evolution and Environment in Trop- ical America. J.B.C. Jackson, A.F Budd, and A.G. Coates, eds., University of Chicago Press, Chicago, pp. 168-204. Budd, A.F., and Kievman, C.M. In press. Coral assemblages in Neogene to Recent cores from the Bahamas platform and their use in paleoenvironmental interpretation. SEPM special publication on the Bahamas Drilling Project. Budd, A.F., and McNeill, D.F. 1998. Zooxanthellate Scleractinian Corals from the Bowden Shell Bed, SE Jamaica. Contributions to Tertiary and Quaternary Geology, vol. 35, pp. 49-65. Budd, A.F., Petersen, R.A., and McNeill, D.F. 1998. Stepwise faunal change during evolutionary turnover: a case study from the Neogene of Curagao, Netherlands An- tilles. Palaios, vol. 13, pp. 167-185. Budd, A.F., Stemann, T.A., and Johnson, K.G. 1994a. Stratigraphic distributions of genera and species of Neo- gene to Recent Caribbean reef corals. Journal of Paleon- tology, vol. 68, pp. 951-977. Collins, L.S., Coates, A.G., Jackson, J.B.C., and Obando, J.A. 1995. Timing and rates of emergence of the Limon and Bocas del Toro Basins: Caribbean effects of Cocos Ridge Sub- duction? in Geologic and tectonic development of the Ca- ribbean Plate Boundary in Southern Central America. P. Mann, ed., Geological Society of America Special Paper 295, Boulder, CO, pp. 263-289. Done, T.J. 1983. Coral zonation: its nature and significance. in Perspectives on coral reefs. D.J. Barnes, ed., Australian Institute of Marine Science, Brian Clouston Publisher, Manuka, A.C.T., pp. 107-147. Foster, A.B. 1986. Neogene paleontology in the northern Dominican Repub- lic. 2. The family Poritidae (Anthozoa: Scleractinia). Bul- REEF CORALS: letins of American Paleontology, vol. 90, no. 325, pp. 47— 123, pls. 15-38. 1987. Neogene paleontology in the northern Dominican Repub- lic. 4. The genus Stephanocoenia (Anthozoa: Scleractinia: Astrocoeniidae). Bulletins of American Paleontology, vol. 93, no. 328, pp. 5—22, pls. 1-7. Foster, A.B., Johnson, K.G., and Schultz, L.L. 1988. Allometric shape change and heterochrony in the free- living coral Trachyphyllia bilobata (Duncan). Coral Reefs, vol. 7, pp. 37-44. Gaston, K.J. 1994. Rarity. Chapman & Hall, London, 205 pp. Gauch, H.G., Jr. 1982. Multivariate analysis in community ecology. Cambridge University Press, Cambridge, 298 pp. Geister, J. 1983. Holozane westindische Korallenriffe: Geomorphologie, Okologie und Fazies. Facies, vol. 9, pp. 173-284. Goreau, T.F. 1959. The ecology of Jamaican coral reefs. Part I, Species com- position and zonation. Ecology, vol. 40, pp. 67-90. Goreau, T.F., and Wells, J.W. 1967. The shallow-water Scleractinia of Jamaica: Revised list of species and their vertical distribution range. Bulletin of Marine Science, vol. 17, pp. 442-453. Graus, R.R., and Macintyre, I.G. 1989. The zonation pattern of Caribbean coral reefs as con- trolled by wave and light energy input, bathymetric set- ting and reef morphology: Computer simulation experi- ments. Coral Reefs, vol. 8, pp. 9-18. Hayek, L.C. 1994. Analysis of amphibian biodiversity data. in Measuring and monitoring biological diversity: standard methods for amphibians. W.R. Heyer, M.A. Donnelly, R.W. Mc- Diarmid, L.C. Hayek, and MLS. Foster, eds., Smithsonian Institution Press, Washington, D.C., pp. 207-269. Jackson, J.B.C. 1994. Community Unity? Science, vol. 264, pp. 1412-1413. Jackson, J.B.C., and Budd, A.F. 1996. Evolution and environment: introduction and overview. in Evolution and Environment in Tropical America. J.B.C. Jackson, A-.F Budd, and A.G. Coates, eds., University of Chicago Press, Chicago, pp. 1—20. Jackson, J.B.C., Budd, A.F., and Pandolfi, J.M. 1996. The shifting balance of natural communities? in Evolu- tionary Paleobiology. D. Erwin, D. Jablonski, and J. Lipps, eds., University of Chicago Press, Chicago, pp. 89-122. James, N.P. 1984. Reefs. in Facies Models. R.G. Walker, ed., Geological As- sociation of Canada, Toronto, pp. 229-244. Johnson, K.G., and Budd, A.F. 1996. Three-dimensional landmark techniques for the recogni- tion of reef coral species. in Advances in Morphometrics. L.F Marcus, M. Corti, A. Loy, D. Slice, and G. Naylor, eds., NATO ASI Series, v. A284, Plenum, New York, pp. 345-353. Johnson, K.G., Budd, A.F., and Stemann, T.A. 1995. Extinction selectivity and ecology of Neogene Caribbean reef corals. Paleobiology, vol. 21, pp. 52-73. Johnson K.G., and McCormick, T. 1995. STATPOD: Statistical Analysis of Palaeontological Oc- BUDD EFT AL. 151 currence Data, Version 0.1. Computer Program distributed by the Department of Geology and Applied Geology, Uni- versity of Glasgow, UK. Knowlton, N., Weil, E., Weigt, L.A., and Guzman, H.M. 1992. Sibling species in Montastraea annularis, coral bleaching, and the coral climate record. Science, vol. 255, pp. 330— 333. McCune, B., and Mefford, M.J. 1995. PC-ORD, Multivariate Analysis of Ecological Data, Ver- sion 2.0. MJM Software Design, Gleneden Beach, Oregon, 126 pp. MeNeill, D.F., Budd, A.F., and Borne, P.F. 1997. An Earlier (Late Pliocene) First Appearance of the Reef- building Coral Acropora palmata: Stratigraphic and evo- lutionary implications. Geology, vol. 25, pp. 891-894. McNeill, D.F., Coates, A.G., Budd, A.F., and Borne, P.F. In press. Stratigraphy of Late Neogene reefs and siliciclastics of Limon, Costa Rica: A coastal emergence record of the Central American Isthmus. Geological Society of Amer- ica Bulletin. Pandolfi, J.M. 1996. Limited membership in Pleistocene reef coral assemblag- es from the Huon Peninsula, Papua New Guinea. Paleo- biology, vol. 22, pp. 152-176. Ricklefs, R.E. 1990. Ecology, third edition. W.H. Freeman, New York, 896 pp. Saunders, J.B., Jung, P., Geister, J., and Biju-Duval, B. 1982. The Neogene of the south flank of the Cibao Valley, Do- minican Republic: a stratigraphic study. Transactions of the 9th Caribbean Geological Conference (Santo Domin- go, 1980), vol. 1, pp. 151-160. Scoffin, T.P. 1987. An Introduction to Carbonate Sediments and Rocks. Blackie, Glasgow, 274 pp. Shi, G.R. 1993. Multivariate data analysis in palaeoecology and palaeo- biogeography—a review. Palaeogeography, Palaeoclima- tology, Palaeoecology, vol. 105, pp. 199-234. Stemann, T.A. 1991. Evolution of the reef-coral family Agariciidae (Anthozoa: Scleractinia) in the Neogene through Recent of the Carib- bean. Unpublished Ph. D. dissertation, University of lowa, Iowa City, Iowa, 321 pp. In press. Neogene Paleontology in the Northern Dominican Re- public. The Family Agariciidae (Anthozoa: Scleractinia). Bulletins of American Paleontology. Stemann, T.A., and Johnson, K.G. 1992. Coral assemblages, biofacies, and ecolgical zone in mid- Holocene reef deposits of the Enriquillo Valley, Domini- can Republic. Lethaia, vol. 25, pp. 231-241. Swedberg, J.L. 1994. Systematics and distribution of the scleractinian coral Madracis in the Miocene to Pleistocene of Tropical Amer- ica. Unpublished M.S. thesis, University of Iowa, Iowa City, IA, 114 pp. Vaughan, T.W. 1919. Fossil corals from Central America, Cuba, and Porto Rico with an account of the American Tertiary, Pleistocene, and Recent coral reefs. U.S. National Museum Bulletin, vol. 103, pp. 189-524, pls. 68-152. Wells, J.W. 1973. New and old scleractinian corals from Jamaica. Bulletin of Marine Science, vol. 23, pp. 16-58. APPENDIX 1 SPECIES OCCURRENCES Species identified at PPP sites. Data are at the PPP internet site, http://www. fiu.edu/~collinsl/. 639 646 715 719 771 772 942 943 948 949 962 963 1100 1101 1102 1103 1104 1105 Acropora cervicornis 1 1 Acropora palmata Acropora sp. Agaricia grahamae 3 25 Agaricia lamarcki 3 Agaricia undata 1 Antillia dentata Antillophyllia sawkinst 1 Archohelia limonensis 1 1 Caulastraea portoricensis Colpophyllia amaranthus Colpophyllia natans 1 Colpophyllia sp. A Dichocoenia caloosahatcheensis 1 Dichocoenia eminens Dichocoenia stokesi 2 1 1 1 Dichocoenia stellaris 1 Dichocoenta tuberosa Diploria clivosa Diploria labyrinthiformis Diploria sarasotana Diploria strigosa 1 Eusmilia fastigiata 1 1 1 Eusmilia sp. A Favia fragum Goniopora imperatoris Helioseris cucullata Isophyllastrea sp. B Madracis asperula 1 Madracis decactis 4 Madracis mirabilis 1 7 Madracis pharensis 1 Maadracis sp. A 1 Manicina areolata 1 1 Manicina mayori 1 Manicina puntagordensis 2 Meandrina braziliensis Meandrina meandrites 1 Meandrina sp. A 1 Millepora complanata 1 Millepora sp. 1 Montastraea canalis Montastraea cavernosa-2 1 1 Montastraea cavernosa-3 1 1 Montastraea cylindrica 1 4 1 3 4 1 Montastraea faveolata 1 1 1 1 Montastraea franksi 1 1 1 Montastraea limbata-| Montastraea limbata-2 4 Montastraea sp. A Mussa angulosa 1 Mussismilia aff. M. hartti 1 Mycetophyllia aliciae 1 Mycetophyllia danaana Mycetophyllia ferox 1 Mycetophyllia lamarckiana 1 Mycetophyllia reesi 1 2 2 1 1 1 2 Mycetophyllia sp. A Pocillopora crassoramosa Porites astreoides 1 1 1 Porites baracoaensts Porites branneri 5 1 1 Porites colonensis 3 Porites furcata 1 Porites porites Porites portoricensis Porites waylandi 1 1 1 Porites sp. Placocyathus trinitatis Placocyathus variabilis Scolymia cubensis Scolymia lacera Siderastrea radians Siderastrea siderea 1 1 1 Solenastrea bournont 1 Stephanocoenia intersepta 1 4 Stephanocoenia duncani 6 Stephanocoenia spongiformis Stylophora affinis Stylophora granulata Stylophora minor Stylophora monticulosa Thysanus corbicula Thysanus sp. A Undaria agaricites 12 29 Undaria crassa Undaria pusilla Undaria sp. agariciid Total specimens 28 1 NN Ny Nr a oon as) one — ho mt) Nee wes BNR i) wWew nN Noe is) b NN eye aS NV Nu Nv Nn N Nv KH wp N ty i) NN Ne Ne fw Nw oO hw eo) 16 6 2 3 tN eN w es i=} So = 00 Ww an w w N an nN w oS 28 2 1 146 96 55 7 21 11 22 21 REEF CORALS: BUDD ET AL. 153 Appendix 1|.—Continued. LOGS UO 7;, S108 AUTOS ST OFF A a2 IS AS) SG 7 SE SS) 20 1121) 122) 138s aneA ais 3 3 1 10 3 2) 1 2 2 5 5 1 4 1 5 3 1 2 2 2 l 1 18 1 1 l 2 l r 3 1 2 2 l 1 2 1 1 1 1 2 l l 2 3 3 4 2 1 1 2 1 4 1 2 I l 1 7 l 1 1 1 1 1 I 1 1 3 5 1 5 2 2 5 6 1 2 2 2 2 1 2 2 2 1 2 2 1 3 1 3 1 1 3 1 1 1 1 2 8 2 l 1 6 l 1 2 l 1 1 1 1 2} 14 1 1 1 1 1 2 1 2 1 2 1 3 1 1 1 3 1 3 1 1 1 1 4 4 2 3 l 3 3 2 5 1 2 3 1 2 1 3 1 1 1 1 1 2 5 5 1 5 2 4 3 1 12 1 3 9 7 1 1 2 10 154 BULLETIN 357 Appendix 1|.—Continued. I 1126 1127 1251 1260) 1285. 1310! 1316 1331 11332 11333) 13834 "11335, 1336) 1337) 1388 Acropora cervicornis 1 1 Ul 1 1 Acropora palmata 2 1 1 1 1 Acropora sp. Agaricia grahamae Agaricia lamarcki Agaricia undata Antillia dentata Antillophyllia sawkinst Archohelia limonensis Caulastraea portoricensis 1 3) 1 Colpophyllia amaranthus Colpophyllia natans 1 1 1 2 3 1 Colpophyllia sp. A 1 1 Dichocoenia caloosahatcheensis Dichocoenia eminens 1 Dichocoenia stokesi 1 Dichocoenia stellaris 1 Dichocoenia tuberosa 1 Diploria clivosa Diploria labyrinthiformis 2 Diploria sarasotana Diploria strigosa 3 1 1 1 3 4 Eusmilia fastigiata Eusmilia sp. A 1 1 1 1 Favia fragum Goniopora imperatoris Helioseris cucullata 3} Isophyllastrea sp. B Madracis asperula Madracis decactis 1 Madracis mirabilis Madracis pharensis Madracis sp. A Manicina areolata 1 Manicina mayori 1 1 1 Manicina puntagordensis 1 2 Meandrina braziliensis Meandrina meandrites 1 1 Meandrina sp. A 1 1 3 Millepora complanata Millepora sp. Montastraea canalis 1 Montastraea cavernosa-2 1 1 Montastraea cavernosa-3 Montastraea cylindrica 1 4 3 Montastraea faveolata 3 1 Montastraea frankst 1 5 4 Montastraea limbata-| Montastraea limbata-2 4 1 3 1 1 Montastraea sp. A 5 1 1 Mussa angulosa 1 Mussismilia aff. M. hartti 1 Mycetophyllia aliciae Mycetophyllia danaana 1 1 1 Mycetophyllia ferox Mycetophyllia lamarckiana Mycetophyllia reesi 1 1 Mycetophyllia sp. A Pocillopora crassoramosa 1 Porites astreoides 1 Porites baracoaensis Porites branneri 1 Porites colonensis Porites furcata 1 Porites porites Porites portoricensis 1 1 Porites waylandi Porites sp. Placocyathus trinitatis Placocyathus variabilis 8 Scolymia cubensis Scolymia lacera 1 Siderastrea radians Siderastrea siderea 1 1 2 1 1 NNw Ann w a) i) Nu we i) i) te i) i) ey = is) - & tN na N N oN i) ‘= a i) nN Solenastrea bournoni Stephanocoenia intersepta Stephanocoenia duncani 1 Stephanocoenia spongiformis Stylophora affinis 1 Stylophora granulata 4 4 Stylophora minor Stylophora monticulosa 2 1 Thysanus corbicula 1 Thysanus sp. A 1 Undaria agaricites 16 1 2 4 7 1 5 Undaria crassa 2 Undaria pusilla 1 Undaria sp. agariciid Total specimens NS nN i) n nN n Ww wn 21 12 6 45 41 16 33 19 30 22 26 14 13 REEF CORALS: BUDD ET AL. 155 Appendix 1.—Continued. 1339 1340 1341 1342 1343 1344 1345 1346 1347 1348 1349 1350 1351 1352 1353 1355 1357 1358 1359 1360 1362 10 4 2 1 2 3 3 1 2 1 1 1 1 l 2 1 1 1 1 2 1 4 1 1 1 1 1 1 2 2 1 1 4 1 1 1 2 1 1 1 1 1 3 3 1 1 1 1 1 1 1 1 2, 1 2 1 1 1 1 1 1 2 5 1 1 1 1 1 1 1 2 1 1 1 1 1 1 2 1 2 1 1 2 1 2 1 1 5 1 4 1 1 2 7 1 1 1 1 1 1 1 1 1 1 1 1 9 1 1 2; 3 1 1 1 3 3 1 1 3 1 1 1 1 3 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 3 5 1 1 3 1 1 1 2 5 1 1 1 1 1 1 1 1 1 1 3 3 2 1 6 4 5 2 2 3 1 1 2 1 1 1 1 1 5 4 21 35 21 69 9 C) 3 20 12 2 36 6 7 4 @) 13 6 1 2) 156 BULLETIN 357 Appendix 1.—Continued. ee 1363 1364 1365 1366 1367 1370 1372 1373 1374 1375 1381 1384 1385 1386 1387 Acropora cervicornis 3 2 3 4 2 Acropora palmata 1 2 3 Acropora sp. 1 Agaricia grahamae 3 Agaricia lamarcki 1 1 Agaricia undata 1 1 Antillia dentata Antillophyllia sawkinst Archohelia limonensis Caulastraea portoricensis 4 3 3 3 1 1 Colpophyllia amaranthus Colpophyllia natans 4 1 Colpophyllia sp. A 1 Dichocoenia caloosahatcheensis Dichocoenia eminens 1 Dichocoenia stokesi 1 Dichocoenia stellaris 1 Dichocoenia tuberosa Diploria clivosa 1 1 Diploria labyrinthiformis Diploria sarasotana 1 Diploria strigosa 3 1 1 Eusmilia fastigiata Eusmilia sp. A 5 Favia fragum 1 1 1 Goniopora imperatoris 1 Helioseris cucullata 4 1 Isophyllastrea sp. B Madracis asperula 1 Madracis decactis Madracis mirabilis 1 Madracis pharensis Madracis sp. A Manicina areolata 3 Manicina mayor 1 1 1 Manicina puntagordensis 1 1 Meandrina braziliensis 1 Meandrina meandrites 1 Meandrina sp. A Millepora complanata 1 Millepora sp. 1 Montastraea canalis 1 1 Montastraea cavernosa-2 1 1 Montastraea cavernosa-3 Montastraea cylindrica Montastraea faveolata 1 1 Montastraea franksi 1 1 Montastraea limbata-\ Montastraea limbata-2 Montastraea sp. A Mussa angulosa Mussismilia aff. M. hartti Mycetophyllia aliciae Mycetophyllia danaana Mycetophyllia ferox Mycetophyllia lamarckiana Mycetophyllia reesi Mycetophyllia sp. A Pocillopora crassoramosa 17 1 1 Porites astreoides Porites baracoaensis Porites branneri Porites colonensis Porites furcata 2 Porites porites Porites portoricensis 2 1 Porites waylandi 1 Porites sp. 1 Placocyathus trinitatis Placocyathus variabilis Scolymia cubensis Scolymia lacera Siderastrea radians Siderastrea siderea 1 6 Solenastrea bournoni 3 9 Stephanocoenia intersepta 1 1 Stephanocoenia duncani 1 3 5 1 Stephanocoenia spongiformis Stylophora affinis 2 Stylophora granulata Stylophora minor 4 Strylophora monticulosa 1 2 Thysanus corbicula Thysanus sp. A 3 1 Undaria agaricites 1 1 Undaria crassa 2) 2 2 Undaria pusilla 1 Undaria sp. agariciid Total specimens 2 6 2 35 23 2 20 34 18 7 57 66 12 10 10 SSS eS 5858608000 nN w Nw — is) w is) N i) rey -N N w i) ws) Nv Ww Ne nN i) a N eal fo.) aN w Now w REEF CORALS: BUDD ET AL. SSI Appendix 1.—Continued 1388 1389 1390 1410 1412 1413 1414 1415 1416 1423 1424 1425 1426 1427 1428 1499 1500 1971 1972 2002 14 1 1 1 1 1 1 2 1 6 1 4 2 1 2 2 6 1 1 1 1 1 2 2 5 1 1 1 1 1 1 1 1 1 2 1 3 1 1 1 1 1 2 3 1 1 1 1 1 3 1 1 1 1 1 1 5 1 1 1 1 2 1 1 1 1 1 1 1 2 1 6 2 1 1 1 3 1 2 2 1 1 1 1 1 1 2 2 1 1 1 3 1 1 1 1 1 1 4 1 1 1 2 1 1 1 1 5 3 2 1 1 3 1 1 1 2 1 2 3 3 1 7 1 1 1 1 1 1 1 1 4 3 1 2 1 2 2 1 1 1 1 4 1 2 1 2 1 1 1 1 1 2 1 1 1 2 1 1 2 2 2 1 1 1 1 1 3 3 3 4 1 1 2 1 1 1 1 1 1 1 1 1 3 1 3 1 3 3 2 3 1 1 1 1 1 2 5 3 1 1 5 3 1 2 2 1 2 2 3 3 1 1 1 37 63 11 11 15 14 14 19 24 28 5) 18 9 ) 9 17 12 24 24 6 158 BULLETIN 357 Appendix 1.—Continued. 2005 2006 2007 2008 2009 2010 2011 Acropora cervicornis Acropora palmata Acropora sp. Agaricia grahamae 1 1 1 Agaricia lamarcki 1 Agaricia undata 1 1 1 1 1 Antillia dentata Antillophyllia sawkinsi Archohelia limonensis Caulastraea portoricensis Colpophyllia amaranthus Colpophyllia natans 1 1 Colpophyllia sp. A 1 1 Dichocoenia caloosahatcheensis Dichocoenia eminens Dichocoenia stokest Dichocoenia Stellaris Dichocoenia tuberosa 1 Diploria clivosa Diploria labyrinthiformis Diploria sarasotana Diploria strigosa Eusmilia fastigiata 1 1 Eusmilia sp. A Favia fragum Goniopora imperatoris Helioseris cucullata 1 Isophyllastrea sp. B Madracis asperula 2 1 Madracis decactis 2 Madracis mirabilis 1 Madracis pharensis Madracis sp. A Manicina areolata 1 1 Manicina mayori 1 Manicina puntagordensis Meandrina braziliensis Meandrina meandrites Meandrina sp. A Millepora complanata Millepora sp. Montastraea canalis Montastraea cavernosa-2 Montastraea cavernosa-3 Montastraea cylindrica Montastraea faveolata Montastraea franksi Montastraea limbata-\ Montastraea limbata-2 1 Montastraea sp. A Mussa angulosa Mussismilia aff. M. hartti Mycetophyllia aliciae Mycetophyllia danaana 1 Mycetophyllia ferox Mycetophyllia lamarckiana Mycetophyllia reesi 3 Mycetophyllia sp. A Pocillopora crassoramosa Porites astreoides 1 1 Porites baracoaensis Porites branneri 1 Porites colonensis Porites furcata Porites porites Porites portoricensis Porites waylandi Porites sp. Placocyathus trinitatis Placocyathus variabilis 1 1 1 1 Scolymia cubensis Scolymia lacera Siderastrea radians Siderastrea siderea 1 1 1 Solenastrea bournoni Stephanocoenia intersepta 3) Stephanocoenia duncani 1 1 22 1 Stephanocoenia spongiformis 1 Stylophora affinis Stylophora granulata Stylophora minor Stylophora monticulosa Thysanus corbicula Thysanus sp. A Undaria agaricites 1 5) 1 1 3} 1 3 Undaria crassa Undaria pusilla Undaria sp. agariciid Total specimens 18 24 6 10 15 9 21 UU tttIttIIIEIISIIISSSSSSSSSSSSSStStSttSa nN ES i) nN NV tN i) nN CHAPTER 8 NEOGENE CHEILOSTOME BRYOZOA OF TROPICAL AMERICA: COMPARISON AND CONTRAST BETWEEN THE CENTRAL AMERICAN ISTHMUS (PANAMA, COSTA RICA) AND THE NORTH-CENTRAL CARIBBEAN (DOMINICAN REPUBLIC) ALAN H. CHEETHAM Department of Paleobiology National Museum of Natural History, Smithsonian Institution Washington, D.C. 20560-0121, U.S.A. JEREMY B. C. JACKSON Smithsonian Institution, Smithsonian Tropical Research Institute Washington, D.C. 20560-0580, U.S.A., and University of California at San Diego, Scripps Institution La Jolla, California 92093-0244, U.S.A. JOANN SANNER Department of Paleobiology, National Museum of Natural History Smithsonian Institution Washington, D.C. 20560-0121, U.S.A. YIRA VENTOCILLA Smithsonian Institution, Smithsonian Tropical Research Institute Washington, D.C. 20560-0580, U.S.A. INTRODUCTION The Neogene (Miocene—Pleistocene) deposits of tropical America have long been known to contain a rich fossil record of cheilostome Bryozoa (Canu and Bassler, 1918, 1919, 1923, 1928), but detailed quan- titative study of its diversity, spatial and temporal dis- tribution, and evolutionary significance has only re- cently become possible. Extensive collections (Text- fig. 1) from detailed stratigraphic sequences in Panama and Costa Rica (Panama Paleontology Project, PPP; Coates et al., 1992; Jackson et al., 1996; Coates, this volume), and from the Dominican Republic (DR; Saunders et al., 1982; Saunders et al., 1986) provide a new basis for this undertaking. The opportunity now exists to document species ranges in space and time more precisely, and thus to explore the relationships between the cheilostome fauna and major environmen- tal changes, such as those associated with the shoaling and final emergence of the Central American isthmus and closure of the isthmian seaway (Duque-Caro, Atlantic Ocean 2 DR SS BE. S Caribbean Sea ning: South America ae Pacific Ocean Text-figure 1—Sketch map of the Caribbean and adjacent areas showing general locations of the PPP and DR collecting sites. 160 BULLETIN 357 Table 1.—Diversity and abundance of cheilostome bryozoans (total and by growth form) in Panama Paleontology Project (PPP) collections containing six or more species each. Number of species Abundance Collection Age Paleodepth Free- Free- number (Ma) (m) Total Encrusting Erect living Total Encrusting Erect living Canal Basin PPP 222 11.6 275 6 6 0 0 15 15 0 0 PPP35 9.6 2iES 8 2 6 0 35 2 33 0 PPP 162 8.6 25.0 22 13 2 iT 40 13 2 25 Bocas del Toro Basin PPP’ 391 Sii/ 150.0 9 4 1 4 18 4 1 13 PPP 60 4.3 60.0 9 6 1 2 9 6 1 2 PPP 201 4.3 60.0 6 1 0 5) 15S 1 0 14 PPP 203 4.3 42.5 8 2 0 6 251 11 0 240 PPP 204 4.3 42.5 16 9 0 7 169 18 0 151 PPP 205 4.3 42.5 28 20 1 7 271 47 1 223 PPP 206 4.3 42.5 21 13 2 6 471 58 2 411 PPP 207 4.3 42.5 14 6 1 7 527 6 1 520 PPP 208 4.3 42.5 7 1 0 6 241 1 0 240 PPP 419 4.3 50.0 7 2 0 5 34 11 0 23 PPP 422 4.3 50.0 13 5 1 7 166 5 1 160 PPP 423 4.3 50.0 8 0 1 Y 152 0 10 142 PPP 425 4.3 50.0 6 0 0 6 51 0 0 SI PPP 426 4.3 50.0 6 0 0 6 420 0 0 420 PPP 64 3.6 60.0 14 7 2 5 41 7 2 32 PPP 65 3.6 60.0 45 29 7 9 243 74 16 153 PPP 66 3.6 60.0 33 23 2 8 627 23 2 602 PRPs6)/ 3.6 60.0 14 6 0 8 617 0 611 PPP 294 3.6 60.0 25 15 2 8 97 15 2 80 PPP 295 3.6 60.0 28 16 4 8 415 52 13 350 PPP 298 3.6 60.0 27 1S 4 8 432 60 22 350 PPP 306 3.6 60.0 14 6 0 8 356 6 0 350 PPP 307 3.6 60.0 20 11 1 8 443 20 1 422 PPP 308 3.6 60.0 25 14 1 10 83 50 1 532 PPP 311 3.6 60.0 9 3 0 6 234 3 0 231 PPP SIZ 3.6 60.0 13 5 2 6 40 5 2 33 PPP 326 3.6 60.0 16 8 2 6 52 8 2 42 PPP 334 3.6 60.0 27 14 4 9 531 167 22. 342 PPP 335 3.6 60.0 20 10 1 9 191 37 1 153 PPP 340 3.6 60.0 25 13 3 9 234 51 3 180 PPP 341 3.6 60.0 16 6 2 8 74 17 2 S55 PPP 345 3.6 42.5 19 9 3 7 109 27 12 70 PPP 346 3.6 42.5 18 8 3 7 81 17 3 61 PPP 348 3.6 42.5 19 7 3} 9 91 16 12 63 PPP 349 3.6 42.5 14 2 4 8 77 2 13 62 PPP 350 3.6 42.5 29 14 4 11 677 113 22 542 PPP 352 3.6 42.5 40 27 3 10 751 189 12 550 PPP 354 3.6 42.5 13 6 1 6 40 6 1 33 PPP'355) 3.6 42.5 16 7 2 Y 439 7 2 430 PPPS 3.6 11 1 1 9 164 1 1 162 PPP 365 3.6 125.0 12 1 4 7 75 10 13 52 PPP 367 3.6 125.0 27 11 7 9 342 56 16 270 PPP 368 3.6 125.0 24 9 4 11 114 18 13 83 PPP 370 3.6 125.0 19 6 4 9 190 15 4 171 PPPS 355 li 1 2 4 16 10 2 4 PPP 55 3.5 8 4 0 4 26 22 0 4 PPPISi/ 3.5 30.0 10 0 1 9 154 0 1 153 PPP 63 35 30.0 11 4 1 6 236 4 1 231 PPP 68 3.5 48 23 15 10 1488 482 285 721 PPP 69 335 36 21 5 10 1107 273 nS Wel CHEILOSTOME BRYOZOA: CHEETHAM ET AL. 161 Table 1.—Continued. Number of species Abundance Collection Age Paleodepth Free- Free- number (Ma) (m) Total Encrusting Erect living Total Encrusting Erect living PPP 71 3hD) 7 1 0 6 25 1 0 24 PPP 72 Sh) i 0 1 6 52 (0) 1 51 PPP 74 3.5 12 1 2 g) 345 10 2 333 PPP 193 335 30.0 11 3 1 7 344 3 10 331 PPP 194 3.5 30.0 14 5) 1 8 365 5 10 350 PPP 195 35) 30.0 9 1 1 7 315 1 1 313 PPP 196 3.5 30.0 10 4 0 6 46 4 0 42 PPP 197 35 30.0 8 1 1 6 53 1 10 42 PPP 198 35) 30.0 18 7 1 10 380 16 10 354 PPP 210 3)5) 45 23 13 9 873 365 247 261 PPP 211 35 20 8 10 2 263 17/ 136 110 PPP 212 3 31 15 8 8 445 60 35 350 PER S379 325) 175.0 9 3 0 6 63 3 0 60 PPP 358 2.8 125.0 7 1 1 5 25 1 1 23 PPP 361 2.8 125.0 8 3 1 4 35 5 10 22 PPP 362 2.8 125.0 18 4 2 12 288 13 2 273 leled2) IU 7/ 2.1 125.0 6 2 0 4 15 2 0 13 PPP 178 Da 125.0 14 5 2 7 77 5 2 70 PPP 214 2.0 10 4 2 4 136 4 2 130 Limon Basin PPP 679 3.5 30.0 16 2 8 6 475 20 134 321 PPP 683 355 30.0 i 1 2 4 124 1 2 121 PPP 695 35) 30.0 6 1 0 5 33 1 0 32 PPP 704 3)5) 30 16 Wl 7 201 34 34 133 PPP 705 35) 13 5 3 5 49 5 3 41 PPP 708 335 25 6 11 8 484 15 119 350 PPP 709 Be) 30.0 53 Sil 13 9 1196 202 364 630 PPP 720 3.5 30.0 48 20 18 10 696 56 279 361 PPP 722 35 30.0 44 19 15 10 1079 2 465 442 PPP 723 3.5 30.0 42 19 13 10 1347 172 454 721 PPP 932 3.5 12.5 14 2 6 6 239 2 24 213 PPP 933 3.5 12.5 10 3 2 5 226 3 2 221 PPP 935 3.5 12.5 6 0 2 4 42 0 11 31 PPP 937 B15) 12S 17 8 3 6 179 26 3 150 PPP 939 35) 9 1 3 5 126 1 3 122 PPP 940 35 8 0 3 5 26 0 3 23 PPP 697 3) 30.0 11 2 3 6 164 2 12 150 PPP 663 3)-I 5.0 27 12 12 3 378 21 345 12 PPP 668 Sal 30.0 11 3 1 7 146 3 1 142 PPP 669 Syl 30.0 8 2 0 6 44 2 0 42 PPP 670 3.1 30.0 18 10 1 7 243 10 1 232 PPP 671 Bal 30.0 8 4 1 3 7 4 1 lp PPP 672 Sal 30.0 7 3} 1 3 25 3 1 21 PPP 689 3.1 30.0 7 0 2 5 43 0 2 41 PPP 691 Bal 30.0 9 1 2 6 234 1 2 231 PPP 180 2.8 125.0 21 8 6 7 183 35 6 142 PPP 634 Nog 61.5 45 31 9 5 378 94 153 131 PPP 635 e7/ 61.5 41 24 9 8 860 186 234 440 PPP 639 ey 61.5 49 37] 4 8 283 190 13 80 PPP 640 -7/ 61.5 44 29 8 7 1052 29 233 610 PPP 642 7/ 61.5 39 26 8 5 S61 323 233 5 PPP 644 ley 61.5 26 15 6 5 44 24 15 5 PPP 645 i7/ 61.5 30 20 7 3 102 47 52 3 PPP 710 Ne7/ 61.5 31 20 6 >) 94 56 6 32 PPP 653 1.6 200.0 25 13 8 4 97 31 35 31 PPP 738 1.6 29 18 8 3 164 27 125 12 PPP 943 1.6 61.5 35 21 10 4 224 66 136 22 Table 1.—Continued. BULLETIN 357 Number of species Abundance Collection Age Paleodepth Free- Free- number (Ma) (m) Total Encrusting Erect living Total Encrusting Erect living PPP 944 1.6 61.5 2 33 12 7 1645 1059 246 340 PPP 948 1.6 61.5 38 20 11 u 497 191 254 52 PPP 949 1.6 61.5 59 45 10 4 608 171 433 4 PPP 950 1.6 61.5 58 42 10 6 958 546 361 51 PPP 962 1.6 61.5 D2 33 13 6 484 159 274 51 PPP 963 1.6 61.5 56 40 11 5 551 283 245 23 PPP 631 1.6 75.0 48 28 11 9 1578 514 254 810 PPE w2: 1.6 20.0 20 11 5 + 200 56 slat} 31 Burica region PPP 47 1.8 32 30 0 2 50 30 0 20 PPP 86 1.8 op) 49 4 2 253 220 22 11 PPP 137 1.8 17 14 1 2 17 14 1 . PPP 144 1.8 47 40 4 3 128 103 22 3 PPP 146 1.8 40 35 2 3 85 53 11 21 PPP 148 1.8 37 35 0 2 82 71 0 1 PPP 156 1.8 34 29 3} 2 43 38 3 2 Nicoya Peninsula PPP 832 1.0 10 ] 3 3h7/ 6 10 21 PPP 833 1.0 7 3 1 25 12 1 12 1990; Coates et al., 1992; Coates and Obando, 1996). Equally importantly, the abundant, often well pre- served cheilostome material has provided a new re- source for applying finer-scale, quantitative morpho- logic approaches to taxonomic distinctions (Jackson and Cheetham, 1990, 1994). Previous work on cheilostomes from the PPP and DR collections has focused on evolutionary patterns in two genera, Metrarabdotos and Stylopoma (Cheet- ham, 1986, 1987; Cheetham and Hayek, 1988; Jackson and Cheetham, 1994; Cheetham ef al., 1994; Chee- tham and Jackson, 1995, 1996). Here we consider the fauna as a whole, even though a sizable proportion of the species remains undescribed. The need for addi- tional taxonomic splitting will no doubt become ap- parent as the fauna is studied in more detail. This paper is thus a general survey, the purpose of which is to provide initial estimates of: (1) the diversity of the PPP and DR cheilostomes and their affinities with the liv- ing and fossil fauna of the Caribbean; (2) the adequacy with which the PPP and DR collections reflect the di- versity and affinities of this fauna; and (3) the spatial and temporal distribution of species, their abundance, and their colony growth forms within and across areas. Preliminary study of 204 PPP collections (each of which was obtained from a roughly 10-kg sediment sample) has yielded 179 cheilostome taxa identified to species, 70 (39%) of which remain undescribed. Un- like other studies in this volume, ours incorporates 26 PPP collections from the Pacific side of the isthmus (with a total of 73 species), in addition to 124 DR collections (with a total of 132 cheilostome species). An even greater proportion of the DR species is un- described (59%). Inclusion of these other collections allows us to estimate the minimum diversity of the Neogene tropical American cheilostome fauna at 250 species (53% of which are undescribed), just slightly under the 273 species estimated to be present in the living cheilostome fauna of the tropical western Atlan- tic (Schopf, 1973). The number of species is likely to grow significantly as both the PPP and DR collections are studied in more detail, and as new PPP collections from both sides of the isthmus are included. However, the same is probably true for the living Caribbean fau- na. Recent studies have shown that correspondence be- tween morphologic and genetic differences is maxi- mized by splitting morphospecies to the limits of sta- tistical significance; thus, many widely distributed, morphologically variable “‘species”’ probably represent suites of genetically distinct species (Jackson and Cheetham, 1990, 1994). We have attempted to use these findings as a guideline for the initial analysis of the PPP and DR collections by recognizing even the smallest observed morphologic differences as tentative species distinctions, pending more detailed (morpho- metric and statistical) study. ACKNOWLEDGMENTS We thank A. G. Coates for age estimates and L. S. Collins for paleodepth estimates for the PPP collec- CHEILOSTOME BRYOZOA: CHEETHAM ET AL. 163 tions; P. Jung and J. Saunders for material, stratigraph- ic documentation, advice, and hospitality during study of the DR material; D. P. Gordon for current infor- mation on the revision of family-level and higher chei- lostome systematics; and EF K. McKinney and P. D. Taylor for comments on an earlier version of the man- uscript. This work was supported by grants from the Scholarly Studies, Research Opportunities, and Wal- cott Funds of the Smithsonian Institution; the Marie Bohrn Abbott Fund of the National Museum of Nat- ural History; the Smithsonian Tropical Research Insti- tute; and the National Geographic Society. DISTRIBUTION, AGE, AND ADEQUACY OF COLLECTIONS The 204 PPP collections represent five of the major Panamanian-Costa Rican regions listed in the PPP Da- tabase (Kaufmann, this volume) and shown on maps of the PPP collection sites (Appendix A): (1) Bocas del Toro Basin (Bocas del Toro Province, Caribbean Panama), 87 collections; (2) Canal Basin (Caribbean Panama), 10 collections; (3) Limon Basin (Caribbean Costa Rica), 66 collections; (4) Darien (Darien Prov- ince, Pacific Panama), 15 collections; and (5) the Pa- cific coasts of Panama and Costa Rica (Burica, Osa, Nicoya, and Golfo Dulce), 26 collections. Each bryo- zoan collection comprises specimens picked from a bulk sample (approximately 10 kg) of unconsolidated sediment. A collection contains 1—59 species (median 8; mean 14.4; CV [coefficient of variation] 103%). Sixty-two percent (126) of the 204 collections com- prise 6 or more species each; the median number of species in these collections increases to 17 (mean 21.8) and CV decreases to 67%. These collections are from all the major regions except the Darien (Table 1). The DR collections comprise 124 bulk samples, each consisting of 2 to more than 20 liters of sediment, and 63 sets of individually collected specimens (Saun- ders et al., 1986). The collections from bulk samples include 1—40 species each (median 19, mean 17.7, CV 50%); the individually collected sets include a median number of 2 species (mean 3.2, CV 96%). Species from the non-bulk collections were combined with those from stratigraphically equivalent bulk samples in the same section; there was virtually no change in the median or mean number of species, or in the CV (18, 17.6, and 49%, respectively) from those of the bulk collections. The DR collections represent four of the nine major areas in the Cibao region of the northern Dominican Republic (Saunders ef al., 1986, text-fig. 3): Rio Cana, 22 collections; Rio Gurabo, 61 collec- tions; Rio Mao, 21 collections; and Rio Yaque del Norte, 20 collections. A total of 117 collections (94%) include 6 species or more; these collections represent all four areas (Table 2). The average number of species in these collections (median 19, mean 18.5) is about the same as that for the 124 PPP collections with 6 or more species (median 17, mean 21.8), but is less var- iable (CV 44% compared to 67%) and has a smaller maximum (41 species compared to 59). The greater variability in apparent diversity (i.e., the number of species recovered) of the PPP collections is probably at least partly a function of preservation. Many collections were made from units that are not obviously fossiliferous, and the quality of bryozoan specimens is correspondingly variable. However, the most abundant and ubiquitous bryozoans in the PPP, even in the collections with the lowest diversity, are species with generally less preservable aragonitic skel- etons. Thus, the numbers of species likely vary with other (e.g., environmental) factors as well. The temporal distribution of PPP and DR bryozoan collections and their abundance is shown in Text-figure 2, together with the distribution of numbers of species represented in each area and occurring in both. Abun- dance totals more than 39,000 for the PPP and more than 21,000 for the DR, but these are minimum values based on counts of colonies and colony fragments con- verted to a scale coded as follows: 1—9 = | (rare), 10— 99 = 10 (common), and 100+ = 100 (abundant). For each of the DR collections, counting was stopped at 100 for any given species, whereas actual counts were recorded in the PPP Database. Basing calculations on the coded scale reduces over-dominance by the most abundant species (especially in the PPP collections) and helps increase comparability of the PPP and DR data. Taken together, the PPP and DR collections span an interval from late Early or early Middle Miocene (cal- careous nannoplankton zones NN 4—6, approximately 17—13 Ma) to Pleistocene (zone NN 19, approximately 0.5—2 Ma), with the two sets of collections overlapping in age by about 8 m.y., or half the approximately 16- m.y. interval (Text-fig. 2). However, the PPP collec- tions (median age 3.5 Ma, range 11.6—1 Ma) are gen- erally younger than those from the DR (median age 7.1 Ma, range 17—3 Ma); 84% of the PPP collections are concentrated in the interval younger than 5 Ma, whereas 87% of those from the DR are older than 5 Ma. The contrast is even more pronounced in terms of abundance, with 98% of PPP abundance concentrated in the younger interval and 93% of DR abundance from the older. The 8—7 Ma (Late Miocene, exclusive- ly DR) and 4—3 Ma (Early Pliocene, chiefly PPP) in- tervals are especially well sampled. The paucity of col- lections in both areas in the long interval between 15 and 8 Ma (Middle and early Late Miocene) stands in marked contrast. The situation with respect to the number of species recovered is less uneven (Text-fig. 2C). Of the 179 164 BULLETIN 357 Table 2.—Diversity and abundance of cheilostome bryozoans (total and by growth form) in Dominican Republic project (DR) collections containing six Or more species each. Number of species Abundance Collection Age Free- Free- number (Ma) Total Encrusting Erect living Total Encrusting Erect living Rio Yaque del Norte NMB 17283 16.2 11 2 6 3 119 2 6 111 NMB 17285 16.1 9 1 6 2 54 1 33 20 NMB 17286 16.0 12 2 7 3 255 11 43 201 NMB 17287 15.9 itil 1 7 3 164 1 52 111 NMB 17288 15.8 14 3 8 3 356 3 143 210 NMB 17289 15.8 12 2 7 3 147 2 25) 120 NMB 17290 15.7 12 2 7 3 174 20 34 120 NMB 17327 157 13 2 8 3} 247 2 224 21 NMB 17184 15.7) 25 4 18 3 493 13 369 111 NMB 16935 15.6 iT 7 17 3 288 16 161 111 Olsson 179 15:5 32 10 19 3 383 19 253 111 NMB 17265 15.5 19 6 11 2 82 15 56 11 NMB 16936 15.5 18 6 10 2 459 105 244 110 NMB 16938 15.4 20 6 11 3 290 15 164 111 NMB 17190 15.4 14 2 10 2 41 2 28 11 NMB 16942 15.4 18 5 11 2 261 23 137 101 NMB 17278 5.6 6 2 4 0 6 2 4 0 NMB 17268 4.6 25 9 13 3 70 9 31 30 USGS 8702 3.9 31 10 17 4 139 10 98 31 Rio Mao USGS 8525 8.0 27 10 13 4 378 19 346 13 NMB 17269 8.0 22 9 10 3 121 63 37 21 NMB 16913 8.0 22 9 10 3 202 27 154 21 NMB 16912 7.9 6 1 2 3 33 1 11 21 NMB 16922 Te) 15 7 5) 3 204 52 32 120 NMB 16927 7.8 22 9 10 3 292 36 226 30 NMB 16923 7.8 10 4 3 3 55 13 21 21 NMB 16917 7.8 14 7 4 3 149 16 22 111 NMB 16916 7.8 DH, 13 12 2 360 94 246 20 NMB 16915 7.8 18 8 7 3 279 26 43 210 NMB 16926 7.8 22. 9 10 3 103 27 46 30 NMB 16924 7.8 18 6 9 3 387 24 243 120 NMB 16918 7.8 2a 7 11 3 489 43 236 210 NMB 16928 W/ 26 15 9 2 656 114 432 110 NMB 16929 7.6 25 11 11 3 430 155 254 21 NMB 16932 7.6 10 4 3 3 55) 4 21 30 NMB 16914 7.6 13 i 3 3 85 25 30 30 NMB 16930 7.5 12 3 6 3 147 3 114 30 NMB 16802 TS 9 3 3 3 144 3 21 120 NMB 16910 7.4 22 S 13 4 211 23 67 121 NMB 17175 7.4 21 6 10 5 102 24 46 32 Rio Gurabo NMB 15915 7.9 20 10 7 3 173 28 115 30 NMB 15914 7.9 18 10 5 3 243 19 104 120 NMB 15912 7.9 20 9 8 3 164 18 125 21 NMB 15911 7.8 21 10 8 3 174 19 125 30 NMB 15910 7.8 11 2 6 3 47 20 6 21 NMB 15903 7.8 10 3 4 3 28 3 4 21 NMB 16192 77 10 5 2 3 28 5 2 21 NMB 15900 Dell 20 6 11 3 83 15 38 30 NMB 15901 7.7 13 3 7 3} 58 3 34 21 NMB 16191 Tesi 6 1 2 3 15 1 2; 12 NMB 15904 ev 9 2 4 3 36 11 4 21 NMB 15907 sil 9 2 4 3 45 11 13 21 NMB 15906 7.6 15 4 8 3 294 2D 62 210 Table 2.—Continued Collection number NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB 15897 15896 16186 15890 15882 15881 16167 15878 15876 15874 15873 16810 15865 15871 15869 15864 15863 15860 15849 16811 15854 15851 15853 15846 15842 15840 15837 15838 15836 15835 15805 15962 15804 15934 15815 15964 15814 15823 15828 16103 15829 15832 15833 Rio Cana NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB NMB 16857 16856 16855 16995 16844 16842 16841 16839 16838 16837 16836 16835 16834 (Ma) 6.5 5.5 6.5 Number of species Total Ne ooonn Ne ANE DONATI WW OO WNN = Ne ie) Encrusting WN NY WORWNHRK KB KWON ~) 0 Nn Mm MN WwW wert nownne £ Wn w BSS Erect WNNOWKDMRANE MANDEPDANWAHUNAN SF Oo ~ Aan vy — iy a — Or ODKN FR KH DWH wOr~A Free- living WhWW WWW WWW WwW WONNNOOFKF OCOFP RRP HRWA RH Nn ann Mnnn wWONnNrF ON WNOWNN — WW NNN WwW CHEILOSTOME BRYOZOA: CHEETHAM ET AL. Abundance Encrusting Erect 2 1 1 2 2 13 3 112 36 323 16 234 16 54 5 325 3 11 1 2 1 12 1 13 14 124 111 23 22 13 3 26 5 165 25 196 90 158 51 24 35 24 53 122 81 141 67 176 28 77 5 13 14 38 22 46 5 32 23 46 13 8 38 231 > 30 17 77 18 57 5 27 7 39 4 68 3 34 3 2 7 2 3 131 31 23 29 133 37 26 21 1 36 125 52 74 30 47 2 4 57 160 69 155 139 33 406 334 31 155 59 12, 165 Free- living 120 300. Ww NONRK ONKN tO Se Ne NN ON O - om) Nu NeEneHHKHnowr 166 BULLETIN 357 Table 2.—Continued. Number of species Collection Age number (Ma) Total Encrusting Erect NMB 16833 39) 39 20 17 NMB 16832 59 31 17 12 NMB 16828 Dil) 37 16 17 NMB 16818 5.0 37 18 16 NMB 16817 5.0 37] 15 19 NMB 16860 4.5 8 3 2 NMB 16865 3.8 22 15 3 NMB 17023 3.1 32 14 16 Abundance Free- Free- living Total Encrusting Erect living 2 237 92 125 20 2 148 71 66 11 4 397 79 296 22 3 271 54 187 30 3 307 78 208 21 3 17 12 2 3 4 49 33 3 13 2 284 14 268 2 species that occur in the PPP, 134 (75%) are found in the 4—3 Ma (Early Pliocene) interval, which also con- tains 78 (59%) of the 132 species that occur in the DR. The numbers of species in other 1-m.y. intervals, except that between 5 and 4 Ma, are more biased to- ward one area or the other (Text-fig. 2C). However, the cumulative fauna in the 3-m.y. interval from the latest Miocene to mid-Pliocene (6—3 Ma) includes ful- ly 72% (85) of the 118 species found only in the PPP, 66% (47) of the 71 present only in the DR, and 97% (59) of the 61 occurring in both areas. By this crite- rion, the 6—3 Ma at least appears well sampled. For a more detailed estimate of sampling adequacy (i.e., the probability that the numbers of species re- covered represent a major proportion of those present), we plotted cumulative numbers of species recovered as a function of the numbers of collections examined in each area (Text-fig. 3). In both plots, collections were added region by region (PPP) or section by sec- tion (DR), from oldest to youngest within each region or section. Although the DR has fewer species and collections than the PPP, the overall rate of increase in species is the same in the two plots, and the curves are similar in shape, rising in a series of steps corre- sponding to the different areas sampled. Overall, a slight flattening of the curves is apparent (thus indi- cating at least the beginning of an approach to “‘true”’ diversity): distinctly more than 50% of the species were recovered after 50% of the collections were tal- lied (124 species, or 69%, in the PPP, and 93 species, or 70% in the DR). Flattening is somewhat more ap- parent in the DR curve, with recovery of 50% of the total number of species requiring only 16% of the col- lections, compared with 30% for the PPP. If sampling is somewhat better for the DR than the PPP, as suggested by the collecting curves in Text- figure 3, PPP diversity can ultimately be expected to exceed that for the DR by even more than the 36% reflected in the total numbers of species so far recov- ered. This may well be related to the greater size and heterogeneity of the area sampled by the PPP (Text- fig. 1); the DR collections are all from a single sedi- mentary basin, the Cibao Basin of the northern Do- minican Republic, littke more than 100 km across (Saunders er al., 1986, text-fig. 2), whereas the area sampled for the PPP is nearly 1000 km long. However, comparison of curves for individual well-sampled PPP and DR areas (Text-fig. 4) suggests that PPP diversity consistently exceeds that for the DR even on scales of tens of km. The same is true for time intervals (Text- fig. 4). Moreover, similar plots of diversity against cu- mulative abundance (not shown) have virtually the same form as those plotted against numbers of collec- tions. Thus, the differences in total numbers of chei- lostome species appear to reflect real differences in “true”’ diversity. Even though some of the PPP collections contain up to 44% more species and more than twice the abun- dance of the richest DR collections, the proportion of PPP collections characterized by few species and low abundance is much greater than that in the DR (Text- fig. 5). As noted above, 38% of PPP collections but only 6% of those from the DR include fewer than six species each. Indeed, the modal number of species per DR collection slightly exceeds the median number (20 versus 19), whereas the modal number in the PPP is just one species (Text-fig. 5) compared to a median of 8. The preponderance of low diversity collections in the PPP, together with the higher total number of spe- cies, illustrates why the estimation of “‘true’’ diversity is even less certain than for the DR. ABUNDANCE AND DISTRIBUTION OF SPECIES AND GROWTH FORMS GROWTH FORMS AND THE ADEQUACY OF COLLECTIONS Species were assigned to three major colony growth-form categories (Table 3): encrusting (EN), erect (EE erect flexible; ER, erect rigid), and free-liv- ing (FL). Examples of PPP and DR species with these growth forms are illustrated in Cheetham and Jackson (1999). We used this generalized approach, rather than CHEILOSTOME BRYOZOA: CHEETHAM ET AL 167 Number of collections Abundance (thousands) Number of species Time interval (Ma) Text-figure 2.—Temporal distribution of 204 PPP and 124 DR collections, the abundance of bryozoan specimens (total 60,757; see text for counting method), and the numbers of species (total 250) represented in each interval. Ages of collections are medians of ranges; collections are grouped in l-m.y. “bins” ending in whole units, e.g., 3.1—4.0 Ma. a more detailed classification such as that of Hageman et al. (1998), for two reasons. First, the great majority of species, 67%, show relatively little variation in pos- sessing single- to multilayered sheetlike or platelike colonies. With few exceptions, species in the encrust- ing category occur as small, usually rare colony frag- ments. Thus, the scope for recognizing subsets of mor- phologies in this group is restricted. Secondly, al- though erect and free-living species show many, much 150 100 nw o ® ro 0) ¢2) oe re) — a 2 f= 0 =) iS o > = ro = £ =>) O 0 50 100 150 200 Cumulative number of collections Text-figure 3.—Cumulative curves (collecting curves) of species recovered with increasing numbers of collections examined (total 204 for PPP, 124 for DR). For each area, collections were added region by region (PPP) or section by section (DR), from oldest to youngest in each. 100 Gurabo Cumulative number of species 0 50 100 0 50 Cumulative number of collections Text-figure 4—Cumulative collecting curves for PPP and DR col- lections, each grouped by 3-m.y. interval and by region or section. 168 BULLETIN 357 Table 3.—Growth forms, abundances, and numbers of PPP and DR collections in which each occurs (= occurrences) of the 71 most common cheilostome bryozoan species (cumulative abundance > 100) arranged by rank in total abundance. Growth Rank in Abundance Abundance Rank in Occurrences Occurrences Species form abundance in PPP in DR occurrences in PPP in DR Cupuladria biporosa BL 1 4410 2543 1 133 113 Discoporella n. sp. 6 FL 2 4451 0 5 121 0 Mamillopora tuberosa FL 3 2974 861 2 112 96 Cupuladria n. sp. 1 aff. C. biporosa Re 4 2742 0 14 77 0 Cupuladria n. sp. 4 aff. C. canariensis FL S) 2205 0 7 108 0 Nellia tenella EF 6 870 1250 4 60 71 Discoporella n. sp. | lal’, 7 0 1889 13 0 80 Thalmoporella biperforata ER 8 8 1865 8 8 92 Tremogasterina mucronata EN ©) 1382 477 6 41 Ue Discoporella n. sp. 5 FE 10 1706 0 9 86 0 Reteporellina evelinae ER 11 1550 125 30 38 8 Discoporella n. sp. 7 FL 12 1436 0 9 86 0 Discoporella n. sp. 4 Be 13 0 1340 84 0 17 Mamillopora cavernulosa EE 14 1284 0 35 42 0 Metrarabdotos colligatum ER 15 11 1153 39 2 37 Gemelliporella punctata ER 16 389 720 15 29 45 Corynostylus labiatus EF 17 13 989 39 4 35 Cellaria mandibulata EF 18 824 65 34 23 20 Biflustra savartit EN 19 602 281 3} 89 74 Cupuladria n. sp. 8 aff. C. canariensis EE 20 875 0 28 47 0 Discoporella n. sp. 3 FL 21 828 13} 38 36 4 Celleporaria albirostris EN 22 834 10) 47 33 0 Vibracellina laxibasis EN 23 726 0 35 42 0 Semihaswellia sinuosa EF 24 159 519 39 15S 24 Metrarabdotos n. sp. 10 ER 25 444 214 52 12 16 Schizoporella magniporosa ER 26 55 235) 18 10 58 Metrarabdotos auriculatum ER 27 25 484 64 | 16 Schedocleidochasma n. sp. 3 EN 28 501 0 23 51 0 Celleporaria brunnea EN 29 310 160 19 49 16 Schedocleidochasma cleidostoma EN 30 458 0 24 49 0 Scrupocellaria regularis EF 31 446 0 37 41 0 Steginoporella parvicella EN 32 26 398 16 8 65 Skylonia dohmi EF 33 0 403 85 0 16 Scrupocellaria n. sp. | EF 34 142 248 27 7 41 Mamillopora n. sp. | BE 35 384 0 89 15 0 Bracebridgia subsulcata ER 36 Sil 0 73 20 0 Buskea n. sp. aff. B. dichotoma ER 37 365 0 92 14 0 Adeonellopsis n. sp. 3 ER 38 355 0 101 13 0 Celleporaria magnifica EN 39 54 258 17 18 51 Cupuladria n. sp. 5 aff. C. canariensis lab 40 287 0 55 26 0 Stylopoma spongites EN 41 110 176 46 11 23 Petraliella bisinuata EN 42 283 0 67 22 0 Adeonellopsis deformis ER 43 0 265 20 0 58 Margaretta buski EF 44 261 0 45 36 0 Canda simplex BE 45 190 57 24 28 21 Cigclisula porosa ER 46 122 117 9 41 45 Labioporella miocenica EN 47 0 238 24 0 49 n. gen. B n. sp. y Scolaro ER 48 23 198 48 5 27 Hippoporella gorgonensis EN 49 122 96 12 50 33 Ditaxipora n. sp. 2 EF 50 0 202 173 0 4 Turbicellepora n. sp. EN 51 201 0 39 39 0 Gemelliporella? n. sp. ER 2 81 114 60 18 6 Steginoporella n. sp. 1 EN 53 0 187 85 0 16 Margaretta n. sp. 1 EF 54 0 172 21 0 55 Microporella umbracula EN 54 172 0 52 28 0 Vincularia n. sp. EF 56 0 158 92 0 14 Cellaria aff. C. bassleri BE 57 43 111 118 7 3 Pasythea n. sp. ER 58 0 149 64 0 23 Gemelliporidra multilamellosa EN 59 147 0 69 21 0 CHEILOSTOME BRYOZOA: CHEETHAM ET AL. 169 Table 3.—Continued. Growth Rank in Species form abundance Schedocleidochasma porcellanum EN 60 Characodoma contractum EN 61 Metrarabdotos lacrymosum ER 61 Lagenicella atf. L. mexicana EN 63 Parasmittina parsevaliformis EN 64 Trematooecia vaughani EN 65 Schizoporella cornuta EN 66 Antropora leucocypha EN 67 Hippaliosina rostrigera EN 67 Metrarabdotos n. sp. 4 ER 67 Stylopoma n. sp. 11 EN 70 Steginoporella magnilabris EN 71 more obvious differences in morphology, for example, in branch shapes and jointing (erect forms) or in per- manent or intermittent nonattachment (free-living forms), they represent much smaller numbers of spe- cies which have similar patterns of occurrence and abundance (a possible exception is Nellia tenella, the only erect species occurring in more than 50 collec- tions in both the PPP and DR). Overall, the encrusting growth form comprises the majority of species in both the PPP (73%) and the DR collections (56%), but only a minority in abundance for both sets of collections (21% in the PPP, 18% in the DR). Consequently, collecting curves for encrust- ing species (Text-fig. 6) closely reflect the curves for the cheilostome faunas as a whole (Text-fig. 3), i.e., rising stepwise with only a slight tendency for overall flattening. In contrast, the curves for erect and free- living species tend to level off, and to about the same extent for both the PPP and DR collections (Text-fig. 6), implying similarly adequate sampling for species of these growth forms in both areas. However, there Number of collections ; 20 40 O 500 Species per collection Abundance per collection oO Text-figure 5.—Frequency distributions of numbers of species and abundance per collection for PPP and DR collections. Abundance Abundance Rank in’ Occurrences Occurrences in PPP in DR occurrences in PPP in DR 4 141 30 4 42 134 0 22 53 0 11 123 33 2 42 130 0 101 13 0 70 49 44 25 13 0 113 64 0 23 110 0 28 47 0 Si, 72 30 28 18 109 0 52 28 0 0 109 16) 0 19 107 0 130 8 0 99 2 50 27 2 are major differences in relative diversity and abun- dance between the PPP and DR. Erect forms make up 39% of the species and 51% of the abundance in the DR collections, but only 19% of the species and 18% of the abundance in those from the PPP (Text-fig. 6). Even more striking is the difference in the free-living components of the two faunas: 8% of the species and 61% of the abundance in the PPP versus 5% of the species and 31% of the abundance in the DR (Text- fig. 6). The virtually complete reversal of relative abundance of free-living and erect species is the most conspicuous difference in the cheilostome faunas of the two areas. Free-living species are almost three times as abundant as either erect or encrusting species in the PPP, whereas erect species are almost twice as abundant as each of the other growth forms in the DR (Text-fig. 6). These patterns probably reflect both tem- poral and environmental differences between the two areas. 150 15 - 3 Encrusting 2 Erect 1 Free-living n a) oO o a w a {e) Se = 0 100 Saeco j= o 60 2 w 40 FS a 7 ae (r oO oe 0 50 100 0 5 10 Cumulative abundance (thousands) Cumulative collections Text-figure 6—PPP and DR collecting curves for cheilostome species of three growth forms plotted against numbers of collections and abundance. 170 BULLETIN 357 PPP 1201 <3 Ma 3 |1201< 3 Ma 80 80 40 es 2 o a= 1 oO 2 0 20 40 60 100 n” ae 6-3 Ma 3 oO 80 ® 60 a —E 40 >} 5 20 1 2 0 | w 0) 50 100 z a 20 15 10 = 2 Erect 5 Yo 0 ieee) 1 Free-living 0 10 20 30 0 0.2 0.4 Cumulative abundance (thousands) Cumulative collections Text-figure 7.—PPP collecting curves for cheilostome species of three growth forms in each of three 3-m.y. intervals. The relative diversities of encrusting, erect, and free-living cheilostomes are maintained in about the same rank order in each of the three 3-m.y. time in- tervals represented by the PPP collections (Text-fig. 7). However, the number of encrusting species increases from 55% of the total in the oldest interval (>6 Ma) to 65% and finally 75% in the youngest (0O—3 Ma). In the youngest interval, the percent share in abundance for the encrusting species more than doubles to 33% (from 13% and 11% in the successively older inter- vals), partly but not entirely because of the 75% re- duction in abundance of free-living species. These changes are probably related to the increased number of reef-associated stratigraphic units sampled in the Late Pliocene—Pleistocene interval in the Bocas and Limon Basins. In the DR (Text-fig. 8), the relative diversities and abundances of the three growth forms are more uni- form, especially for the 6—3 Ma and 9—6 Ma intervals. The oldest (>9 Ma), however, shows a deficit of en- crusting species relative to younger intervals, possibly a sampling artifact. Erect species are dominant in abundance in all three time intervals, comprising 50% or more of the total abundance in each. Frequency distributions of numbers of species and abundance per collection for each growth form (Text- figs. 9, 10) are similar to those for the faunas as a whole (Text-fig. 5). Modal numbers of species are all 6-3 Ma ee) 1 1D 0G | A eee 3 0 10 20 2 60, 9-6 Ma 3] 60 3 9-6 Ma {e) — ® 40 40 a = 2 20 EG i o 0 20 40 60 =} >9Ma 2 2 3 O 45 10 2 1 0 0 5 10 15 0 1 2 Cumulative abundance (thousands) Cumulative collections Text-figure 8.—DR collecting curves for cheilostome species of three growth forms in each of three 3-m.y. intervals. one for the PPP and more than one for the DR, with abundances showing less difference between the two areas. These relationships suggest that sampling adequacy is good to fair for species of different growth forms, and that the quality of sampling is similarly good for the two younger 3-m.y. intervals in each area, and only slightly reduced for the oldest interval in each. PPP DR Encrusting wn = £ 10 20 oO A) Te} 15) io. 50 = o 2 = 0 E 0 20 40 0 10 20 = 501 Free-living 50 Free-living 0 0 20 40 ) 10 20 Number of species per collection Text-figure 9.—Frequency distributions of numbers of species per collection of three growth forms in the PPP and DR. CHEILOSTOME BRYOZOA: CHEETHAM ET AL. 171 PPP DR Encrusting iz) [= 7 2 0.2 0.4 3 Erect To) Oo cS to) re} VERO £ 0.2 0.4 a z Free-living Na 0 0.2 0.4 Abundance per collection (thousands) Text-figure 10.—Frequency distributions of abundance per collec- tion of three growth forms in the PPP and DR. ABUNDANCE AND OCCURRENCE OF SPECIES Largely but not entirely because of its superabun- dance in the PPP collections, free-living growth char- acterizes the 5 most abundant species in the combined PPP-DR database, and 13 of the 25 most abundant species (Table 3). The same free-living species, Cu- puladria biporosa, ranks first or second in abundance and number of occurrences in both the PPP and the DR, and is one of only two species to rank in the top 10 in both areas. (The other, Mamillopora tuberosa, is also classed as free-living on the basis of morphology illustrated in Cheetham and Jackson, 1999.) The 19 free living species have a median rank in abundance of 14 among the 250 cheilostome species, compared to 79 for the 64 erect species and 152.5 for the 167 encrusting ones. Four free-living species occur in 100 or more collections, in contrast to only two erect and two encrusting ones (Text-fig. 11). Further differences in the patterns of distribution of the three growth forms are evident in the relationship between the abundance of a species and the number of collections in which it occurs (occurrences; Text- fig. 11). Although abundance is highly significantly correlated with occurrences for all three growth forms (Spearman rank-order correlation 0.87—0.93, P < 0.001 in all cases), the rate at which abundance in- creases with occurrences for free-living species is about 50% greater than the rate of increase for erect species and 100% greater than for encrusting ones (Text-fig. 11). Unsurprisingly, the most frequently oc- curring and abundant species in each group are those that occur in both the PPP and the DR, although other less common species in each group also occur in both areas (Text-fig. 11). oO no) (= ae = % Free-living species 2 N= 19 fe) £ = r, = 0.93 @ 0 50 100 150 200 250 Erect species ie N= 64 Te) r, = 0.87 xt : 0 50 100 150 200 250 Encrusting species N = 167 ra=0102 0 50 100 150 200 250 Number of occurrences Text-figure 11.—Relationship between total numbers of occur- rences ( = number of collections in which each species occurs) and total abundance for 250 cheilostome species in 204 PPP and 124 DR collections. Regressions of abundance on occurrences and Spearman rank-order correlations (all with P < 0.001) are shown for each of three growth forms. For comparison of slopes of regres- sion lines, the line for erect species (dashed) is shown on each of the three plots. Differences among the three growth forms are also apparent in frequency curves showing the numbers of species with different numbers of occurrences (Text- fig. 12). Encrusting species have strongly unimodal (“hollow’’) distributions, with 46% occurring in five PPP DR Encrusting 0 A 0 0 il 0 20 40 60 480 0 20 40 60 80 100 °5- 2040 60 80 100 Number of species Free-living AA A A oUu_M Ld 0 20 40 60 80 0 20 40 60 80 100 Number of collections per species Text-figure 12.—Frequency distributions of numbers of collec- tions per species for each of three growth forms in PPP and DR. 172 BULLETIN 357 Encrusting a ©) Erect WB Free-living ” 2 oO ® rot ® 100 = ° — © a g 3 50 0 no Encrusting 2 oO ® rol 7) — ° ae (= © oO — © a Time interval (Ma) Text-figure 13.—Temporal distribution of numbers (A) and per- cent (B) of species of three growth forms in PPP and DR collections combined. Collections are “binned” in 1-m-.y. intervals as in Text- figure 2. or fewer collections (i.e., fewer than 2% of the 328 PPP and DR collections). Only two encrusting species, Tremogasterina mucronata and Biflustra savartii, are among the 20 most abundant species in the combined dataset, and both are present in both the PPP and the DR collections (Table 3). In marked contrast, free-liv- ing species show ‘“‘flat’’ frequency curves, i.e., with equal numbers of species having few and many oc- currences (Text-fig. 12). The curves for erect species are intermediate in pattern, distinctly less hollow than those for encrusting ones (Text-fig. 12). Only two erect species, Nellia tenella and Thalamoporella biperfora- ta, both present in both the PPP and DR collections, are among the top ten in abundance in the combined dataset (Table 3). Frequency curves for the same growth form are strikingly similar between the PPP and DR collections (Text-fig. 12), suggesting that these are inherent prop- erties of species having these growth forms rather than effects of environmental or preservational differences between the two areas. Frequency curves plotted against abundance (not shown) are very similar to those plotted against occurrences (Text-fig. 12), be- £2 Encrusting © Erect MB Free-living Abundance/collections Free-living . ® ° ry . ry ry ° ry ry . . . . Percent abundance 15 10 5 0 Time interval (Ma) Text-figure 14.—Temporal distribution of abundance (A) and per- cent abundance (B) of three growth forms in PPP and DR collections combined. Abundance is normalized to the number of collections in each l-m.y. interval. Collections are “binned” as in Text-figure 2. cause of the tight correlation between abundance and number of occurrences (Text-fig. 11). TEMPORAL DISTRIBUTION OF SPECIES AND GROWTH FORMS CHANGES IN RELATIVE IMPORTANCE OF GROWTH FORMS The distribution of growth forms in the combined PPP-DR database shows two apparent sets of temporal relationships: (1) a rise in numbers of encrusting spe- cies and decline in numbers of erect species (Text-fig. 13; Cheetham and Jackson, 1996), and (2) a comple- mentary fluctuation in the relative abundances of erect and free-living species (Text-fig. 14). Confidence in both sets of relationships, however, is affected by the incomplete overlap in the ages of the PPP and DR collections and the small numbers of collections rep- resenting several of the 1-m.y. intervals (Text-fig. 2). In addition, the questions of sampling adequacy dis- cussed above make it difficult to judge just how close- ly differences in numbers of species recovered from different intervals track changes in “‘true”’ diversity. The rise in number of encrusting species and decline CHEILOSTOME BRYOZOA: CHEETHAM ET AL. 173 in erect forms show considerable fluctuation, much of which appears related to small numbers of samples (Text-fig. 13). The number of free-living species also fluctuates, although with no obvious relationship to the other growth forms (Text-fig. 13). The decline in num- ber of erect species seems fairly steady beginning at 8 Ma, with the sharpest decline beginning at 3 Ma and coinciding with the peak in numbers of encrusting spe- cies. Because encrusting diversity may be significantly less adequately sampled than that of other growth forms, much of the apparent fluctuation in the numbers of encrusting species could be sampling artifact. Free- living diversity is much better sampled, and thus fluc- tuation in species numbers for this growth form should be more meaningful. Although changes in the abundance of free-living species appear to be complementary to those of erect species (Text-fig. 14), they are also random in time, and appear unrelated to changes in diversity of any of the three growth forms. Despite the strong disparity in abundance of free-living species in the PPP and DR (Text-figs. 7 and 8), the greatest temporal change in free-living abundance occurs within the PPP, between the well-sampled intervals younger and older than 3 Ma (Text-fig. 14). TEMPORAL DURATION OF SPECIES For the 250 species in the combined PPP-DR da- tabase, first and last occurrences and estimated ranges are listed in Table 4. First occurrences of 230 species (92%) fall within the database, as do last occurrences of 141 species (56%). With two exceptions (Metra- rabdotos auriculatum, and Thalamoporella chubbi), species whose ranges extend to stratigraphic levels younger than the collections included in the database are still living. Prominent among the still living species not previously known from the fossil record are: Bracebridgia subsulcata, Semihaswellia sinuosa, Ste- ginoporella connexa, Stylopoma projecta, and Tetra- plaria dichotoma, all widely distributed Caribbean species; Metrarabdotos pacificum and M. unguicula- tum, encrusting species of an otherwise erect genus; and Parasmittina crosslandi, P. fraseri, and ‘‘Stegi- noporella” cornuta, common eastern Pacific species. As expected, given the difference in median age be- tween the PPP (3.5 Ma) and DR faunas (7.1 Ma), the proportion of still living species in the PPP (99 of 179, or 55%) is greater than that in the DR (46 of 132, or 35%); the difference is significant statistically (chi- square = 12.78, P < 0.005, 1 df). The median observed stratigraphic range of the 250 species in the PPP-DR database is 3.6 m.y. (Table 4). As an estimate of the actual median duration (‘‘true” range) of the cheilostome fauna as a whole, this value is likely to be biased both by preservation failure at range limits, tending to truncate ranges, and failure of taxa with the shortest durations to be preserved at all, tending to lengthen average range (Foote and Raup, 1996). An additional truncating effect could be ex- pected because of the large number of species ranging to the present, 7.e., not yet extinct. However, the me- dian range of the 107 still living species (5.7 m-.y.) is actually significantly longer than that of the 143 ap- parently extinct species (2.1 m.y.) (Mann-Whitney U = 3992, P = 0.0000); thus preservation failure at range limits appears to be a much more significant factor. Given the larger proportion of still living spe- cies in the PPP fauna than in the DR, one might expect median ranges of species in the two areas to be dif- ferent (Text-fig. 15). However, the median observed ranges of species that occur in one area but not the other are not significantly different either among all collections (Mann-Whitney U = 3814, P = 0.3026) or among collections from the well-sampled, 6—3 Ma in- terval (U = 2345.5, P = 0.8465). These results suggest that, however significant the biases on species dura- tions may be, they are unlikely to apply to the faunas in the two areas differently. The median stratigraphic range of species occurring in both the PPP and DR collections, however, is highly significantly different from those for species occurring in only one area or the other (Text-fig. 15). With the data for all collections, the median range of the species from both areas (7.8 m.y.) is at least three times that for species from one area alone (2.7 m.y., U = 745, P = 0.0000, for the PPP; 2.0 m.y., U = 587, P = 0.0000, for the DR). The comparisons are much the same for the well-sampled, 6—3 Ma interval (8.0 m.y. for spe- cies in both areas; 3.6 m.y., U = 874.5, P = 0.0000, for the PPP; 4.0 m.y., U = 587, P = 0.0000, for the DR). These results are consistent with the correlation between stratigraphic and geographic range noted in Metrarabdotos and Stylopoma (Cheetham and Jack- son, 1996), i.e. a lowered probability of extinction with increased geographic range (Jackson, 1974). In marked contrast, the small differences in median stratigraphic ranges among species with different growth forms (Text-fig. 16) are all nonsignificant (P = 0.0780—0.7470 in Mann-Whitney U tests). Deletion of the two species with extremely long ranges, Trypos- tega venusta and Nellia tenella, changes calculated values slightly but leaves the statistical tests un- changed. Thus, stratigraphic ranges (i.e., extinction probabilities) appear to depend much less on the marked differences in patterns of occurrence and abun- dance among species of different growth forms noted above (Text-figs. 8—11) than on the extent of a species’ geographic range. 174 Table 4.—Stratigraphic ranges and oldest and youngest occurrences of cheilostome bryozoan species present in PPP and DR collections. BULLETIN 357 References in parentheses are for fossil occurrences outside the PPP-DR database. Range Species (m.y.) Oldest occurrence Youngest occurrence Adeonellopsis deformis 4.4 NMB 16913, 17269; USGS 8525 NMB 15829 Adeonellopsis n. sp. 1 7.4 TU 1293 Living Adeonellopsis n. sp. 2 0.0 NMB 17273 NMB 17273 Adeonellopsis n. sp. 3 1.9 PPP 334, 367, 368 PPP 710 Aimulosia palliolata 5.3. NMB 15964 Living Alderina smitti 3.6 PPP 370 Living Amphiblestrum pustulatum 3.1 NMB 17023 Living Antropora granulifera 3:67 PPPJ65 Living Antropora leucocypha 16.2 (Scolaro, 1968) Living Antropora typica Saf) SPPPM83" 391 Living Arthropoma cecilii 4.3 PPP 205 Living n. gen. B n. sp. y 12.7. (Scolaro, 1968) PPP 708, 709, 720, 722, 723 Bellullopora bellula 3:5)" RPP709 Living Biflustra denticulata 1.8 PPP 86 Living Biflustra savartii 16.0 NMB 17285 Living Bracebridgia subsulcata 4.3. PPP 345 Living Buskea n. sp. 1 aff. B. dichotoma 5) SPPPIG63 PPP 644, 653, 943, 944, 948-950, 962, 963 Caberea sp. 1.9 PPP 68, 720 PPP 738 Calpensia sp. 0.0 PPP 86, 137, 144, 146, 156 PPP 86, 137, 144, 146, 156 Calyptooecia insidiosa 6.6 NMB 15849 Living Canda simplex 16.0 NMB 16935 Living Canda n. sp. 1.9 NMB 16817 NMB 17023 Cauloramphus atf. C. brunea 0.0 PPP 47, 86, 144, 146 PPP 47, 86, 144, 146 Cellaria bassleri 3.6 PPP 294, 295, 370 Living Cellaria aff. C. bassleri 2.2 USGS 8702 PPP 710 Cellaria mandibulata 7.7 _NMB 15901 Living Celleporaria albirostris 4.3 PPP 346, 352 Living Celleporaria brunnea 16.0 NMB 16935 Living Celleporaria magnifica 16.2 (Scolaro, 1968) Living Celleporaria n. sp. OOS SPER 352 PPP 352 Chaperia condylata 16.0 NMB 17288 Living Characodoma contractum 16.2 (Scolaro, 1968) Living Cigclisula porosa 14.4 NMB 17283 PPP 631, 943, 944, 948-950, 962, 963 Coleopora aff. C. americana 1.6 PPP 644, 949, 963 Living Coleopora granulosa 4.3. NMB 17175 NMB 17023 Copidozoum planum 4.3. PPP 60 Living Copidozoum aff. C. tenuirostre 1.3. NMB 15903 NMB 15851 Corynostylus labiatus 12.5 NMB 17283 PPP 932 Crassimarginatella aff. C. corbula 1.7 PPP 642 Living Crepidacantha longiseta 1.6 PPP 949, 950 Living Crepidacantha poissonii 4.3. PPP 60 Living Cupuladria biporosa 20.0 (McGuirt, 1941) Living Cupuladria n. sp. | aff. C. biporosa 10.0 PPP 4, 10 PPP 631, 644, 717, 718, 944, 948-950, 962 Cupuladria n. sp. 2 aff. C. biporosa 22) EPPIGI BRP 77; Cupuladria n. sp. 3 aff. C. biporosa 720) SEPP162 PPP 631, 644, 653, 712, 943, 944, 948-950, 962, 963 Cupuladria n. sp. 4 aff. C. canariensis 2.7 PPP 348, 349 PPP 631, 644, 653, 712, 944, 948-950, 962 Cupuladria n. sp. 5 aff. C. canariensis 2. PPPF35053525422 PPP 631, 944, 948, 950, 963 Cupuladria n. sp. 6 8.8 PPP 1 PPP 180, 362 Cycloperiella rubra 6.8 PPP 162 PPP 47, 86, 144 Cycloperiella n. sp. 0.7 PPP 205, 346, 348, 350, 352 PPP 65, 294, 295, 298, 306, 307, 334, 335 Discoporella n. sp. 1 4.3. NMB 15916, 17269 NMB 16865 Discoporella n. sp. 2 4.1 NMB 16856 NMB 17024 Discoporella n. sp. 3 S28) DUP1225 PPP 631, 962 Discoporella n. sp. 4 0.7 NMB 17284 NMB 16942 Discoporella n. sp. 5 726) ERPV162 PPP 822 Discoporella n. sp. 6 8.9 PPP 898 PPP 781, 819-824, 832, 833 Discoporella n. sp. 7 8:5) PPP; 4) 10 PPP 663, 668, 570, 691 Ditaxipora n. sp. | 6.4 NMB 16913; USGS 8525 PPPe iil Ditaxipora n. sp. 2 0.0 NMB 15851, 15853 NMB 15851, 15853 Table 4.—Continued. CHEILOSTOME BRYOZOA: CHEETHAM ET AL. Range Species (m.y.) Oldest occurrence Drepanophora tuberuculatum 7.8 NMB 16916 Electra biscuta 4.3 PPP 206, 352 Escharella? sp. 0.0 PPP 210 Escharina pesanseris 6.7 NMB 15860 Escharina porosa 7.1. NMB 16856 Escharoides costifer 16.2 (Scolaro, 1968) Escharoides n. sp. 2.0 NMB 15865 Exechonella cf. E. antillea 0.8 NMB 15846 Fedora aff. F. nodosa 4.6 TU 1294 Floridina antiqua 1.8 PPP 86, 148, 156 Floridina minima 1.4 (Canu and Bassler, 1923) Floridinella parvula 3.6 PPP 306 Gemellipora n. sp. 1 0.8 NMB 15823 Gemellipora n. sp. 2 0.8 NMB 15823 Gemelliporella glabra 238) IPPPs36il Gemelliporella punctata 8.0 PPP 35 Gemelliporella? n. sp. 3.8 NMB 15962 Gemelliporidra magniporosa 1.8 PPP 47 Gemelliporidra multilamellosa 3.6 PPP 65 Gemelliporidra? sp. 4.1 NMB 15911 Gephyrophora cf. G. rubra 6.0 NMB 16834 Gigantopora fenestrata 6.5 NMB 15851 Hiantopora intermedia 1.6 PPP 949 Hincksina sp. 0.0 Olsson 179 Hippaliosina rostrigera 16.2. (Scolaro, 1968) Hippaliosina n. sp. 0.8 NMB 16915 Hippomenella? fissurata 2.0 PPP 65, 298 Hippomenella? atf. H.? fissurata 0.6 NMB 16995 Hippopetraliella cf. marginata 2.1 NMB 16856 Hippopleurifera mucronata 16.2 (Scolaro, 1968) Hippopleurifera n. sp. 1 0.9 NMB 16833 Hippopleurifera n. sp. 2 0.1 PPP 710 Hippopodina cf. H. bernardi 1.6 NMB 15914 Hippopodina feegeensis 1.7. PPP 635 Hippopodina aff. H. feegeensis 9.7 NMB 16935 Hippoporella costulata 1.9 PPP 68, 69, 210, 212, 704, 709, 720 Hippoporella gorgonensis 16.2 (Scolaro, 1968) Hippoporella aff. H. rimata 2.7 PPP 205, 206, 352 Hippoporidra edax 11.6 PPP 10 Hippoporina aculeata 8.6 PPP 162 Hippoporina n. sp. | 0.7 PPP 60 Hippoporina n. sp. 2 1.8 PPP 66 Hippoporina n. sp. 3 0.0 PPP 86, 137, 144 Jaculina sp. x 0.7. (Scolaro, 1968) Labioporella aff. L. dumonti 0.0 PPP 668 Labioporella miocenica 12.2. NMB 17286 Labioporella aft. L. miocenica 0.1 PPP 634 Lagenicella marginata 1.7. PPP 635, 642 Lagenicella aft. L. mexicana 2.6 PPP 312, 367 Lagenicella n. sp. 19) SERB 10 Lagenipora sp. 0.1 PPP 640 Mamillopora cavernulosa 2.7. PPP 345, 350, 352, 354 Mamillopora tuberosa 15.2 (Scolaro, 1968) Mamillopora n. sp. 1 2.0 PPP 365, 367, 368, 370 Mamillopora n. sp. 2 4.1 PPP 187 Margaretta buski 16.2 (Scolaro, 1968) Margaretta n. sp. 1 12.4 Olsson 179 Margaretta n. sp. 2 0.1 NMB 17290 Membraniporella? sp. 1.4 PPP 670 Metrarabdotos auriculatum 5.7 NMB 16186 Youngest occurrence Living Living PPP 210 Living Living Living NMB 16818 NMB 15838 NMB 15833 Living PPP 631 Living NMB 17023 NMB 17023 Living PPP 653 PPP 644, 943, 944, 948-950, 962, 963 Living Living NMB 16865 Living Living Living Olsson 179 Living NMB 16855 PPP 653, 943, 944, 948, 950, 962, 963 NMB 16834 NMB 16817 Living NMB 16818 PPP 949 NMB 16835 Living NMB 16832 PPP 631, 738, 944, 950, 962, 963 Living PPP 962 Living Living PPP 294, 298 PPP 86, 137, 146 PPP 86, 137, 144 Olsson 179 PPP 668 USGS 8702 PPP 949 Living PPP 781, 835 PPP 949, 963 PPP 712. PPP 738, 940, 944, 950 PPP 832, 833 PPP 631 PPP 362 Living NMB 17023 NMB 17184 PPP 634 (Cheetham, 1968) 175 176 Table 4.—Continued. BULLETIN 357 Range Species (m.y.) Oldest occurrence Youngest occurrence Metrarabdotos colligatum 4.2 TU 1294 USGS 8702 Metrarabdotos lacrymosum 5.1 NMB 15915 PPP 362 Metrarabdotos pacificum 3.6 PPP 66 Living Metrarabdotos tenue 4.6 NMB 17268 Living Metrarabdotos unguiculatum 5.9 NMB 16833 Living Metrarabdotos n. sp. | 0.5 NMB 17284 NMB 17265 Metrarabdotos n. sp. 2 0.3. NMB 17184 NMB 16942 Metrarabdotos n. sp. 3 3.9 NMB 16191 USGS 8702 Metrarabdotos n. sp. 4 22) DUWA293 NMB 15814 Metrarabdotos n. sp. 5 1.4 NMB 15911 NMB 16986 Metrarabdotos n. sp. 6 0.6 NMB 16191 NMB 15878 Metrarabdotos n. sp. 7 22 EU 1293 NMB 15814 Metrarabdotos n. sp. 8 8:0) (PPP'35 PPP 653 Metrarabdotos n. sp. 9 3.5 NMB 17005 USGS 8702 Metrarabdotos n. sp. 10 4.5 NMB 16844 PPP 214 Micropora coriacea 8.6 PPP 162 Living Microporella cf. M. ciliata 3.4 NMB 15851 NMB 17023 Microporella normant 8.9 PPP 1152 PPP 773, 832 Microporella umbracula Sy SPBP39) Living Mollia? sp. 0.0 NMB 17023 NMB 17023 Monoporella nodulifera 4.3 PPP 352 Living Mychoplectra? sp. 0.0 NMB 16995 NMB 16995 Nellia tenella 65.0 (Winston and Cheetham, 1984) Living Nellia cf. N. tenuis 8.4 NMB 17286 NMB 16929 Odontoporella adpressa 3.6 PPP 66 Living Onychocella aff. O. angulosa 5.5 NMB 16811 PPP 833 Onychocella n. sp. 3.5 NMB 16856 NMB 15829 Parasmittina aff. P. areolata 8.0 NMB 17269 Living Parasmittina crosslandi 4.3 PPP 206, 350, 532 Living Parasmittina fraseri 4.3 PPP 60, 205 Living Parasmittina hastingsae 3.5 PPP 68, 69 Living Parasmittina aff. P. murarmata 8155 JPPP7085937 Living Parasmittina parsevaliformis 15.5 Olsson 179 Living Parasmittina spathulata NEG PRP 222 Living Parasmittina n. sp. 1 10.0 PPP 222 PPP 644, 944 Parasmittina n. sp. 2 0.9 NMB 17269 NMB 16856 Parasmittina n. sp. 3 15.5 NMB 17265 Living Parasmittina n. sp. 4 0.3. NMB 16913 NMB 16928 Parasmittina n. sp. 5 0.0 NMB 15842 NMB 15842 Parasmittina n. sp. 6 0.0 NMB 15851 NMB 15851 Parasmittina n. sp. 7 0.0 PPP 631 PPP 631 Parasmittina n. sp. 8 4.3 PPP 205 Living Parellisina curvirostris 1.7 PPP 639, 640 Living Parkermavella punctigera 1.6 PPP 631, 949 Living Pasythea tulipifera 3.6 PPP 64, 65 Living Pasythea n. sp. 12.8 NMB 17327 USGS 8702 Petraliella bisinuata 3.5 PPP 68, 69, 210-212, 704 Living Pleurocodonellina sp. 0.0 PPP 68, 210 PPP 68, 210 Poricellaria n. sp. | 11.8 NMB 17184 NMB 15823 Poricellaria n. sp. 2 0.0 NMB 16935 NMB 16935 Puellina innominata Bebe 09 Living Puellina radiata 4.3 PPP 205, 348, 352 Living Puellina n. sp. aff. P. radiata 1.9 PPP 69, 708, 709, 723 PPP 653, 943, 949, 950, 962, 963 Reptadeonella bipartita 43) JPPP12055352 Living Reptadeonella hastingsae 15.4 NMB 16942 Living Reptadeonella tubulifera 4.3 PPP 201 Living Reptadeonella n. sp. 2.9 NMB 15849 NMB 15829 Reteporellina evelinae 9.6 PPP 35 Living Retevirgula tubulata 3.1 NMB 17023 Living 0.0 PPP 148 PPP 148 Rhynchozoon aff. R. phyrnoglossum a la eae a oc tS ee cies ES Table 4—Continued. CHEILOSTOME BRYOZOA: CHEETHAM ET AL. 177 Species Rhynchozoon rostratum Rhynchozoon verruculatum Savignyella sp. savignyellid? sp. Schedocleidochasma cleidostoma Schedocleidochasma porcellanum Schedocleidochasma n. sp. | Schedocleidochasma n. sp. 2 Schedocleidochasma n. sp. 3 Schizoporella cornuta Schizoporella floridana Schizoporella magniporosa Scrupocellaria maderensis Scrupocellaria pusilla Scrupocellaria regularis Scrupocellaria aff. S. unguiculata Scrupocellaria n. sp. | Scrupocellaria n. sp. 2 Semihaswellia sinuosa Skylonia dohmi Smittina? n. sp. 1 Smittina? n. sp. 2 Smittipora aff. S. acutirostris Smittipora levinseni Smittoidea maleposita Smittoidea pacifica Smittoidea prolifica Steginoporella magnilabris Steginoporella parvicella Steginoporella n. sp. 1 Steginoporella n. sp. 2 Steginoporella n. sp. 3 aff. S. connexa Steginoporella n. sp. 4 aff. S. connexa “Steginoporella”’ cornuta Stylopoma informatum Stylopoma minutum Stylopoma projectum Stylopoma spongites Stylopoma n. sp. 3 Stylopoma n. sp. 4 Stylopoma n. sp. 5 Stylopoma n. sp. 6 Stylopoma n. sp. 7 Stylopoma n. sp. 11 Stylopoma n. sp. 13 Stylopoma n. sp. 14 Tetraplaria dichotoma Thalamoporella biperforata Thalamoporella chubbi Thalamoporella n. sp. Thalamoporella n. sp. Thalamoporella n. sp. Thalamoporella n. sp. Thalamoporella n. sp. Thalamoporella n. sp. 6 Trematooecia aviculifera Trematooecia cf. T. hexagonalis Trematooecia turrita AkWN Tremagasterina vaughani Tremogasterina mucronata Range (m.y.) 4.3 1.8 12.6 0.2 8.6 16.2 2.6 1.9 Pied] 9.9 4.3 5.6 3.6 35) 16.2 0.0 12.4 12.4 7.4 0.7 0.0 0.0 5.6 16.2 13.7 1.6 1.8 9.6 6.3 1.5 0.1 10.0 0.1 dell 8.0 7.8 SES) 16.2 Ss) il 0.1 15:5 6.5 3.6 Te 6.9 eal 12.4 0.7 1.5 10.6 10.6 0.0 0.0 0.1 7.8 3.4 6.6 4.0 16.2 Oldest occurrence Youngest occurrence PPP 206, 207, 352, 355, 422 PPP 145, 146, 156 NMB 17184 NMB 17184 PPP 162 (Scolaro, 1968) PPP 345, 350, 352 PPP 937 PPP 204, 205, 345, 348-350, 352, 354 PPP 1071 PPP 419, 422 USGS 8525 PPP 367 PPP 68 (Scolaro, 1968) PPP 68, 69, 210 Olsson 179 Olsson 179 NMB 16910 NMB 17283 PPP 823, 833 PPP 944, 963 NMB 15878 (Scolaro, 1968) Olsson 179 PPP 653, 949 PPP 86 PPP 35 USGS 8528 NMB 16844 NMB 16838 PPP 222 PPP 640, 710 NMB 16856 USGS 8525 NMB 16916 PEP 227 23) (Scolaro, 1968) PPP 720 NMB 15851 NMB 16928 Olsson 179 NMB 16842 PPP 367 NMB 16928 NMB 15863 NMB 16856 Olsson 179 PPP 295 NMB 16995 NMB 16935 NMB 16935 PPP 35 PPP 346 PPP 370 NMB 16916 NMB 15854, 16811 NMB 15849 NMB 16916 (Scolaro, 1968) Living Living NMB 17023 Olsson 179 Living Living PPP 634 PPP 949 PPP 738, 944, 948, 950, 962, 963 Living Living PPP 691; NMB 17023 Living Living Living PPP 68, 69, 210 NMB 17023 NMB 17023 Living NMB 16942 PPP 823, 833 PPP 944, 963 PPP 738, 949, 962, 963 Living PPP 86, 137, 144, 146, 156 Living Living Living PPP 634 NMB 16817 NMB 16836 PPP 944, 948-950, 962, 963 PPP 631, 738 Living Living Living Living Living Living NMB 15962 NMB 16929 Living Living Living Living Living Living PPP 672 (Lagaaij, 1959) NMB 16817 NMB 16818 NMB 16817 PPE 35 PPP 346 PPP 708, 709, 720, 722, 723, 937 Living NMB 17023 Living USGS 8702 Living 178 BULLETIN 357 Table 4.—Continued. Range Species (m.y.) Oldest occurrence Youngest occurrence Tremoschizodina lata 1:6) JPPRiG3 1738 Living Triporula stellata 3.1 PPP 663 Living Trypostega venusta 35.0 (Canu and Bassler, 1920) Living Trypostega sp. 1.4 NMB 16836 NMB 16817 Turbicellepora n. sp. 2.7 PPP 205, 352, 422 PPP 631, 944, 948, 962, 963 Vibracellina laxibasis 231 PPPs352 PPP 631, 644, 653, 738, 943, 944, 948-950, 962, 963 Vibracellina aff. V. laxibasis 12.6 NMB 17184 NMB 17023 Vincularia n. sp. 0.7 NMB 17285 NMB 16942 Vittaticella sp. 9.8 NMB 17184 NMB 16833 Watersipora subovoidea 1.8 PPP 148 Living TEMPORAL DISTRIBUTION OF FIRST OCCURRENCES The apparent durations of species in the PPP-DR database are affected by incompleteness of temporal overlap between the PPP and DR collections (Text-fig. 2) and by the relatively small proportion of species occurring in both faunas (Text-fig. 15). Fully 66% of the PPP fauna (118 of 179 species) and 54% of the DR fauna (71 of 132 species) do not occur in the other area (the comparable values based only on species in the well-sampled, 6-3 Ma interval are similar: 92 of 139 PPP species, or 66%, and 47 of 99 DR species, All collections 6-3 Ma PPP only N= 118 median = 2.7 m.y. PPP only N=92 median = 3.6 m.y| DRonly N=52 median 4.0 my. DR only N=71 median = 2.0 m.y. Number of species median = 8.0 m.y. median = 7.8my. | 10} N=47 0 5 10 15 20 Stratigraphic range (m.y.) Text-figure 15.—Frequency distributions of observed stratigraphic ranges of cheilostome species occurring only in the PPP or DR, or in both areas. Distributions are shown for all collections (250 spe- cies) and for the 6—3 Ma interval that is well-sampled in both areas (191 species). Two extreme outliers, Trypostega venusta (35 m.y., PPP only) and Nellia tenella (65 m.y., both areas), are not shown. or 47%). Nearly half the combined PPP-DR fauna (107 of 250 species, or 43%) range to the present, so the problem of incompleteness is more acute for the distribution of first occurrences than for last occur- rences. Despite unevenness of sampling, first occurrences that fall within the PPP-DR database are distributed virtually throughout the approximately 16 m.y. repre- sented by the collections (Text-fig. 17). For species occurring in only one area or the other (Text-fig. 17A), significant numbers of first occurrences are distributed 6-3 Ma Encrusting N= 121 median = 4.3 m.y. All collections Encrusting N = 167 median = 3.5 m.y. Erect median N=52 50my. Erect N =64 median = 4.0 m.y. Number of species 6| Free-living N=18 median = 5.2 m.y. 6| Free-living N=19 median = 4.6 m.y. 0 0 5 oS ao eaB 2h Stratigraphic range (m.y.) Text-figure 16.—Frequency distributions of observed stratigraphic ranges of cheilostome species of three growth forms. Distributions are shown for all collections (250 species) and for the 6-3 Ma in- terval that is well-sampled in both areas (191 species). Two extreme outliers, Trypostega venusta (35 m.y., encrusting) and Nellia tenella (65 m.y., erect), are not shown. CHEILOSTOME BRYOZOA: CHEETHAM ET AL. 179 Species in PPP or DR only cI Ppp Mi OR ” o Oo = © = oO oO fe} 2 iL Species in both PPP and DR w oO Oo iS o 5 oO Oo (2) 2 iL ro c 2 = £ Ww BO) (5) o a 0) Time interval (Ma) Text-figure 17.—Temporal distribution of first occurrences of cheilostome species occurring only in the PPP or DR (A, total 189), or in both areas (B, total 61); and the number of species recovered from both areas that were found in each l-m.y. interval (C). “Bin- ning” is as in Text-figure 2. through most of the interval from 9 to 3 Ma in both the PPP and DR. More than 60% (73 of 120) of these first occurrences are in the PPP. In contrast, 93% (42 of 45) of the first occurrences of species found in both areas in the 9—3 Ma interval are in the DR (Text-fig. 17B). Despite their strong bias toward first occurrence in the DR, the species that occur in both areas do so in each of the 1-m.y. intervals from 9 Ma to 3 Ma (Text-fig. 17C), suggesting that the biased distribution is not just a sampling artifact. It is tempting to con- clude that the bias toward DR first occurrences for these species indicates preferentially westward migra- tion, consistent with the apparent origination of most of the species of Metrarabdotos and many of those of Stylopoma in the central Caribbean (Cheetham and Jackson, 1996), and with the prevalent direction of near-surface Caribbean circulation. Almost half the species with first occurrences between 9 Ma and 3 Ma in the DR (42 of 89 species, or 47%) are found in the PPP, while only 4% (3 of 76 species) with first occur- rences in the same interval in the PPP are found in the DR (Text-fig. 17). The peaks in apparent originations (Text-fig. 17) at about 8 Ma in the DR (41 species) and at about 4 Ma in the PPP (44 species) coincide approximately with the peaks in origination inferred for Stylopoma and Metrarabdotos (Cheetham and Jackson, 1996). How- ever, more data are needed to test hypotheses of rates of origination and directions of migration, especially data filling the apparent gap between about 8 Ma and 16 Ma. COMPARISON OF PPP AND DR FAUNAS GENERAL CHARACTERISTICS As described above, the PPP fauna is largely young- er than that of the DR (Text-fig. 2), richer in species both within areas and for all collections combined (Text-figs. 3, 4), and characterized by greater abun- dance and diversity of free-living species and lower diversity of erect species (Text-figs. 6-8). Inclusion of collections from the Pacific coast of Panama and Costa Rica (Burica, Golfo Dulce, Nicoya, and Osa regions) appears to be a relatively minor factor in the greater diversity of the PPP collections. Only 10 of 75 species (13%) from the Pacific side of the PPP do not occur on the Caribbean side, compared with the 54% of the DR species (71 of 132) not present in the PPP collec- tions (53% for collections of comparable age in the two areas, i.e., the 6-3 Ma interval). Moreover, only two of the Pacific PPP species, Smittoidea prolifica and Watersipora subovoidea, that are absent from the Caribbean PPP collections appear to be ‘‘true” Pacific species. Others, such as ‘‘Steginoporella”’ cornuta and Metrarabdotos pacificum, formerly known only living in the eastern Pacific, are present in the Caribbean PPP collections. Thus, the percentage of cheilostome spe- cies present on both sides of the isthmus since its emergence at about 3.5 Ma may even exceed the 87% shared in the PPP collections; closure of the isthmian seaway appears to have had relatively little evolution- ary effect on the cheilostome fauna. The most significant difference between the PPP 180 BULLETIN 357 and DR faunas is in their contrasting diversity, abun- dance, and number of occurrences of free-living spe- cies (Text-figs. 6-8). In the PPP, such species comprise 58% (7 of 12) of those that occur in 50 or more col- lections, compared with only 25% (3 of 12) in the DR (Table 3). Although 93% of both PPP (189 of 204) and DR (115 of 124) collections include free-living species, the median abundance of free-living species is 83% of total cheilostome abundance per collection in the PPP but only 28% in the DR. Only 3 of the 7 most abundant free-living species (Cupuladria bipo- rosa, ranked 1, Mamillopora tuberosa, ranked 3, and Discoporella n. sp. 1, ranked 7) occur in the DR, whereas all but one (Discoporella n. sp. 1) are present in the PPP collections (Table 3). The 2-to-1 difference in diversity of free-living species (15 in the PPP versus 7 in the DR) appears to be real. Despite our attempt to employ the same “‘splitting philosophy” for collec- tions from both areas, we were able to recognize only 1 species of Cupuladria, 4 of Discoporella, and 1 of Mamillopora in the DR versus 7, 4, and 4, respective- ly, in the PPP. The ubiquity, abundance, and diversity of free-liv- ing species in the PPP fauna are similar to the domi- nance of such species in bryozoan faunas on the con- tinental margins of North and South America (Marcus and Marcus, 1962; Maturo, 1968; Cadée, 1975; Win- ston and Hakansson, 1986). However, the morpholog- ically distinctive group of species typified by Cupu- ladria doma appears to be entirely unrepresented in the PPP and DR collections, even though these species are among the most abundant bryozoans living on the continental shelf of the southeastern United States (Winston and Hakansson, 1986), and also occur exten- sively in Neogene deposits in that area (Cook, 1965; Spencer and Campbell, 1987). The importance of the erect growth form in the DR fauna stands in marked contrast (Text-fig. 8) to the dominance of free-living forms in the PPP. Almost half (42%, or 5 of 12) of the species that occur in 50 or more collections in the DR are erect, compared to only 8% (1 of 12) of such species in the PPP (Table 3). Only 4 of 124 (3%) collections from the DR lack erect species compared to 86 of 204 (42%) from the PPP. However, erect species are not typically this important in the modern bryozoan fauna of the central Carib- bean, where reef-associated species are predominantly encrusting (Jackson and Winston, 1982; Winston and Jackson, 1984). A number of erect genera in the DR collections (Adeonellopsis, Cigclisula, Gemelliporella, and Metrarabdotos) show abundant evidence of hav- ing grown on seagrasses (Cheetham and Jackson, 1996), which were more abundant in the Neogene than they are in the Caribbean today (Ivany ef al., 1990). Moreover, recent collecting in the Miocene and Plio- cene of Venezuela suggests that faunas dominated by erect species of Metrarabdotos and Schizoporella, similar to those in the Dominican Republic, may also characterize some Neogene mainland assemblages in sedimentary environments more like those in the Neo- gene of the DR than the PPP (Jackson and Cheetham, unpublished data). QUANTITATIVE DIFFERENCES BETWEEN FAUNAS To explore the effects of temporal and spatial influ- ences on the abundance, diversity, and species com- position of the PPP and DR cheilostome bryozoans, we used the abundance data from collections (Tables 1, 2) containing six or more species (each of which occurs in at least two collections) in a series of ordi- nation analyses. Separate analyses were made with the PPP (Text-fig. 18) and DR data (Text-fig. 19), and with the two sets of data combined (Text-fig. 20). In each case, detrended correspondence analysis (DCA) was used to relate the differences in species abundances to independent axes of decreasing variation. This tech- nique is a nonparametric analogue of principal com- ponents analysis with eigenvectors extracted from a matrix of chi-square distances between collections (McCune and Medford, 1995). Pachut et al. (1995) found DCA to be more effective than cluster analysis in ordering abundance data for living Caribbean reef bryozoan assemblages in congruence with water depth. We used PC-ORD version 3.0 to calculate all ordina- tions; in contrast to earlier versions of detrended cor- respondence analysis programs, results are no longer dependent on input order of collections or species (McCune and Medford, 1997). In both the PPP (Text-fig. 18A, B) and the DR (Text-fig. 19A, B) ordinations, collections from the different constituent regions or sections overlap exten- sively on all three DCA axes. However, there is some separation on DCA axis 2 between PPP collections from the Pacific coast and those from the Caribbean basins (Text-fig. 18A). With the PPP and DR data combined (Text-fig. 20A, B), separation between PPP Pacific and Caribbean collections disappears as the dif- ference between the PPP and DR is emphasized. These results are consistent with percentages of species shared among the three sets of collections, noted above. To interpret the results of the PPP and DR ordina- tions, we calculated Spearman rank-order correlations (Table 5) between scores on the DCA axes and (1) the abundance and diversity of colony growth forms in each collection, and (2) the ages and estimated paleo- depths for each collection where available (Tables 1, 2). For the DR collections, ages were estimated by CHEILOSTOME BRYOZOA: CHEETHAM EFT AL. 181 400 B Bocas Basin C Canal Basin L Limon Basin P Pacific PPP 300 200 DCA axis 2 100 0 100 200 300 400 500 DCA axis 1 400, 300} 200) DCA axis 3 100 0 100 200 300 400 500 DCA axis 1 DCA axis 1 12 6 8 Age (Ma) Text-figure 18.—Ordination plots (A, B) of 126 PPP collections (Table 1) on three axes obtained by detrended correspondence anal- ysis (DCA) of abundances (counted as explained in text) of 160 cheilostome species; and (C) relationship between the ages of col- lections and their scores on DCA axis 1. linear interpolation based on stratigraphic thickness between biostratigraphic markers (Saunders ef al., 1986; Cheetham, 1986). For the PPP collections, mid- points of age range estimates are from Coates (in Jack- son et al., Appendix 1, this volume; written commun., 1998); paleodepths are midpoints of depth ranges based on benthic foraminiferal data (Collins et al., 500 C Rio Cana G Rio Gurabo 400 M Rio Mao Y Rio Yaque 300 200 DCA axis 2 0 200 400 600 DCA axis 1 400 300 200 100) DCA axis 3 600 600 400 DCA axis 1 200 10 Age (Ma) Text-figure 19.—Ordination plots (A, B) of 117 DR collections (Table 2) on three axes obtained by detrended correspondence anal- ysis (DCA) of abundances (counted as explained in text) of 120 cheilostome species; and (C) relationship between the ages of col- lections and their scores on DCA axis 1. 1995; Collins et al., this volume; Collins in Jackson et al., Appendix 1, this volume). Highly significant correlations with age (Table 5) suggest that the arrangement on DCA axis | in both cases is strongly controlled by temporal changes in species composition and abundance (Text-figs. 18C, 19C). However, the correlation is far stronger for the DR (-0.83) than for the PPP (-0.49; with the three 182 BULLETIN 357 600 x Caribbean PPP p Pacific PPP oDR 400 N 2 x oO . Coates, A.G., Jackson, J.B.C., Collins, L.S., Cronin, T.M., Dow- sett, H.J., Bybell, L.M., Jung, P., and Obando, J.A. 1992. Closure of the Isthmus of Panama: the near-shore marine record of Costa Rica and western Panama. Geological So- ciety of America Bulletin, vol. 104, pp. 814-828. Coates, A.G., and Obando, J. 1996. The geologic evolution of the Central American isthmus. in Evolution and environment in tropical America. J.B.C. Jackson, A.F Budd, and A.G. Coates, eds., University of Chicago Press, Chicago, pp. 21—S6. Collins, L.S., Coates, A.G., Jackson, J.B.C., and Obando, J. 1995. 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N. Eldredge and S.M. Stanley, eds., Springer Ver- lag, New York, pp. 257-265. Winston, J.E., and Hakansson, E. 1986. The interstitial bryozoan fauna from Capron Shoal, Flor- ida. American Museum Novitates, no. 2865, pp. 1—50. Winston, J.E., and Jackson, J.B.C. 1984. Ecology of cryptic coral reef communities. [1V. Commu- nity development and life histories of encrusting cheilo- stome Bryozoa. Journal of Experimental Marine Biology and Ecology, vol. 76, pp. 1-21. APPENDIX OF SPECIES OCCURRENCES In the following listing, species are arranged alphabetically in each of the suborders of the Cheilostomatida. A complete systematic listing for these species, following the classification of Gordon (1984, 1986, 1989), along with illustrations of most of the species, is available on the internet, at , the homepage for ““Neogene Marine Biota of Tropical America” (NMITA) (Cheetham er al., 1998). For each species, occurrences are arranged numerically under PPP and DR groupings. For the DR occurrences, NMB (Naturhistorisches Museum Basel, Switzerland), TU (Tulane University), and USGS (U. S. Geological Survey) lo- calities are listed and described in Saunders et al. (1986); the Olsson localities are in Vokes (1979). The occurrence data are available at the PPP internet site http://www.fiu.edu/collinsl/. As systematic studies are completed, all specimens will be deposited in the Na- tional Museum of Natural History, Washington, D. C. Order CHEILOSTOMATIDA Busk, 1852 Suborder MALACOSTEGINA Levinsen, 1902 Biflustra denticulata Smitt, 1873: PPP 86, 712. Biflustra savartii (Audouin, 1826): PPP 34, 47, 58, 63-66, 68, 69, 74, 86, 144, 146, 148, 156, 162, 188, 194, 197, 198, 203, 205— 207, 210-212, 214, 294, 295, 298, 306-308, 312, 326, 334, 335, 340, 341, 345, 346, 350, 352, 354, 355, 365, 370, 379, 419, 422, 631, 640, 671-673, 677, 679, 681, 683-685, 691, 695, 697, 704, 708, 709, 712, 720, 722, 723, 820, 822, 824, 832, 833, 835, 896, 907, 932, 933, 937, 939, 946, 975, 1142, 1163, 1164. DR NMB 15804, 15814, 15815, 15835-15838, 15842, 15865, 15869, 15878, 15882, 15896, 15897, 15900, 15901, 15903, 15904, 15906, 15907, 15910-15912, 15914-15916, 15934, 15962, 16191, 16192, 16802, 16818, 16828, 16838, 16839, 16842, 16844, 16855, 16857, 16910, 16912-16918, 16922-16924, 16926-16930, 16932, 16935, 16936, 16942, 16995, 17184, 186 BULLETIN 357 17265, 17268, 17269, 17285-17290, 17327; Olsson 179; TU 1294, 1379; USGS 8525. Corynostylus labiatus Canu and Bassler, 1919; PPP 326, 896, 898, 932. DR NMB 15903, 15906, 15907, 15910, 16913, 16915— 16918, 16922-16924, 16926-16929, 16935, 16936, 16938, 16942, 17184, 17190, 17265, 17269, 17283, 17285-17290, 17307, 17327; Olsson 179; TU 1294, 1379; USGS 8525. Electra biscuta Osburn, 1950: PPP 144, 148, 206, 352, 933. Mychoplectra? species: DR NMB 16836-16839. Suborder FLUSTRINA Smitt, 1867 Alderina smitti Osburn, 1950: PPP 47, 148, 370. Amphiblestrum pustulatum (Canu and Bassler, 1928): DR NMB 17023. Amphiblestrum? species: DR NMB 17023. Antropora granulifera (Hincks, 1880): PPP 65, 86, 146, 148, 738. Antropora leucocypha (Marcus, 1937): PPP 47, 66-69, 86, 137, 144, 146, 148, 162, 198, 206, 207, 294, 295, 298, 306, 346, 352, 354, 379, 631, 639, 642, 705, 709, 710, 950. DR NMB 15835, 15851, 15874, 15881, 15882, 15916, 16817, 16818, 16828, 16832, 16833, 16835-16838, 16842, 16857, 16865. Antropora typica (Canu and Bassler, 1928): PPP 47, 63, 65, 66, 68, 69, 86, 144, 148, 183, 193, 204, 205, 210, 211, 295, 307, 334, 352, 391, 631, 634, 639, 640, 653, 704, 705, 709, 712, 722, 723, 738, 943, 944, 948-950, 962, 963. Caberea species: PPP 68, 631, 720, 738. calloporid species: PPP 824, 1163, 1164. Calpensia species: PPP 86, 137, 144, 146, 156. Canda simplex Busk, 1884: PPP 35, 68, 86, 210-212, 334, 349, 631, 634, 635, 642, 653, 663, 705, 709, 710, 720, 722, 723, 738, 943, 944, 948-950, 962, 963. DR NMB 15805, 15823, 15827, 15828, 15837, 15934, 15962, 16817, 16818, 16832-16835, 16927, 16929, 16935, 17023, 17268; Olsson 179; USGS 8525, 8702. Canda new species [small zooids]: DR NMB 15823, 16817, 17023. Cauloramphus aff. C. brunea Canu and Bassler, 1930: PPP 47, 86, 144, 146. Cellaria bassleri Hastings, 1947: PPP 68, 178, 180, 210, 294, 295, 358, 362, 370, 639, 640, 663, 704, 738, 943, 944, 950, 963. Cellaria aff. C. bassleri Hastings, 1947: PPP 367, 708, 709, 710, 720, 722, 723. DR NMB 15832, 15833; USGS 8702. Cellaria mandibulata Hincks, 1882: PPP 211, 295, 298, 350, 634, 635, 640, 642, 644, 649, 653, 663, 709, 720, 722, 723, 943, 944, 948, 949, 950, 962, 963. DR NMB 15804, 15805, 15814, 15828, 15837, 15842, 15846, 15849, 15860, 15863, 15901, 15934, 15962, 15964, 16810, 16817, 16818, 16828, 16842, 17023. Chaperia condylata Canu and Bassler, 1930: PPP 709. DR NMB 16832, 16833, 17023, 17288; Olsson 179. Copidozoum planum (Hincks, 1880): PPP 60, 66, 86, 144, 146, 156. Copidozoum aff. C. tenuirostre (Hincks, 1880): DR NMB 15851, 15897, 15903, 15906, 16928. Crassimarginatella aff. C. corbula (Hincks, 1880): PPP 642. Crassimarginatella species: PPP 156, 709, 950, 963. Cupuladria biporosa Canu and Bassler, 1923: PPP 1, 42, 57, 58, 61, 62, 65—69, 71, 72, 74, 86, 137, 144, 161, 162, 184, 187-189, 191, 193-208, 210, 212-214, 294, 295, 298, 306-308, 311, 312, 326, 334, 335, 340, 341, 345, 346, 348-350, 352, 354, 355, 357, 362, 367, 368, 370, 379, 391, 419, 422, 423, 425, 426, 631, 634, 635, 639, 668-672, 675-679, 681-684, 688, 689, 691, 695, 697, 704-709, 712, 720, 722, 723, 738, 773, 820, 822, 832, 833, 907, 908, 916, 931, 932-935, 937, 939, 940, 943, 962, 963, 1134, 1137, 1142, 1145, 1163, 1171. DR NMB 15804, 15805, 15814, 15815, 15823, 15829, 15833, 15835, 15836, 15838, 15840, 15842, 15849, 15854, 15860, 15863-15865, 15869, 15871, 15873, 15874, 15876, 15878, 15881, 15882, 15890, 15895— 15897, 15900, 15901, 15903, 15904, 15906, 15907, 15910— 15916, 15929, 15934, 15936, 15937, 15939, 15947, 15962, 15964, 15966, 15982, 16103, 16167, 16186, 16191, 16192, 16802, 16810, 16817, 16818, 16824, 16828, 16832-16839, 16842, 16844, 16854-16857, 16862, 16865, 16910, 16912— 16918, 16922-16924, 16926-16930, 16932, 16935, 16936, 16938, 16942, 16961, 16973, 16985, 16995, 17023, 17175, 17184, 17190, 17265, 17268, 17269, 17283-17290, 17322, 17327; Olsson 179, 180; TU 1225, 1227A, 1293, 1294; USGS 8525, 8702. Cupuladria new species 1 aff. C. biporosa Canu and Bassler, 1923 [medium zooids]: PPP 4, 10, 34, 57, 60, 61, 63-69, 74, 162, 178, 194-196, 198, 204, 207, 210, 212, 214, 294, 295, 298, 306— 308, 311, 312, 326, 334, 335, 340, 341, 345, 346, 348-350, 352, 354, 355, 357, 358, 361, 362, 365, 367, 368, 370, 423, 631, 635, 639, 640, 642, 644, 663, 668, 683, 684, 688, 709, 710, 717, 718, 720, 722, 723, 944, 948-950, 962. Cupuladria new species 2 aff. C. biporosa Canu and Bassler, 1923 {small to medium, flat colonies; small zooids]: PPP 52, 53, 57, 61, 63, 65, 66, 71, 177, 193. Cupuladria new species 3 aff. C. canariensis (Busk, 1859) [large zooids]: PPP 53, 57, 63, 64, 66-72, 74, 162, 177, 178, 180, 187, 193-198, 200-208, 210, 212, 213, 294, 295, 298, 306-308, 311, 326, 334, 335, 340, 341, 345, 346, 348-350, 352, 354, 355, 357, 358, 362, 365, 367, 368, 370, 379, 391, 419, 422, 423, 425, 426, 631, 634, 635, 639, 640, 642, 644, 653, 669, 670, 676, 679, 689, 695, 697, 704-706, 708-710, 712, 720, 722, 723, 738, 931-935, 937, 940, 943, 944, 948-950, 962, 963. Cupuladria new species 4 aff. C. canariensis (Busk, 1859): PPP 57, 68, 69, 74, 178, 180, 193, 194, 197, 198, 210, 334, 340, 348, 349, 357, 361, 362, 365, 367, 368, 370, 631, 634, 635, 639, 640, 642, 644, 645, 653, 678, 695, 697, 704, 708-710, 712, 720, 722, 723, 944, 948-950, 962. Cupuladria new species 5 aff. C. canariensis (Busk, 1859) [colony with large basal pores; medium zooids]: PPP 68, 69, 198, 210, 212, 306, 308, 350, 352, 361, 362, 367, 368, 370, 422, 631, 635, 639, 640, 720, 722, 723, 944, 948, 950, 963. Cupuladria new species 6 [colonies with very few basal pores; large zooids]: PPP 1, 68, 69, 180, 350, 362. Discoporella new species 1 [growth determinate, regeneration rare; zooids large; cryptocyst with central pores; opesia with pair of small denticles]: DR NMB 15804, 15805, 15809, 15812, 15814, 15815, 15823, 15835, 15836, 15842, 15846, 15860, 15863- 15865, 15869, 15871, 15873, 15874, 15876, 15878, 15881, 15882, 15890, 15895-15897, 15900, 15901, 15903, 15904, 15906, 15907, 15910-15912, 15914-15916, 15929, 15933, 15934, 15936-15944, 15946, 15947, 15952, 15962, 15964, 15965, 15968, 15969, 16167, 16186, 16191, 16192, 16802, 16810, 16817, 16818, 16824, 16827, 16828, 16837, 16838, 16842, 16854, 16856, 16857, 16859, 16860, 16862, 16865, 16879, 16910, 16912-16918, 16922-16924, 16926-16930, 16932, 17175, 17268, 17269; TU 1225, 1227A, 1293, 1294; USGS 8525, 8702. Discoporella new species 2 [growth determinate, regeneration rare; colony base concave; zooids small; cryptocyst without central pores; opesia smooth]: DR NMB 15804, 15805, 15809, 15812, 15814, 15815, 15835, 15838, 15860, 15864, 15865, 15869, 15874, 15934, 15944, 15946, 15952, 15962, 15964, 16810, 16854, 16856, 17024. Discoporella new species 3 [like D. n. sp. 2, but with filled colony base]: PPP 55, 65, 67—69, 74, 178, 180, 205, 210, 212, 295, 298, 308, 335, 340, 341, 345, 348-350, 352, 362, 365, 367, 368, 370, CHEILOSTOME BRYOZOA: CHEETHAM ET AL. 187 631, 668-670, 676, 678, 708, 937, 962. DR NMB 15835, 16865; TU 1225; USGS 8702. Discoporella new species 4 [growth indeterminate, regeneration common; zooids small; cryptocyst rarely with central pores; ope- sia with many fine denticles]: DR NMB 16935, 16936, 16938, 16942, 17184, 17190, 17265, 17283-17290, 17322, 17327; Ols- son 179. Discoporella new species 5 [colonies flat; basal surface smooth; zo- oids small]: PPP 65-67, 71, 72, 74, 146, 162, 194, 195, 198, 200-208, 212-214, 294, 295, 298, 307, 308, 326, 334, 335, 340, 341, 345, 346, 348-350, 352, 355, 362, 368, 376, 379, 391, 419, 422, 423, 425, 426, 668-672, 675-677, 679, 681—685, 688, 689, 691, 695, 697, 705, 706, 708, 709, 720, 722, 723, 822, 931-935, 937, 939, 948. Discoporella new species 6 [growth determinate; zooids large; cryp- tocyst without central pores]: PPP 47, 53, 55, 57, 58, 61, 63-69, 71, 72, 74, 148, 156, 162, 177, 178, 180, 187, 188, 193-199, 202-208, 210-212, 294, 295, 298, 306-308, 311, 312, 326, 334, 335, 340, 341, 345, 346, 348-350, 352, 354, 355, 357, 358, 362, 365, 367, 368, 370, 379, 391, 419, 422, 423, 425, 426, 631, 634, 635, 639, 640, 642, 644, 645, 649, 668-670, 678, 679, 691, 697, 704-706, 708-710, 712, 720, 722, 723, 773, 781, 818-824, 832, 833, 898, 932, 937, 939, 940, 943, 944, 948-950. Discoporella new species 7 [zooids medium; basal surface finely pitted; cryptocyst with few central pores]: PPP 1, 4, 10, 34, 39, 42, 57, 64-69, 72, 74, 89, 91, 144, 146, 162, 193-198, 201-208, 210, 211, 294, 295, 298, 306-308, 311, 312, 326, 334, 335, 340, 341, 345, 346, 348-350, 352, 355, 357, 368, 379, 422, 423, 425, 426, 663, 668, 670, 679, 691, 704, 708, 709, 720, 722, 723, 932, 933, 939, 940, 1137, 1139, 1142, 1145, 1155, 1163, 1164, 1171. Floridina antiqua (Smitt, 1873): PPP 86, 148, 156. Floridina minima Canu and Bassler, 1923: PPP 631. Floridinella parvula Canu and Bassler, 1928: PPP 47, 86, 156, 210, 308, 635, 663, 670, 818, 944, 950, 962, 963. Hiantopora intermedia Kirkpatrick, 1890: PPP 949. Hincksina species: DR Olsson 179. Labioporella aff. L. dumonti (Canu and Bassler, 1928): PPP 668. Labioporella miocenica (Canu and Bassler, 1919): DR NMB 15814, 15842, 15849, 15860, 15863, 15869, 15871, 15873, 15876, 15878, 15882, 15890, 15900, 15911, 15912, 15914, 15915, 15934, 15962, 15964, 16167, 16802, 16817, 16818, 16828, 16832-16839, 16842, 16855-16857, 16910, 16916, 16926— 16929, 16983, 16989, 17005, 17175, 17268, 17269, 17286; Ols- son 180; TU 1293; USGS 8702. Labioporella aff. L. miocenica (Canu and Bassler, 1919): PPP 634, 949, Micropora coriacea (Johnston, 1847): PPP 137, 162, 207, 307, 653. Mollia? species: DR NMB 17023. Monoporella nodulifera Hincks, 1881: PPP 65, 352, 720. Nellia tenella (Lamarck, 1816): PPP 53, 57, 63-66, 68, 69, 144, 162, 171, 178, 180, 193-195, 197, 198, 206, 207, 210-212, 295, 298, 312, 326, 334, 340, 345, 346, 348-350, 354, 355, 357, 365, 367, 370, 631, 634, 635, 653, 679, 697, 705, 709, 712, 720, 722, 723, 908, 932-934, 937, 939, 948, 962. DR NMB 15814, 15815, 15837, 15842, 15846, 15849, 15860, 15876, 15878, 15900, 15901, 15903-15907, 15910-15912, 15934, 15962, 16167, 16186, 16817, 16818, 16828, 16832-16839, 16842, 16844, 16856, 16857, 16913, 16916-16918, 16922, 16924, 16926— 16930, 16935, 16936, 16938, 16942, 16995, 17184, 17190, 17265, 17268, 17269, 17283, 17285-17290, 17307, 17327; Ols- son 179, 180; TU 1294; USGS 8525, 8702. Nellia cf. N. tenuis Harmer, 1926: DR NMB 15914, 15915, 16916, 16929, 16935, 17184, 17286, 17327. Onychocella aff. O. angulosa Reuss, 1847: PPP 47, 67, 86, 137, 144, 146, 156, 634, 635, 639, 644, 645, 710, 833, 943, 948-950, 962, 963. DR NMB 15851, 15853, 16811. Onychocella new species 1 [small zooids]|: DR NMB 15823, 15829, 15851, 15853, 15962, 16811, 16856. Parellisina curvirostris (Hincks, 1862): PPP 639, 640, 944, 949, 950. Poricellaria new species 1 [P. aff. P. ratoniensis (Waters, 1887) of Cheetham, 1973]: DR NMB_ 15823, 16817, 16935, 16942, 17184; Olsson 179. Poricellaria new species 2: DR NMB 16935. Retevirgula tubulata (Hastings, 1930): PPP 47, 86, 144, 146, 156, 949. DR NMB 17023. Scrupocellaria maderensis Busk, 1860: PPP 367, 948, 962. Scrupocellaria pusilla (Smitt, 1872): PPP 68, 631, 943, 944, 948, 949, 962, 963. Scrupocellaria regularis Osburn, 1940: PPP 53, 65, 66, 68, 86, 137, 144, 146, 156, 180, 210, 211, 295, 298, 312, 341, 350, 352, 370, 631, 634, 635, 679, 705, 708, 709, 720, 722, 723, 738, 773, 832, 833, 932, 933, 937, 939, 943, 944, 949, 962. Scrupocellaria aff. S. unguiculata Osburn, 1950: PPP 68, 69, 210. Scrupocellaria new species 1 [large zooids]: PPP 35, 65, 211, 212, 720, 722, 723. DR NMB 15815, 15823, 15828, 15837, 15838, 15842, 15849, 15860, 15864, 15900, 15901, 15903, 15904, 15910-15912, 15914, 15915, 15962, 16817, 16818, 16828, 16832, 16833, 16836, 16856, 16857, 16865, 16916, 16918, 16924, 16935, 16936, 16942, 17023, 17184, 17190, 17268, 17269; Olsson 179; USGS 8525. Scrupocellaria new species 2 [small zooids]: DR NMB 15823, 15828, 15838, 15964, 16817, 16818, 16828, 16833, 16838, 16935, 17023, 17184; Olsson 179. Skylonia dohmi (Sandberg, 1962): DR NMB 16935, 16936, 16938, 16942, 17184, 17190, 17265, 17283, 17285-17290, 17327; Ols- son 179. Smittipora aff. S. acutirostris (Canu and Bassler, 1928): PPP 47, 86, 144, 710, 738, 949, 962, 963. DR NMB 15878, 16855. Smittipora levinseni (Canu and Bassler, 1917): PPP 50, 65, 66, 86, 144, 146, 148, 156, 162, 205, 294, 295, 298, 308, 334, 340, 352, 634, 670, 709, 723, 944. Steginoporella magnilabris (Busk, 1854): PPP 35, 65, 66, 68, 69, 210-212, 222, 367, 653, 663, 685, 704, 708-710, 712, 720, 722, 723, 933, 948-950, 962, 963. DR NMB 16855, 16857. Steginoporella parvicella (Canu and Bassler, 1919): PPP 203, 308, 334, 335, 340, 345, 634, 679. DR NMB 15804, 15805, 15814, 15815, 15829, 15835, 15837, 15838, 15842, 15846, 15849, 15854, 15860, 15864, 15865, 15869, 15871, 15876, 15878, 15881, 15882, 15900, 15906, 15911, 15912, 15914, 15915, 15934, 15962, 15964, 16167, 16186, 16192, 16810, 16811, 16817, 16818, 16828, 16832-16834, 16842, 16855-16857, 16865, 16910, 16913-16918, 16922-16924, 16926-16930, 17175, 17269; Olsson 180; TU 1379; USGS 8525. Steginoporella new species 1: DR NMB 16817, 16818, 16828, 16832-16839, 16842, 16844, 16856, 16857, 16995. Steginoporella new species 2: DR NMB 16836-16838. Steginoporella new species 3 aff. S. connexa (Harmer, 1900): PPP 222, 634, 635, 639, 642, 645, 944, 948-950, 962, 963. Steginoporella new species 4 aff. S. connexa (Harmer, 1900): PPP 631, 640, 710, 738. “Steginoporella” cornuta Osburn, 1950: PPP 47, 86, 257, 704, 720. DR NMB 16856. Thalamoporella biperforata (Canu and Bassler, 1919): PPP 74, 672, 684, 704, 708, 709, 720, 722. DR NMB 15804, 15805, 15814, 15815, 15832, 15835, 15842, 15846, 15849, 15860, 15863— 15865, 15869, 15873, 15876, 15878, 15881, 15882, 15890, 15895-15897, 15900, 15901, 15903, 15904, 15906, 15907, 188 BULLETIN 357 15910-15916, 15934, 15962, 15964, 16167, 16186, 16192, 16810, 16817, 16818, 16828, 16832-16834, 16836-16839, 16841, 16842, 16844, 16846, 16852, 16855-16857, 16879, 16910, 16912-16918, 16922-16924, 16926-16930, 16932, 16935, 16936, 16938, 16942, 16959, 16961, 16971, 16972, 16988, 16995, 17004, 17175, 17184, 17190, 17265, 17268, 17269, 17278, 17288, 17327; Olsson 179; TU 1379; USGS 8525, 8702. Thalamoporella chubbi Lagaaij, 1959: PPP 212, 295. Thalamoporella new species 1 [similar to T. biperforata, but avi- cularia smaller and sibling zooids highly torqued (see Soule et al., 1987, for terminology)]: DR NMB 15815, 16817, 16818, 16828, 16832-16836, 16838, 16839, 16842, 16844, 16995. Thalamoporella new species 2 [like 7. chubbi, but avicularia slight- ly asymmetrical]: DR NMB 16818, 16833, 16836, 16838, 16914, 16917, 16923, 16927, 16932, 16935, 16938, 17184, 17265, 17268; Olsson 179. Thalamoporella new species 3 [small, pointed avicularia]: DR NMB 15900, 15903, 15904, 15906, 15907, 15910-15912, 15914, 15915, 16192, 16817, 16828, 16856, 16857, 16935; Olsson 179. Thalamoporella new species 4 [large zooids; rounded avicularia; sibling zooid slightly torqued]: PPP 35. Thalamoporella new species 5 [small, crescent-shaped avicularia; sibling zooid not torqued]: PPP 346. Thalamoporella new species 6 [rounded avicularia similar to zooids in size; sibling zooid moderately torqued]: PPP 370, 669, 708, TOON I2ON 22 23) 93i- Vibracellina laxibasis Canu and Bassler, 1928: PPP 65, 66, 68, 69, 162, 178, 180, 205, 210-212, 295, 306, 308, 352, 362, 367, 368, 370, 631, 634, 635, 639, 640, 642, 644, 653, 663, 704, 709, 720, 722, 723, 738, 937, 943, 944, 948-950, 962, 963. Vibracellina atf. V. laxibasis Canu and Bassler, 1928: DR NMB 15805, 15815, 15829, 15832, 15833, 15835, 15836, 15900, 15901, 15934, 15962, 16103, 16192, 16817, 16818, 16832, 16833, 16835, 16865, 16936, 16938, 16942, 17023, 17175, 17184; TU 1294. Vincularia new species: DR NMB 16935, 16936, 16938, 16942, 17184, 17190, 17265, 17285, 17287-17290, 17327; Olsson 179. Suborder ASCOPHORINA Levinsen, 1909 Adeonellopsis deformis (Canu and Bassler, 1919): DR NMB 15804, 15805, 15814, 15815, 15829, 15836-15838, 15840, 15854, 15860, 15873, 15874, 15878, 15881, 15882, 15896, 15900, 15901, 15906, 15912, 15934, 15962, 15964, 16167, 16186, 16191, 16192, 16802, 16817, 16818, 16828, 16832, 16833, 16835, 16860, 16865, 16910, 16913-16916, 16918, 16922, 16924, 16926-16930, 16932, 16975, 17268, 17269, 17273, 17307; Olsson 180; TU 1225, 1227A, 1294; USGS 8525, 8702. Adeonellopsis new species 1 [A. sp. of Cook, 1973, p. 252]: DR NMB 15838, 15840, 15842, 15846, 15849, 15860, 15863, 15865, 15871, 16910, 17175; TU 1227A, 1293. Adeonellopsis new species 2 [spiramen multiporous]: DR NMB 17273. Adeonellopsis new species 3: PPP 334, 367, 368, 679, 683, 697, 704, 708-710, 720, 722, 723. Aimulosia palliolata (Canu and Bassler, 1928): PPP 47, 69, 86, 144, 146, 148, 156, 204, 308, 639, 642, 645, 670, 710, 720, 738, 944, 949, 950, 962, 963. DR NMB 15828, 15829, 15964, 16103. Arthropoma cecilii (Audoin, 1826): PPP 205. new genus B new species y Scolaro, 1968: PPP 708, 709, 720, 722, 723. DR NMB 15823, 15828, 15835, 15838, 15842, 15846, 15849, 15851, 15853, 15854, 15860, 15863, 16811, 16856, 16910, 16935, 16936, 16938, 16942, 17023, 17184, 17190, 17265, 17278, 17283, 17284; Olsson 179; TU 1293; USGS 8702. Bellullopora bellula (Osburn, 1950): PPP 709, 950. Bracebridgia subsulcata (Smitt, 1873): PPP 68, 210, 212, 345, 631, 634, 635, 640, 642, 645, 663, 710, 712, 738, 940, 944, 948, 950, 962, 963. Buskea new species aff. B. dichotoma (Hincks, 1862): PPP 635, 639, 640, 642, 644, 653, 663, 943, 944, 948, 949, 950, 962, 963. Calyptooecia insidiosa Winston, 1984: PPP 639, 644, 949, 950, 962, 963. DR NMB 15838, 15840, 15846, 15849, 15853, 16865; USGS 8702. Celleporaria albirostris (Smitt, 1873): PPP 53, 68, 69, 71, 86, 144, 177, 210, 212, 346, 352, 361, 367, 631, 634, 635, 639, 640, 642, 644, 645, 653, 663, 712, 738, 832, 943, 944, 948-950, 962, 963. Celleporaria brunnea (Hincks, 1884): PPP 47, 64—66, 68, 86, 137, 148, 194, 198, 204-206, 210, 212, 214, 294, 295, 298, 308, 334, 335, 340, 341, 345, 346, 348-350, 362, 368, 391, 631, 634, 635, 639, 640, 642, 704, 709, 720, 722, 943, 944, 948-950, 962, 963. DR NMB 15804, 15815, 15832, 15833, 15837, 15846, 15849, 15853, 15865, 15871, 15934, 16103, 16935, 16936, 16938, 17175. Celleporaria magnifica (Osburn, 1914): PPP 148, 178, 180, 334, 367, 368, 634, 635, 639, 640, 642, 645, 704, 705, 709, 943, 950, 963. DR NMB 15842, 15846, 15849, 15860, 15878, 15881, 15901, 15911, 15912, 15914, 15915, 15962, 16167, 16192, 16817, 16818, 16828, 16832-16839, 16842, 16844, 16856, 16857, 16865, 16879, 16913-16918, 16924, 16926-16929, 16942, 16975, 16976, 16983, 16984, 16995, 17023, 17268, 17269, 17278, 17283; Olsson 179; TU 1294, 1379; USGS 8525, 8702. Celleporaria new species: PPP 352. Characodoma contractum (Waters, 1899): PPP 47, 60, 65, 68, 69, 86, 137, 144, 146, 148, 156, 162, 178, 180, 193, 194, 196, 204— 207, 307, 311, 326, 340, 346, 350, 352, 354, 355, 357, 367, 368, 422, 631, 634, 635, 639, 640, 642, 645, 709, 710, 720, 722, 723, 943, 944, 948-950, 962, 963. Cigclisula porosa (Canu and Bassler, 1919): PPP 65, 68, 69, 74, 205, 206, 210, 294, 298, 307, 308, 340, 341, 345, 346, 348, 365, 391, 631, 634, 635, 639, 640, 642, 645, 663, 670, 679, 697, 704, 708, 709, 720, 722, 723, 932, 944, 949, 950, 962, 963. DR NMB 15805, 15842, 15846, 15851, 15854, 15860, 15863-15865, 15871, 15934, 16828, 16834-16836, 16842, 16844, 16910, 16913, 16915, 16916, 16918, 16924, 16926-16928, 16932, 16935, 16938, 16942, 17184, 17190, 17265, 17268, 17269, 17283, 17285-17290, 17327; Olsson 179; TU 1294. Coleopora aft. C. americana Osburn, 1940: PPP 644, 949, 962. Coleopora granulosa Canu and Bassler, 1928: PPP 205. DR NMB 15823, 15828, 15829, 15832, 15833, 15838, 15840, 15849, 15851, 15853, 15962, 16811, 16818, 17023, 17175. Crepidacantha longiseta Canu and Bassler, 1928: PPP 639, 949, 950. Crepidacantha poissonii (Audouin, 1826): PPP 60, 67, 86, 144, 949. Cycloperiella rubra Canu and Bassler, 1923: PPP 47, 65, 86, 144, 162, 340, 350, 352, 722, 723. Cycloperiella new species: PPP 65, 205, 294, 295, 298, 306, 307, 334, 335, 346, 348, 350, 352. Ditaxipora new species 1: PPP 65, 349, 355, 708, 711, 720, 722, 723. DR NMB 15823, 15838, 15849, 15900, 15911, 15912, 15914, 15915, 16817, 16818, 16828, 16833, 16836, 16838, 16844, 16856, 16913, 16918, 16929, 17023; USGS 8525. Ditaxipora new species 2: DR NMB 15837, 15846, 15851, 15853. Drepanophora tuberculatum (Osburn, 1914): PPP 631. DR NMB 15842, 15846, 15849, 15853, 16811, 16828, 16832, 16856, 16865, 16915, 16916, 16926, 16928, 16929; USGS 8702. Escharella? species: PPP 210. CHEILOSTOME BRYOZOA: CHEETHAM ET AL. 189 Escharina pesanseris (Smitt, 1873): PPP 639, 663, 709, 738, 948— 950. DR NMB 15846, 15853, 15860, 17023. Escharina porosa (Smitt, 1873): PPP 634, 635, 639, 640, 642, 944, 948-950, 963. DR NMB 15838, 15851, 15853, 15962, 16811, 16855, 16856. Escharoides costifer (Osburn, 1914): PPP 949. Escharoides new species [like E. costifer, but with avicularia curved and directed distolaterally]: DR NMB 15853, 15865, 16818. Exechonella ct. E. antillea (Osburn, 1927): DR NMB 15838, 15846. Fedora aft. F. nodosa Silén, 1947: DR NMB 15804, 15805, 15814, 15815, 15829, 15833, 15835, 15836, 15863, 15934, 15962, 15964, 16828, 17175; Olsson 180; TU 1293, 1294. Gemellipora new species 1 [small zooids]: DR NMB 15823, 15832, 15833, 17023. Gemellipora new species 2 [large zooids]: DR NMB 15823, 15832 15833, 17023. Gemelliporella glabra Smitt, 1873: PPP 361, 710, 712. Gemelliporella punctata Canu and Bassler, 1919: PPP 35, 68, 69, 72, 162, 180, 183, 210-212, 214, 340, 365, 367, 368, 370, 390, 422, 645, 653, 663, 668, 679, 683, 689, 932, 935, 937, 940. DR NMB 15804, 15814, 15815, 15835, 15836, 15842, 15849, 15878, 15881, 15882, 15890, 15900, 15912, 15962, 16167, 16817, 16818, 16828, 16832, 16833, 16835, 16836, 16837, 16839, 16842, 16844, 16857, 16860, 16913, 16915, 16916, 16918, 16924, 16926, 16927, 16928, 16929, 16995, 17023, 17175, 17268, 17269; TU 1225, 1293, 1379; USGS 8525, 8702. Gemelliporella? new species [colony adeoniform]: PPP 365, 634, 640, 642, 644, 645, 663, 704, 720, 722, 935, 943, 944, 948-950, 962, 963. DR NMB 15804, 15814, 15815, 15823, 15962, 16103. Gemelliporidra magniporosa (Canu and Bassler, 1923): PPP 47, 639, 949. Gemelliporidra multilamellosa (Canu and Bassler, 1923): PPP 55, 65, 631, 634, 635, 639, 640, 642, 644, 645, 663, 709, 710, 712, 943, 944, 948-950, 962, 963. Gemelliporidra? species: DR NMB 15815, 15846, 15849, 15851, 15853, 15854, 15878, 15911, 15962, 15964, 16811, 16817, 16818, 16855, 16856, 16865, 17268, 17278. Gephyrophora ct. G. rubra Osburn, 1940: DR NMB 16834. Gigantopora fenestrata (Smitt, 1873): PPP 639, 644, 710, 949, 963. DR NMB 15851, 17023. Hippaliosina rostrigera (Smitt, 1873): PPP 65, 68, 69, 86, 205, 210, 212, 295, 298, 326, 631, 634, 635, 639, 640, 642, 645, 653, 709, 720, 722, 723, 944, 948-950, 962, 963. Hippaliosina new species: DR NMB 16855, 16915. Hippomenella? fissurata (Canu and Bassler, 1928): PPP 65, 298, 634, 635, 639, 640, 645, 653, 943, 944, 948, 950, 962, 963. Hippomenella? atf. H.? fissurata (Canu and Bassler, 1928): DR NMB 16834, 16835, 16838, 16839, 16995. Hippopetraliella cf. H. marginata (Canu and Bassler, 1928): DR NMB 15842, 15846, 15854, 16817, 16828, 16832, 16833, 16836-16839, 16842, 16856, 15849. Hippopleurifera mucronata (Smitt, 1873): PPP 47, 86, 144, 146, 148, 352. Hippopleurifera new species 1: DR NMB 16818, 16828, 16833. Hippopleurifera new species 2: PPP 710, 949. Hippopodina ct. H. bernardi Lagaaij, 1963: DR NMB 15911, 15914, 16835, 16836. Hippopodina feegeensis (Busk, 1884): PPP 635. Hippopodina aft. H. feegeensis (Busk, 1884): DR NMB 15849, 15851, 15853, 15915, 16811, 16832, 16935, 16936. Hippoporella costulata Canu and Bassler, 1923: PPP 68, 69, 86, 144, 156, 210, 212, 631, 634, 635, 639, 645, 704, 709, 710, 720, 738, 944, 950, 962, 963. Hippoporella gorgonensis Hastings, 1930: PPP 47, 64—66, 68, 86, 89, 137, 144, 146, 148, 156, 162, 193-196, 198, 204-208, 294, 295, 307, 311, 312, 326, 345, 350, 352, 355, 379, 631, 639, 640, 644, 653, 670, 709, 937, 943, 949, 950, 962, 963, 1171. DR NMB 15809, 15814, 15836, 15837, 15842, 15846, 15849, 15851, 15853, 15854, 15860, 15863, 15865, 15881, 15900, 15964, 16802, 16811, 16833-16836, 16839, 16844, 16855-16857, 16860, 16865, 16914, 16995, 17268; USGS 8702. Hippoporella att. H. rimata Osburn, 1952: PPP 47, 148, 205, 206, 307, 352, 962. Hippoporidra edax (Busk, 1859): PPP 10, 47, 65, 66, 86, 137, 144, 146, 148, 156, 180, 188, 212, 354, 639, 697, 704, 709, 720, 722, 723, 832, 950, 1142. Hippoporina aculeata (Canu and Bassler, 1928): PPP 65, 68, 69, 144, 148, 156, 162, 204, 206, 210, 291, 326, 340, 345, 350, 352, 355, 631, 640, 642, 645, 704, 708, 710, 738, 944. Hippoporina new species 1: PPP 60, 294, 298. Hippoporina new species 2: PPP 66, 86, 137, 146. Hippoporina new species 3: PPP 86, 137, 144. Hippoporina? species: DR NMB 15840, 15934, 16828, 17175; Ols- son 179. Jaculina species x (Scolaro, 1968): DR NMB_ 16935, 17184, 17285-17289, Olsson 179. Lagenicella marginata (Canu and Bassler, 1930): PPP 635, 642, 644, 738, 949, 962. Lagenicella aft. L. mexicana (Osburn, 1952): PPP 68, 86, 137, 144, 146, 156, 211, 312, 367, 634, 781, 835, 950. Lagenicella new species: PPP 86, 144, 146, 210, 645, 949, 963. Lagenicella species: DR NMB 15832, 16817, 16818, 16828, 16833, 16836, 16841, 16842, 16928, 17269. Lagenipora species: PPP 640, 712. Mamillopora cavernulosa Canu and Bassler, 1928: PPP 55, 57, 65, 197, 198, 294, 295, 298, 306-308, 312, 334, 335, 340, 345, 350, 352, 354, 357, 358, 635, 639, 640, 653, 663, 688, 689, 691, 695, 704, 709, 710, 720, 722, 723, 738, 937, 940, 944, 950. Mamillopora tuberosa Canu and Bassler, 1918: PPP 10, 47, 55, 57, 60, 63-69, 71, 72, 74, 86, 137, 144, 146, 148, 156, 162, 178, 180, 183, 193-196, 198, 203-208, 210, 214, 294, 306, 307, 311, 312, 334, 335, 341, 345, 346, 348, 350, 352, 354, 355, 357, 362, 365, 367, 368, 370, 379, 419, 422, 423, 425, 426, 631, 634, 635, 639, 640, 642, 644, 645, 653, 668-673, 675-679, 683, 685, 689, 691, 697, 704, 705, 708, 709, 720, 722, 723, 832, 833, 908, 931— 933, 935, 939, 943, 944, 948, 962, 963, 1145, 1171. DR NMB 15805, 15814, 15815, 15833, 15835, 15836, 15840, 15846, 15860, 15863-15865, 15869, 15871, 15873, 15874, 15876, 15878, 15881, 15882, 15890, 15895-15897, 15900, 15901, 15903, 15904, 15906, 15907, 15910-15912, 15914, 15915, 15934, 15942, 15962, 15964, 15965, 15968, 15969, 15974, 16167, 16186, 16191, 16192, 16802, 16810, 16817, 16818, 16824, 16827, 16828, 16832-16839, 16842, 16844, 16854, 16856, 16857, 16860, 16865, 16910, 16912-16915, 16917, 16918, 16922-16924, 16926, 16927, 16929, 16930, 16932, 16935, 16938, 16961, 16970, 16971, 16973, 16975, 16978, 16985, 16995, 17005, 17175, 17184 17268, 17269, 17283, 17286-17290, 17322, 17327; Olsson 179, 180, TU 1225, TU 1293, 1294; USGS 8525, 8702. Mamillopora new species 1 [large, pointed avicularia on basal side]: PPP 53, 171, 175, 177, 178, 180, 212, 358, 361, 362, 365, 367, 368, 370, 631. Mamillopora new species 2 [tiny, conical colonies]: PPP 61, 62, 63, 187, 200, 201, 308, 362. Margaretta buski Harmer, 1957: PPP 35, 65, 68, 86, 144, 146, 156, 210, 211, 350, 367, 631, 634, 635, 640, 642, 644, 645, 663, 679, 190 BULLETIN 357 708, 709, 710, 712, 720, 738, 932, 939, 940, 943, 944, 948, 949, 950, 962, 963. Margaretta new species 1: DR NMB 15804, 15814, 15815, 15837, 15838, 15840, 15842, 15846, 15849, 15851, 15853, 15854, 15860, 15863, 15864, 15871, 15878, 15881, 15882, 15900, 15914, 15915, 15934, 15962, 15964, 16167, 16811, 16817, 16818, 16828, 16832, 16833, 16835-16838, 16842, 16856, 16857, 16910, 16913, 16916, 16918, 16926, 16927, 16929, 16936, 16938, 16942, 17023, 17190, 17265, 17268; Olsson 179; USGS 8702. Margaretta new species 2: DR NMB 17184, 17290. Membraniporella? species: PPP 634, 670. Metrarabdotos auriculatum Canu and Bassler, 1923: PPP 68, 210, 211, 212, 671, 689, 691. DR NMB 15876, 15878, 15881, 15882, 16167, 16186, 16817, 16818, 16824, 16828, 16836, 16837, 16839, 16842-16844, 16846, 16852, 16857, 16858, 16959, 16961, 16962, 16970-16972, 16984, 16986, 16988, 16989, 16993, 16995, 17005, 17012, 17019, 17026. Metrarabdotos colligatum Canu and Bassler, 1919: PPP 191, 423. DR NMB 15869, 15876, 15878, 15881, 15890, 15895-15897, 15900, 15904, 15906, 15907, 15910-15912, 15914, 15915, 16167, 16191, 16802, 16844, 16852, 16857, 16910, 16912— 16918, 16922-16924, 16926-16930, 16932, 17175, 17268, 17269; TU 1293, 1294, 1379; USGS 8525, 8702. Metrarabdotos lacrymosum Canu and Bassler, 1919: PPP 212, 362. DR NMB 15804, 15814, 15815, 15833, 15835, 15846, 15849, 15854, 15860, 15863-15865, 15873, 15874, 15878, 15881, 15882, 15890, 15911-15913, 15915, 15934, 15962, 15964, 16167, 16802, 16817, 16818, 16824, 16828, 16832-16839, 16842, 16844, 16858, 16930, 16984, 16988, 16989, 16995, 17286; TU 1225. Metrarabdotos pacificum (Osburn, 1952): PPP 66, 86, 144, 146, 156, 257, 271. Metrarabdotos tenue (Busk, 1884): DR NMB 17268. Metrarabdotos unguiculatum Canu and Bassler, 1928: PPP 211, 723. DR NMB 16833. Metrarabdotos new species 1 Cheetham, 1986: DR NMB 17265, 17284, 17286-17290; Olsson 179. Metrarabdotos new species 2 Cheetham, 1986: DR NMB 16935, 16936, 16938, 16942, 17184, 17265; Olsson 179. Metrarabdotos new species 3 Cheetham, 1986: DR NMB 15837, 15838, 15840, 15842, 15846, 15860, 15863, 15864, 15900, 16191, 16910, 17175; TU 1293; USGS 8702. Metrarabdotos new species 4 Cheetham, 1986: DR NMB 15804, 15814, 15835, 15836, 15838, 15840, 15842, 15846, 15849, 15860, 15863-15865, 15869, 15871, 15934, 15962, 16810, 16811, 16833; Olsson 180; TU 1225, 1293. Metrarabdotos new species 5 Cheetham, 1986: DR NMB 15900, 15901, 15903, 15906, 15910, 15911, 16839, 16844, 16986, 16988, 16989, 16993, 16995. Metrarabdotos new species 6 Cheetham, 1986: DR NMB 15878, 15881, 15882, 15904, 15906, 16191. Metrarabdotos new species 7 Cheetham, 1986: DR NMB 15804, 15805, 15814, 15815, 15835, 15836, 15842, 15849, 15934, 16910, 17175; Olsson 180; TU 1293. Metrarabdotos new species 8 Cheetham, 1986: PPP 35, 60, 68, 210, 211, 352, 653. DR NMB 15860, 15863, 15864, 15869, 15873, 16810; USGS 8702. Metrarabdotos new species 9 Cheetham, 1986: DR NMB 16817, 16824, 16828, 16832-16834, 16838, 16879, 16959, 16961, 16970-16973, 16975, 16976, 16978, 17005, 17268; USGS 8702. Metrarabdotos new species 10 Cheetham, 1986: PPP 214, 335, 348, 349, 352, 663, 679, 708, 709, 720, 722, 723. DR NMB 15804, 15815, 15934, 15962, 16817, 16818, 16824, 16828, 16832— 16835, 16837, 16838, 16842, 16844, 16961, 16976, 16983, 16984; USGS 8702. Microporella cf. M. ciliata (Pallas, 1766): DR NMB 15851, 15853, 17023. Microporella normani Canu and Bassler, 1928: PPP 86, 144, 148, 156; 162; 773,832; 1152: Microporella umbracula (Audouin, 1826): PPP 47, 65, 86, 144, 146, 148, 180, 352, 354, 367, 391, 635, 639, 640, 642, 653, 663, 709, 720, 723, 943, 944, 948, 950, 962, 963. Odontoporella adpressa (Busk, 1854): PPP 47, 66, 86, 144, 146, 148, 156, 832, 949, 950. Parasmittina aff. P. areolata (Canu and Bassler, 1927): PPP 65, 204, 206, 210, 294, 307, 326, 334, 335, 634, 639. DR NMB 16828, 16832-16834, 16865, 16913-16918, 16922, 16926— 16929, 16932, 16935, 17269; TU 1294; USGS 8525. Parasmittina crosslandi (Hastings, 1930): PPP 47, 68, 86, 144, 146, 156, 196, 206, 257, 271, 292, 307, 350, 352. Parasmittina fraseri Osburn, 1952: PPP 47, 60, 66, 86, 144, 146, 148, 156, 205, 271, 291, 292. Parasmittina hastingsae Soule and Soule, 1973: PPP 68, 69, 963. Parasmittina aff. P. murarmata (Kirkpatrick, 1888): PPP 670, 671, 708, 937, 963. Parasmittina parsevaliformis Soule and Soule, 1973: PPP 65, 137, 205, 210, 212, 222, 294, 298, 308, 340, 631, 634, 635, 639, 642, 644, 645, 704, 722, 943, 944, 948, 950, 962, 963. DR NMB 15934, 16817, 16818, 16828, 16832-16834, 16836, 16838, 16842, 16865, 17268; Olsson 179. Parasmittina spathulata (Smitt, 1873): PPP 222, 352, 640, 642, 644, 710, 943, 944, 949, 950, 962, 963. DR NMB 15815, 15849, 15851, 15878, 15881, 15882, 15912, 15914, 16167, 16811, 16836, 16837. Parasmittina new species 1: PPP 65, 222, 634, 635, 644, 944. DR NMB 15881, 15915, 16828, 16833, 16834, 16836-16839, 16856, 16916, 16995; USGS 8702. Parasmittina new species 2: DR NMB 16856, 16913, 16915-16918, 16922, 16926-16929, 17269; TU 1294; USGS 8525. Parasmittina new species 3: PPP 640, 642, 645, 709, 710, 720, 944, 949, 950, 962. DR NMB 15849, 16818, 17265. Parasmittina new species 4: DR NMB 16913, 16916, 16922, 16928; TU 1294; USGS 8525. Parasmittina new species 5: DR NMB 15842. Parasmittina new species 6: DR NMB 15851, 15853. Parasmittina new species 7: PPP 631. Parasmittina new species 8: PPP 86, 148, 205. Parkermavella punctigera (MacGillivray, 1883): PPP 631, 949. Pasythea tulipifera (Ellis and Solander, 1786): PPP 64, 65. Pasythea new species: DR NMB 15900, 15901, 15906, 15907, 15934, 15962, 16833, 16835, 16836, 16838, 16916, 16926, 16928, 16935, 16936, 16938, 16995, 17184, 17190, 17265, 17307, 17327; Olsson 179; TU 1294; USGS 8525, 8702. Petraliella bisinuata (Smitt, 1873): PPP 68, 69, 210-212, 361, 362, 631, 634, 635, 639, 640, 642, 645, 663, 704, 712, 738, 943, 944, 950, 963. Pleurocodonellina species: PPP 68, 210. Puellina innominata (Couch, 1844): PPP 86, 144, 146, 709. Puellina radiata (Moll, 1803): PPP 148, 205, 210, 348, 352, 631, 640, 670, 738, 944, 949, 950, 963. Puellina new species aff. P. radiata (Moll, 1803): PPP 69, 639, 642, 645, 653, 708-710, 723, 943, 949, 950, 962, 963. Puellina species: DR NMB 15823, 15853, 16811, 16860, 17023. Reptadeonella bipartita (Canu and Bassler, 1928): PPP 47, 65—69, 86, 144, 146, 148, 156, 205, 271, 352, 670, 937. DR NMB 15842, 15864, 16855, 16856; TU 1227A; USGS 8702. CHEILOSTOME BRYOZOA: CHEETHAM ET AL. 191 Reptadeonella hastingsae Cheetham and Sandberg, 1964: DR NMB 15878, 16167, 16856, 16865,16942. Reptadeonella tubulifera (Canu and Bassler, 1930): PPP 47, 66, 86, 144, 146, 148, 156, 201, 214. DR NMB 17023. Reptadeonella new species [similar to R. joloensis (Bassler, 1935) and other Pacific species]: DR NMB 15823, 15828, 15829, 15840, 15849, 15851, 15853, 15854, 16811. Reteporellina evelinae Marcus, 1958: PPP 35, 68, 69, 86, 144, 156, 180, 183, 210-212, 367, 368, 631, 634, 635, 639, 640, 642, 644, 645, 653, 663, 704, 708, 709, 712, 720, 722, 723, 738, 943, 944, 948-950, 962, 963. DR NMB 15823, 15828, 15832, 15833, 15837, 15853, 16856, 17023. Rhynchozoon aff. R. phyrnoglossum Marcus, 1932: PPP 148. Rhynchozoon rostratum (Busk, 1856): PPP 47, 63, 65, 66, 86, 144, 206, 207, 294, 352, 355, 422, 634, 639, 642, 644, 943, 944, 949, 950, 962. Rhynchozoon verruculatum (Smitt, 1873): PPP 146, 148, 156. Rhynchozoon species [at least three species]: DR NMB 15842, 15869, 15871, 15878, 15881, 15882, 15906, 15911, 15912, 15914, 15915, 15962, 16167, 16186, 16810, 16817, 16818, 16828, 16832-16836, 16839, 16842, 16844, 16855-16857, 16913-16916, 16918, 16922, 16926-16930, 16935, 16936, 16938, 16942, 17184, 17265, 17269, 17288-17290; Olsson 179; TU 1294, 1379. Savignyella species: DR NMB 15823, 15837, 16817, 17023, 17184: Olsson 179. savignyellid? species [new genus with biserial branches]: DR NMB 17184; Olsson 179. Schedocleidochasma cleidostoma (Smitt, 1873): PPP 47, 66, 68, 69, 86, 144, 162, 180, 210, 212, 294, 295, 298, 308, 334, 335, 340, 341, 345, 348, 350, 361, 362, 367, 368, 370, 634, 635, 639, 640, 642, 644, 653, 663, 668, 669, 671, 704, 709, 710, 720, 722, 723, 943, 944, 948-950, 963. Schedocleidochasma porcellanum (Busk, 1860): PPP 86, 146, 148, 156. DR NMB 15833, 15846, 15849, 15851, 15853, 15854, 15863, 15882, 15911, 15912, 15914, 15915, 15934, 16811, 16817, 16818, 16832-16836, 16842, 16855, 16856, 16865, 16913, 16915, 16916, 16918, 16922, 16924, 16926-16930, 16936, 16938, 17190, 17265, 17269; TU 1294; USGS 8525. Schedocleidochasma new species 1: PPP 64, 65, 198, 308, 340, 341, 345, 350, 352, 634. Schedocleidochasma new species 2: PPP 937, 949. Schedocleidochasma new species 3: PPP 64—66, 68, 69, 178, 180, 204, 205, 210, 212, 214, 294, 295, 298, 306-308, 312, 326, 334, 335, 340, 341, 345, 348-350, 352, 354, 358, 367, 368, 370, 631, 634, 635, 639, 640, 642, 663, 668, 670, 672, 710, 738, 944, 948, 950, 962, 963. Schizoporella cornuta (Gabb and Horn, 1862): PPP 60, 63-69, 86, 144, 146, 148, 156, 162, 188, 194, 196, 198, 204, 206, 212, 294, 295, 298, 308, 311, 312, 326, 334, 346, 350, 352, 355, 635, 639, 640, 642, 705, 709, 722, 723, 835, 944, 949, 950, 963, 1171. Schizoporella floridana Osburn, 1914: PPP 419, 422. Schizoporella magniporosa (Canu and Bassler, 1923): PPP 334, 368, 679, 691, 704, 708, 709, 720, 722, 723. DR NMB 15804, 15805, 15814, 15815, 15829, 15835-15838, 15840, 15842, 15846, 15849, 15853, 15860, 15863-15865, 15869, 15878, 15881, 15882, 15900, 15911, 15934, 15962, 15964, 16103, 16167, 16186, 16810, 16811, 16817, 16818, 16828, 16832, 16833-16835, 16842, 16865, 16910, 16913, 16915, 16916, 16918, 16924, 16926-16930, 17023, 17175, 17268, 17269; TU 1293, 1379; USGS 8525, 8702. Semihaswellia sinuosa (Canu and Bassler, 1928): PPP 68, 210, 631, 644, 645, 653, 663, 738, 943, 944, 948-950, 962, 963. DR NMB 15804, 15814, 15815, 15823, 15828, 15837, 15838, 15842, 15846, 15849, 15854, 15860, 15863-15865, 15871, 15934, 15962, 15964, 16811, 16910, 17023, 17175; USGS 8702. Smittina? new species 1: PPP 823, 833. Smittina? new species 2: PPP 944, 963. Smittoidea maleposita (Canu and Bassler, 1923): PPP 86, 137, 144, 146, 156, 180, 335, 350, 367, 368, 709. DR NMB 16938; Olsson 179. Smittoidea pacifica Soule and Soule, 1973: PPP 653, 949. Smittoidea prolifica Osburn, 1952: PPP 86. Stylopoma informatum (Lonsdale, 1845): DR TU 1294; USGS 8525. Stylopoma minutum (Canu and Bassler, 1923): PPP 55, 65, 66, 67, 68, 69, 144, 146, 148, 205, 206, 307, 334, 335, 340, 346, 350, 352, 355, 464, 704, 709, 937. DR NMB 15882, 16837, 16916. Stylopoma projectum (Canu and Bassler, 1923): PPP 631, 634, 712, (22123. Stylopoma spongites (Pallas, 1766): PPP 210, 222, 368, 464, 639, 645, 709, 944, 948, 962, 963. DR NMB 15912, 15914, 15915, 16817, 16818, 16828, 16832, 16834-16837, 16839, 16844, 16856, 16857, 16913, 16914, 16916, 16928, 16929, 16932, 16995, 17268, 17269; TU 1294; USGS 8525. Stylopoma new species 3 Jackson and Cheetham, 1994: PPP 631, 634, 635, 640, 712, 720, 943, 949, 950. Stylopoma new species 4 Jackson and Cheetham, 1994: DR NMB 15838, 15851, 15962; Olsson 180. Stylopoma new species 5 Jackson and Cheetham, 1994: DR NMB 16928, 16929. Stylopoma new species 6 Jackson and Cheetham, 1994: DR NMB 16832, 16833, 16913, 16916, 16924, 16928, 17265, 17269; Ols- son 179. Stylopoma new species 7 Jackson and Cheetham, 1994: DR NMB 16811, 16833, 16835, 16838, 16842, 16865, 17268. Stylopoma new species 11 Jackson and Cheetham, 1994: PPP 367, 634, 712, 943, 944, 949, 950, 962, 963. Stylopoma new species 13 Jackson and Cheetham, 1994: PPP 294, 295, 298, 710, 722. DR NMB 15869, 16928. Stylopoma new species 14 Jackson and Cheetham, 1994: PPP 639, 640. DR NMB 15863, 16910. Tetraplaria dichotoma (Osburn, 1914): PPP 180, 631, 720. DR NMB 15805, 15823, 15827, 15835, 15838, 15846, 15849, 15851, 15853, 15860, 15964, 16811, 16828, 16833, 16856, 17268. Trematooecia aviculifera (Canu and Bassler, 1923): PPP 55, 210, 634, 710, 738, 949, 962, 963. DR NMB 15849, 15962, 16916, 16928, 16929; USGS 8702. Trematooecia cf. T. hexagonalis (Canu and Bassler, 1930): DR NMB 15805, 15814, 15838, 15854, 16811, 17023; USGS 8702. Trematooecia turrita (Smitt, 1873): PPP 55, 64, 65, 86, 146, 148, 156, 645, 720, 944, 949, 950, 963. DR NMB 15804, 15814, 15815, 15838, 15840, 15849, 15962, 16828, 16860, 16865. Trematooecia vaughani (Canu and Bassler, 1919): DR NMB 15836, 15838, 15840, 15846, 15878, 15882, 15890, 16167, 16817, 16818, 16828, 16832-16839, 16842, 16844, 16855, 16856, 16916, 16928, 16929, 16984, 16995; USGS 8702. Tremogasterina mucronata (Smitt, 1873): PPP 68, 69, 198, 210, 211, 294, 295, 298, 308, 334, 335, 340, 341, 345, 348, 368, 631, 634, 635, 639, 640, 642, 644, 645, 653, 685, 704, 709, 710, 720, 722, 723, 738, 932, 943, 944, 948-950, 962, 963. DR NMB 15804, 15814, 15815, 15836-15838, 15842, 15846, 15849, 15851, 15853, 15854, 15860, 15863, 15864, 15876, 15878, 15881, 15882, 15890, 15911, 15912, 15914, 15915, 15934, 15962, 15964, 16167, 16811, 16817, 16818, 16828, 16832, 16833-16839, 16842, 16844, 16857, 16865, 16910, 16917, 16918, 16922-16924, 16926-16929, 16935, 16936, 16938, 16942, 16983, 16988, 16989, 16993, 16995, 17023, 17175, 192 BULLETIN 357 17184, 17190, 17265, 17268, 17269, 17288, 17289, 17290, 17327; Olsson 179; TU 1293. Tremoschizodina lata (Smitt, 1873): PPP 631, 738. Triporula stellata (Smitt, 1873): PPP 137, 631, 639, 640, 663, 943, 950, 963. Trypostega venusta (Norman, 1864): PPP 34, 47, 60, 66, 69, 86, 144, 146, 148, 150, 156, 348, 391, 634, 709, 720, 722, 723. Trypostega species: DR NMB 16817, 16836. Turbicellepora species: PPP 47, 65, 66, 86, 144, 146, 148, 1 177, 178, 205, 210, 212, 295, 298, 306, 308, 334, 352, 422, 63 639, 640, 642, 663, 670, 671, 672, 704, 705, 709, 720, 722, 7 738, 944, 948, 962, 963. Vittaticella species: DR NMB 16833, 16935, 16938, 17184; Olsson 179; USGS 8525. Watersipora subovoidea (d’Orbigny, 1852): PPP 148. CHAPTER 9 DIVERSITY AND ASSEMBLAGES OF NEOGENE CARIBBEAN MOLLUSCA OF LOWER CENTRAL AMERICA JEREMY B. C. JACKSON Smithsonian Institution Smithsonian Tropical Research Institute Washington, D.C. 20560-0580, U.S.A. Scripps Institution University of California at San Diego La Jolla, California 92093-0244, U.S.A. JONATHAN A. TODD Department of Palaeontology Natural History Museum London SW7 5BD, United Kingdom HELENA FORTUNATO Smithsonian Institution Smithsonian Tropical Research Institute Washington, D.C. 20560-0580, U.S.A. PETER JUNG Naturhistorisches Museum Basel Augustinergasse 2 CH-4051 Basel, Switzerland INTRODUCTION Our goal is to document and understand paleobio- logical patterns and trends for tropical American mol- lusks in marine coastal environments in relation to the rise of the lower Central American isthmus and global climate change. The time frame is the Neogene defined by the new Cenozoic chronology as the last 23.7 mil- lion years (Berggren et al., 1995a, 1995b) when most modern clades of mollusks diversified. To this end, we have attempted to sample the cells of a 3-dimensional matrix whose axes are gradients in time, spatial scale and environment. Recent molluscan faunas differ greatly among geographic regions and environments (Sanders, 1968; Jackson, 1972, 1974; Rex, 1981; Roy et al., 1996, 1998), and the same was true in the past (Valentine and Jablonski, 1993; Koch 1995,1996; Roy et al., 1995). Therefore, it is necessary in paleobiolog- ical surveys to sample many geographic locations and environments throughout the entire time interval in question, and to know something about the environ- ments of deposition of each sample. Otherwise, it is impossible to establish whether differences among fau- nas of different ages represent temporal trends or are artifacts of sampling different regions or environments over time. In addition, it is necessary to sample rig- orously and consistently every cell of the 3-dimen- sional matrix so that observed differences between ages, places and environments are not artifacts of dif- ferential sampling effort (Koch 1987, 1995, 1996; Koch and Morgan, 1988; Sepkoski and Koch, 1995). For all these reasons, hypotheses of major evolution- ary events (Petuch, 1995) or the division of paleobi- ogeographic provinces (Petuch, 1988) based on a few new taxa are only speculation. Neogene mollusks of tropical America have been studied extensively during the past century, with de- tailed monographic descriptions of faunas ranging from Trinidad to Ecuador in the south, to Florida and Chiapas in the north. Some of these published faunas are very diverse (Text-fig. 1, Table 1), with the record held by Woodring’s (1925-1928) systematic mono- graphs of the Late Pliocene Bowden Formation in Ja- maica with 347 genera or subgenera and 610 species. However, most of the other paleontological collections were made by petroleum geologists for the purpose of 194 BULLETIN 357 Caribbean Sea G & if 2 “3 COSTA RICA A 11,12 5 ES ae 19 16-18 7 4-10 2 0 4 - 20,21 « a VENEZUELA Eastern Facific COLOMBIA 22,23 Text-figure 1.—Location of Neogene molluscan faunas listed in Table 1. Table 1.—Numbers of genera and subgenera reported in the major monographs of Neogene and Quaternary molluscan faunas of southern tropical America from Trinidad to Costa Rica to Ecuador. Taxa were counted as listed by the authors with no attempt to reconcile taxonomic usage over the years. G/SG = genera and subgenera. Blank spaces for scaphopods mean that they were not described rather than absent. Bivalves Scaphopods Gastropods Total mollusks Formation or fauna G/SG Species G/SG Species G/SG Species G/SG Species 1. Bowden, Jamaica 117 187 10 20 220 406 347 610 2. Grand Bay, Carriacou 18 20 1 2 70 87 89 109 3. Belmont, Carriacou 8 8 1 1 34 44 43 53 4. Brasso, Trinidad 21 28 18 23 39 51 5. Manzanilla, Trinidad 24 35 16 22 40 57 6. Springvale, Trinidad 25 35 31 47 56 92 7. Springvale, Trinidad 46 56 2 3 61 95 119 154 8. Melajo, Trinidad 45 SW 1 1 88 110 134 168 9. Coubaril, Trinidad 45 55 1 1 37 40 83 96 10. Matura, Trinidad 47 58 3 3 82 92 132 160 11. Mare, Venezuela 65 82 99 144 164 226 12. Malquetia, Venezuela 42 53 65 82 107 135 13. Cantaure, Venezuela 37 49 2 Z 67 95 106 146 14. “Miocene,” Colombia 19 29 1 1 25 44 45 74 15. “‘Miocene,’’ Colombia 20 49 24 43 44 92 16. Lower Gatun, Panama 66 73 4 4 119 170 189 247 17. Middle Gatun, Panama 107 134 7 8 145 217 259 359 18. Upper Gatun, Panama 3{3) 64 6 8 92 98 151 170 19. Limon Basin, Costa Rica 79 120 65 147 144 267 20. Armuelles, Panama and Costa Rica 42 51 1 1 49 62 92 114 21. Charco Azul, Panama and Costa Rica 36 50 1 1 55 79 92 130 22. Angostura, Ecuador 29 32 48 66 77 98 23. ‘““Esmeraldas,’’? Ecuador 33 35 2 3 84 120 119 158 References: 1, Woodring (1925, 1928); 2-3, Jung (1971); 4-6, Maury (1925); 7, Rutsch (1942); 8-10, Jung (1969); 11-12, Weisbord (1962, 1964); 13, Jung (1965); 14, Weisbord (1929); 15, Barrios (1960); 16-18, Woodring (1957-1982); 19, Olsson (1922); 20-21, Olsson (1942); 22-23, Olsson (1964). MOLLUSKS: JACKSON ET AL. 195 stratigraphic and facies reconnaissance based on com- mon taxa. Due to this more limited sampling, the me- dian numbers of taxa for the 23 studies listed in Table 1 are only 107 subgenera and 135 species. Failure to consider the limited and inconsistent sam- pling among these studies, coupled with imprecise and sometimes faulty stratigraphy, has led to highly erro- neous interpretations of patterns and trends of mollus- can diversity in space and time throughout the region. Numerous authors concluded, for example, that num- bers of molluscan taxa declined dramatically in the tropical western Atlantic during the Pliocene (Woodr- ing, 1966; Vermeij, 1978; Stanley and Campbell, 1981; Jones and Hasson, 1985; Stanley, 1986; Vermeij and Petuch 1986). Furthermore, they attributed these supposed changes to oceanographic consequences of the rise of the Isthmus of Panama or intensification of glaciation in the Northern Hemisphere. However, their data were strongly biased by much greater sampling of Miocene and Early Pliocene compared with Late Pleistocene to recent faunas. More recent and exten- sive sampling of younger faunas demonstrates that di- versity did not decrease (Allmon et al., 1993, 1996; Roy et al., 1995, 1998) and may have increased (Jack- son et al., 1993). Rates of extinction and origination intensified greatly towards the end of the Pliocene, but these processes were roughly balanced so that total numbers of taxa effectively stayed the same. This paper describes the analysis of 245 collections of fossil mollusks from the Limon Basin of Costa Rica and the Bocas del Toro and Panama Canal basins of Panama that range in age from approximately 11.6 to 1.4 million years. The PPP occurrence data are avail- able at the internet site http://www.fiu.edu/~collinsl/. Our goals are to: 1. describe in detail how the collections were made and assess possible biases in sampling, processing, identification and analysis; 2. compare collections broken down by basins of de- position, age, and environment to determine how well we sampled the total diversity at any age, place or water depth, and the adequacy of these data to assess trends in diversity over time; 3. identify common taxa in the collections and their patterns of association in space and time based on ordination analyses; and 4. use correlations of ordination scores with age and water depth to estimate the relative importance of age and environment to variations in faunal com- position over 10 million years. Throughout the paper, we emphasize problems of sam- pling and taxonomy to demonstrate what we believe is required to rigorously establish faunal patterns and trends. The amount of work required is enormous and much still remains to be done. Nevertheless, it is al- ready possible to recognize consistently common taxa and draw conclusions about the stability of diversity over time. Future papers will build on these results to examine changes in rates and selectivity of extinction and origination, taxonomic composition, body size, shell thickness and ornamentation, incidence of pre- dation, and the proportions of different functional groups defined on the basis of life habits and diet. ACKNOWLEDGEMENTS Tony Coates measured the sections and developed the entire stratigraphic framework for the PPP upon which this study is based. Age determinations were provided by Marie-Pierre Aubry, Bill Berggren, Laurel Bybell, Harry Dowsett and Don McNeill, and coor- dinated by Laurie Collins, who also provided the pa- leodepth information. Tony Coates, Laurie Collins, Tim Collins, Antoine Heitz, René Panchaud, Jorge Ob- ando, David West, Yira Ventocilla and a great many others helped with the fieldwork. Magnolia Calderon, Antoine Heitz, Karl Miiller, and Yira Ventocilla pro- cessed the samples. René Panchaud and Antoine Heitz curated and managed the collections at the Naturhis- torisches Museum (NMB) in Basel, Switzerland. Felix Wiedenmayer created the database for all of the NMB Venezuelan and Trinidadian collections. Winifred and Jack Gibson-Smith assisted greatly in the original identifications. Xenia Guerra prepared figures 1—5. Discussions with Ann Budd and Alan Cheetham helped to organize our thoughts about so many taxa, and reviews by Lee-Ann Hayek, Carl Koch and Geerat Vermeij greatly improved the manuscript. This work was supported by grants from the Kuglerfonds of the NMB, National Geographic Society, Scholarly Studies and Walcott Funds of the Smithsonian Institution, Schweizerischer Nationalfonds Forschung (Grant Numbers 21-36589.92 and 20-43229.95), U. S. Na- tional Science Foundation (Grant Numbers BSR90- 06523, DEB-9300905, DEB-9696123, and DEB- 9705289), the NMB, and the Smithsonian Tropical Re- search Institute. STRATIGRAPHY, COLLECTIONS AND TAXONOMY Any study of the distribution and abundance of fos- sil taxa depends upon the quality and consistency of four basic factors: (1) stratigraphic control in space and time, (2) independent paleoenvironmental analy- sis, (3) methods of collection and processing of sam- ples, and (4) identification of taxa. 196 BULLETIN 357 b “Nicaragua “i a N afi Caribbean Sea Text-figure 2.—Map of Panama and Costa Rica showing the locations of the four areas sampled for this paper. LIMON (37) (38) (36) (35) Mar Mbr. |= has Empalme = Pueblo Nuevo Cemetery East Buenos Aires Mbr. BRIBRI (27) Text-figure 3—Summary of the stratigraphy of the Limon Basin for all sections sampled for this paper. Numbers in parentheses above stratigraphic columns are section numbers (Coates, this volume, Chapter 1). The stratigraphic position of each molluscan faunule is shown by numbers in boldfaced italics. STRATIGRAPHY The 245 collections analyzed in this paper come from three small, adjacent basins along the Caribbean coast of southeastern Costa Rica to central Panama (Text-fig. 2; Coates et al., 1992; Coates and Obando, 1996; Coates, Chapter 1, Appendix A, this volume). Most ages used for this paper are medians of age rang- es that were defined using planktic foraminifera, cal- careous nannofossils and (for the Limon Basin only) paleomagnetics (App. 1; Bybell, Chapter 2, this vol- ume; Cotton, Chapter 3, this volume; Aubry and Berg- gren, Appendix | in Chapter 1, this volume; McNeill et al., in press). Ages of two sets of undated collections from Isla Popa and Rio Tuba were assumed to be equivalent to nearby dated horizons at Cayo Agua and Rio Sand Box respectively based on stratigraphic po- sition. In addition, very approximate ages (“Late Pli- ocene”’ and ‘‘Late Miocene”) were arbitrarily assigned to three sets of undated collections from Ground Creek, Rio Calzones and Miguel de la Borda based on inferred field relationships. Medians of all these in- ferred ages are given in brackets to emphasize their uncertainty. The 103 collections from the Limon Basin range in age from late Late Miocene (7.7 Ma) at Rio Sand Box to near the Plio—Pleistocene boundary (1.6 Ma) at Lo- mas del Mar (Text-fig. 3; Appendix B, this volume). However, the great majority of the collections come from the late Early to early Late Pliocene Rio Banano Formation, the earliest Late Pliocene Quebrada Choc- olate Formation and the basal Pleistocene Moin For- mation. The 96 collections from the Bocas del Toro Basin also range from late Late Miocene (6.9 Ma) at MOLLUSKS: Fish Hole Reef Mor. CAYO AGUA ISLAND Cayo Agua Fm | | | | | t | | | | | t | | | | | r | | | | | a Pa e Soe eSsaaSs> (19) (20) (17) Piedra Roja Tiburon Point Point West JACKSON ET AL. 197 ESCUDO DE VERAGUAS ISLAND VALIENTE PENINSULA Shark Hole Point Fm Nancy Point Fm Nancy PointFm Tobabe Sst. Tobabe Ss} Text-figure 4—Summary of the stratigraphy of the Bocas del Toro Basin for all sections sampled for this paper. Section numbers and faunule numbers shown as in Text-figure 3. Finger Island on the Valiente Peninsula to Early Pleis- tocene (1.4 Ma) at Swan Cay; but the collections are more evenly distributed in age than in the Limon Basin (Text-fig. 4; Appendix B, this volume). The 46 collec- tions from the Canal Basin are all Late Miocene (Text- fig. 5), ranging from the lower Gatun Formation at the Martin Luther King housing development (11.6 Ma) to the middle Gatun Formation at Isla Payardi (9.0 Ma). There are no collections so far from the mollusk- rich Rio Indio facies (sensu Collins et al., 1996) of the Late Miocene (mostly 6.4—5.8 Ma) Chagres Forma- tion. Finally, the three collections from the north-cen- tral coast of Panama are as yet undated but are very probably Late Miocene. The stratigraphy of all three basins and the strengths and weaknesses of the age dating are discussed in de- tail elsewhere in this volume. Here we only consider possible problems of particular relevance to the mol- lusks. In the Limon Basin, the entire Rio Banano For- mation, including the section from Quitaria through La Bomba, is now considered to be 3.6—2.9 Ma (McNeill et al., in press). Previously, La Bomba had been dated at 2.4—2.5 Ma (Coates et al., 1992), but it is now con- sidered to be 3.1 Ma. In addition, the sequence of coral reef tracts, flank deposits and inter-reef basins extend- ing westward from Limon is now known to range from 1.7-1.5 Ma at Lomas del Mar to 3.6—3.3 Ma at Que- brada Chocolate, and there is also a patch reef within the Rio Banano sequence at Brazo Seco tentatively dated at 5.2—-4.3 Ma (Budd et al., 1996, this volume; Budd and Johnson, 1997; McNeill et al., in press. Im- mediately west of Limon, the new Quebrada Chocolate Formation replaces the uppermost part of the Rio Ban- ano Formation sensu Coates et al. (1992). The Que- 198 BULLETIN 357 NORTH COAST IN SENISES (Undated Upper Miocene) (6) Calzones River Change of Scale Cretaceous | 77.7 Volcanics oN Sabanita- Text-figure 5—Summary of the stratigraphy of the Canal Basin and central north coast of Panama for all sections sampled for this paper. Section and faunules numbers shown as in Text-figure 3. brada Chocolate Formation overlaps in age with the Rio Banano Formation at the latter’s type locality at Bomba. Finally, the age and stratigraphic position are still uncertain for the mollusk-rich deposits at Pueblo Nuevo, the Cementerio General and the Progressive Baptist Church, although they are probably the same age as Lomas del Mar. Ages of formations in the Bocas del Toro Basin have remained more stable, and the principal devel- opments are the inclusion of numerous new, mollusk- rich horizons that fill important gaps in the sequence. The most important included here are the Late Plio- cene to Early Pleistocene horizons at Fish Hole, Ground Creek and Swan Cay in the northwest and Late Miocene Finger Island in the southeast. Finally, the age of the important Gatun Formation in the Pan- ama Canal Basin now extends back to the uppermost Middle Miocene (11.8—11.4 Ma) (Collins et al., 1996) instead of 8.2 Ma (Coates et al., 1992). PALEOENVIRONMENTAL ANALYSIS It is essential to identify environments independent- ly of the mollusks to avoid circular reasoning. The most complete paleoenvironmental data available so far are for water depth, based primarily on benthic foraminifera, but also ostracodes, otoliths and aher- matypic corals (Appendix 1; Collins et al., Chapter 4). These data are available for 29 of the 37 depths used herein. The remainder were assigned conservatively based on regional stratigraphic and facies relationships pending analysis of benthic foraminifera. Detailed sed- imentary facies analyses have not yet been attempted. COLLECTIONS Our basic sampling unit is a collection, which we define as the sum total of all the samples of fossils collected at one time from some stratigraphically well defined horizon at a single location. Subsequent col- lections from exactly the same site are given a new collection number. Most of our collections from the Limon Basin come from river banks and new construc- tion sites; from the Bocas del Toro Basin they are pri- marily from sea cliffs; and from the Panama Canal Basin they are from new construction sites. In the lat- ter case, the great majority of Woodring’s (1957-1982) original localities are gone. The process of assembling a collection is a long and commonly iterative process. The problem is that no one method is suitable for collecting all the mollusks at a site, primarily because of differences in size and preservation. We therefore collect two kinds of sam- ples, which we call “specimen” and “bulk.” Speci- men samples comprise all of the visible shells at the outcrop that can be collected individually in place from the outcrop or as float at the immediate base of the outcrop. Bulk samples are typically 10-kg sacks of sediment that are excavated for future processing at the laboratory. Specimen and bulk samples are given numbers in the field according to the individual inves- tigator’s system, but these subsequently are assigned PPP numbers, which are used hereout for convenience and accuracy of comparison. Specimen and bulk samples are subject to numerous sources of bias, which we try to avoid through use of standard methods. Factors, which affect both types of sample, are the condition of the material and the amount of time and number of collectors available. For example, to make one collection at a single small, but rich, site may require four experienced collectors an entire afternoon. Moreover, as we shall see, no single collection contains more than an indeterminate small MOLLUSKS: JACKSON ET AL. 199 fraction of the total fauna at a site as can be observed after repeated collecting. Two additional factors that affect the usefulness of the specimen collections are care in individually wrap- ping and packing the specimens at the outcrop or in camp and investigator bias. Specimens are wrapped in tissue, packed, sealed in cans or other rigid containers, and then repacked for eventual shipment to Basel. In addition, comparison of our own and earlier collec- tions from the same sites repeatedly demonstrates that many earlier collections were biased in favor of large, pretty snails at the expense of bivalves, smaller snails and unattractive or fragile fossils in general. The most important additional factor affecting the quality of the bulk samples is the depth of excavation before taking a sample. Most of our collections come from volcaniclastic silty sands to sandy silts that are gray to dark brown at the weathered, outcrop surface but a highly distinctive slate blue-gray when fresh. It is commonly necessary to excavate at least 0.5 m be- neath the outcrop surface to encounter fresh material, except after fortuitous floods or storms excavate riv- erbanks and coastal cliffs en masse. Digging “‘to the blue’’ may require an hour or more per sample, but the resulting numbers of taxa collected in a single bulk sample may increase more than two-fold, accordingly. Finally, the number of bulk samples made at any site depends on available time and energy as well as the apparent richness of the material. One bag of bulk sed- iment is routinely collected at most sites, but 3 to 10 bags are collected where the material is rich in fossils. Processing of specimen samples is mostly a matter of routine museum curation. All collections are as- signed a Naturhistorisches Museum of Basel (NMB) number but are stored separately in cabinets arranged by sedimentary basin and PPP number. In contrast, processing of bulk samples is more complex and time consuming. The same bulk samples are processed for corals, bryozoans, otoliths, fish and shark teeth, echi- noderms and brachiopods, as well as for mollusks. Therefore, three sizes of sieve openings are used: 2000, 500 and 125 wm. Our goal is to process and pick mollusks from the 2000-j.m (2-mm) fraction from at least one bulk sample from every collection. How- ever, this task was not completed for two thirds of the collections for inclusion in this paper. The 500-1m fraction contains many micromollusks and the larval shells of larger species. These micromollusks are com- monly extremely abundant and diverse, but thousands of additional hours would be required just to pick them, and they are a separate study in themselves. Washing and diasaggregation (processing) of the bulk samples is done as gently as possible. Sometimes it is unnecessary to do more than soak the sample in water before sieving, but other times it is necessary to use detergent, hydrogen peroxide or to heat and freeze the sample. The latter treatments inevitably damage some specimens, but the condition and diversity of the material so far obtained suggests little more damage occurs than is sustained through wet sieving. All pick- ing is done at 10 magnification using a Wild M-5 stereomicroscope. Fossils from the bulk samples are curated and stored in the same drawers as the corre- sponding specimen samples. TAXONOMY It cannot be overemphasized that the superspecific taxonomy and systematics are poorly resolved for the majority of Neogene and Quaternary mollusks of trop- ical America. Diagnostic characters of many of the commonest genera or subgenera are not consistently stated or applied, and only a handful of taxa have been analyzed cladistically to help clarify relationships. Sim- ilar problems apply to species, which are commonly assigned names uncritically based on comparison with monographs of other faunas, without examining the types, a practice that produces misinformation rather than precision (Robinson, 1993; Waller, 1993). We conservatively estimate that more than half of the species in our collections are undescribed, and that it would require a decade or more to describe them prop- erly group by group. We base this estimate on results of recent and ongoing studies of the few common groups so far examined in detail throughout the region. These include the Strombina Group (Jung, 1989; Jack- son et al., 1996; Fortunato and Jackson, unpublished data); Muricidae (Vokes, 1989; D. Miller, unpublished data), Turridae (J. Todd, unpublished data), and Tuce- tona (P. Tschudin, unpublished data). We therefore adopted the following pragmatic policy for the prelim- inary identification of taxa for faunal lists: 1. All identifications are at the generic or subgeneric level. 2. Great effort is devoted to establishing lists of char- acters for the consistent recognition of genera and subgenera, with the greatest emphasis paid to the roughly 200 genera or subgenera that make up more than 90% of the specimens (Appendix 2). Many of these taxa may prove to be polyphyletic in subsequent systematic study, but they will have been identified consistently so that the data will be useable should future study of strict monophyla be desirable. 3. A reference collection is being established for com- mon taxa that includes diagnostic characters and specimens from each of the sedimentary basins and ages where the genus or subgenus occurs (broken 200. BULLETIN 357 down as Late Miocene, Pliocene, and Pleistocene from each of the three basins). Digital images and diagnostic characters of the common taxa are being placed on the World Wide Web site of the Neogene Marine Biota of Tropical America (NMITA) taxo- nomic database system (http://nmita.geology. uiowa.edu). 4. No attempt was made to revise (even informally) numerically important problematic taxa such as Turritella, Anadara (Rasia), and most of the turrids due to lack of time, even though they almost cer- tainly include numerous, unrecognized subgenera or even genera. This lack of resolution inevitably reduces our initial estimates of diversity. On the other hand, about one third of the taxa (almost all of them rare) are questionably identified due to problems of preservation or inadequate published descriptions, which inflates our estimates because questionable identifications are listed separately in the database. Following these guidelines, specimens in each col- lection were identified to genus or subgenus and count- ed. Because of the difference in effort and time in- volved, identifications and tabulations of taxa were usually completed for the specimen samples from a given locality long before the bulk samples were pro- cessed from the same locality. Mollusks from the bulk samples of only one third of the 245 collections have been included with material from the specimen sam- ples in the following analyses. DISTRIBUTION, AGE, AND ADEQUACY OF COLLECTIONS The numbers of collections and specimens upon which this paper is based are broken down by sedi- mentary basins and age in Text-figure 6. There are two clear biases in these data. First, the sampling effort is unevenly distributed through time because most of the collections were made in the early stages of the PPP before we were confident of the stratigraphy, and be- cause some ages are better represented than others in the areas studied. Second, space and time are con- founded because all the collections younger than 6 Ma come from the two western basins whereas all those older than 8 Ma come from the Canal Basin. There are similar sampling problems with depths and environments of deposition determined using ben- thic foraminifera (Appendix 1). Late Miocene mol- lusks sampled so far from the Canal Basin (11.6—-8.6 Ma) were all deposited in only 15-40 m water depth, whereas those from the western basins (7.7—5.7 Ma) range from 60—200 m. Pliocene and Pleistocene de- 0 Limon Bocas del Toro mw Panama Canal and North Coast i rep) oo S Oo Oo Oo S Number of Collections 30 pe) oO oO Thousands of Specimens = abil F 12: AlN Gie Ba, Te Ou- Ora 2aI20 Age (Ma) Text-figure 6—Numbers of collections and thousands of speci- mens per million years for each sedimentary basin. Each collection was assigned to one age interval using the median of the estimate of the age. MOLLUSKS: JACKSON ET AL. 201 all basins combined 800 600 400 pe) Oo oO 0 TIONS SS 7 (605.4 C2710 0 Limon 800 7 & Bocas del Toro m Panama Canal and North Coast 600 400 Number of Genera or Subgenera 200 > K> <] iS Ro “as Cex 0 Text-figure 7.—Numbers of genera or subgenera per million years for all sedimentary basins combined and for each separate basin. The totals are larger for the separate basins because many taxa occur in more than one basin. sa All Mollusks All Mollusks Early Pleistocene Early Pleistocene 6007 | > 5 De) oO oO 4 ise} Q o g 8 E Late Miocene Cumulative Number of Genera or Subgenera : T =T T 0 30 60 90 0 15 30 45 60 75 Number of Collections Thousands of Specimens Text-figure 8.—Cumulative numbers of genera and subgenera col- lected as a function of both the numbers of collections made and numbers of specimens accumulated for three time intervals (Late Miocene, Pliocene, and Early Pleistocene). L = Limon basin, B = Bocas del Toro Basin, C = Canal Basin and Panama north coast, T = total for all basins combined. posits from both of the western basins include a better mix of environments ranging from about 10—200 m. Numbers of genera and subgenera collected closely parallel the sampling effort (Text-fig. 7), which strong- ly suggests that a large fraction of taxa from each age interval is uncollected (Koch, 1987). This is confirmed by the steep increase in cumulative numbers of taxa as a function of the numbers of collections or speci- mens, whether broken down by age and basin (Text- fig. 8), or for all 245 collections combined (Text-fig. 9). The reasons for these daunting results are that most collections contain comparatively few specimens or taxa (Text-fig. 10) and, as expected in the tropics (Sanders, 1969), most taxa are extremely rare (Text- fig. 11). Half of the 1021 genera or subgenera are rep- 202 BULLETIN 357 — genera [a == +) aS ae —aplls | 0 On OO. 150 200 | 250 Cumulative Number of Collections — S S) Oo +4 (o0) S S Cumulative Number of Genera or Sub = @ B 0 25 60 75 100 125 150 Cumulative Number of Specimens (thousands) Text-figure 9.—Cumulative numbers of genera or subgenera col- lected as a function of both the numbers of collections made and numbers of specimens accumulated for all 245 collections combined. B = bivalves, G = gastropods, T = total mollusks (includes bi- valves, gastropods and scaphopods). resented by fewer than five specimens from fewer than three collections. Nevertheless, subgeneric to generic diversity per basin per million years in our PPP col- lections (Text-fig. 7) generally exceeds that obtained in most of the studies listed in Table 1. COMPOSITION OF THE FAUNA There are a total of 149 families and 1021 genera or subgenera of mollusks in the 245 collections that Total Mollusks mean=48.4 mean-623.2 4 median=42 median=285 std dev=33.5 std dev=605.3 range=1-151 range=1-3744 mean=3 17.5 median=149 std dey=418.3 range=1-3213 Number of Collections Bivalves mean= 15.1 mean=205.2 median= 14 median=101 std dev=102 std dev=128.0 range=1-56 range=1-2142 + T 0 40 80 120 160 O 1 2 8 4 Number of Genera Thousands of or Subgenera Specimens Text-figure 10.—Frequency distributions of numbers of taxa and specimens per collection for total mollusks (including scaphopods), and for gastropods and bivalves each taken alone. break down as summarized in Table 2. The proportions of genera or subgenera for gastropods, bivalves and scaphopods (0.66:0.32:0.02) are broadly similar to those reported in the faunas listed in Table 1. This suggests that the PPP collections are not biased some- how in the representation of these three major taxa. Only 156 out of the total 1021 genera and subgenera are represented by 100 or more specimens in the 245 collections (listed in descending order in Appendix 2). These include 96 gastropods, 52 bivalves and 8 sca- phopods. The abundance of some of these, such as Turritella and Anadara (Rasia), is clearly artificially high due to taxonomic lumping. In contrast, abun- dances of most of the smaller taxa (including the top three bivalves Crassinella, Caryocorbula and Varicor- bula) are too low because specimens from bulk sam- Total Mollusks mean=11.6 median=3 std dev=23.0 4 range=1-175 4 | mean= 1256 median=6 MOLLUSKS: JACKSON ET AL. Table 2.—Composition of the molluscan fauna from 245 collec- tions of fossils. Numbers of Numbers of genera std dev=558.1 Taxa families and subgenera range=1-10296 Gastropoda 89 675 Bivalvia 54 326 Scaphopoda 7 19 Cephalopoda 1 1 Totals 149 1021 ples have not been processed for two thirds of the mean=1130 collections. eee As expected for such data (Williams, 1964; Hanski range=1~10296 et al., 1993; Hayek and Buzas, 1997), there is a strong Tees soealcoeg nla nmi lunen nae eae Get wo 4 = a | = ® 2h > Gastropods Bel C 1000 mean=11.8 | median=3 © 8004 std dev=236 4| range=1-175 © 600 ae & 400 ® 4 pe ed oO a 7] eer Ima Da oe —t T ; gx Bivalves 5 1000 + mean= 10.7 a 4 median=2 800 std dev=21.1 range= 1-160 0 380 60 90120 160 180 0 2 4 Number of Collections positive correlation between the abundance of these 156 commonest taxa, as measured by numbers of spec- imens (range 100 to 10,296), and their frequency of occurrence in the different collections (range 3 to 175) (Spearman rank-order correlation, r = +0.669, P < 0.000; Text-fig. 12). Nevertheless, there is considerable variation in frequency. Many taxa of closely similar wee abundance exhibit a 5- to 10-fold range in the numbers std dev-6 109 of collections in which they occur. These differences range=1-5727 6 8 10 Thousands of Specimens Text-figure 11.—Frequency distributions of numbers of collec- tions and numbers of specimens per genus or subgenus for total mollusks, gastropods and bivalves. reflect the relative eurytopy or stenotopy of taxa (Jack- son, 1974). For example, 407 Volvulella (Volvulella) occur in 95 collections, whereas the scaphopod Gad- ilopsis occurs as 435 specimens in only 11 collections. Likewise, 1104 Polystira are distributed among 134 collections, whereas the next most abundant taxon Sin- cola (Sincola) is represented by 1085 specimens from only 34 collections. In both cases, the more narrowly distributed taxa are limited to shelf envionments whereas more widespread taxa occur in a greater di- versity of environments. The156 most common gastropods, bivalves and sca- 10000 J (cS (ad) = © (ad) ion Y 4000-4 © ab) 1@) = Za 100 4 10 100 Number of Collections 1000 Text-figure 12.—Logarithmic plot of numbers of specimens versus numbers of collections for all genera or subgenera represented by 100 or more specimens in all collections combined as listed in Appendix 2. Filled circles = gastropods, triangles = bivalves, and squares = scaphopods. 204 BULLETIN 357 Table 3.—List of 37 faunules (molluscan taxa from a single horizon at a single outcrop or closely grouped outcrops) and descriptive statistics used for the ordination analyses. Taxa are genera or subgenera. Lists of PPP numbers for each faunule are given in Appendix 3. Documentation for ages and depths are given in Appendix 1. Estimated ages and depths placed in brackets. L = Limon Basin, B = Bocas del Toro Basin, NC = North Coast of Panama, C = Panama Canal Basin. Number Number Faunule Section Median Median _ of col- of Number Fisher’s number Faunule name Basin number age depth lections specimens of taxa alpha ] Swan Cay B 25 1.4 100 1 1,418 135 36.691 2 Cemetery Pueblo Nuevo L 35 1.6 75 1 452 67 21.744 3 Upper Lomas del Mar east (reef) IB, 36 1.6 75 12 5,986 219 44.637 4 Empalme IG 34 1.6 20 5 2,188 143 34.508 5 Cangrejos Creek L 37 1.6 200 5 828 116 36.734 6 Lower Lomas del Mar east (non- L 36 ea WS 10 6,458 304 66.229 reef) 7 Northwest Escudo de Veraguas B 10 2 125 4 433 49 14.206 8 Fish Hole B 22/23 2.6 70 3 331 114 61.516 9 Ground Creek B [2.6] [S50] 2 1,723 90 20.283 10 North central Escudo de Veraguas B 10 2 125 8 5,019 227 48.916 11 Rio Limoncito L 3.0 [30] 1 148 39 17.269 12 Chocolate Buenos Aires r 33 Sul [SO] 3) 1,011 45 9.657 13 Bomba Ie, 29 3.1 30 34 18,181 285 47.980 14 Agua 1, 29 3.3 30 2 841 53 12.565 15 Bruno Bluff B 12 3:5 175 4 1,310 133 35.822 16 Cayo Agua: west side Punta Norte B 16 355) 30 8 2S) 139 31.238 17 Quitaria IL, 29 35 30 7 12,690 179 29.508 18 Rio Vizcaya L 39 3.5 25 yf 979 47 10.296 19 Santa Rita 1, 32 355 30 2 497 81 27.462 20 Northeast Escudo de Veraguas B 10 3.6 125 4 2,588 175 42.847 21 Southeast Escudo de Veraguas B 11 3.6 125 9 2,215 166 41.888 22 Cayo Agua: Punta Tiberon B 19 3.6 60 9 4,001 270 65.368 23 Cayo Agua: Punta Nispero west B 19) 3.6 60 6 F339 122 32.648 24 Cayo Agua: southeast Punta B 20 3.6 60 qT 3,307 75) 39.562 Nispero 25 Isla Popa B [4.3] [60] i 2,445 101 22.431 26 Cayo Agua: Punta Norte east B 19 4.3 60 6 2,185 124 28.663 Dil Cayo Agua: Punta Piedra Roja west B 17 4.3 43 6 6,640 275 57.881 28 Quebrada Brazo Seco 1 4.8 {SO} 3} 240 S7 23.632 29 Shark Hole Point B 12 SE7/ 150 7 432 57 17.586 30 Finger Island B 14 6.9 80 3 1,817 165 44.354 31 Rio Sand Box and Hone Creek IL, Dil Tell 175 6 697 65 17.534 32 Rio Tuba IL, [7.7] [175] 5 91 40 27.279 33 Rio Calzones NC 9 [8.3] [25] 2 185 43 18.598 34 Miguel de la Borda NC 6 [8.3] 25 1 699 97 30.580 35 Isla Payardi Cc 1 9.0 28 14 14,627 172 27.376 36 Mattress Factory (e 1 9.0 28 16 11,957 236 41.677 37 Martin Luther King Jr. Ee 1 11.6 28 11 9,242 155 26.455 phopods also differ in the median numbers of collec- tions in which they occur. The median for the 96 gas- tropod taxa is 46.5 collections, the median for the 52 bivalve taxa is 36 collections, and the median for the eight scaphopod taxa is 28 collections (Kruskal-Wallis test, chi-square = 4.89, df. = 2, P = 0.087). These marginally significant results suggest that common gastropods were more eurytopic than bivalves, which were, in turn, more eurytopic than scaphopods. FAUNAL ASSEMBLAGES The median numbers of specimens and taxa per col- lection are small (285 and 42, respectively), and there is enormous variation in the richness of individual col- lections (Text-fig. 10). We therefore assembled the 245 collections into 37 groups called ‘‘faunules” to try to decrease the effects of sample size for the analysis of patterns and trends in diversity and composition of faunal assemblages (Table 3; Text-figs. 3-5). The groupings were made based on age and location. Faun- ules correspond to a single fossiliferous stratigraphic horizon at a single outcrop (e.g., Swan Cay or Finger Island; Text-fig. 4). Due to pervasive bioturbation at the great majority of sites, bedding could not usually be observed. Therefore, packages of lithologically identical sediment, typically amounting to a few me- MOLLUSKS: JACKSON ET AL. 205 1000 i Number of Genera or Subgenera 3 Ss 2 0 aa 100 oo $7 13 13 1&6 ry ee ae ay Fe ‘40 12 it Tl 1000 10000 Number of Specimens Text-figure 13—Numbers of genera or subgenera versus numbers of specimens for the 37 faunules listed by the same numbers in Table 3. ters O1 section, were treated as a single horizon. Near- by but physically separate outcrops of the same strati- graphic horizon were treated as separate faunules for purposes of replication in the analyses. Examples of replicate faunules include exposures on different head- lands of the 3.5—3.6 Ma horizon of the Cayo Agua Formation (Faunules 16, 22, 23 and 24) or the 9.4— 8.6 Ma horizon of the Gatun Formation (Faunules 35 and 36) (Text-figs. 4, 5). The mollusks contained in the 37 faunules range from a minimum of 91 to a maximum of 18,181 spec- imens, and from 39 to 304 genera or subgenera (me- dians: 1723 specimens and 124 subgenera per faunule; Table 3). These numbers are large enough to include all of the faunules in analyses of faunal patterns in space and time. Nevertheless, most of this variation in numbers of specimens and taxa is still due to differ- ences in sampling effort, as demonstrated by the high- ly significant positive correlation between numbers of taxa and specimens among the 37 faunules (Spearman rank order correlation, r = +0.889, P < 0.000; Text- fig. 13). DIVERSITY OF FAUNULES We used Fisher’s alpha as the best single index of diversity because of great differences in sample size among both the faunules (Text-fig. 13) and the one- million-year intervals (Text-figs. 6—7) (Magurran, 1988; Rosenzweig, 1995; Hayek and Buzas, 1997). Use of alpha is based on the assumption that the abun- dances of species fit a log-series distribution (Fisher et al., 1943), in which case alpha is independent of the number of specimens. Alpha also can be used as a measure of diversity when the log series is not a good statistical fit to the data if the ratio of numbers of spec- imens to taxa (N/S) > 1.44 (Hayek and Buzas, 1997). This was always true for our data. Estimates of alpha for faunules and one-million-year intervals with fewer than 5000 specimens were obtained from Appendix 4 in Hayek and Buzas (1997). Estimates for those with more than 5000 specimens were kindly provided by Lee-Ann Hayek (written commun., 1997, 1998). Values of alpha for the 37 faunules range more than six-fold, from a low of 9.7 at Chocolate Buenos Aires to a high of 67 at lower Lomas del Mar (median for the 37 faunules = 30.6; Table 3). Alpha is positively correlated with the number of specimens in the faun- ules (Spearman rank-order correlation, r = +0.549, P = 0.000), but not so strongly as the number of genera or subgenera with the number of specimens (Text-fig. 13). Alpha also increases nonsignificantly with in- creasing depth (r = +0.216, P = 0.100 for 1-tailed test. The |-tailed test is appropriate because molluscan diversity increases with depth in Recent seas from nearshore to bathyal environments (Rex, 1981). The faunules are listed in decreasing order of Fish- er’s alpha in Table 4, along with information gleaned from the stratigraphic sections (Appendix B, this vol- ume) about the presence of corals. Diversity of mol- luscan faunules from horizons where corals were com- mon is 50% higher than for faunules without corals (median alphas 41.9 versus 27.4, Mann-Whitney Test, P < 0.01). The type of coral does not appear to be particularly important, since diversity is high regard- less of whether corals are cemented or free-living, sol- itary or colonial, or with or without symbiotic zoo- xanthellae. Abundance of free-living, zooxanthellate corals suggests the presence of extensive seagrass beds 206 BULLETIN 357 Table 4.—Faunules listed in descending order of Fisher’s diversity index alpha. Presence of corals as common to abundant based on descriptions of stratigraphic sections in Appendix B (this volume). Asterisk indicates azooxanthellate corals. Fisher’s Faunule Occurrence Alpha number of corals 66.2 6 reef-building 65.4 22 free-living 61.5 8 reef-building Sie) 2 free-living 48.9 10 dense thicket* 48.0 13 44.6 3 reef-building 44.4 30 42.9 20 horn* 41.9 21 horn* 41.7 36 39.6 24 free-living 36.7 25 reef-building 36.7 5 35.8 15S 34.5 4 reef-building 32H 23 31.2 16 free-living 30.6 34 293 17 28.7 26 Dales 19 27.4 B95 DAES 32 26.5 37 23.6 28 reef-building 22.4 25 PIEA 2 20.3 9 18.6 33 17.6 29 7A) 31 17.3 11 14.2 7 horn* 12.6 14 10.3 18 OFF 12 reef-building (Johnson et al., 1995), which is consistent with the high diversity of molluscan faunas in Recent, Carib- bean seagrass environments (Jackson, 1972, 1973). In contrast, abundant deep burrows were absent from most horizons where corals were common (Appendix B, this volume). The implications of this very prelim- inary analysis are that comparatively stable sediments, as inferred from the abundance of corals and absence of deep burrows, supported higher molluscan diversity than unstable sediments, just as in the Recent. In spite of all this environmental heterogeneity, there is no significant change in alpha of the 37 faun- ules over the 10.2 million years for which we have collections (Text-fig. 14, top; Spearman rank-order correlation, r = —0.138, P = 0.415). This apparent stability strongly supports an earlier conclusion that southern Caribbean molluscan diversity did not decline during the past 12 million years (Jackson ef al., 1993; Jackson, 1994) despite faunal turnover at the end of the Pliocene. Moreover, alpha increases significantly towards the Recent using the data for one-million-year intervals (Text-fig. 14, bottom; Spearman rank order correlation, r = —0.751, P = 0.012). This result is obviously biased by the much greater number of faun- ules in the younger intervals, and thus the much great- er variety of environments sampled. Regardless, di- versity definitely did not decrease as had been claimed previously (Woodring, 1966; Stanley, 1986; Vermeij and Petuch, 1986). ORDINATION OF FAUNULES We used detrended correspondence analysis (DCA) as a measure of similarities in molluscan faunal com- position of the 37 faunules based on the occurrence of common taxa. To examine possible trends in faunal composition, we calculated Spearman rank-order cor- relations between scores on the DCA axes and the ages and paleodepths of each faunule as presented in Table 2. DCA is a non-parametric procedure analogous to principal components analysis in relating differences in generic or subgeneric occurrences to independent axes of decreasing variation by the eigenanalysis of a matrix of chi-square distances between collections (McCune and Medford, 1995). To calculate the ordi- nations, we used the DCA option of the PC-ORD pro- gram (ibid.). The ordinations were repeated using both binary (presence-absence) and ranked abundance data for the occurrence of the 25 most abundant molluscan genera or subgenera in each of the 37 faunules (Appendix 3). When there were ties in the abundance of the 25th most abundant genus or subgenus in a faunule, we included all of the taxa that were tied. In these cases the numbers of genera or subgenera in a faunule is greater than 25. We used ranked abundance (0 speci- mens = 0, 1-9 specimens = 1, 10—99 specimens = 2, 100-999 specimens = 3, = 1000 specimens = 4) rath- er than raw abundance because the latter is more sen- sitive to sampling bias. The choice of the 25 commonest taxa from each faunule resulted in lists of 254 unique molluscan genera or subgenera. Eleven of the 156 genera or subgenera represented by >100 specimens (Text-fig. 12, App. 2) are not included among the 254 unique mollusk taxa in Appendix 3 because they were not among the top 25 taxa in any faunule. In contrast, 109 genera and sub- genera among the 256 unique mollusks are represented by fewer than 100 specimens in all of the collections combined. These are listed in order of numerical abun- MOLLUSKS: JACKSON ET AL. 207 80 Faunules 6 22 60 4 ‘ 27 1043 38 30 386 40 + 18 x 4 26 32 eat oe 37 5 95. 28 20 al oan 81' (33) 7 1248 av) 1 OS ay T T T T T T =a O- 100 : e ; ae <{ One-million-year 80 + intervals e e 60 + e e 40 4 ° ° e 20 5 O | T — T tT T T ial T OG 1 2 © ® 7 FF © TH BW te AQ e (Ma) Text-figure 14.—Variation in Fisher’s diversity index alpha with age for all mollusks in the 37 faunules listed in Table 3 and for the one- million year intervals in Text-figures 6—7. dance at the end of Appendix 2. Each of these 109 taxa occurs in very few faunules where, however, they are relatively abundant. Thus it should be possible to dis- criminate faunules dominated by otherwise rare taxa that would be excluded from any list based solely on total abundance for all the collections combined. Results of the DCA analyses are illustrated separately for binary and ranked abundance data in Text-figure 15. Simple inspection suggests that faunule composition strongly varies with age and depth along the ordination axes 2 and | respectively. Moreover, the separation of faunules by age along axis 2 is clearer for the analysis using ranked abundance data than for binary data. These impressions are confirmed by highly significant correlations of the ordination scores on axis 2 with age (r = —0.623, P < 0.001) and of the scores on axis 1 with depth (r = 0.740, P < 0.001) for analyses based on ranked abundances (Text-fig. 16). In contrast, cor- relations of scores on axis 2 with age were only mar- ginally significant (r = 0.380, P = 0.020) for analyses based on binary data, but were significant for scores on axis 1 with depth (r = 0.688, P < 0.001). These results appear to support the use of ranked abundances rather than simple binary data in paleoeco- logical analyses and studies of macroevolutionary trends (Jackson and McKinney, 1990). However, the number of faunules is not great. Thus, as the number of faunules increases, differences between the results of ordination analyses based on binary and ranked abundance data are likely to decrease (Hayek and Buzas, 1997). DISCUSSION The diversity of our collections exceeds that of all previous studies of Caribbean Neogene mollusks (compare Table 1 and Text-fig. 8). Numbers of genera or subgenera in our collections from one-million-year intervals of the Late Pliocene exceed those for any other Neogene Caribbean collection. In addition, num- bers of taxa for all but two of the remaining one-mil- lion-year intervals sampled equals or exceeds that of the faunas listed in Table 1 except for the Bowden and middle Gatun formations. Similar claims can be made for about half of the 37 faunules in Table 3. Never- theless, our collections are still subject to sampling biases in age, environment and geography (Text-figs. 6-11, 13). Some of these problems should be at least 208 BULLETIN 357 presence/absence 300 200 DCA Axis 1 3 800 200 100 0) 100 200 300 400 DEA Axis 2 Text-figure 15.—Plots of detrended correspondence analyses (DCA) for the 37 faunules listed by the same numbers as in Table 3. Numbers increase from youngest (1) to oldest (37). Faunules from estimated water depths > 75 m indicated in boldface. Data for the ordinations are occurrences (presence-absence or ranked abundance) of the 254 molluscan genera or subgenera listed in Appendix 3. partially resolved when we incorporate extensive new collections from the same three sedimentary basins, and from the transisthmian Chuqunaque Basin in Dar- ien, that are now being processed. The differences in numbers of taxa between PPP and earlier collections reflect greater collecting effort, and are not unique to the three basins sampled. For example, Jung’s (1965) description of the early Middle Miocene Cantaure Formation was based on oil com- pany reconnaissance collections that produced 109 genera and subgenera and 146 species. In contrast, re- 110 155 200 Depth (m) 20 65 O12 8 oS G 7 OO DW Age (Ma) Text-figure 16.—DCA axis scores versus age and depth of water for the 37 molluscan faunules as numbered in Table 3. Data for the ordination analyses were ranked abundances of the 254 genera or subgenera listed in Appendix 3. peated collecting over more than ten years by Jack and Winifred Gibson-Smith from the same localities at Cantaure produced 471 genera and subgenera and 737 species now at the Naturhistorisches Museum in Basel (unpubl. PPP-NMB taxonomic database compiled by Felix Wiedenmayer). Likewise, published faunas from the Mare Formation (Weisbord, 1962, 1964) include 164 genera and subgenera and 226 species, versus 352 genera and subgenera and 531 species in the Gibson- Smith collections (ibid.). Despite all these caveats about sampling, a number MOLLUSKS: JACKSON ET AL. 209 of apparently robust patterns have emerged. Slopes of the cumulative diversity curves for different ages and basins (Text-fig. 8) appear to be stable with the repeated addition of new collections. This suggests that the slopes of the cumulative curves can be used reliably to infer the comparative diversity of our faunas, much like Fisher’s diversity index alpha (Fisher ef al., 1943; Ma- gurran, 1988; Hayek and Buzas, 1997). In addition, the list of the most abundant taxa in our collections (Ap- penidx 2) is unlikely to change much with further col- lecting, except for reasons of taxonomic refinement, be- cause most taxa are extremely rare (Text-fig. 11). Most of the abundant genera and subgenera have very long stratigraphic ranges in our collections (Ap- pendix 2), which suggests that they were eurytopic. Restriction of a minority of abundant taxa to relatively few collections (Text-fig. 12) may be due to rapid evo- lutionary turnover or stenotopy (Jackson, 1974), or to environmental or stratigraphic biases in our collec- tions. In addition, difficulty of collection (e.g., large, fragile bivalves) and diagenesis (Koch and Sohl, 1983) may be important factors. For example, shells are typ- ically leached and the matrix sandy at all the collection sites in the Gatun and Rio Banano formations. Lastly, large, attractive, or rare taxa, such as many of the tur- rid gastropods, inevitably attract more attention in the field, and are therefore likely to be over-represented relative to their actual frequency of occurrence. Molluscan diversity generally increases while abun- dance generally decreases with increasing depth in Re- cent seas (Sanders, 1968, 1969; Jackson, 1972, 1974; Rex, 1981). Abundance of fossils per volume of sedi- ment also decreases dramatically with depth in our col- lections (Table 3). Abundance is especially low at out- crops of deep continental shelf to continental slope de- posits including Cangrejos Creek, Bruno Bluff, Shark Hole Point, and Rio Sand Box. Moreover, faunules from other deeper water horizons, such as the Nancy Point Formation (Coates et al., 1992; Coates, this vol- ume), could not be included in the ordination analyses because the collections were too small. Sediments de- posited in water depths =150 m are therefore very dif- ficult to sample, and estimates of diversity are accord- ingly uncertain compared to more fossiliferous deposits. Much more collecting is required to determine whether diversity increases with depth as in the Recent. The apparent stability or increase of molluscan di- versity in the southern Caribbean over ten million years (Text-fig. 14) is consistent with data for Pliocene to Recent molluscan diversity in Florida (Allmon et al., 1993, 1996). Both of these results contrast with the abrupt decline in diversity of the Strombina-group and other so-called paciphile mollusks in the Late Pli- ocene about 2—3 Ma. (Woodring, 1966; Vermeij, 1978; Jung, 1989; Jackson et al., 1993, 1996). Total Carib- bean reef coral diversity also declined precipitously at the end of the Pliocene despite an extended burst of origination throughout most of the Late Pliocene (Budd et al., 1996; Budd and Johnson, 1997). The ordination analyses effectively discriminate among faunules based on age and water depth. Dif- ferences in the composition of faunules separated by only 100 m of depth may be as great as differences among faunules separated by ten million years. Simi- larly, large differences in faunal composition between water depths have been reported for ostracodes from the continental slope versus the deep sea ranging in age from Cretaceous to Pleistocene (Benson, 1979). Much more detailed analyses of sedimentary facies of the different faunules are required to understand the environmental basis of faunal variation. SUMMARY Our study is the largest yet attempted to describe molluscan diversity and faunal composition from the Neogene of tropical America. Problems of sampling have been largely overcome, although older collec- tions are limited to a few, exclusively very shallow- water deposits. Local diversity at the level of the faun- ule varies more than six-fold, and is generally highest where corals (and perhaps seagrasses) were present. As in Recent seas, molluscan abundance decreases greatly with depth. In contrast, diversity does not in- crease with depth as in the Recent, most probably be- cause of problems of finding enough fossils in deeper water deposits. Diversity as estimated by Fisher’s al- pha was constant, or may even have increased, over the ten million years studied. This stability clearly lays to rest the earlier view of mass extinction and de- creased diversity after the Early Pliocene. Ordination using DCA separates faunules along axes highly sig- nificantly correlated with age and depth, which pro- vides a good first step towards separating the evolu- tionary and ecological bases for changes observed. 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NO = Nw BULLETIN 357 APPENDIX 1 AGES AND PALEOBATHYMETRIES OF FAUNULES These are ages and paleobathymetries for the 37 faunules in Table 2. Ages were provided by A. G. Coates (written commun., 1998; see also Aubry and Berggren in Coates this volume; Bybell, this volume; Cotton, this volume; McNeill er al., in press). Most water depths are based on analyses of benthic foraminifera (Laurel Collins, written communs., 1996, 1998; Collins, 1993; Collins et al., 1995; Collins er al., 1996). Exceptions indicated by footnotes were based on stratigraphic relationships. Section numbers of Appendix B, this volume. il. why ie 18. NnNny WwW n in) ON Faunule (with section #) Swan Cay (#25) . Cemetery Pueblo Nuevo (#35) . Upper Lomas del Mar East (reef) (#36) . Empalme (#34) . Cangrejos Creek (#37) . Lomas del Mar East (non-reef) (#36) . Northwest Escudo de Veraguas (#10) . Fish Hole (#22/23) . Ground Creek (no section) . North-central Escudo de Veraguas (#10) . Rio Limoncito (no section) . Chocolate Buenos Aires (#33) . Bomba (#29) . Agua (#29) . Bruno Bluff (#12) . Cayo Agua: West side of Punta Norte (#16) Quiteria (#29) Rio Vizcaya (#39) . Santa Rita (#32) . Northeast Escudo de Veraguas (#10) . Southeast Escudo de Veraguas (#11) . Cayo Agua: Punta Tiburon (#19) . Cayo Agua: Punta Nispero West (#19) 4. Cayo Agua: Punta Nispero Southeast (#20) . Isla Popa (no section) . Cayo Agua: Punta Norte East (#19) Age (Ma) 1.6-1.2 a an eal nn N | in 1.6-1.5 Depth (m) 80-120! 50-100 50-100 150-250 50-100 100-150 75-100 (up- per mud- stone) 40-100 (lower reef conglomerate) <50? 100-150 20-40? <50? 20-40 20-40 150-200 20—40 20-40 <25 20—40 100-150 100-150 40-80 40-80 40-80 <50? 40-80 Abundant diagnostic taxa Amphistegina gibbosa, Cassidulina curvata, Epon- ides antillarum, Eponides repandus, Pararotalia rosea, Planulina ariminensis var. exorna, Quin- queloculina lamarckiana, Siphonina pulchra based on lithostratigraphic relation to faunule #3 C. curvata, Elphidium discoidale, P. ariminensis var. exorna, Sigmoilina tenuis, Spirillina vivipara E. discoidale, Fursenkoina pontoni, Nonionella at- lantica, Pararotalia magdalenensis, Sagrina pulchella Bulimina aculeata, Bulimina marginata, Cassiduli- na minuta, Gyroidina regularis, Planulina fov- eolata, Trifarina eximia based on lithostratigraphic relation to Faunule #3 Bolivina paula, B. marginata, C. curvata, C minu- ta, G. regularis, Hanzawaia concentrica, Melon- is barleeanum, Reussella spinulosa, S. tenuis, S. pulchra, Uvigerina laevis, Uvigerina peregrina B. marginata, E. antillarum, P. ariminensis vat. exorna, T. eximia, U. peregrina, A. gibbosa, E. discoidale, E. antillarum, Nodobaculariella cas- sis, P. ariminensis var. exorna, Q. lamarckiana, S. pulchra estimate based on sediments and mollusks same as Faunule #7 based on apparent stratigraphic relationship to Faunule #’s 13 and 17-19 Based on lithostratigraphic position between reef trends Ammonia decorata, P. magdalenensis, P. sarmien- toi, Rotorbinella umbonata, S. tenuis Based on stratigraphic relations to Faunule #’s 13, 17-19 B. marginata, C. curvata, C. minuta, C. norcrossi australis, T. eximia, U. peregrina E. discoidale, E. antillarum, F. pontoni, H. con- centrica, N. cassis, N. atlantica, Quinqueloculi- na compta, Q. lamarckiana Same as Faunule #’s 13 and 19 Ammonia beccarii, A. gibbosa, Buccella hannai, N. atlantica, P. magdalenensis, Trifarina occi- dentalis A. gibbosa, E. antillarum, Hauerina fragillissima, N. cassis, P. ariminensis var. exorna, R. umbon- ata Same as Faunule #7 Same as Faunule #7 Cassidulina subglobosa, E. discoidale, E. antillar- um, F. pontoni, H. concentrica, N. atlantica, P. ariminensis, R. spinulosa, S. tenuis Same as Faunule #22 Same as Faunule #22 Based on apparent stratigraphic equivalence and proximity to older Cayo Agua Fm. Same as Faunule #22 MOLLUSKS: JACKSON ET AL. APPENDIX 1.—Continued. Abundant diagnostic taxa Faunule (with section #) Age (Ma) Depth (m) 27. Cayo Agua: Punta Piedra Roja West (#17) 5.0-3.5 10-75 28. Quebrada Brazo Seco (no section) 5.2—4.3 <50? 29. Shark Hole Point and top of Nancy Point (#12) S56 100—200 30. Finger Island (#14) 8.2-5.6 60-100 31. Rio Sand Box (#27) 8.7-7.2 150-200 32. Rio Tuba (no section) 8.2-7.24 150-200 33. Rio Calzones (#9) 25? 34. Miguel de la Borda (#6) 251) 35. Isla Payardi (#1) 15—40 36. Mattress Factory (#1) 9.4-8.6 15—40 37. Martin Luther King (#1) 11.8-11.4° 15—40 A. gibbosa, Cancris sagra, E. discoidale, E. antil- larum, Quinqueloculina spp. Based on stratigraphic position between reef tracts and Rio Banano Fm. Bolivina barbata, Bolivina imporcata, N. atlantica, P. ariminensis, U. peregrina A. gibbosa, B. barbata, C. curvata, E. antillarum, H. concentrica, Hanzawaia isidroensa, Lenticu- lina calcar, P. ariminensis, Quinqueloculina seminulum, S. pulchra, U. peregrina B. imporcata, Bolivina lowmani, Bolivina mexi- cana, C. minuta, N. atlantica, P. magdalenensis, R. umbonata Assumed equivalent to Faunule #31 based on ap- parent stratigraphic position assumed equivalent to Faunule #34 A. beccarii, Bolivina merecuani, Bolivina vaugh- ani, B. hannai, P. magdalenensis, R. spinulosa, R. umbonata same as Faunule #35 same as Faunule #35 ' We used the maximum rather than the median depth because the sediments are a reef talus slump deposit. > Assumed Late Pliocene age based on inferred stratigraphic position. * Assumed equivalent to older Cayo Agua Fm. * Assumed equivalent to nearby Rio Sandbox. > Assumed Late Miocene (Collins et al., 1996). a oTl x OI 6b TS8L8 6£8 D oupl[[Ours.ieyy AAYIPUINYIG YjNIISAad vl LL xX 0T v9 €1OL8 898 9) dePIATO uoss[O (D]JaNUIsIDP) P1]2A1O oT OTT x £T 8L Sr198 C88 a oeplipre +) H&G CVS) Bipsvoom0si4y oT 96 x LT 66 O9TS8 C68 HY) ae pr[oursseyy UISUBULLIOF] Winundd oT O11 x Ie 6ll 89£P8 c06 () SePIOIUOJIITYIV SUIpPOY VIIUOJJaNYIAW tl OIL x O¢ Sol 99re8 L06 S sepryeuod 6( US) wniypoluagd VT oIT x cI tp 6SST8 066 ) aePIAYO uOss|O (DAT]OIUIP) P]]9AO ol OIT xe CT 6L 69S18 IcOl dq dePLIauaA, APD 2 lueIy (Viuvidydayp) vIsyjvIOLID ES OT 5% Il PE 8rSO8 S801 Dy aeprlyjaquinjo) UOSIGIRH{ 2 UOSS[O (Yj/OIUIG) DJOIUIg tl O1l x [AS rel t9OroLl Fort 9) oepluny, Suupoom vuus< Ol TE SSEle OI8P i) seplAllo BULIPOOM (P//APIAIIVG) YI12ANO vl 96 >,¢ LT L8 6£S9T 90TS da aepriynqio+) SIPD 2 JURID VjINGLOIUDA oT OTT x Or 9TI £ELIT OLES qd aeptyngio+) Jaupiey vjngsov0Kiny ale aa ax 6 laa €7ZO9I LTLS d oepl[[olesseiD Addny vjjauissp1D vl Tl xX O£ TSI 96701 96701 D sept [[ouny DI] AUAANT, (RWW) (RW) suoy safnuney = suondaT[oo suountoads suauoads sseID Aprurey snuasqns pue snuan 99udLINIDO = 99Ua.IND90 -BUIpIO jo jo jo Joquinu jo sey] ISI ul pos) JOquinyy dJoquinyy sanenuiny Joquinyy “Apmis soyziny Sumber dnoss paaposas Ajsood Apuasaid ev 10} aureu parjdde Ajasoo] 10 uoNRUSIsSap [BULIOJUL ue soyeorpul (x) YSL9}se UY “PXP} oneuaqoid UOWIUIOS UIBDUOS SaJOU SIULOUOXe) PpetsquInny “¢ xipuoddy UI poySI] SopNuUNey Le 234) JO Yyoro 4Ioy BXP] CT do} oy suowe pepny[out BXP] jeuonippe 6OT Ie snjd ‘suountoads OOl= Aq pajuasaida BXP] QC] [[P Sepnpour 4ST] sy peurquios SUOTJI9T[OD [[B UT suauntoads jo toquinu ye101 Aq sourpungde jo Jgpi0 Ul peyst] ore BXeL = VUANADENS YO VAANAD NOWWOD c XIGNHddV : JACKSON ET AL. 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UT saN[NOYJIP (q “UOTSTASI onewaiscs paystqnd Apuases ev Jo yor] (ev :Jo sduaNbasuos vw udaq sey sty, ‘Apmis s9y1iny Surpuad sasAyeue ay) 10J exe) auIOs dnosd 0) sn pasioj sey AiISS99aN :SA.LON OINONOXV.L ‘penunuo)\— ~ XIGNHddV Number of Number of Genus and subgenus Class Specimens Genus and subgenus Class Specimens 1. Swan Cay (Bocas del Toro Basin). PPP: 1995. Turritella G 1194 Olivella (Minioliva) G 367 Conis G 397 Limopsis B 110 Mitra G 291 psi d Nassarius (s.1.) G 87 Antillophos (Antillophos) G 233 Granula G 71 Argopecten B 214 Granulina G 64 Dimya B z 12 Antillophos (Antillophos) G 41 More G 207 Saccella B 32 u olvarina G 187 Volvarina G 32 Siliquarta G 171 rissoine G 31 Velvaring ? At G 152 Subcancilla ? G 27 Olivella (Macgintiella) G 131 Crassinella B 22 Haustellum ae G 123 Alvania G 2] Olivella (Minioliva) G 106 Barleeia G 21 Polystira G 97 Syntomodrillia ? G 20 Naccella B 93 Conus GS 19 Knefastia G 93 Alvania ? G 18 Diodora G 80 Teno stonia G 18 Barbatia (Barbatia) B 75 Olivella (Macgintiella) G 15 INGSSATEES (s.L.) G 74 Polystira G 15 Latirus G 70 Hyotissa B 15 Dentalium (s.1.) S 66 turbonilline G 13 Rueuona ? B 65 Metula G 13 Hindsiclava G 62 Arene (Arene) G 13 Gorabliophila G 55 Arcopsis B 12 Flabellipecten B 51 Cerithiopsis G 12 4. Empalme (Limon Basin). PPP: 715, 718, 719, 759, 1987. 2. Cemetery Pueblo Nuevo (Limon Basin). PPP: 631. Argopecten B 310 Caryocorbula B 125 Luria G 286 Argopecten B 51 Caryocorbula B 248 Gouldia B 45 Bulla G 168 Anadara (Rasia) B 29 Macrocypraea G 85 Limopsis B 17 Crassostrea B 81 Petaloconchus G 14 St FDS G 716 Crassinella B 13 Zonaria . G 47 Varicorbula B 11 Parvanachis G 41 Solariorbis G 10 Conus G 36 Diodora G 7 cypraeid G 32 Nassarius (s.1.) G 6 turbonilline G 31 Cyclopecten B 6 Nassarius (s1) eee G 27 Nea B 6 pace rocaiised (Megapitaria) B 26 Gortopsis G 6 in goniocardia (s.1.) B 25 Arene G 6 LSAT B 23 Jupiteria B 6 Plicatula B 22 Nucula (Lamellinucula) B 5 triphorid G 21 Chama B 5 Cc adulus Ss 21 ISaecelia B 4 J SEGEEES (Glycymerella) B 20 Sulcoretusa G 4 Oliva (Oliva) G 20 Heliacus G 4 Apadara (Cunearca) ? B 17 triphorid 'G 3 Pitar . B 17 Leionucula B 3 Barbatia (Barbatia) B 16 caecid G 3 Saccella B 16 Parvilucina (s.1.) B 3 5. Cangrejos Creek (Limon Basin). PPP: 648, 649, 651, 653, 657. Tucetona B 3 Desh = aa Mearns G 3 Dentalium (s.1.) S 64 3. Upper Lomas del Mar East (reef) (Limon Basin). PPP: Cadulus S 56 rissoine G 45 MOLLUSKS: JACKSON ET AL. APPENDIX 3 Most ABUNDANT TAXA Listed are the 25 most abundant taxa per faunule, based on numbers of specimens obtained from each of the 37 faunules listed in Table 2. 468,469, 628, 942, 943, 944, 948, 949, 950, 951, 962, 963. NO NO a i) tO BULLETIN 357 APPENDIX 3.—Continued. Number of Number of Genus and subgenus Class Specimens Genus and subgenus Class Specimens Olivella (Macgintiella) G 42 Anadara (Rasia) B 16 Olivella (Minioliva) G 37 Haustellum G 14 Nassarius (s.1.) G 35 Argopecten B 13 Alvania G 34 Agladrillia (Agladrillia) G 12 Kurtziella (Kurtziella) G 28 Crassispira (Crassispira) G 11 Styliola G 27 Tucetona B 10 Limopsis B Pal Conus G 10 Anachis G 19 Volvarina G 9 Jupiteria B 13 Leionucula B 8 Glyphostoma (Glyphostoma) G 12 Dentalium (s.1.) S 6 turbonilline G 12 Miraclathurella ? G 5 Kurtziella (Cryoturris) G 12 Natica (Naticarius) G 5 Teinostoma G 11 Metula G 4 Carinodrillia G 11 Serpulorbis G 3 Saccella B 10 Sincola (Sinaxila) G 3 Stigmaulax G 10 Strombus G 3 Trigonulina B 10 Cosmioconcha G 3 Diacria G 9 Polystira G 2 Ringicula (Ringiculella) G 9 Chlamys B 2 turrid G 2) Miraclathurella G 2 Polinices G 7 Jupiteria B 2 Euchelus (Mirachelus) G 7 eulimid G 2 Thelecythara G 7 Subcancilla G 2 . Lower Lomas del Mar East (non reef) (Limon Basin). PPP: 464, OliveRONST) S 2 465, 466, 467, 634, 635, 710, 757, 1982, 1988. 8. Fish Hole (Bocas del Toro Basin). PPP: 1254, 1256, 1304. Caryocorbula B 853 Olivella (Minioliva) G 30 Siliquaria G 519 Conus G 21 Turritella G 292 Anadara (Rasia) B 16 Conus G 285 Voluta G 15 Volvarina G 255 Tucetona B 12 Olivella (Minioliva) G 206 Saccella B 8 Nassarius (s.1.) G 176 carditesine B 7 Argopecten B 174 Nassarius (s.1.) G 4 scaphopod S 145 Polystira G 6 Dentalium (s.1.) S 131 Dentimargo G 6 Vermicularia G 116 Syntomodrillia G 6 Crassinella B 116 Knefastia G 6 triphorid G 96 Volvarina G 6 Voluta G 91 Antillophos (Antillophos) G 5) Gouldia B 88 Sconsia G 5 Dimya B 88 Granulina G 5 Antillophos (Antillophos) G 82 Oliva (Oliva) G 5 Cadulus S 78 Axinactis (Glycymerella) B 4 turrid G 71 Rhinoclavis (Ochetoclava) G 4 turbonilline G 69 Solenosteira G 4 Alabina G 62 Cassis G 4 Granulina G 58 rissoine G 4 Myrtea B 54 triphorid G 4 Polystira G 54 Crassinella B 4 Diodora G 533) Acar B 4 . Northwest Escudo de Veraguas (Bocas del Toro Basin). PPP: 176, 9. Ground Creek (Bocas del Toro Basin). PPP: 1285, 1286. 177, 178, 1974. Ghione B 450 Olivella (Minioliva) G 97 Nucula (Nucula) B 155 Saccella B sill Bulla G 117 Antillophos (Antillophos) G 36 Saccella B 74 Oliva (Oliva) G 34 Lucinisca B 66 Olivella (Macgintiella) G 27 Tagelus (Mesopleura) B 60 Cadulus S 20 Macrocallista (Megapitaria) B 59 10. Nucula (Lamellinucula) MOLLUSKS: JACKSON EFT AL. APPENDIX 3.—Continued. Number of Genus and subgenus Class Specimens Genus and subgenus Eurytellina B 51 Polystira caecid G 50 Cyclopecten Varicorbula B 42 Trachycardium (s.1.) Argopecten B 42 Conus Parvilucina (s.1.) B 42 Nucula (Nucula) Meioceras G 36 Oliva (Oliva) Alabina G 29 Architectonica Stigmaulax G 24 Sulcoretusa Dosinia B 24 Saccella Noetia (Noetia) B 93 Dentalium (s.1.) Angulus B 2] Hindsiclava Tricolia G 19 Cadulus Laevicardium B 19 NOE Anadara (Cunearca) B 19 Buridrillia ? Volvulella (Volvulella) G 18 Srgmaulax Strombus G 17 Acteocina turbonilline G 15 Agaronia Acteocina G 14 Knefastia Conus G 14 Voluta North Central Escudo de Veraguas (Bocas del Toro Basin). PPP: 179, 180, 358, 359, 361, 362, 363, 364. Olivella (Macgintiella) Nassarius (s.1.) Gadilopsis Antillophos (Antillophos) Dentimargo Volvarina Kelliella Saccella Cavolinia Sulcoretusa rissoine Cadulus Crassinella Sincola (Sinaxila) ? Pectunculina Styliola Strioterebrum Caryocorbula Acteocina Alabina Gouldia Turritella Yoldia Abra Eulimella (Ebalina) Tucetona Turritella scaphopod Parvilucina (s.1.) caecid Limacina Anadara (Rasia) Crassinella Ringicula (Ringiculella) Strombus ADDADNAADWAARPOAWVNNAXADWANAIMYAA . Rio Limoncito (Limon Basin). PPP: 463. WOAWMWWHAAWNOD 443 366 195 171 160 147 145 131 125 111 110 95 89 87 86 84 DNDNANDAY~ =e Wn £& 0O Nan nan wok HHH EUNDOONH Rhinoclavis (Ochetoclava) Careliopsis ? Atys Meioceras Anadara (Cunearca) ? Petaloconchus Caryocorbula Nucula (Nucula) Chionopsis Alabina Cadulus Acteocina scaphopod Ostreola Ringicula (Ringiculella) Vermicularia Anadara (Rasia) Cardiomya Saccella Olivella (Olivella) Dentalium (s.1.) Pitar Bulla Eurytellina Olivella (Minioliva) Macrocallista (Megapitaria) Eulimastoma Ithycythara Dendostrea Anadara (s.1.) Epitonium (Asperiscala) Teinostoma Cylichnella eulimid pyramidelline triphorid Nassarius (s.1.) Merisca Ondina Class QADWAANAANANANANAADP*ZO*TDOAANDWOTWA ADWNANANANAAADPAHAAMAHOWOMNOADMDAWAADNVNODAVNYOAWOYD tw tO Ww Number of Specimens eR ON ON ON OY ON ON tt . Chocolate Buenos Aires (Limon Basin). PPP: 1083, 1772, 1773. 286 254 111 56 46 33 29 Man owWoO oO oO wwe tk WWW wWww vs) 224 BULLETIN 357 APPENDIX 3.—Continued. Number of Specimens Number of Genus and subgenus Class Specimens Genus and subgenus Class 13. Bomba (Limon Basin). PPP: 451, 452, 455, 456, 457, 458, 459, 15. Bruno Bluff (Bocas del Toro Basin). PPP: 376, 379, 381, 1975. 460, 461, 462, 668, 669, 672, 678, 686, 691, 758, 1726, 1727, 1728, 1729, 1730, 1731, 1732, 1733, 1764, 1983, 1984, 1986. NMB: 13836, 17477, 17478, 17479, 17480. Crassinella Olivella (Dactylidella) Anadara (Cunearca) Caryocorbula Turritella Olivella (Olivella) Strioterebrum Parvilucina (s.1.) Tucetona Alabina caecid Prunum Acteocina Persicula Stigmaulax Natica (Naticarius) Varicorbula Nassarius (s.1.) Sincola (Sincola) Strombus Dentalium (s.1.) Macrocallista (Megapitaria) Conus Limacina Voluta . Agua (Limon Basin). PPP: 696, 697. Olivella (Dactylidella) Persicula Tucetona Conus Laevidentalium Stigmaulax pyramidelline Prunum Strioterebrum caecid Acteocina Sincola (Sincola) Niso Turritella Sincola (Sinaxila) Mitra Strombus Parvilucina (s.1.) Terebra Lirophora Nassarius (s.1.) turrid Polinices Anadara (Cunearca) ? Oliva (Oliva) Voluta Cancellaria (s.1.) Eontia QW QANADWNNAADPAAAAANNMNDPDAAOTD DAANDDPANNDAWARHAAANIAAAMAAAMAYOWOAOA Ww wWnn ~~) oc oO NNNNNY WW Varicorbula Caryocorbula Anadara (Rasia) Parvilucina (s.1.) Gouldia Olivella (Niteoliva) Alabina Saccella naticid Moerella pyramidelline Turritella Gadilopsis Linga (Bellucina) Trigoniocardia (s.1.) Crassinella Agladrillia (Agladrillia) Calyptraea Tucetona Acteocina Polystira Meioceras Bathoxiphus Dimya Argopecten Varicorbula Tucetona Caryocorbula Argopecten Cadulus Alabina Turritella Cyclopecten Petaloconchus Crassinella turbonilline Plicatula Trigoniocardia (s.1.) Saccella Cavolinia Acteocina Anadara (Rasia) Chama Volvarina caecid scaphopod Crenella Flabellipecten Xenophora Sulcoretusa 1985. Olivella (Dactylidella) Turritella Olivella (Olivella) WMWNDOHNAWOANDADAANHADWOHAAHNADDOIIYD QANDWANODOODDDAHOWDADWDHOWDODHOAVDIOSD G G G 225 90 77 72 45 40 39 36 27 22 21 21 19 19 17 16 16 16 16 15 1S 14 13 1S} 13 . Cayo Agua: West side Punta Norte (Bocas del Toro Basin). 195, 196, 197, 198, 470, 471, 472, 473. 436 S79) 343 145 109 104 103 87 83 71 51 49 45 40 40 35 32 32 23 19 17 16 16 1S 14 2295 950 885 PPP: . Quitaria (Limon Basin). PPP: 449, 450, 679, 695, 1734, 1735, . Rio Vizcaya (Limon Basin). PPP: 925, 9 MOLLUSKS: JACKSON ET AL. APPENDIX 3.—Continued. Genus and subgenus Class Crassinella B Sincola (Sincola) Nassarius (s.1.) Alabina Anadara (Cunearca) Caryocorbula caecid Tucetona Macrocallista (Megapitaria) Stigmaulax Persicula Anadara (Potiarca) Parvilucina (s.1.) Ringicula (Ringiculella) Strioterebrum Agladrillia (Agladrillia) Strombus Volvarina Cylichnella turrid Prunum AAADAAANDANAARPBHAADWDOAWDWOHAAA Acteocina ee) \o ee) tN 1082. Olivella (Dactylidella) Anadara (Potiarca) Prunum Natica (Naticarius) Anadara (Rasia) Turritella Conus Axinactis (Glycymerella) Stigmaulax Strioterebrum Anadara (Cunearca) Noetia Sincola (Sinaxila) Volvarina Dosinia Caryocorbula Hindsia Strombus Voluta Chionopsis Polinices Cancellaria (s.1.) Oliva (Oliva) Eurytellina Volvarina ? Tucetona Strombinophos Antillophos (Antillophos) QAADWADRAANADPAANADPBHAADBAOADHDAWAAROH . Santa Rita (Limon Basin). PPP: 709, 723. Tucetona Cyclopecten Eucrassatella (Eucrassatella) Myrtea Varicorbula Daonmnww i) tw Nn Number of Number of Specimens Genus and subgenus Class Specimens 840 Volvarina G 9 702 Conus G 9 602 Sconsia G 8 602 Arcopsis B 6 575 carditesine B 6 486 Crassinella B 6 455 Anadara (Rasia) B 6 243 Flabellipecten B 5 208 Trachycardium (s.1.) B 5 207 Gouldia B 5 176 Laevidentalium S 5 158 Arene G 5 146 Crucibulum (Crucibulum) G 5 143 Anomia B 5 143 Barbatia (Barbatia) B 4 143 Plicatula B 4 135 Dentalium (s.1.) S 4 119 Spathochlamys B 4 111 Strombus G 3 110 Architectonica G 3 105 Voluta G 3 99 triphorid G 3 933, 935. 937 Marginella G 3 ce) die lata Nassarius (s.1.) G 3 Nucula (Lamellinucula) B 3 439 Phalium G 3 i 20. Northeast Escudo de Veraguas (Bocas del Toro Basin). PPP: 48 365, 366, 367, 368. 41 Olivella (Niteoliva) G 136 36 Nassarius (s.1.) G 127 23 Crassinella B 105 18 Volvarina G 94 17 Anadara (Rasia) B 88 15 Caryocorbula B 82 15 Saccella B 74 13 Kelliella B 71 12 rissoine G 71 12 Cavolinia G 64 11 Gadilopsis S 59 10 Dentimargo G 58 9 Sulcoretusa G 53 8 Tellina (Scissula) B 50 8 Gouldia B 49 8 Antillophos (Antillophos) G 47 5 Polinices G 44 4 Alabina G 42 4 Argopecten B 42 3 Strioterebrum G 41 3 Marginella ? G 38 3 Parvilucina (s.1.) B oi B Abra B 36 3 Pectunculina B 34 Styliola G 33 176 21. Southeast Escudo de Veraguas (Bocas del Toro Basin). PPP: 72 168, 170, 431, 478, 479, 480, 481, 482, 483. 32 Antillophos (Antillophos) G 199 14 Gadilopsis S 149 11 Cadulus s 132 NO i) tN N Ww BULLETIN 357 APPENDIX 3.—Continued. Genus and subgenus Saccella Nassarius (s.1.) Eulimella (Ebalina) Dentalium (s.1.) Volvarina Conus Bathoxiphus Olivella (Minioliva) rissoine Strioterebrum Leionucula Anadara (Rasia) Dentimargo Microgaza Parvilucina (s.1.) Architectonica Olivella (Macgintiella) turbonilline Styliola Haustellum Granulina ? Volvulella (Volvulella) Compsodrillia ? 296, 297, 335, 337, 339, 340, 341. Olivella (Dactylidella) Caryocorbula Tucetona Turritella Crassinella Argopecten Acteocina Oliva (Oliva) Anadara (Rasia) Varicorbula Gouldia Cadulus Parvilucina (s.1.) Persicula Alabina Strombus Nucula (Lamellinucula) Olivella (Macgintiella) Volvarina Ringicula (Ringiculella) Strioterebrum Natica (Natica) turbonilline Stigmaulax Ervilia 323, 325, 326, 330, 1303. Turritella Plicatula Tucetona Oliva (Oliva) Caryocorbula Varicorbula Number of Class Specimens 131 98 73 69 65 64 64 60 ANAAANAADWAAWDWAADNAN*NAYVOAYD WWWW DArNe NWA OSO NNN NN NY - NM Ww 559 420 262 160 155 117 DAADAAADAADAPAANADYVZDADWHAOADWOWWOHA a N 34 471 65 65 53 52, 33 BwWawDO . Cayo Agua: Pt Tiburon (Bocas del Toro Basin). PPP: 294, 295, . Cayo Agua: Pt Nispero West (Bocas del Toro Basin). PPP: 318, Number of Genus and subgenus Class Specimens Saccella B 26 Axinactis (Glycymerella) B 25 Tellina (Phyllodina) B 23 scaphopod S 23 Strombus G 20 Nucula (Nucula) B 19 Prunum G 17 Eucrassatella (Eucrassatella) B 17 Strioterebrum G 19/ Haustellum G 15 turbonilline G 14 Lunarca ? B 13 Parvilucina (s.1.) B 12 Ondina G 11 Dendostrea B 11 Alabina G 11 Eulimastoma G 11 Petaloconchus G 10 Lirophora B 10 . Cayo Agua: Southeast Pt Nispero (Bocas del Toro Basin). PPP: 307, 308, 310, 311, 313, 476. Varicorbula B 1494 Tucetona B 242 Caryocorbula B 215 Crassinella B 168 Argopecten B 110 Gouldia B 77 Alabina G 49 Ringicula (Ringiculella) G 39 Alvania G 34 Chama B 32 Petaloconchus G 31 Acteocina G 27 turbonilline G 26 Volvulella (Volvulella) G 23 Sulcoretusa G 21 Meioceras G 21 Strombus G 20 Turritella G 20 Anadara (Rasia) B 20 Cyclopecten B 19 Cavolinia G 19 marginellid G 18 Parvilucina (s.1.) B 18 Tellidorella B 17 Conus G 16 Natica (Naticarius) G 16 . Isla Popa (Bocas del Toro Basin). PPP: 422, 426, 427, 1276, 1277, 1283, 1284. Varicorbula B 1498 Caryocorbula B 174 Tucetona B 116 Meioceras G 86 Alabina G 48 Lirophora B 33 Gouldia B 27 Fissidentalium S 26 Macrocallista (Megapitaria) B 24 21: Genus and subgenus Saccella Oliva (Oliva) Stigmaulax Trigoniocardia (s.1.) Argopecten Conus Petaloconchus Polystira Crenella Eucrassatella (Eucrassatella) Prunum Anomia Arcinella Lucina (Lepilucina) Dendostrea ? Cadulus Strombus Voluta 217, 373, 475, 1203. Antillophos (Antillophos) Anadara (Rasia) Turritella Polystira Conus Strioterebrum Hindsiclava Prunum Cancellaria (s.1.) Sconsia Petaloconchus Stigmaulax Oliva (Oliva) Crassinella Voluta Serpulorbis Solenosteira Strombus Tucetona Olivella (Macgintiella) Argopecten Lirophora Fasciolaria (Fasciolaria) Architectonica Calliostoma Cayo Agua: Pt. Piedra Roja West (Bocas del Toro Basin). PPP: 204, 345, 346, 348, 350, 1188. Varicorbula Tucetona Caryocorbula Alabina Argopecten Crassinella Acteocina Gouldia Anadara (Rasia) Oliva (Oliva) Strombus MOLLUSKS: JACKSON ET AL. APPENDIX 3.—Continued. Class Number of Specimens QANYM*MBAWDAOWDWDANADPWIAAD QAAARDWABDAANANAWAARDAAAADANAAATRA QAAWDOHAWAWBDOWWYD WW NNN NN NN Woy . Cayo Agua: Pt Norte East (Bocas del Toro Basin). PPP: 200, 201, 538 296 143 106 103 79 78 76 66 58 S7 47 44 44 36 24 23 19 18 14 14 13 11 11 10 870 569 528 396, 1976; 1977, 1978. Fissidentalium Amarophos Polystira Argopecten Volvarina Stigmaulax Bathygalea (Miogalea) Conus turrid Oliva (Oliva) Natica (Naticarius) Antillophos (Antillophos) Polinices . Shark Hole Point (Bocas del Toro Basin). QAANAAANADAAARAAY PPP: 390, 391, 119 75 41 36 iI) 17 11 9 fA NO 227 Number of Genus and subgenus Class Specimens Turritella G 151 Moerella B 139 Macrocallista (Megapitaria) B 97 Cadulus S 94 Hyotissa B 94 Ringicula (Ringiculella) G 90 carditesine B 70 Conus G 65 Cyclopecten B 56 Natica G 35) Dentalium (s.1.) S 53 Trachycardium (Phlogocardia) B 48 Spondylus B 45 caecid G 45 Crenella B 45 - Quebrada Brazo Seco (Limon Basin). PPP: 1775, 1776, 1777. Varicorbula B 37 Turritella G 33 Caryocorbula B 20 Volvarina G 9 Natica (Naticarius) G 8 Stigmaulax G 1 pyramidelline G 7 Dentimargo G 7. Ancilla (Eburna) G 6 Lirophora B 5 Marginella G 5 Strombinophos G 5 Natica G 5 Dendostrea ? B 5 Olivella (Dactylidella) G 5) Prunum G 4 Anomia B 4 Persicula G 3 Polystira G 3 Kurtziella (Cryoturris) G 3 Saccella B 3 Kurtziella (Kurtziella) G 3 Sincola (Sinaxila) G 3 Sincola (Sincola) G 3 Solariella G 3 Nassarius (s.1.) G 3 Volvulella (Volvulella) G 3 Olivella (Macgintiella) G 3 31. Number of Number of Genus and subgenus Class Specimens Genus and subgenus Class Specimens Anadara (Rasia) B 4 Natica (Naticarius) G 11 Homalopoma (Leptothyropsis) G 4 Strombinophos G 9 Carinodrillia G 4 Volvarina ? G 9 Compsodrillia G 3 Cancellaria (s.1.) G 9 Saccella B 3 Prunum G 7 Crucibulum (Crucibulum) G 3 Conus G 7 Strioterebrum G 3 Caryocorbula B 6 Fusiturricula G 3 Terebra G 6 Anomia B 3 Voluta G 6 typhine G 3 Microgaza ? G 6 Miraclathurella G 3 Dentalium (s.1.) S 5 Natica (Natica) ? G 2 Chiodrillia ? G 4 use ladrallia (Agladriliia) Se a 32. Rio Tuba (Limon Basin). PPP: 1765, 1766, 1768, 1769, 1770. Architectonica G 2 Cancellaria (s.1.) G 2 scaphopod 2 a Macrocallista (Megapitaria) B 2 Teinostoma G 8 turbonilline G 5 30. Finger Island (Bocas del Toro Basin). PPP: 191, 477, 1996. G@adilus S 5 Conus G 231 pyramidelline G 4 Tellidorella B 172 Fissidentalium S 4 Polystira G 156 Subcancilla G 4 Strioterebrum G 79 Dentalium (s.1.) S 4 Subcancilla G 50 Eulimastoma ? G 3 Tesseracme S 49 Meioceras G 3 Dentalium (s.1.) S 49 Natica (Naticarius) G 8 Cancellaria (s.1.) G 47 Saccella B 3 Sconsia G 46 Turritella G 2 Ervilia B 36 caecid G 2 Scobinella G 36 Acteocina G 2 Crassinella B 35 Polinices G 2 Natica (Naticarius) G 34 Prunum G 2 Saccella B 33 Arcinella B 2 Ficus G 33 Dentimargo G 1 Hindsiclava G 31 Tucetona B 1 Anadara (Rasia) B 31 Cyclostremiscus (Ponocyclus) G 1 Syntomodrillia G 29 Cosmioconcha G 1 Antillophos (Antillophos) G 29 triphorid G 1 Architectonica G 28 Varicorbula B 1 Acila B 25 Macrocallista (Megapitaria) B 1 Carinodrillia G 21 Balcis G 1 Polinices G 19 Bellaspira G 1 Antillophos (Antillophos) ? G 18 Antillophos ? G 1 Linga (Pleurolucina) B 17 Volvulella (Volvulella) G 1 Solenosteira G i) B 1 G 1 G 1 B 1 G 1 B 1 G 1 G 1 G 1 B 1 G 1 Rio Sand Box and Hone Creek (Limon Basin). PPP: 453, 454, 1736, 1737, 1774, 1989. Turritella Strombina (Strombina) ? Polystira Olivella (Macgintiella) Sconsia Natica Antillophos (Antillophos) Nassarius (s.1.) Acila Cadulus Oliva (Oliva) Polinices turrid QAQAYRWNAAANAAAAA BULLETIN 357 APPENDIX 3.—Continued. 295 33 32 29 28 24 24 21 20 14 13 13 11 Argopecten ? Alabina Architectonica Chama Carinodrillia Eurytellina Cancellaria (s.1.) Strioterebrum Agladrillia ? Lamelliconcha Oliva (Oliva) Argopecten Tucetona Ervilia Flabellipecten Axinactis (Glycymerella) 33. Rio Calzones (North Coast). PPP: 162, 163. Dom ww © MOLLUSKS: JACKSON ET AL. 229 APPENDIX 3.—Continued. Number of Number of Genus and subgenus Class Specimens Genus and subgenus Class Specimens Acteocina G 4 Gadilopsis Ss 7 Nucula (Nucula) B 4 Nita B 7 Gouldia B 3 Anadara (Rasia) B 6 Conus G 3 Mitrella G 6 Parvilucina (s.1.) B 3 Cyclostremiscus (Ponocyclus) G 6 Eucrassatella (Eucrassatella) B 3 A ; Architectonica G 2 25. Isla Payardi (Panama Canal Basin). PPP: 34, 225, 226, 487, 488, Witrinellan?, G 2 489, 1077, 1079, 1080, 1081, 1086, 1087, 1307, 1308. Trus (Irus) ? B 2 Turritella G 3136 Terebra G 2 Anadara (Rasia) B 1661 Crassinella B 2 Lirophora B 1069 Oliva (Oliva) G 2 Strombina (Strombina) G 1048 Solariella G 5) Oliva (Oliva) G 800 Spondylus B 2 Cadulus S 676 Transennella B 1 Antillophos (Antillophos) G 483 neogastropod G 1 Cancellaria (s.1.) G 448 Calliostoma ? G 1 Conus G 386 Cantharus G 1 Chama B 365 Cyclopecten B 1 Hyotissa B 356 Alabina G 1 Natica (Naticarius) G 344 naticid G 1 Polinices G 324 Macrocallista (Megapitaria) B 1 Polystira G 284 Tricolia G 1 Architectonica G 253 Chama B 1 Olivella (Niteoliva) G 225 Hiatella ? B 1 Stigmaulax G 208 Polystira G 1 Petaloconchus G 184 Anomia B 1 Nassarius (s.1.) G 154 Petaloconchus G 1 Strioterebrum G 145 carditesine B 1 Ervilia B 132 Meioceras G 1 Caryocorbula B 132 corbulid B 1 Cylichnella G 128 Hyotissa B 1 Terebra (Panaterebra) G 110 Ringicula (Ringiculella) ? G 1 Flabellipecten B 83 Lirophora B l 36. Mattress Factory (Panama Canal Basin). PPP: 224, 227, 229, cnlimid G l 230, 484, 485, 486, 1030, 1031, 1032, 1033, 1034, 1035, 1078, STS G l 1305, 1306. 34. Miguel de la Borda (North Coast). PPP: 1973. Turritella G 1523 Crassinella B 75 Ervilia B 1470 Varicorbula B 72 Cymatophos G 949 Saccella B 44 Alabina G 526 Lirophora B 43 Alveinus B 487 Nassarius (s.1.) G 34 Olivella (Niteoliva) G 471 caecid G 30 Polinices G 447 Eurytellina ? B 26 Strioterebrum G 423 Parvilucina (s.l.) B 21 Cadulus S 385 Macrocallista (Megapitaria) B 19 Nassarius (s.1.) G 280 Natica (Naticarius) G 17 Anadara (Rasia) B 268 Caryocorbula B 16 Cancellaria (s.1.) G 259 Cadulus Ss 15 Stigmaulax G 212 Anadara (s.1.) B 15 Leptopecten B 194 Acteocina G 15 Caryocorbula B 187 Tellina (Scissula) B 15 Oliva (Oliva) G 163 Lamelliconcha B 15 Trigoniocardia (s.1.) B 145 Acila B 14 Cylichnella G 144 Turritella G 13 Anadara (Cunearca) B 140 Crepidula G 10 Natica (Naticarius) G 138 eulimid G 9 Acteocina G 138 Cylichnella G 9 Strombina (Strombina) G 136 Oliva (Oliva) G 8 Eucrassatella (Hybolophus) B 134 APPENDIX 3.—Continued. Genus and subgenus BULLETIN 357 Crassinella Crucibulum (Crucibulum) 221, 222, 223, 231, 232, 233, 490, 1075. Cymatophos Turritella Strioterebrum Cancellaria (s.1.) Olivella (Niteoliva) Ervilia Trigoniocardia (s.1.) Polinices Anadara (Rasia) Crassinella Architectonica Macrocallista (Megapitaria) Natica (Naticarius) Caryocorbula Leptopecten Oliva (Oliva) Conus Nassarius (s.1.) Eupleura Cadulus Solenosteira Agladrillia (Agladrillia) Semele Chionopsis Neverita (Glossaulax) QWWDAAYVNOHANDADPAOWOWDDAOAWDWDOAAIAADA Number of Class Specimens B 117 G 105 . Martin Luther King (Panama Canal Basin). PPP: 218, 219, 220, 2124 1022 467 435 420 318 314 286 264 245 234 210 206 184 181 166 146 133 111 101 87 76 76 70 66 CHAPTER 10 NEOGENE-QUATERNARY OSTRACODA AND PALEOENVIRONMENTS OF THE LIMON BASIN, COSTA RICA, AND BOCAS DEL TORO BASIN, PANAMA PAMELA FE. BORNE Louisiana Sea Grant College Program Louisiana State University Baton Rouge, Louisiana 70803, U.S.A. THOMAS M. CRONIN U.S. Geological Survey 926 National Center Reston, Virgina 20192, U.S.A. JOSEPH E. HAZEL Department of Geology and Geophysics Louisiana State University Baton Rouge, Louisiana 70803, U.S.A. INTRODUCTION Tropical marine ostracodes from Neogene and Qua- ternary sediments of the Central American Caribbean region have been the subject of biostratigraphic (Bold, 1988), ecological (Krutak, 1971), taxonomic (Teeter, 1975), and evolutionary studies (Cronin, 1988; Cronin and Schmidt, 1988). As part of the Panama Paleon- tology Project (PPP), Neogene and Quaternary ostra- codes are being studied from the Central American region. The overall goal of this research is to evaluate the impact of the emergence of the Central American Isthmus as a land barrier between the Caribbean/trop- ical Atlantic and the Pacific oceans on marine ostra- code biodiversity and the oceanic environments in which extant ostracodes evolved. Due to the ecological specificity of many living tropical ostracode species, they are ideally suited for reconstructing paleoenviron- ments on the basis of their occurrence in fossil assem- blages, which in turn can lead to a better understanding of the tropical climatic and tectonic history of Central America. The principal aims of this chapter are: (a) to docu- ment the composition of the ostracode assemblages from the Limon Basin of Costa Rica and the Bocas del Toro Basin of Panama, two areas yielding extensive ma- rine ostracode assemblages; (b) to describe the environ- ments of deposition within these basins; and (c) to doc- ument the stratigraphic distribution of potentially age- diagnostic ostracode species in the Limon and Bocas del Toro basins in order to enhance their use in Central American biostratigraphy. A secondary, but none-the- less important goal is to assemble a database on the distribution of modern ostracode species in the Carib- bean and adjacent areas as a basis for comparison with fossil assemblages. Although the ecological, biostrati- graphic and paleoenvironmental conclusions presented here will improve as additional material is studied, these fossil and modern ostracode databases constitute the foundation for future evolutionary and geochemical studies of tropical Caribbean and eastern Pacific Ocean ostracodes. Moreover, we present here evidence that major faunal and oceanic changes occurred in the west- ern Caribbean over the last 4 million years, probably related to changes in ocean circulation due to the emer- gence of the Isthmus as well as other climatic events. ACKNOWLEDGMENTS The authors extend their sincere thanks to A. G. Coates, J. B. C. Jackson, L. S. Collins, A. F Budd, D. E McNeill and the staff at the Smithsonian Tropical Research Institute for assistance and field support, to 232 BULLETIN 357 Table 1.—Limon Basin, Costa Rica, and Bocas del Toro Basin, Panama, PPP samples used herein. Section numbers and names from Appendix B. Samples (PPP numbers) Formation Section 59-62, 293, 298, 300, 306, 307, 334, 335, 337, 371-374 Cayo Agua 19. North Point to Tiburon Point 63 Cayo Agua 16. North Point, Western Side 168-169 Escudo de Veraguas 11. Southeastern Coast 175, 358, 360-369 Escudo de Veraguas 10. Northern Coast 389 Shark Hole Point 12. Bruno Bluff to Plantain Cays, Valiente Penin- sula 410 Nancy Point 12. Bruno Bluff to Plantain Cays, Valiente Penin- sula 634-638, 645, 953-955, 959-960, 1480-1483 Moin 36. Lomas del Mar, Eastern Sequence 647-658, 1357, 1375-1376, Moin 37. Lomas del Mar, Western Reef Flank Sequence 668, 670, 672, 678, 679, 682-684, 685-688, 690 Rio Banano 29. Rio Banano 710 Moin * Los Laureles 712, 1369-1371, 1392-1406, 1435, 1436, 1438, 1442, 2003 Moin 34. Empalme 720-721 Moin 32. Santa Rita 1433-1434 Moin * Avenida Barracuda * = jsolated outcrop. A. FE Budd for support (National Scientific Foundation Grant EAR-9219138), to B. K. Sen Gupta and W. van den Bold for taxonomic help, and to P. A. Fithian for use of her dissertation data. We also thank our review- ers, Mervin Kontrovitz and Robin Whatley, for helpful comments. MATERIAL AND METHODS Table 1 lists the formations, section names, and sample numbers associated with each outcrop section that we studied for ostracodes. Sample numbers are keyed to the Panama Paleontology Project Database described by Kaufmann and Fortunato (this volume). Most samples described here were taken for ostracode study and were processed and/or picked for ostracodes at the U. S. Geological Survey in Reston, Virginia, or the Department of Geology and Geophysics, Louisiana State University, Baton Rouge, Louisiana. We pro- cessed sediment first by soaking 50-gram (dry weight) samples for one to two days in water in which a small amount of sodium bicarbonate had been dissolved. In rare cases, we disaggregated sediment using Varsol. Sediment was then washed through a 63 pm sieve and the size fraction greater than 150 wm was picked for ostracodes. Because of the large number of samples, we focused on key sections from each region to obtain data on the preservation, abundance and diversity of the assemblages. In most cases, samples were com- pletely picked of ostracodes; abundances ranged from seven to 1501 valves, with an average of 229 valves in 104 samples. In samples where ostracodes were pre- sent in great numbers, standard micro-splitting tech- niques were used so that at least 200-300 individual valves were obtained in order to perform quantitative analyses of assemblages. Table 2 contains fossil ostracode census data con- sidered in this chapter. Because the study of the tax- onomy of tropical marine ostracodes from the Central American Caribbean region is uneven, the taxonomic categories used in our analyses reflect this unevenness in that we were obliged to use both genus- and species- level categories in the census data. Figured specimens shown in Plates 1, 2 and 3, have been reposited at the United States National Museum of Natural History. A large modern ostracode database was assembled in or- der to provide a basis for paleoenvironmental interpre- tations. The fossil and modern databases are available electronically from the first author and the fossil data are at http://www.fiu.edu/ “collinsl/. Two sets of semi-quantitative cluster analyses were executed using presence-absence data: one analysis of 57 Moin Formation samples using 41 taxa and a sec- ond set of analyses of 44 taxa from selected samples from the Limon and Bocas del Toro Basins. The Jac- card coefficient provides a commonly used binary co- efficient that tends to slightly emphasize the differenc- es between two assemblages more than some other co- efficients (Cheetham and Hazel, 1969). We used pres- ence-absence as opposed to relative abundances because the wide variation in ostracode abundance in samples from different lithologies makes it unattrac- tive to compare samples using species proportions. Second, the specific ecology of many tropical ostra- code species is such that dominant species characterize one or two of the tropical biofacies encountered. Jac- card coefficients were calculated for the presence-ab- sence matrix and a cluster analysis of the resulting matrix was performed using the average linkage meth- od (Text-fig. 1) and complete linkage method (Text- fig. 2a and 2b). OSTRACODES: BORNE ET AL. MODERN OSTRACODES FROM THE CARIBBEAN SEA Maddocks (1974) and Bold (1983, 1988) give ex- cellent comprehensive introductions to the taxonomy, ecology and biostratigraphy of Caribbean ostracodes. Our goal was to construct from the large but scattered and somewhat uneven literature a database (Modern Ostracode Database) on modern ostracode ecology and zoogeography for species living in the tropical seas of the Caribbean and adjacent regions. This database pro- vides a reasonable basis for making paleoenvironmen- tal inferences from fossil assemblages occurring in the Central American region. We chose to include selected species and/or generic census data from five primary sources (Cronin and Dowsett, 1990; Fithian, 1980; Kontrovitz, 1976; Krutak, 1982; Teeter, 1975) for three reasons. First, they all provided excellent illustrations or references to illustrations, allowing taxonomic con- sistency across different geographical areas. Secondly, they span a wide spectrum of ecological habitats, from restricted lagoons to the continental slope, and include all biofacies encountered in Neogene and Quaternary sediments. Finally, each reference contains species census counts (not just presence/absence data) that give us information on the relative frequency of dom- inant and rare taxa and which will allow future quan- titative comparisons between fossil and Recent assem- blages. It is important to emphasize that the paleoen- vironmental reconstructions given below for Neogene and Pleistocene ostracode biofacies are based heavily on the ecological data contained in the Modern Ostra- code Database, deemed adequate for reconstructing past environments of deposition. OSTRACODES FROM LIMON BASIN, COSTA RICA Ostracode species occurrence data for the Moin and Rio Banano formations of the Limon Basin and for the Bocas del Toro Basin are given in Table 2. The fol- lowing are short summaries of the ostracodes from each formation and the environmental significance of the biofacies. MOIN FORMATION We identified ostracode biofacies in the Moin For- mation, Limon Basin, to determine the late Pliocene- early Pleistocene environmental history of the region and to provide regional biostratigraphic correlation be- tween the Limon and the Bocas del Toro basins. The lithostratigraphy followed here is based on work by Taylor (1975), Cassell and Sen Gupta (1989), Coates et al. (1992), and Coates (this volume). The Moin For- mation consists of several distinct lithofacies and mac- rofaunal biofacies, most notably a fine-grained facies i) Ww WwW (originally described from an unnamed creek that flows through the Cangrejos community) and a coral reef facies (Appendix A, Map 11, Inset B). There ap- pear to be at least three distinct coral reef trends in the region of the city of Limon (Taylor, 1975; McNeill et al., 1996; McNeill et al., in press; Budd er al., this volume). Although the stratigraphic relationships be- tween the Cangrejos creek and reefal facies are not yet firmly established, we were able to distinguish the youngest trend from the older trends on the basis of the ostracode assemblages and provide a preliminary interpretation of their significance. We were also able to make a preliminary comparison to ostracode assem- blages from the Quebrada Chocolate section (Appen- dix A, Map 11, Inset A; associated with the oldest trend and now considered part of the Quebrada Choc- olate Formation). A total of 57 samples from two measured sections (Cangrejos creek and Lomas del Mar) and several smaller exposures of the Moin Formation, and one measured section (CTA Fence) currently considered part of the Rio Banano Formation (Appendix A, Map 11, Inset C) form the basis of this analysis (Table 1, 2). The cluster analysis of the ostracode data (57 sam- ples, 41 taxa; Text-fig. 1) revealed five groups of sam- ples, A—E, referred to here as biofacies (Hazel, 1971, 1988), each being characteristic of a faunal assemblage and indicative of distinct environments of deposition. Biofacies A This biofacies is represented by Cluster A (Text-fig. 1) and consists of 12 samples, nine of which come from the type section of the Moin Formation exposed along Cangrejos creek west of the city of Limon. The dominant taxa in order of their mean percentage oc- currence in a sample are: Krithe spp. (12%), Cyther- opteron wardensis Puri, 1954 (12%), Argilloecia spp. (14%), Bradleya aff. B. acceptabilis Liibimova and Sanchez-Arango, 1974 (7%), Loxocorniculum spp. (6%). Other taxa averaging 5% include Radimella con- fragosa (Edwards, 1944), Cytherella spp. and Echin- ocythereis madremastrae Bold, 1988. Bradleya aff. B. acceptabilis is closely related to the living species B. normani (Brady, 1866), which Bold (1968) suggested inhabits the Caribbean today. This biofacies represents a mixture of shallow, warm-water, carbonate-platform taxa (Loxocornicu- lum, and to a lesser extent, Radimella), taxa typical of outer shelf/upper slope, cooler-water habitats in the At- lantic/Caribbean (Cytherella, Echinocythereis), and typically deep colder-water taxa (Krithe, Bradleya). Krithe is the predominant genus in the world’s deep- sea environments (Coles eft al., 1994: Van Harten. 1996), usually comprising 40-70% of assemblages in NO Ww & BULLETIN 357 Table 2.—Ostracode census data by PPP number from the Limon Basin, Costa Rica, and Bocas del Toro Basin, Panama. Data are available at internet site http://www.fiu.edu/~collinsl/. ou uoxe| Taxa Actinocythereis gomillionensis (Howe and Ellis, 1935) 6S 09 19 c9 e9 891 691 GLb E62 862 00€ 90€ Z0€ PEE Gee > Ss > n no] js) =. =a > TS, WONMDNHL WMH —! Basslerites minutus Bold, 1953 1 2 1 3 4 Basslerites sp. 2 Sr4 Bradleya aff. B. acceptabilis Lubimova and Sanchez-Arango, 1974 6 10 Buntonids Bythoceratina spp. 1.3 1 Caribbella puseyi Teeter, 1975 24 Cativella sp. Cativella navis Coryell and Fields, 1937 ae th 7 ee ZA) 10 4 dh Slee eee Cativella pulleyi Teeter, 1975 9 27 Caudites aff. C. rectangularis (Brady, 1869) Caudites aff. C. highi Teeter, 1975 Caudites medialis Coryell and Fields, 1937 2 “bp 8 a) Caudites nipeensis Bold, 1946 35 5 Caudites symmetricus Bate, Whittaker and Mayes, 1981 Caudites spp. 3 1 2) Se 4) Teel Coquimba fissispinata Benson and Coleman, 1963 Costa stokesae Bold, 1967 icentg Cushmanidea sp. 1 Cytherella spp. SB 2 2 10) Ze 4K! Cytherelloidea spp. Cytheropteron spp. 23), 16) 54) 15245 4 45 Cytheropteron (Lobosocytheropteron) palton Bold, 1966 1 1 Cytherura spp. “ul © if 1 iil 2 & Echinocythereis madremastrae Bold, 1988 Eucytherura spp. 3 et Gangamocytheridea plicata Bold, 1968 Heinia spp. 1 2 Hemicytherura spp. Wh cop) > eas for) nm w a =n = a WONA=B SSNIS (p) an Nm Wor lop) = Ss > D cat a (2) Seat LS) Kangarina spp. Krithe spp. Loxoconcha spp. Loxocorniculum spp. Macrocyprids 2 4 9 10 7 Mw MmNm— NR > an 45 |Megacythere johnsoni (Mincher, 1941) 46 |Microcythere sp. 47 |Munseyella bermudezi Bold, 1966 1 48 |Neocaudites scottae Teeter, 1975 42a Seo ete site 76) al re Awe 4 49 |Costa cf. Costa variabilicostata recticostata Bold, 1970 2 50 |Occultocythereis angusta Bold, 1963 51 | Orionina boldi Cronin and Schmidt, 1981 §2 |Orionina vaughani (Ulrich and Bassler, 1904) 3 1 410) Neha det gv od 1 2 53 |Palmoconcha spp. 54 |Paracyprids 1 10 55 |Paracytheridea spp. 1 Ws 1 POX PH aK) 720) SAN 56 |Paradoxostoma spp. 2 2 1 2 57 |Parakrithe alta Bold, 1988 58 |Parakrithella sp. 59 |Pellucistoma spp. v ap 6b 4 B ih eh 4 16 1 60 |Perissocytheridea spp 2 2 4 i 2 40) 61 |Phlyctocythere spp. 1 1 62 |Pontocyprids 7 63 |Propontocyprids 2 5 3 64 |Pseudoceratina droogeri Bold, 1965 65 |Pseudocythere caudata Sars, 1865 66 |Pseudosammocythere spp. 2A eee, (lame A CE | 1 67 |Pterygocythereis spp. 2 3 7 68 |Pumilocytheridea sp. 1 4 2 A) il ee 1 1 69 |Puriana spp. 77 88 70 |Reussicythere reussi (Brady, 1869) 71 |Quadracythere howei (Puri, 1953) Sid eZ 06 DP AK0 72 |Quadracythere producta (Brady, 1868) 6 73 |Radimella cf. R. confragosa (Edwards, 1944) 18 3 8 74 |Radimella ovata Bold, 1988 10) “3 8 Sms 3 Woe 42°46 75 |Radimella aff. R. ovata Bold, 1988 10 76 |Radimella wantlandi (Teeter, 1975) 4 15 77 |Semicytherura spp. 78 |Touroconcha lapidiscola (Hartmann, 1959) 4 | th) 2 7 se A ie 25 o 3 SB 79 | Triangulocypris laeva (Puri, 1960) 80 |Triebelina spp. 1 81 |Uroleberis sp. 82 |Xestoleberis spp. ie) al 5 GS} a! Bil eh Al aa be SN aS} 83 |Ostracode A 84 | Other (unidentified) Total specimens 196 106 201 347 178 135 257 425 109 424 155 14 164 159 328 OSTRACODES: BORNE ET AL Table 2—Continued. 1 2 3 4 5 6 7 8 9 23 123) 4 3 1 1 204 5 AS 2 9 30 1 3 cle ra 2 7 7) {3} 6 Wes 22 332 33 Ww 16 47 on 44 164 uo NOM — Orn 15 133 One ENS, fw nme 98 —=N 14 26 74 On 12 95 mw 23 0 12 33 121 NW Con 21 76 Rw No -oOo © 6 24 No 13 igel 1 ial 96 103 14 82 24 34 16 26 104 343 590 422 281 183 67 494 655 219 166 168 2 1 —-wW 2 18 Mr ~ Sf Say Cort aS} wn fo: oo Pw lok ine) Pro oen-9 76 1 NOM ine} Onna Nh Ls) —NMOnr NR wo ined On Nw > wrnw Sr — NPN nm o-=8 20 aoa om 22 21 woud [o,Kop) -f 11 ar-O-®D On wn wn as. Demme 30 2022200 al HP Sh BG 9A ab 236 1 2 3 4 5 6 7 8 9 =—=hM NoS fon 2 MN WOnNNr on 1 ON Wwnr ~ DWWG 20 20 =~ wo On 63 10 10 of. Nyeo— 7 12 27 30 91 63 16 108 1 31 14 10 9 1 65 111 138 145 344 340 312 335 234 57, NOM -D 12 144 37 90 44 —NM 82 BULLETIN 357 Table 2.—Continued. 15 122 11 2 198 132 113 41 353 121 357 175 307 236 oa 16 12 1 CB ey 4 oy a a 22 27 1 Ze) Oil 11 28 123 of —-Mhm NO Noo 87 27 3 wo Mon 48 27 76 3 WP 21 1 on 32 36 15 10 23 ENS) 87 0 3 2 2 8 12 4 3 32 14 ss 59 WhnNw 2506 Gon wo a As i GY 119 26 91 Se) i) 2 288 4 3 st3 1 51 90 149 295 297 563 373 662 tN WwW ~ OSTRACODES: BORNE ET AL. Table 2.—Continued. 1 2 3 4 5 6 7 8 9 OD =NG MwO=D oNN= =) een 39 21 2 48) 21 57 206 157 16) “51/36 4) 1114 81 3 11 42 | 16 1 43 iS € iW 49 ay fe G2 22 sh) Bil Ge Ze) 2 16 44 133) <4) ~2 44 | 11 (Sid 2 7 3. 41 «13 > oOo ns = uo for wo > Le) > oa —P 282. 4 heal uo oS = = = —M 121 98 75 98 62 53 81 72 8 444 43 6 4 i 7 ass 62 17 10 66 | 80 12 4 8 2 6 71 42 16 ; 2 73} 2 5 1 334 68 73 144 68 29 33 33 16 365 2 EAR OU SEO aa i) > NN uo > > w w > 80 7 2 |\25) gh 2 <3 4) 4). OG Aes Cy Ge allt 14 24 2; 2 5) 18 160 8 1 1. 3 5 Umer an 509 527 102 889 333 313 745 441 179 243 406 46 1501 13 68 24 32 55 215 8 51 7 102 145 238 86 894 239 24 BULLETIN 357 PLATE 1 1—2. Radimella confragosa (Edwards, 1944), X118. 1. LV, KE Locality PPP 678, Rio Banano Formation, USNM 490889. 2. RV, E Locality PPP 369, Escudo de Veraguas Formation, USNM 490890. 3—4. Radimella wantlandi Teeter, 1975, *118. 3. LV, E Locality PPP 638, Moin Formation, USNM 490891. 4. RV, E Locality PPP 638, Moin Formation, USNM 490892. OSTRACODES: BORNE ET AL 239 water deeper than 2000 m, and as fossil, 50-90% of North Atlantic Ocean Pliocene assemblages (Rodri- guez-Lazaro and Cronin, in press). Species of Krithe rarely inhabit shallow-water environments; the best known shallow-water species is K. praetexta (Sars, 1866), which lives in shelf and slope areas off northern Europe (Athersuch et al., 1989). A few sighted Krithe also live in shallow-water around Australia (Whatley et al., 1983). In the Gulf of Mexico and Caribbean Sea, Krithe is also usually found on the continental slope and deeper waters (Morkhoven, 1972), with the important exception of the Paria-Trinidad-Orinoco Shelf (see below and Fithian, 1980, App. 1). Most spe- cies of Bradleya are also most common in deep-sea environments (Benson, 1972; Whatley et al., 1983), with the exceptions of B. normani (Brady, 1866), which inhabits continental slopes and fjords of south- ern South American (Shuckstes, 1995; Whatley ef al., in press), and B. andamanae Benson, 1972, which lives in depths of 70—500 m in the northeastern Indian Ocean (Benson, 1972). The database of modern ostracodes from the Carib- bean and Gulf of Mexico allows us to identify modern environments in which the mixture of species found in Biofacies A occurs today. The only place identified in the Caribbean where Krithe and Bradleya occur to- gether with shallow-water taxa is the Paria-Trinidad- Orinoco Shelf off northern Venezuela (Fithian, 1980; Bold, 1978a), a region of modern tropical coastal up- welling water. A similar anomalous shallow-water colonization of Krithe and Bradleya occurs in the southern part of the Magellan Straits, where these and other normally psy- chrospheric, blind taxa occur together with the indig- enous shallow-water assemblage (Whatley ef al., in press). Whatley ef al. (in press) interpreted the occur- rence of these taxa in shallow-water as due to cold water temperatures and upwelling. A relatively shallow paleodepth for the Cangrejos Creek and lower part of the Lomas del Mar localities of the Moin Formation is suggested by the strong de- velopment of the eye tubercle in Echinocythereis mad- remastrae (Pl. 3, Fig. 2). It is well known that ostra- (= 5—6. Radimella ovata Bold, 1988, X118 5. LV, FE Locality PPP 68, Isla Solarte, USNM 490893. 6. RV, F Locality PPP 68, Isla Solarte, USNM 490894. 7-8. Radimella aff. R. ovata Bold, 1988, X118. 7. LV, FE Locality PPP 631, Moin Formation, USNM 490895. 8. RV, E Locality PPP 631, Moin Formation, USNM 490896. 9-10. Radimella ovata Bold, 1988 *109. codes living in deep water below the euphotic zone (< about 100-200 m) do not develop prominent eye tu- bercles as do most of those living in shallow-water (Benson, 1975; Howe and Bold, 1975). Moreover, Kontrovitz and Myers (1988) quantified the relation- ship between ostracode ocular structures and ambient downwelling sunlight in seawater and concluded that the biconvex eyespot-tapetum structure typical of po- docopid ostracodes like Echinocythereis would be of no use below depths of about 280 m in clear ocean water and 85 m in more turbid coastal water. The combination of ecological data for colder water conditions and morphological evidence for shallow- water paleoenvironments (<100—200 m), leads us to postulate that Biofacies A represents an outer shelf to upper slope environment, perhaps similar to that off northern Venezuela where Krithe and Bradleya live today. The upwelling of cool, nutrient-rich water that characterizes the Venezuela shelf, and elsewhere, ap- pear to be the typical oceanographic conditions that allow Krithe and Bradleya to migrate upslope into rel- atively shallow-water habitats. Benthic foraminiferal assemblages from the type Moin section support the interpretation of a deposi- tional environment at the continental shelf edge (150— 250 m of water depth), as they include a mixture of shelf edge and nearshore taxa (Cassell, 1986; Collins et al., 1995a). Collins et al. (1995a) interpreted the foraminiferal assemblages as evidence for downslope transport of nearshore taxa; Collins et al. (1995b) sug- gested benthic foraminifera assemblages were not rep- resentative of upwelling assemblages. Nonetheless, oceanographic parameters such as dissolved oxygen, nutrients, food, temperature, light, rather than simply abstract water depth, are the critical factors that influ- ence ostracode species’ ecology and their depth distri- butions. For example, oceanographic factors have been shown to be especially important for deep-sea and mid-depth taxa living along the continental slope, where bottom water temperatures in the thermocline limit the shallowest depth for many species (Dingle and Lord, 1990; Dingle et al., 1989; Rodriguez-Lazaro and Cronin, in press). Off southwest Africa (south of 9. LV, F Locality PPP 678, Rio Banano Formation, USNM 490897. 10. RV, F Locality PPP 678, Rio Banano Formation, USNM 490898. 240 3-4. 9. 10. BULLETIN 357 PLATE 2 Caudites medialis Coryell and Fields, 1937, *120. 1. LV, Locality PPP 668, Rio Banano Formation, USNM 490899. 2. RV Locality PPP 668, Rio Banano Formation, USNM 490900. Caudites aff. C. rectangularis (Brady, 1869), «120. 3. LV, Locality PPP 606, Rio Bartolo, USNM 490901. 4. RV, Locality PPP 768, Rio Banano Formation, USNM 490902. . Caudites cf. C. asymmetricus Bate et al., 1980, *120. 5. LV, Locality PPP 63, Cayo Agua, USNM 490903. 6. RV, Locality PPP 63, Cayo Agua, USNM 490904. Caudites medialis Coryell and Fields, 1937, 120. LV, Locality PPP 62, Cayo Agua Formation, USNM 490905. Caudites nipeensis Bold, 1946, * 120. LV, Locality PPP 369, Escudo de Veraguas Formation, USNM 490906. Caudites rectangularis (Brady, 1869), * 120. LV, Locality PPP 655, Moin Formation, USNM 490907. Pterygocythereis sp., 94. LV, Locality PPP 653, Moin Formation, USNM 490908. OSTRACODES: BORNE ET AL. 241 -1.000 DISSIMILARITIES 0.000 PPP 1400 PPP 1401 PPP 1398 PPP 1394 PPP 1397 PPP 1393 PPP 1395 PPP 1392 PPP 1399 PPP 1396 PPP 1370 PPP 1442 D PPP 1438 PPP 634 PPP 635 PPP 953 B PPP 954 PPP 955 PPP 1480 PPP 1481 PPP 636 PPP 1371 A PPP 655 PPP 649 PPP 653 PPP 651 PPP 650 PPP 658 PPP 959 PPP 656 PPP 652 PPP 654 PPP 647 PPP 1376 PPP 1433 C PPP 645 PPP 1357 PPP 710 PPP 960 PPP 638 PPP 1482 PPP 1375 PPP 1483 PPP 657 PPP 648 PPP 1369 E PPP 712 Text-figure 1—Q-mode dendrogram of cluster analysis, average linkage method, of Jaccard coefficients, based on 57 localities from the Moin and Rio Banano formations, Limon Basin, Costa Rica. Letters A—-E refer to ostracode biofacies that are characteristic of faunal assemblages and indicative of distinct environments of de- position. See text for further explanation. 28°S), Dingle (1992a, 1992b) found that shelf upwell- ing and intrusion of shelf currents, especially the cold, low-salinity, nutrient-rich Antarctic Intermediate Wa- ter, controls the distribution of ostracode species. Wha- tley (1991) also demonstrated the influence of low ox- ygen in slope environments for certain platycopid os- tracodes. In summary, we favor an “oceanographic” interpretation of the depositional environment for Bio- facies A of the Moin Formation—that the ostracodes indicate cool water (12—15° C) on the outer shelf/upper slope, perhaps due to upwelling of cold, deep nutrient- rich waters. Biofacies B Biofacies B is represented by Cluster B (Text-fig. 1) and is composed of five samples from the Lomas del Mar locality containing assemblages that are similar to those of Biofacies A in that they also contain signifi- cant numbers of Krithe spp. (7%), although in smaller numbers than for Biofacies A. Biofacies B is also dis- tinguished from Biofacies A by higher proportions of neritic ostracodes, including Pseudosammocythere sp. (average 32%), Munseyella bermudezi Bold, 1966 (21%), Radimella confragosa (5%) and Cytherella spp. (4%). The in situ coral lenses from which the microfossil samples were collected indicate a water depth of less than 40 m (A. E Budd, 1996, written communication). If the interpretation of the deposi- tional environment for Biofacies A is correct, then the presence of Krithe and other normally deeper-dwelling ostracodes in the Lomas del Mar reef sediments sug- gest upwelling of cooler waters occurred during the deposition of this unit. Biofacies C Biofacies C is represented by samples from Cluster C (Text-fig. 1) and consists of 13 samples that are primarily from Lomas del Mar and Avenida Barracuda outcrops. Biofacies C appears to be transitional be- tween the cooler upwelling assemblages of Biofacies A and the shallow (<30 m) carbonate platform assem- blages of Biofacies D (see below). The dominant taxa that clearly indicate a relatively shallow, primarily warm-water, tropical environment are the bairdiids (16%; i.e., Bairdoppilata, Neonesidea, Paranesidea), Xestoleberis spp. (10%), Jugosocythereis pannosa (Brady 1869) (5%), Loxocorniculum spp. (8%) and Radimella confragosa (11%). If the interpretation of the occurrence of typical deep-water taxa Krithe and Bradleya (14% and 9%, respectively, in this biofacies) presented above is correct, then there was at least pe- riodic upwelling of cooler deep-water during the de- position of Biofacies C, although there may be more time-averaging than occurred in Biofacies A. Biofacies D Cluster D (Text-fig. 1) consists of 15 samples, most of which are from a single outcrop (CTA Fence lo- cality, Rio Banano Formation), and represents Biofa- cies D. Four taxa dominate this assemblage: Radimella confragosa (18%), Loxocorniculum spp. (16%), Par- acytheridea tschoppi Bold, 1946 (20%), and Jugoso- cythereis pannosa (14%). Other important occurrences 242 BULLETIN 357 PLATE 3 1. Bradleya aff. B. acceptabilis Liibimova and Sanchez-Arango, 1974, x99. LV, FE, Locality PPP 657, Moin Formation, USNM 490909. 2. Echinocythereis madremastrae Bold, 1988, 90. LV, E Locality PPP 653, Moin Formation, USNM 490910. OSTRACODES: BORNE ET AL. 243 include Caudites nipeensis Bold, 1946, Orionina vaughani (Ulrich and Bassler, 1904), Gangamocyther- idea? plicata Bold, 1968, Quadracythere howei (Puri 1953), and Perissocytheridea subrugosa (Brady, 1870). This assemblage is found in the lower part of the CTA Fence section and signifies a shallow-water carbonate assemblage (water depth <10 m; Teeter, 1975) distinct from assemblages in Biofacies A—C. Biofacies E Only two samples cluster in this group (Cluster E) and represent Biofacies E (Text-fig. 1). They are dom- inated by Reussicythere reussi Teeter, 1975, described from the nearshore areas and main lagoon of Belize. This species accounts for an average of 74% of the total valve count in each sample. Also present are Pur- iana aff. P. matthewsi Teeter, 1975, (11% average be- tween the two samples), Cytherura sp. (5%) and Pel- lucistoma howei Coryell and Fields, 1937 (3%). The low diversity and the high dominance of only a few taxa suggest that this biofacies represents an assem- blage living in an environmentally restricted nearshore environment. Spacial and Temporal Relationships of Limon Area Samples The biofacies of the Moin Formation described above reflect spatially and/or temporally complex en- vironments of deposition. Analysis of these fossil os- tracode assemblages has provided insight as to the po- tential stratigraphic relationships among several strati- graphic sections of the Moin Formation and a more detailed understanding of the late Pliocene and early Pleistocene environmental history in the Limon Basin. If we assume the Bradleya- and Krithe-bearing finer- grained sediments (Biofacies A and B) stratigraphi- cally underlie the reef facies (Biofacies C) at the Lo- mas del Mar site (Text-fig. 3a), our results indicate a deep to shallow environmental change in the Moin Formation and/or a diminished upwelling towards the upper part of the section. An alternative interpretation (McNeill, written communication, 1995) is that the up- welling assemblages (Biofacies A) represent a tran- gression that postdates the deposition of the reef facies at the Lomas del Mar site. Such a scenario implies that these sediments were draped along the flanks of the Lomas del Mar reef and also in the low-lying Cangre- jos creek area (Text-fig. 3b). Given the absence of core data for this geographic area, clarification of the strati- graphic relationships for this portion of the Moin For- mation will depend on additional geologic mapping and refinement of the ages based on biostratigraphic and magnetostratigraphic work. The distinct clustering (Biofacies D) of many CTA Fence samples suggests that this section represents a reefal facies different from, and probably older than that at Lomas del Mar, an interpretation that is consis- tent with the field relationships of the two sections (Coates, this volume; Budd et al., this volume). Pre- vious geologic mapping has assigned the CTA Fence and Quebrada Chocolate sections to the Rio Banano Formation (Taylor, 1975; Cassell and Sen Gupta, 1989; Coates et al., 1992). McNeill et al. (in press) have now described the Quebrada Chocolate Formation, which includes these older trend outcrops. The Quebrada Chocolate section is located 2 km west of the CTA Fence section and contains ostracodes typical of the older Rio Banano Formation samples (Basslerites spp., Perissocytheridea spp., Puriana spp.), supporting the hypothesis that the Quebrada Chocolate section is part of a reefal trend older than the Lomas del Mar trend. Although this reefal unit has a similar carbonate plat- form ostracode assemblage to that of the Lomas del Mar reefal facies, it is distinguished from the younger 3. Jugosocythereis pannosa (Brady, 1869), * 108. LV, FE Locality PPP 710, Moin Formation, USNM 490911. 4. Quadracythere howei (Puri, 1953), * 162. LV, FE Locality PPP 710, Moin Formation, USNM 490912. 5. Hermanites hornibrooki Puri, 1960, * 162. LV, E, Locality PPP 710, Moin Formation, USNM 490913. 6. Cativella navis Coryell and Fields, 1937, 108. LV, FE Locality PPP 682, Rio Banano Formation, USNM 490914. 7. Actinocythereis gomillionensis (Howe and Ellis, 1935), * 108. LV, E, Locality PPP 673, Rio Banano Formation, USNM 490915. 8. Costa aff. C. bellipulex Levinson in LeRoy and Levinson, 1974, 108. LE FE Locality PPP 368, Escudo de Veraguas Formation, USNM 490916. 9. Costa variabilicostata recticostata Bold, 1970, * 108. LV, FE Locality PPP 631, Moin Formation, USNM 490917. 10. Krithe sp. RV, FE internal view, 135, Locality PPP 657, Moin Formation, USNM 490918. 244 BULLETIN 357 0.000 Distances 2.000 PPP 712 PPP 683 A PPP 682 PPP 59 PPP 61 PPP 389 PPP 688 PPP 62 PPP 670 PPP 672 PPP 685 PPP 300 PPP. 337, PPP 298 PPP 335 PPP 686 PPP 293 PPP 636 PPP 307 PPP 638 PPP 710 PPP 645 PPP 635 PPP 631 PPP 634 PPP 637 PPP 668 PPP 361 PPP 358 PPP 366 PPP 678 PPP 363 PPP 721 PPP 720 PPP 334 PPP 364 PPP 365 B PPP 360 PPP 362 PPP 718 PPP 175 PPP 653 PPP 655 PPP 657 PPP 648 PPP 374 PPP 372 PPP 60 PPP 63 PPP 687 Cc PPP 679 PPP 690 PPP 684 PPP 410 PPP 303 PPP 369 PPP 368 PPP 168 PPP 371 PPP 169 Text-figure 2a—Q-mode dendrogram of cluster analysis of Jac- card coefficients, complete linkage method, showing relationship of Limon Basin, Costa Rica, and Bocas del Toro Basin, Panama, os- tracode assemblages. A. Samples primarily from Rio Banano and Caya Agua formations. B. Samples from Moin and Escudo de Ver- aguas formations. C. Samples from Caya Agua, Valiente Peninsula, Escudo de Veraguas (southern coast), Moin and Rio Banano for- mations. 0.000 Distances 2.000 Macrocyprids Occultocythereis Bythoceratina Caribella Neonesidea Propontocyprids 1 Jugosocythereis Hermanites Paranesidea ? Puriana Costa Bairdoppilata Bradleya Echinocythereis Krithe 2 Munseyella Xestoleberis Pterygocythereis Phlyctocythere Loxoconcha Paradoxostoma 3 Cativella Cytheropteron Cytherelloidea Coquimba Hulingsina Lobosocytheropteron Radimella Orionina Paracytheridea 4 Kangarina Cytherura Caudites Pumilocytheridea Loxocorniculum Quadracythere Aurila Touroconcha Neocaudites Puriana Actinocythereis Cytherella 5 Basslerites Pellucistoma Text-figure 2b.—R-mode dendrogram showing relationship of taxa from the same cluster analysis as Text-figure 2a: 1, 4. Taxa representative of carbonate lagoon environments. 2. Outer shelf to upper slope taxa. 3. Taxa representative of lagoon environments, some of which prefer phytal habitats. 5. Taxa that prefer fine- grained, muddy substrates. Lomas del Mar reefal facies by its lack of deeper-water ostracodes. Efforts are underway to revise the geologic map of the geologically complex Limon area and to elucidate the stratigraphic relationships between the reef exposures and intervening clastic deposits (McNeill et al., 1996; McNeill et al., in press). Rio BANANO FORMATION Assemblages from exposures of the Rio Banano Formation south of the city of Limon contain a very distinct assemblage containing Basslerites spp., Cy- therella spp., Cativella navis Coryell and Fields, 1937, OSTRACODES: BORNE ET AL. 245 Text-figure 3.—Spatial and temporal relationship of selected sam- ples from Biofacies A—C (Text-fig. 1) in the Limon area. 3a. Relationship of ostracode Biofacies A—C as proposed herein. Biofacies A = deeper water ostracode assemblage (possibly up- welling related). Biofacies B = transitional assemblage of deeper water ostracodes and shallower carbonate-platform ostracodes. Bio- facies C = carbonate-platform ostracode assemblage with fewer deeper-water ostracodes. The grain-size generally coarsens upward from predominantly claystone in Biofacies A (with some intervals of coarser material) to a silty sandstone in Biofacies C. 3b. Alternative interpretation of relationship of ostracode Biofacies A-C (McNeill, written communication, 1995). Biofacies A postdates or is coeval to Biofacies B and C. Finer-grained sediments have been draped along flanks of Lomas del Mar reef trend. Loxocorniculum spp., Pellucistoma howei, Puriana spp., and Radimella ovata Bold, 1988. The presence of Actinocythereis gomillionensis (Howe and Ellis, 1935) is noteworthy because it is not found in other formations in the Limon area (with the exception of two samples of the Moin Formation) nor in the Bocas del Toro area. The Rio Banano Formation ostracode assemblages are lacking in typical carbonate platform taxa (e.g., Jugosocythereis, Hermanites, Radimella wantlandi Teeter, 1975, Quadracythere spp.). The en- vironment of deposition was probably inner to middle continental shelf, near a delta or estuary that was pro- viding clastic sediment. This interpretation is corrob- orated by the benthic foraminiferal assemblages of the Rio Banano Formation, which indicate deposition in 10—40 m of water in an open marine, inner to middle neritic environment (Cassell, 1986; Cassell and Sen Gupta, 1989; Collins et al., 1995), with a slight deep- ening higher in the section (Cassell, 1986; Cassell and Sen Gupta, 1989). It is possible that there were periods of non-marine deposition and/or depositional hiatuses within the Rio Banano section in areas lacking good exposures. OSTRACODES FROM BOCAS DEL TORO BASIN, PANAMA At present, ostracodes have been studied from only a few samples from the Nancy Point Formation (Ap- pendix A, Map 5) and they contained an outer shelf to upper slope assemblage including the following genera Argilloecia, Ambocythere, Cytherella, Krithe, and Munseyella. Cayo Agua Formation (App. A, Map 6) ostracode assemblages generally contain typical shallow marine, tropical taxa such as Cativella navis, Loxocorniculum spp., Costa variabilicostata recticos- tata Bold, 1970, Orionina vaughani group, Paracy- theridea tschoppi, Quadracythere howei, Radimella ovata, and Touroconcha lapidiscola Hartmann, 1959. Ostracode assemblages such as these are found in in- ner-middle shelf environments today. Ostracodes from the Escudo de Veraguas Formation (Appendix A, Map 4) are quite different from those in Cayo Agua in that there are rare to common occur- rences of the ostracode taxa Ambocythere, Argilloecia, Eucytherura, Kangarina, macrocyprids, paracyprids, Pseudosammocythere, Pterygocythereis, Radimella aff. R. ovata Bold, 1988, and Caudites nipeensis Bold, 1946. In general, these taxa suggest a deeper, outer shelf environment compared to those of Cayo Agua. The ostracode evidence generally supports the inter- pretation of the benthic foraminiferal assemblages from these sections (Collins, 1993; Collins er al., 1995). CORRELATION OF THE BOCAS DEL TORO AND LIMON BASIN In order to provide a comparison between the Bocas del Toro Basin and the Limon Basin ostracode assem- blages, we carried out Q- and R-mode cluster analyses using presence-absence data for 44 taxa in a small number of samples from each formation that contain representative assemblages. The Q-mode cluster anal- ysis (Text-fig. 2a) shows samples clustering together based on shared species and reveals the following groups: Group A consists of samples mainly from the Rio Banano and Cayo Agua formations and represents ostracode assemblages characteristic of the early—mid- dle Pliocene, roughly 2.4—5.0 Ma on the basis of planktonic foraminiferal and calcareous nannofossil biostratigraphy (Coates et al., 1992). Group B consists mainly of samples from the Moin and Escudo de Ver- aguas formations and represents an outer shelf envi- ronment of late Pliocene and perhaps earliest Pleisto- cene age, about 1.6—3.0 Ma. Group C is a mixture of 246 WwW n 6. BULLETIN 357 PLATE 4 . Orionina boldi Cronin and Schmidt, 1981, * 120 LV, E Locality PPP 710, Moin Formation, USNM 490919. . Orionina vaughani (Ulrich and Bassler, 1904), * 120. LV, F, Locality PPP 668, Rio Banano Formation, USNM 490920. Puriana sp. D, 120. LV, FE Locality PPP 671, Rio Banano Formation, USNM 490921. . Reussicythere reussi (Brady, 1869), *120 LV, FE Locality PPP 712, Moin Formation, USNM 490922. . Puriana ct. P. gatunensis (Coryell and Fields, 1937), * 150. LV, FE Locality PPP 59, Cayo Agua Formation, USNM 490923. Coquimba fissispinata (Benson and Coleman, 1963), * 120. LV, FE, Locality PPP 634, Moin Formation, USNM 490924. OSTRACODES: BORNE ET AL. Table 3.—Biostratigraphic distribution of ostracode taxa, Limon Basin, Costa Rica, and Bocas del Toro Basin, Panama. Ages from Coates et al. (1992), Collins et al. (1995) and Cotton (Chapter 2, this volume). Rio Banano Moin Nancy Point Escudo de Caya Agua Fm. Fm. Fm. Veraguas Fm. Fm. 2.4-2.5 to 1S =1e/1to 5.6 to 1.8-1.9 to 2.9 to Species 3.5—3.6 Ma 1.9 Ma 6.5 Ma 3.5-3.6 Ma 4.6-5.0 Ma Actinocythereis gomillionensis x xX Bradleya aff. B. acceptabilis x Xx xX Cativella navis x x Caudites rectangularis xX x x Caudites medialis xX x Caudites nipeensis xX x ».4 Coquimba fissispinata Xx x Costa cf. C. bellipulex x x Echinocythereis madremastrae Xx x Hermanites hornibrooki x Jugosocythereis pannosa xX x Krithe sp. x Costa sp. A x x xX Orionina boldi > 4 Orionina vaughani group xX x xX Puriana convoluta x Puriana gatunensis Xx X x Puriana minuta x x Puriana aff. P. matthewsi x Puriana sp. D x Quadracythere howei xX x Radimella confragosa xX xX Xx Radimella ovata Xx >. 4 Xx Radimella aff. R. ovata Xx x Radimella wantlandi x Reussicythere reussi xX samples from Cayo Agua, Valiente Peninsula, the southern part of Escudo de Veraguas, and part of the Rio Banano Formation. Some of these samples are considered to be early Pliocene in age (about 3.5—5.0 Ma), although there are older samples having similar biofacies. Within each of the main groups A-—C, there are subclusters in which samples from the same out- crop section show a high degree of similarity because they represent similar ostracode biofacies. The R-mode cluster analysis (Text-fig. 2b) grouped taxa that commonly occur together in the fossil assem- blages. Groups | and 4 represent two species assem- blages commonly found today inhabiting carbonate la- goon environments. Group 2 includes Bradleya and Krithe and represents the outer shelf/upper slope as- semblage discussed above (Biofacies A) that may sig- nify coastal upwelling. Group 3 is a lagoonal assem- blage consisting of taxa that may prefer phytal habi- tats. Group 5 consists of the taxa that dominate in the Rio Banano Formation and may signify an assemblage that prefers fine-grained, muddy substrates. REGIONAL OSTRACODE BIOSTRATIGRAPHY Bold (1983) summarized the stratigraphic ranges of marine and brackish water ostracodes in Cenozoic de- posits from the Caribbean region based on his many years of research on assemblages from dozens of lo- cations. More recent work on the Dominican Republic Neogene ostracode assemblages (Bold, 1988) provides additional important biostratigraphic and taxonomic data on Caribbean assemblages. Studies by Bold (1966, 1978b) showed that there are diverse ostracode co 7. Puriana cf. P. convoluta (Edwards, 1944), * 120. LV, — Locality PPP 631, Moin Formation, USNM 490925. 8. Puriana minuta Bold, 1963, * 120. LV, FE Locality PPP 686, Rio Banano Formation, USNM 490926. 9. Puriana aff. P. matthewsi Teeter, 1975, * 120. LV, F, Locality PPP 712, Moin Formation, USNM 490927. 248 BULLETIN 357 assemblages in the Limon area in the Moin and Rio Banano formations (note that Bold (1978b, Table 9) referred to the Rio Banano assemblages as ‘‘Gatun’’) and in Panama near the Panama Canal, but the detailed stratigraphic distributions of the species were not de- termined. Our work builds on Bold’s biostratigraphy as we establish the stratigraphic ranges of species within the last five million years in the western Carib- bean region of Central America. Table 3 lists age diagnostic ostracode species and their occurrences in the Moin and Rio Banano for- mations. By calibrating the first and last stratigraphic occurrences of each species to independent age data provided by nannofossils, planktonic foraminifers, and/or paleomagnetic stratigraphy, we determined that many species are useful age markers in the study area. These diagnostic species are illustrated in Plates 1—4. CONCLUSIONS The ostracodes collected from the Limon and the Bocas del Toro basins indicate that depositional envi- ronments included lagoon, carbonate platform, restrict- ed nearshore, and outer shelf to upper slope, and, in general, the assemblages from the two regions are sim- ilar in that both contain typical extant tropical taxa. One exception to this similarity is the presence in the Moin Formation of Bradleya and Krithe, two predom- inantly deep-sea genera, and an Echinocythereis with a well-developed eye tubercle. We interpret their abun- dance as signifying outer shelf to upper slope environ- ments influenced by the presence of cold, upwelling currents on the Central American continental shelf dur- ing the Late Pliocene and Early Pleistocene. This up- welling biofacies is sometimes associated with typical carbonate platform taxa, but it is not clear from the field relationships or biostratigraphy if it pre- or post- dates the main Lomas del Mar reefal facies. The Lo- mas del Mar reef is clearly younger than the CTA Fence and Quebrada Chocolate reefs. The distinctive Rio Banano Formation ostracode assemblages at the Bomba and Quiteria sections both indicate shallow marine, clastic environments and lack typical reef or upwelling taxa. 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CHAPTER 11 BATHYMETRIC DISTRIBUTION OF MIOCENE TO PLEISTOCENE CARIBBEAN TELEOSTEAN FISHES FROM THE COAST OF PANAMA AND COSTA RICA ORANGEL AGUILERA Centro de Investigaciones Arqueoldgicas, Antropologicas y Paleontologicas Universidad Francisco de Miranda Coro, Estado Falcén, Venezuela DIONE RODRIGUES DE AGUILERA Centro de Investigaciones Arqueoldégicas, Antropolégicas y Paleontologicas Universidad Francisco de Miranda Coro, Estado Falcon, Venezuela INTRODUCTION Otoliths of teleostean fishes and the teeth of sharks and rays are fairly abundant in Neogene sediments of tropical America (Nolf, 1976; Gillette, 1984; Nolf and Stringer, 1992; Schwarzhans, 1993; Nolf and Aguilera, 1998). Sediment samples processed as for large mi- crofossils commonly yield many tens to hundreds of specimens. Otoliths and teeth can be identified to ge- nus in nearly all cases, and often to species. Detailed analyses of selected genera and families, including comparisons to specimens of living species, will result in many more taxa being identified to species. Life habits can be inferred by comparisons to living species and otolith size can be used to estimate the size of the entire fish. Thus, the natural history and food chains of the entire assemblages can be reconstructed with some confidence. Comparison with recent taxa also al- lows reconstruction of the bathymetry of the deposits in which the otoliths occur, which is the major focus of this paper. Despite the great potential, there has been little sys- tematic collecting for otoliths. To this end, we joined the Panama Paleontology Project (PPP) expedition to Bocas del Toro, Panama, in 1995, and made survey collections of most of the formations of the Bocas del Toro Group (Coates et al., 1992; Coates, this volume). Subsequently, we made a preliminary survey of fishes represented in many PPP samples collected previously from the Limon Basin in Costa Rica and the Panama Canal Basin. Most of the collections examined yielded abundant, well-preserved, and diverse faunas of fossil fishes (Tables 1—5S). These collections already have yielded more specimens and taxa than all previous studies from the region. The abundant occurrence and diversity of fossil fishes through time will allow, for the first time, comparison of faunal change of fishes with that of the rich record of marine benthos from tropical America. Here, we present a preliminary analysis of the tele- ostean fish assemblages identified from the Caribbean coast of Panama and Costa Rica, along with their prob- able bathymetric distributions inferred by comparison to the bathymetric ranges of living representatives of each genus. All but two of the fossil genera are alive today in the region. We also briefly consider the oc- currence of recent faunas from Eastern Pacific sedi- ments to evaluate the potential problems of post-mor- tem mixing of faunas from different depths. Finally, we compare our estimates of water depths to those based on benthic foraminifera (Collins, 1993; Collins et al., 1995, 1996; Collins in Jackson ef al., this volume). ACKNOWLEDGMENTS We especially thank Anthony Coates and Jeremy Jackson for inviting us to join the PPP to work on the ichthyofauna. This study would not have been possible without their encouragement and support. We are also grateful to Laurel Collins, Helena Fortunato and Dirk Nolf for their assistance. This work was supported by the National Geographic Society, the Walcott and Scholarly Studies funds of the Smithsonian Institution, the Smithsonian Tropical Research Institute, and the Francisco de Miranda University, Venezuela. MATERIAL AND METHODS SAMPLING Otoliths were obtained individually from surface ex- posures in the field and from bulk samples. Otoliths BULLETIN 357 N N BERT (0990D snooAndpy ‘Jopul ‘ues ‘oepruehny] ZOBI “MED snyisidoyouoT S681 “IED wmipiydodaT I O€ST FIAND snuaso7 I7 I EER ‘Souusiougea sniuojovT E981 “MED snyssidosy 6£61 “UNOg wnydos«y I 678 JaIAND uojniuanyy ‘Jopul ‘usd ‘oeprpnuoRy Japul ‘uad ‘aepuqoy ‘Japul “uas ‘aRplowiayH ZO6| Aasuatnog xtraqoudkyday (OPS ‘SouustougyeA) suapiuay 8P8I 9MOT DI]aPVD SS8I ‘PIBIID 2 plleg snwojsouiong 9 € I 14 LI SI8I ‘anbsouyey smjanbq ‘JOpul ‘uas ‘oepljneisuq I QIS] TaIAND synvsisuq LEQ] ‘uosdwioyy voporysq S SSRI ‘Yooiqjoy wnajoajdiq _ lunsoulcyjyououauiq,, EVR ‘UeIg 2 AaPOOH auajoszig “Jo I I 61 0681 ‘UuRWUAsIq 27 UURUIUASIq snydrig TIBI “MID uo1ssoudD I PTS ‘UkIg 2 I9[MOy DUaDIISOUa]D “Jd ‘Jopul ‘ues ‘oepuisu0D I 608 ‘BWOID snyouiys0ja0D soumoyiodn[a TIBI TOYA sAyyooyuy 898 JayUNDH sijnpssuaiaD ZOR ‘apedase_T snwodosjuad OI8I ‘anbsouyey sndpivD I 6781 ASAND VjNIOLg 98 Or 174 6L 6L II € Org ‘uosdwuoyy sosaonusaig I ‘JOpul “uas ‘oeplpog QL8I ‘auysepulaig sdoiquiag “Jo 1681 “oopy vadnjo io iO Mesopelag ion io iO Bathyal and/or Otophidium indefatigable Paraconger californiensis Paralonchurus dumeniii Pacific Coast of Panama (Depth range: 41 - 102 m BULLETIN 357 Ponchthys margantatus IRhynchoconger nitens Serranus psittacinus Stellifer illecebrosus Syacium ovale 91% 15% 12% Text-figure 1.—Occurrence of otoliths from one dredge sample from the Gulf of Panama (Locality: GC-97-20, depth: 102 m, 41 otoliths) and 8 dredge samples from the Gulf of Chiriqui (GC-97-72, 41 m, 19 otoliths; GC-97-80, 64 m, 132 otoliths; GC-97-10, 65 m, 5 otoliths; GC-97-95, 65 m, 4 otoliths; GC-97-91, 65 m, 19 otoliths; CG-97-57, 65 m, 1 otolith; GC-97-97, 87 m, 67 otoliths; CG-97-79, 87 m, 22 otoliths) along the Pacific coast of Panama. Solid lines indicate the presence of the genera at the depth intervals and dashed lines indicate the nightly occurrence of mesopelagic genera near the surface. The number of genera present in any 100-m or 50-m depth interval is expressed as a percentage of the total number of taxa collected in the sample. analyzed here come from 93 collections, including 18 collections from the Panama Canal Basin (Table 1), 58 collections from the Bocas del Toro Basin (Tables 2— 4), and 17 collections from the Limon Basin (Table 5). Bulk samples were washed using 2-mm and 500-ym sieves. Otoliths from both types of collections were combined for presentation in Table 1. The data of Ta- bles 1-5 are available at the PPP internet site http:// www.fiu.edu/~collinsl/. TERMINOLOGY AND TAXONOMY Terminology and classification follow Nolf’s (1985) review of otolith anatomy, morphology, variability, on- togeny and preservation. Otoliths consist of calcium carbonate, mainly aragonite, and organic matter called otoline. Otoliths are the integral, specialized hard part of the actinopterygian and sarcopterygian acoustico- lateralis system, situated in the membranous laby- rinths. Each labyrinth is located on either side of the brain in the otic capsules of the neurocranium. Within each membranous labyrinth is a different otolith in the utriculus, sacculus and lagena. The term otolith as used in this study refers to saccular otoliths, except for cat- fish that are represented by utricular otoliths. Otoliths can be identified unambiguously only when they are well preserved and common enough for com- parative observations (Plate 1). A complete inventory of the faunas was not possible in this preliminary sur- vey pending more material. In general, we have been able to make generic identifications with high confi- dence, but species identifications will require more de- tailed study and use of extensive reference collections of the otoliths of living species. Some taxa, such as Diaphus, present particular problems in identification (see discussions in Nolf and Steurbaut, 1987; Nolf and Capetta, 1989; Nolf and Stringer, 1992). In other cases, such as the approximately 113 Recent species of trop- ical western Atlantic gobiids (Richards, 1990), the tax- onomy is not sufficiently resolved to allow reliable identifications because of their highly variable otolith morphologies. Furthermore, in taxa such as the sciaen- ids, juvenile otoliths do not exhibit diagnostic features. TAPHONOMY Most otoliths probably enter the sediments through the excreta of predators (Nolf, 1985; Nolf and Brzo- bohaty, 1992). Large quantities of teleostean remains have been found in the stomachs of numerous large marine predators including cetaceans (Fitch and Brow- nell, 1968), sharks (Cortes and Gruber, 1990; Ebert er al., 1992; Hazin et al., 1994; Ellis et al., 1996; Cortes et al., 1996) and batoids (Hess, 1961; Gilliam and Sul- livan, 1993; Ellis et al., 1996). For example, the stom- achs of 17 cetaceans, comprising seven species, con- tained 18,164 otoliths of fishes, of which more than 89 percent were Myctophidae (lanternfish). However, observations of the occurrence of otoliths in Recent sediments are limited. Most otoliths along the east coast of North America occur on the continental slope (400—2,000 m), whereas otoliths are much less abun- dant on the continental shelf (<100 m), possibly be- cause fluctuations in sea level greatly reduce the time TELEOSTEAN FISHES: AGUILERA AND AGUILERA NO Oo eS) Dorsal Margin Dorsal Depression Antirostrum Excisura Sa Rostrum Ostium Ventral Depression Crista Superior Posterior Margin Cauda Crista Inferior 1mm Ventral Margin PLATE | Principal morphological features of the mesial surface of a right otolith (Epigonus denticulatus, Dieuzeide, 1950), PPP 5057, from the Early Pliocene Cubagua Formation, Venezuela. for otoliths to accumulate at shallower depths (Elder et al., 1996). These problems raise questions about how well the occurrence of otoliths in sediments reflects their dis- tributions in life. To begin to address this question, we examined otoliths from nine sediment samples ob- tained by dredging in depths of 41—102 m in the Gulfs of Panama and Chiriqui along the Pacific coast of Pan- ama (Text-fig. 1). All of the genera obtained have rep- resentatives among the fossils collected from the Li- mon, Bocas del Toro, and Panama Canal basins. Oto- liths were identified based on examination of speci- Water Depth (m) Paralonchurus ----------4Dijaphus Neritic or Epipelagic Bathyal and/or Mesopelagic Middle to Late Miocene Gatun Formation (Panama Canal Basin, Panama mens of living species from the area. Data on depth of occurrence and life habits of living fishes (Allen and Robertson, 1994) were compared to the depth of collection and bottom conditions at the sites of collec- tion of the dredge samples (H. Fortunato, oral com- munication, 1998). ESTIMATES OF PALEOBATHYMETRY We used all the samples from the same formation to make one estimate of paleobathymetry, using the method of Nolf and Brzobohaty (1992). The method is based on the assumption that the taxa encountered 97% Text-figure 2—Present-day bathymetric ranges of taxa represented in the Gatun Formation. Solid and dashed lines drawn as in Text-figure 1. 264 BULLETIN 357 lived together in the same environment represented by the sedimentary facies sampled. The assumption ap- pears reasonable, because of the general lithologic uni- formity within the formations sampled and the agree- ment with estimates of paleobathymetries based on benthic foraminifera (Collins, 1993; Collins ef al., 1995, 1996, this volume). All the identifiable otoliths from the samples were identified to genus and depth ranges were assigned to each taxon based on the known depths of living counterparts. We did not at- tempt to estimate water depths for assemblages of less than 15—20 species from any formation. The number of genera in each 100-m depth interval was converted to the percentage of the total genera in the formation. Occasionally it was possible to subdi- vide the first 100-m interval into two based on the occurrence of common, shallow-water taxa. The depth profile of the formation is thus characterized by vary- ing percentages of taxa per depth interval. These per- centages usually peak at a single 100-m depth interval that is then taken as the most likely paleodepth of the formation. However, sometimes two peaks may occur, possibly due to reworking of sediments. In such cases, an estimation of paleodepth is unreliable. THE NEOGENE FAUNA We obtained 7,770 otoliths distributed among 81 taxa, 70 identified to genera, or about ten percent of the 773 western central Atlantic genera alive today (Richards, 1990). The comparatively low fossil diver- sity is due to several factors, including: (1) small num- bers of samples, (2) differential preservation among facies, (3) numerous broken or abraded specimens, (4) presence of juvenile forms without diagnostic features, and (5) abundance of families whose otoliths are un- described or poorly known. The first three problems can be readily addressed by more extensive sampling, but the rest require much more work on the otolith morphology of Recent species. The fauna has strong similarities to the Neogene tel- eostean faunas of the Dominican Republic (Nolf and Stringer, 1992), Trinidad (Nolf, 1976), Jamaica (Stringer, 1998) and Venezuela (Nolf and Aguilera, 1998). The most common taxa are arlids, clupeids, myctophids, gerreids, pomadasids, sciaenids, gobiids and ophiids. Most are neritic with a few mesopelagic or demersal taxa from the upper slope. The presence of Lactarius is of special interest be- cause the one Recent species of the genus, L. lactarius Schneider, is known only from coastal waters of south- ern Asia (Nolf and Bajpai, 1992). Fossil lactariids are known from the Middle Eocene of Barbados, the Pa- leogene of the Gulf of Mexico and Europe, the Aqu- itaine Basin of France, and the Miocene of Portugal and the Dominican Republic (Steurbaut, 1984; Nolf and Stringer, 1992). Lactarius appears to be a Tethyan relict that survived in the southwest Caribbean until at least the Late Pliocene (Rio Banano Formation, Table 5). The Indo-Pacific genus Plotosus is another Tethyan relic that occurs in the Neogene of the Dominican Re- public (Nolf and Stringer, 1992) and Panama (Table 3): TEST OF THE PALEOBATYHMETRY METHOD FOR RECENT FAUNAS Nine dredge samples that were collected between 41-102 m from the Pacific coast of Panama were an- alyzed. They yielded 310 identifiable otoliths of 33 genera (Text-fig. 1). All but five (85%) live exclusively in depths of 100 m or less, over sandy or muddy bot- toms, and 91% occur in depths of 0-50 m (Allen and Robertson, 1994). Most are benthic (e.g., gilbert floun- der Citharichthys gilberti, conger eel Paraconger cal- iforniensis, spinesnout brotula Lepophidium prorates, longtailed jawfish Lonchopisthus sinuscalifornicus) or live in the lower part of the water column above the bottom (e.g., croaker Micropogonias ectenes, serrated grunt Conodon serrifer, banded serrano Serranus psit- tacinus). Five of the taxa are bathyal or mesopelagic. However, only two of these (Neobythites stelliferoides and Coelorhincus aff. C. scaphopsis) are restricted to such depths and therefore were probably transported upwards by predators. Thus, the otolith fauna matches very well the life habits of the species collected. ESTIMATES OF PALEOBATHYMETRY OF NEOGENE FAUNAS PANAMA CANAL BASIN Gatun Formation (12 collections, 874 otoliths) The 34 taxa from the Middle to Late Miocene Gatun Formation are the second most diverse assemblage from this preliminary survey (Text-fig. 2; Table 1). The fauna includes several neritic sciaenids such as the jacknifefish, Equetus, the weakfish, Cynoscion, the shorthead drum, Larimus, the croaker, Ophioscion, and the barbel drum, cf. Ctenosciaena. Also present is the Tethyan relic false trevallies fish, Lactarius. These taxa, together with the anchovy, Anchoa, the ancho- veta, Cetengraulis, the anchovy, Engraulis, the jaw- fish, Lonchopisthus, and the mullet, Mugil, are all mainly shallow-water fishes that live at depths less than 25 m (Text-fig. 2). Living representatives of 94% of the fauna occur in depths less than 50 m, whereas only one third range into bathyal depths. Our paleo- bathymetric estimate of 0—SO m is strongly supported by assemblages of benthic foraminifera that suggest depths of 20—40 m (Collins et al., 1996). TELEOSTEAN FISHES: AGUILERA AND AGUILERA 265 Late Miocene Chagres Formation, Rio Indio facies (Panama Canal Basin) E a 3 a 3 ® < A . $32 5) 7 q o g & = oo”g Ee ' oD ‘© &| 100 H 84% 2 o ' 5 2 1 os H 2 iG} 200 fe 5 g | 300 2s © $1} 400 =) £3 oO & =| 500 Text-figure 3.—Present-day bathymetric ranges of taxa represented in the Chagres Formation. Solid and dashed lines as in Text-figure 1. Late Miocene Nancy Point Formation (Bocas del Toro Basin, Panama) Water Depth (m) Parascombrops Steindachneria Trachurus 75% Epipelagic ise Be Nenitic or B Bathyal and/or Mesopelagic Text-figure 4—Present-day bathymetric ranges of taxa represented in the Nancy Point Formation. Solid and dashed lines drawn as in Text- figure 1. Early Pliocene Shark Hole Point Formation (Bocas del Toro Basin, Panama) Water Depth (m) Parascombrops Steindachneria --------4+Diaphus me 400 o8 ar 52 Zo} 200 60% 300 70% 400 | 70% 500 Bathyal and/or Mesopelagic Text-figure 5.—Present-day bathymetric ranges of taxa represented in the Shark Hole Point Formation. Solid and dashed lines drawn as in Text-figure 1. BULLETIN 357 266 oe = oO Cc iv] a £ 7) is.) a ° — oO = oD no] 8 {eo} 2 i 2 s oO E = fe} Ww oO D < ° > i) Oo o c a o Ss a aI = oO WwW (w) udag sa}eM\ oiGejadida JO ONUBN oiGejadosayy soypue jeAeg Text-figure 6.—Present-day bathymetric ranges of taxa represented in the Cayo Agua Formation. Solid and dashed lines drawn as in Text- figure 1. Late Pliocene Escudo de Veraguas Formation (Bocas del Toro Basin, Panama) euauyrepula}s| (w) wdeg 138328 oi6ejadid3 JO DUAN aibejadosew, jo/pue jeAujeg Text-figure 7.—Present-day bathymetric ranges of taxa represented in the Escudo de Veraguas Formation. Solid and dashed lines drawn as in Text-figure 1. Pleistocene Swan Cay Formation (Bocas del Toro Basin, Panama) sninydwiAs wniaefs} aibejadidg = aiBejadosay JOONUeN Jo/pue jeAujeg Text-figure 8.—Present-day bathymetric ranges of taxa represented in the Swan Cay Formation. Solid and dashed lines drawn as in Text- gure 1. fi TELEOSTEAN FISHES: AGUILERA AND AGUILERA Water Depth (m) Eucinostomus Odontoscion Paralonchurus IRhynchoconger -------4Diaphus Neritic or — Bathyal and/or Mesopelagic Late Pliocene Rio Banano Formation (southern Limon Basin, Costa Rica) 267 Text-figure 9.—Present-day bathymetric ranges of taxa represented in the Rio Banano Formation. Solid and dashed lines drawn as in Text- figure 1. Chagres Formation (6 collections, 1,908 otoliths) The 20 taxa from the Rio Indio facies of the Late Miocene Chagres Formation include a mixture of ne- ritic and bathyal forms, but 80% have living represen- tatives that inhabit depths less than 100 m versus 45— 60% for deeper water (Text-fig. 3; Table 1). The sug- gested paleobathymetric range of 0 to 100 m compares favorably with that based on benthic foraminifera (Collins et al., 1996). BOCAS DEL TORO BASIN Nancy Point Formation (13 collections, 178 otoliths) The Late Miocene Nancy Point Formation yielded only 12 taxa, with no clear indication of paleobathy- metry (Text-fig. 4; Table 2). The estimate of 300-500 m based on foraminifera (Collins, 1993) is consistent with the limited information on otoliths. Water Depth (m) Parascombrops 3] i 2 es Bathyal and/or Mesopelagic Steindachnena Shark Hole Point Formation (8 collections, 51 otoliths) The Early Pliocene Shark Hole Point Formation contained only nine taxa of mixed environmental af- finities (Text-fig. 5; Table 2). The mesopelagic lantern- fish, Diaphus, the codletsfish, Bregmaceros, the cusk eel, Lepophidium, and Steindachneria range widely in depth. Likewise, the conger eel, Ariosoma, includes several neritic Recent species, but the Caribbean A. selenops inhabits the continental slope down to 550 m (Nolf and Brzobohaty, 1992). The only exclusively shallow-water fishes are the estuarine catfish, Geni- dens, and Lactarius. The estimate of 100—200 m based on benthic foraminifera (Collins et al., 1995) is con- sistent with the limited data for otoliths. Cayo Agua Formation (13 collections, 634 otoliths) The 37 taxa from the Early to Middle Pliocene Cayo Agua Formation comprise the most diverse teleostean Late Pliocene to Middle Pleistocene Moin Formation (southern Limon Basin, Costa Rica) 76% 65% Text-figure 10.—Present-day bathymetric ranges of taxa represented in the Moin Formation. Solid and dashed lines drawn as in Text-figure 1. 268 BULLETIN 357 assemblage obtained in this preliminary survey (Text- fig. 6; Table 3). The fauna contains diverse estuarine taxa such as the bonefish, Albula, the snook, Centro- pomus, the mojarra, Eucinostomus, and Larimus, all suggesting a water depth of less than 50 m. Also pres- ent is the catfish Plotosus, which is another Paleogene, western Tethyan relict that lives today in principally neritic, estuarine, or even fresh waters of the Indo- Pacific and Australian realms (Nolf and Stringer, 1992). Taxa whose living representatives occur in depths less than 50 m comprise 84% of the fauna, whereas those found in bathyal depths are less than half to one third the taxa. Our estimate of 0-100 m compares favorably with that of 20-80 m based on benthic foraminifera (Collins, 1993). Escudo de Veraguas Formation (20 collections, 920 otoliths) The 27 taxa from the Late Pliocene Escudo de Ver- aguas Formation comprise a mixed neritic and meso- pelagic ichthyofauna with no obvious peak (Text-fig. 7; Table 4). The general decline in diversity of taxa whose living representatives are known from deeper waters suggests a paleobathymetry of somewhere be- tween 0-300 m. This is compatible with the estimate of 100-150 m based upon benthic foraminifera (Col- lins et al., 1995). Swan Cay Formation (4 collections, 2,289 otoliths) The Early Pleistocene Swan Cay Formation contains an ecologically diverse assemblage of 30 taxa of pre- dominantly shallow-water affinities (Text-fig. 8; Table 3). The estimated depth of 0-100 m is similar to that of 80-120 m based on benthic foraminifera (Collins in Jackson et al., this volume). LIMON BASIN Uscari Formation (3 collections, 8 otoliths) The Early to Middle Miocene Uscari Formation (up- permost part) produced the lowest abundance and di- versity, yielding only eight otoliths of the mesopelagic lanternfish, Diaphus, that exhibits a wide depth range. The locality sampled (Appendix B, this volume, Sec- tion 27) is about 6.0—5.0 Ma (Coates er al., 1992) with a depositional depth of 300-500 m, based on benthic foraminifera (Collins et al., 1995). Rio Banano Formation (7 collections, 77 otoliths) Living representatives of most of the 19 taxa found in the Late Pliocene Rio Banano Formation inhabit depths less than 50 m (Text-fig. 9; Table 5). The sciaenid, Stellifer, lives primarily near the shoreline. The croaker, Umbrina, Ariosoma, Lepophidium, the Atlantic midshipman, Porichthys, and the conger fish, Rhynchoconger, range today from shallow waters to depths of 300 to 500 m. The only oceanic taxon is Diaphus, which today occurs mainly between 200-— 1,000 m. However, individuals migrate each night to the surface where they may be eaten by epipelagic predators which could excrete Diaphus otoliths in the neritic environment (Nolf, 1985; Nolf and Brzobohaty, 1992). Ninety-five percent of the taxa live today in depths of 0-50 m, which agrees very well with the estimate of 20-40 m based on benthic foraminifera (Collins et al., 1995). Moin Formation (7 collections, 762 otoliths) Half of the 17 taxa from the Late Pliocene to earliest Pleistocene Lomas del Mar Member of the Moin For- mation range across neritic to bathyal depths with no clear peak in diversity (Text-fig. 10; Table 5). Only Neobythites and Diaphus are oceanic. The estimated paleobathymetry of 50-100 m based on benthic fora- minifera (Collins et al., 1995) and 40-73 m based on ahermatypic corals (Cairns, this volume) are consistent with the otoliths but appear somewhat deep for the abundant and diverse reef coral assemblage that occurs in place in the same deposit (Coates ef al., 1992; Budd et al., this volume). CONCLUSIONS 1. Preliminary collections of a Neogene ichthyofauna from three Caribbean sedimentary basins of Pana- ma and Costa Rica yielded identifiable otoliths of 70 genera. Most of these are alive today, which permits estimates of paleobathymetry based on the otolith assemblages. 2. Estimates of paleobathymetry based on diverse as- semblages of otoliths were in good agreement with those based upon benthic foraminifera. 3. Reworking of otoliths by physical or biological transport apparently does not badly obscure original patterns of depth distribution. 4. The great abundance and distribution of Neogene otoliths from these sediments constitute a rich re- source for future investigations of paleoecology and systematics. TELEOSTEAN FISHES: AGUILERA AND AGUILERA 269 REFERENCES CITED Allen, G.R., and Robertson, D.R. 1994. Fishes of the tropical eastern Pacific. University of Hawaii Press, Honolulu, pp. 1—332. Coates, A., Jackson, J.B., Collins, L.S., Cronin, T.M., Dowsett, H.J., Bybell, L.M., Jung, P., and Obando, J.A. 1992. Closure of the Isthmus of Panama: The near-shore marine record of Costa Rica and Western Panama. Geological Society of America Bulletin, vol. 104, pp. 814-828. Collins, L.S. 1993. Neogene paleoenvironments of the Bocas del Toro Basin, Panama. Journal of Paleontology, vol. 67, no. 5, pp. 699— 710. Collins, L.S., Coates, A., Jackson, J.B., and Obando, J.A. 1995. Timing and rates of emergence of the Limon and Bocas del Toro basins: Caribbean effects of Cocos Ridges sub- duction? Geological Society of America Special Paper, no. 295, pp. 263-289. Collins, L.S., Coates, A.G., Berggren, W.A., Aubry, M-P., and Zhang, J. 1996. The late Miocene Panama isthmian strait. Geology, vol. 24, no. 8, pp. 687-690. Cortes, E., and Gruber, S.H. 1990. Diet, feeding habits and estimates of daily ration of young lemon sharks, Negaprion brevirostris (Poey). Copeia, 1990, no. 1, pp. 204-218. Cortes, E., Manire, C.A., and Heuter, R. 1996. Diet, feeding habits and diet feeding chronology of the bonnethead shark, Sphyrna tiburo, in southwest Florida. Bulletin of Marine Science, vol. 58, no. 2, pp. 353-367. Ebert, D.A., Compagno, L.J.V., and Cowley, P.D. 1992. A Preliminary investigation of the feeding ecology of squaloid sharks of the West coast of Southern Africa. South African Journal of Marine Science, vol. 12, pp. 601—609. Elder, K.L., Jones, G.A., and Bolz, G. 1996. Distribution of otoliths in surficial sediments of the U.S. Atlantic continental shelf and slope and potential for re- constructing Holocene fish stocks. Paleoceanography, vol. 11, no. 3, pp. 359-367. Ellis, J.R., Pawson, M.G., and Shackley, S.E. 1996. The comparative feeding ecology of six species of shark and four species of ray (elasmobranchii) in the North-East Atlantic. Journal of the Marine Biological Association of the United Kingdom, vol. 76, pp. 89-106. Fitch, J.E., and Brownell, R.L. 1968. Fish otoliths in cetacean stomachs and their importance in interpreting feeding habits. Journal of the Fishery Re- search Board of Canada, vol. 25, no. 12, pp. 2561-2574. Gillette, D.D. 1984. A marine ichthyofauna from the Miocene of Panama and the Tertiary Caribbean faunal province. Journal of Ver- tebrate Paleontology, vol. 4, no. 2, pp. 172-186. Gilliam, D., and Sullivan, K.M. 1993. Diet and feeding habits of the southern stingray Dasyatis americana in the central Bahamas. Bulletin of Marine Science, vol. 52, no. 3, pp. 1007—1013. Hazin, F., Lessa, R., and Chammas, M. 1994. First observations on stomach contents of the blue shark, Prionace glauca, from Southwestern Equatorial Atlantic. Revista Brasileira de Biologia, vol. 54, no. 2, pp. 195— 198. Hess, P.W. 1961. Food habits of two Dasyatid rays in Delawere Bay. Cop- eia, 1961, no. 2, pp. 239-241. Nolf, D. 1976. Les otolithes de Téléostéens Néogénes de Trinidad. Eclo- gae Geologicae Helvetiae, vol. 69, no. 3, pp. 703-742. 1985. Otolithi Piscium. in Handbook of Paleoichthyology. H.-P. Schultze, ed., Gustav Fischer Verlag, Stuttgart, vol. 10, 145 pp. Nolf, D., and Aguilera, O. 1998. Fish otoliths from the Cantaure Formation (Early Miocene of Venezuela). Bulletin de |'Institut Royal des Sciences Naturelles de Belgique, Sciences de la Terre, vol. 68, pp. 237-262. Nolf, D., and Bajpai, S. 1992. Marine middle Eocene fish otoliths from India and Java. Bulletin de I’ Institut Royal des Science Naturelles de Bel- gique, Sciences de la Terre, vol. 62, pp. 195-221. Nolf, D., and Brzobohaty, R. 1992. Fish otoliths as paleobathymeytric indicators. Paleonto- logia 1 Evolucion, no. 24—25 (1992), pp. 255-264. Nolf, D., and Cappetta, H. 1989. Otolithes de poissons du Pliocéne du Sud-Est de la France. Bulletin de |’Institut Royal des Sciences Naturel- les de Belgique, Sciences de la Terre, vol. 58 (1988), pp. 209-271. Nolf, D., and Steurbaut, E. 1987. Description de la premiére faune ichthyologique exclusi- vement bathyale du Tertiaire d'Europe: otolithes de l’Oligocéne Inférieur du gisement de Pizzocorno, Italie septentrionale. Bulletin de I'Institut Royal des Science Naturelles de Belgique, Sciences de la Terre, vol. 57, pp. 217-230. Nolf, D., and Stringer, G.L. 1992. Neogene paleontology in the northern Dominican Repub- lic, 14. Otoliths of teleostean fishes. Bulletins of Ameri- can Paleontology, vol. 102, no. 340, pp. 41-81. Richards, W.J. 1990. List of the fishes of the western central Atlantic and the status of early life stage information. NOAA Technical Memorandum NMFS-SEFC-267, pp. 1-87. Schwarzhans, W. 1993. A comparative morphological treatise of recent and fossil otoliths of the family Sciaenidae (Perciformes). in Pis- cium Catalogus: Otolithi Piscium, H.P. Friedrich, ed., vol.1, 245 pp. Steurbaut, E. 1984. Les otoliths de Téléostéens de 1’Oligo-Miocéne d’ Aquitaine (Sud-Ouest de la France). Paleontographica, ser. A, vol. 186, no.1—6, pp. 1-162. Stringer, G.L. 1998. Otoliths-based fishes from the Bowden Shell Bed (Plio- cene) of Jamaica: Systematics and Palaeoecology. Con- tribution of Tertiary and Quaternary Geology, vol. 35, no. 1—4, pp. 147-160. 5 j Y A v ah fi | , ‘ “a ye i? a i . j 2 mal of wd = ' ] : ~ : ¥ ait di ' at ’ # i a ‘ — "Row eal) a oa Dutt H “ Hie De A yas a . ~ =) 7 ag Tie bef b die co a st Pile w ty - 4 e 4 ihe a : Disney Wee pall) pace a Te eee s,fr ae te oy, wee ae hagftaide’ Gh. GAD. 0 ch Bre! cose Bid a.” ours a Sha vin = i ihe fi — } cimmabeey buy 4) Oar] we @ 3 leighese= patialg rest (Ow Wii —« 2 tar’ “e Pee ae ee ip alt, sal belt Mig ioe a) Oi - jenipaiia wv! lth! quads ately’ ot ee Pee Bc stile hist agua! Sn i aw are. ¥ shit eh ) dos oO =i od ee Si A be re 6 ae ea cpm nerlil on . a Pea le ai commana a od anak Y te port sae " Par Aare: tw i tis ayes ai Se tel tnd @ ol - a owes) zat fay! ven 27" pan ee vhs } ; \) atebttal ie aes ‘ a aa" : Viaywy 447m, Lae } | jad hai ama ge A ye (at > hak all aad i 1 (dg 4 j stents Hy a real nf Luis” Ale pi peel De iia ian |]! (ore he > { > i wi ‘Fe oo wil ST so I 4 bbe gf ee i N { ii i ON . rer i bint! © haa | tee Ge @ ; tea grre eee oben 4 hy eer =Tips i) 7 , \ | Ot ; ’ een | a Z 7 " yi iti? M seege tt as ee al ‘ era nnd se “e ‘wey uni’ saveue!| Semeol hl ap a \ tags fee ist eal eat, 1 Hh Lit iy ! i ft Ne OS tee be ~ q hans (GRY qidital Oe wel - Pes. {; cwEe 4 - oe > Se coronal eee « Vrind hen += hin, Chinese Clin OG = Ce gedaan ere ty eel ea r bv! eee lt Toes 1. Wir Sena IDs 21 CO) © ai” oie ae CHAPTER 12 A DATA MODEL FOR THE PANAMA PALEONTOLOGY PROJECT KARL W. KAUFMANN Smithsonian Institution Smithsonian Tropical Research Institute Washington, D.C. 20560-0580, U.S.A. INTRODUCTION The goal of the Panama Paleontology Project (PPP) is to describe the geological and biological events leading up to the complete emergence, approximately 3 million years ago, of the land bridge connecting North and South America. Field work in support of the project has required numerous collecting trips to remote areas in Panama, Costa Rica, Nicaragua, Ec- uador and Venezuela since 1986. These trips have re- sulted in the collection of almost 5,000 samples of fossil invertebrates. The samples have in turn been di- vided and shipped to over 30 active scientists in 8 countries. There is a substantial interdependence among the collaborators on the results obtained by the others. It is imperative to track accurately the location and status of preparation of the samples; to integrate age analyses, stratigraphy, section measurements, and paleoenvironmental interpretations; and to distribute this information to other collaborators. WHAT IS A DATA MODEL AND WHY MAKE ONE? A data model is an abstract representation of the information used by an organization. The model is ex- pressed as a list of entities and their interactions that the members of the organization deal with. Examples of entities for the PPP are either distinct objects such as SAMPLEs; places, such as collection SITEs; pieces of information such as AGE DETERMINATIONs; or abstract ideas, such as an assignment of a SITE to a CORRELATED SITE. (Note that the names of entities are in capital letters.) Each of these entities has a list of attributes that describe the entity. For example, the entity SAMPLE has Collector ID and Sample Collec- tion Date among its attributes. (Names of attributes have the first letter of each major word capitalized). Individual instances (or occurrences) of an entity are related to individual instances of other entities. These are called relationships and their verbal descriptions form part of the model as well. One reason for making a data model is to determine how best to arrange the data into tables, where each table represents a particular entity as described above. The overriding goal is to reduce redundancy in the database. It is a common mistake to define entities too broadly. For example, suppose that instead of defining a SAMPLE entity and a SUBSAMPLE entity sepa- rately it were decided to combine them into one table. Then, information, such as Sample Collection Date, that relates only to the SAMPLE as a whole, would have to be repeated each time a row is added for an- other SUBSAMPLE extracted from a single SAMPLE. This opens the possibility for a type of error where one SAMPLE could have two different collection dates, one for each SUBSAMPLE extracted from it. Subsequent queries based on Sample Collection Date would then be in error. There are other more subtle ways that duplication of data can occur, and the pro- cess of making a data model eliminates them. Once the data are properly arranged into tables, then entering data, finding and correcting errors, expanding the scope of the database if required, and extracting data is all much easier. There is another equally important reason for mak- ing a data model. The model, and in particular the graphical representation of the model, provides a means of explaining the working of the database and the project itself to the users. The model is supposed to represent real-life things, easily identified by people who have to work with the data. By arranging the data in an organized way and explicitly defining all of the terms and concepts used by the organization, the mod- el provides a common means of communication be- tween the users, who are well versed in the complex- ities of the organization, and the people designing the database, who may know less about the organization but more about relational database technology. Finally, the process of producing a logical data mod- el puts the data in the format required by a relational database. When a relational database is used, queries on the data may be executed using Structured Query Language (SQL), a powerful query language devel- 272 BULLETIN 357 oped specifically for this type of database. Many com- mercial databases support this language, so that SQL commands written for one particular database program may be used unaltered in another. CONVENTIONS USED IN THE PPP DATA MODEL The data model of the PPP is described both graph- ically (Text-figs. 1-2) and verbally (Definitions of En- tities and Attributes below) (Teorey, 1990; Fleming and von Halle, 1989). Each entity in Text-figure 1 is represented by a box. Above each of the boxes is the name of the entity, such as SITE or STRATIGRAPH- IC UNIT. Inside each box are a list of attributes of that entity. When a data model is transformed into a working database, the entities become tables, and the attributes become columns (also called fields) in those tables. An individual instance of an entity is represented by the information in a single row (also called a record) of its table. It may be easier in the following discussion for the reader used to working with a database to think in terms of tables, columns, and rows rather than en- tities, attributes, and instances. At the top of each list of attributes in Text-figure 1, above the line in the box and in bold letters, are one to four attributes with special significance. These at- tributes, taken together, uniquely identify an individual instance of the entity and are called primary keys. There can also be other attributes that uniquely iden- tify instances of an entity and they are called alternate keys. For example, in the entity SAMPLE, the com- bination of Collector ID and Field Code uniquely iden- tifies any one sample. A person wanting to connect his own personal list of samples with the data in the PPP Database might use these like a primary key, but all references from within the PPP database will be to the Project Sample Number, which is therefore considered the primary key. Bullets in each box mark foreign keys. A foreign key is an attribute that is a primary key for some other entity. They do not have to be unique (except in the entity for which it is a primary key), and often are not. For example, in the SITE entity, the attribute Strati- graphic Unit ID is a foreign key because it is a primary key in the STRATIGRAPHIC UNIT entity. In this case, the same Stratigraphic Unit ID may appear many times as an attribute of SITE but will correspond to only one STRATIGRAPHIC UNIT and hence appear only once in the STRATIGRAPHIC UNIT entity. When two attributes together make up a foreign key, their bullets are tied together with a short line. Between the boxes representing entities are lines that indicate how individual instances of one entity are related to individual instances of another. The lines have hash marks, circles, and crow’s feet on them that describe how many of one instance is related to how many of the other. A verbal description of the rela- tionship is also placed near each line. For example, a SITE is found in a STRATIGRAPHIC UNIT, hence the description of the relationship: is found in. A SITE is found in one and only one STRATIGRAPHIC UNIT. The symbols on the line between the boxes rep- resent this aspect of the relationship with two hash marks. The hash mark nearest the STRATIGRAPHIC UNIT box indicates that the maximum number is 1, and the hash mark slightly further away indicates that the minimum number is also one. The symbol indi- cating the maximum is always immediately adjacent to the box, and the one indicating the minimum is always slightly further away. The relationship may also be described from the STRATIGRAPHIC UNIT side of the relationship. A STRATIGRAPHIC UNIT can contain many different SITEs. It is also possible that a STRATIGRAPHIC UNIT has not yet been sampled, and hence contains zero SITEs. These aspects of the relationship are in- dicated by a crow’s foot, indicating that 2 or more SITEs may be in a STRATIGRAPHIC UNIT or by a 0, indicating that no sites are found there. Note that the words describing the relationship, “is found in” and “‘contains”, change with direction. Only the de- scription for the direction of left to right or top to bottom have been placed in Text-figure 1, for brevity. A special kind of entity is a subtype, which repre- sents a subset of another entity called the supertype. A subtype actually represents the same entity as the supertype, and hence has the same primary key, but includes some additional information. AGE DETER- MINATION is a subtype of SUBSAMPLE and in- cludes information about the age of a sample returned by the investigator. This information is placed into a separate entity because not all SUBSAMPLEs have an AGE DETERMINATION, and including the attributes for the age in SUBSAMPLE would otherwise result in the presence of many null values. To indicate that an entity is a subtype, the line indicating the relation- ship is dashed. BRIEF SUMMARY OF THE PPP DATA MODEL The following section summarizes the entities in Text-figure | and the relationships connecting them. A complete definition of each entity and its attributes is found in the last section. The two main entities in the PPP database are SAMPLE and SITE, shown with a thick outline in Text-figure 1. SAMPLEs are the actual SAMPLE 1 Project Sample Number @ Site Visit Number Field Code Collector ID Type of Sample Bags Sample Collection Date Wash Date Lithology Lost Sample? Sample Comment Y/ collected ; on 1 7 SITE VISIT Site Visit Number (PPP Number) @ Site Number Visit Date VY 1S assigned 1S to reported SITE ® 10 to have Site Number 1S @ Section Number assigned @ Stratigraphic Unit ID to @ Locality Number is found in ® Correlated Site Code o Position in Section is located Latitude Longitude Site Comment In 1S interpreted interpreted to have to have AGE OF SITE 2 14 Youngest Age Oldest Age Age Interpreter Date of Age Interpretation Age Interpolated? Age Comment is separated into aC; Former Site Visit Number Obsolete Site Visit Number O-S e@ Referenced Site Visit Number PPP Data MODEL: KAUFMANN SUBSAMPLE 2 e Project Sample Number Subsample Number 4 Taxon Type yP' H 4), A Repository ID Curator ID Transfer Date Preparation Date Barren Subsample? FORMER SITE VISIT NUMBER 8 e Site Visit Number SITE OVERLAP 9 Overlap Documentation Code @ Current Site Visit Number Type of Overlap STRATIGRAPHIC SECTION 11 Section Number Section Name Depositional Basin STRATIGRAPHIC UNIT 12 aeeey | receives F ]|---- receives a list of 273 AGE DETERMINATION 3 Stratigraphic Unit ID LOCALITY 13 Locality Number Locality Name Detailed Locality Country ID Region ID Ocean ID is assigned to ENVIRONMENT OF SITE e Site Number 15 Paleodepth Paleoenvironment Environment Interpreter Date of Environment Interpretation Environment Comment Stratigraphic Group Stratigraphic Member Stratigraphic Sequence Stratigraphic Formation Name | Type Name 1 | Name 2 Is Current Vi named Name 3 TAXON OCCURRENCE Project Sample Number Subsample Number Original Taxon ID Taxon Type Project Sample Number Subsam ple Number Age Zone Determined Age Documentation Code Age Entry Date Age Unreliable? Age Comment ENVIRONMENT DETERMINATION 4 Environment Determined Depth Determined Environment Documentation Code Environment Entry Date Environment Unreliable? Environment Comment Current Taxon ID Taxon Name Comment CORRELATED SITE 16 Correlated Site Code Defined Youngest Age Defined Oldest Age Text-figure 1.—Entity-relationship diagram, the Data Model for the Panama Paleontology Project. 274 BULLETIN 357 Name of relationship Reference number in list of entities 1 Names of attributes 1 ! Name of entity ' 1 1 Primary key (above line) Site Number uniquely identifies each different instance of SITE ~ ~ 10 e Section Number '! e Stratigraphic Unit ID e Locality Number e Correlated Site Code Position in Section / Latitude Longitude Site Comment is found in—> ‘ Foreign key (e«)~ 3 * 4—( contains ) Stratigraphic Unit ID isa foreign key in SITE. It identifies the instance of STRATIGRAPHIC UNIT with which a particular instance of SITE is associated. ' ! PS 1 ! ! ' vie ------~ / many none one and ; only one \ i] Minimum number} of instances \ \ 1 Maximum number . of instances / ‘\ Dashed relationship line Indicates that one entity is a subtype of the other. This means it has the same primary key and the relationship is one to one or one to none. Multiple primary keys, Foreign key symbols tied together Sometimes two or more attributes are needed to uniquely define an entity. This is indicated for primary keys by including two or more entities above the line in the box. For foreign keys, this is indicated by tying together the symbols. ° Alternate key An attribute that uniquely identifies an entity, but is not directly used to access the data in the table. Text-figure 2.—Key to Data Model of Text-figure 1. Connecting line indicates a relationship ' Any single instance of a SITE is found in at least one instance of a STRATIGRAPHIC UNIT and cannot be found in more than one. STRATIGRAPHIC UNIT Stratigraphic Unit ID 12 Stratigraphic Group ye Stratigraphic Formation Stratigraphic Member Stratigraphic Sequence Relationship from opposite point of w (not explicitly shown in fig. 1) Any single instance of a STRATIGRAPHIC UNIT may exist without any instances of a SITE but may contain more than one instance of a SITE.. PPP DATA MODEL: KAUFMANN PEYS) objects collected in the field, and SITEs are the places where the SAMPLEs were collected. Each SAMPLE is separated into one or more SUB- SAMPLEs which are sent to participants in the project for analysis. A SUBSAMPLE consists of a selection of fossils of a particular type, corals or foraminifera, for example. A participant may return an AGE DE- TERMINATION or an ENVIRONMENT DETER- MINATION based on the fossils found in that partic- ular sample. Or, the participant may return a list of TAXON OCCURRENCES, i.e. a species list, for the SAMPLE. This list will be stored as Original Taxon IDs which are unique for each Taxon Type and which will have full scientific TAXON NAMEs, or an infor- mal description of the fossil. A SAMPLE is collected on a SITE VISIT and all SAMPLES collected from exactly the same place on the same trip receive the same Site Visit Number (also called the PPP_number). Early in the project, SAM- PLEs were assigned different identifiers and thus may have a FORMER SITE VISIT NUMBER. In the field, collectors may record that a SAMPLE collected on a particular SITE VISIT is closely asso- ciated with another SITE VISIT. The SAMPLE may have been collected at nearly the same SITE used in a previous trip, it may span or be included in the same SITE, it may be adjacent to the same SITE, or it may be collected at exactly the same site as another SAM- PLE. These observations are recorded as a SITE OVERLAP. A SITE is a location on an outcrop from which one or more SAMPLEs have been taken, perhaps on dif- ferent collecting trips. Hence, repeat visits to a SITE, say three visits in a five year period, will result in three different SITE VISITs, all of which will receive the same Site Number. This code is, by convention, the same as the first Site Visit Number assigned. Each SITE has a LOCALITY, which is a place that can be found on a map. Several SITEs may have the same LOCALITY. SITEs will also be assigned to a STRATIGRAPHIC SECTION, which is a drawn fig- ure showing the local stratigraphy and the relationship of all the sites within a Depositional Basin. SITEs also belong to a particular STRATIGRAPHIC UNIT. From AGE DETERMINATIONs and ENVIRON- MENT DETERMINATIONS returned about SAM- PLEs collected at each SITE, and at nearby SITEs, it is possible to interpret an AGE OF SITE and ENVI- RONMENT OF SITE for each SITE. In some cases the determination is direct, in other cases it is inter- polated from directly dated SITEs above or below the SITE. In either case, a responsible scientist, the Age Interpreter or Environment Interpreter, is needed to in- terpret possibly conflicting reports for AGE DETER- MINATIONs or ENVIRONMENT DETERMINA- TIONs for relevant SAMPLEs. Many SITEs will overlap in upper and lower time boundaries. For example a series of micro samples may be taken through a section where macro fossils were collected. In order to provide macro fossil work- ers with a set of named, non-overlapping SITEs to use in their work, SITEs are grouped into CORRELATED SITES by Coates, the Chief Stratigrapher. These span a broader geographical area and have a coarser divi- sion of geological time. IMPLEMENTING THE MODEL WITH VISUAL FOXPRO This data model does not assume the use of any particular commercial database program for its imple- mentation. Any program capable of working with re- lational tables (which is what the entities represent) could be used. The PPP uses Microsoft Visual Foxpro 6.0 (VFP) for the implementation because of its well developed programming environment which makes setting up the database, entering data, writing reports, and maintaining the program relatively easy. However, just because VFP is used to enter and maintain the data does not mean that executing queries on the da- tabase requires the same program. In fact, as men- tioned above, any database program capable of exe- cuting Structured Query Language (SQL) commands could be used. In implementing the data model with VFP, each en- tity becomes a separate table, and the attributes be- come fields, or columns, in each table. For entering data and enforcing the rules governing the relation- ships, commands specific to VFP, not SQL, are used. An example of one such rule is: a particular SUB- SAMPLE cannot exist without a SAMPLE to which it belongs. VFP has a data dictionary which makes the enforcement of most rules, including this one, auto- matic. Investigators not using copies of the database for SQL queries, receive reports in the form of printed tables or made-to-order VFP tables which they can ac- cess on their own computer. USING STRUCTURED QUERY LANGUAGE TO EXTRACT INFORMATION Structured Query Language (SQL) was developed specifically for working with relational databases. One of the commands, the SELECT command, is particu- larly adept at combining information from many dif- ferent tables and placing it in a single report or another table. For example, with just one (rather complex) SQL command, a list could be prepared of all samples collected by a particular collector (found in SAMPLE), and in a particular Stratigraphic Formation (found in 276 BULLETIN 357 STRATIGRAPHIC UNIT) along with the interpreted Youngest Age and Oldest Age (found in AGE OF SITE) for the SITE to which it belongs. This infor- mation could be presented on the screen as if it were in a single table, or placed in a separate table and viewed on a remote computer where it could be im- ported into a favorite spreadsheet, or it could be put into a printed report. VFP, as well as earlier versions of Foxpro and other commercial databases, support the SQL SELECT command. Scientists learning SQL will be able to quickly extract specialized information be- yond the prepared reports distributed to them. DEFINITIONS OF ENTITIES AND ATTRIBUTES The following contains precise descriptions of each entity and its attributes. The entity names are in upper case letters and are preceded by a number correspond- ing to the numbers on the boxes in Text-figure 1. En- tity names are followed by a description and then a short list of one or more attributes which provides the primary key for the entity. Then the attributes, starting with the primary key, are listed after each entity name, along with their descriptions. The domain is the set of possible values an attribute may have. The statement “not null” indicates that the field must have some val- ue other than blank or zero, e.g., a default value. To refer to an entity’s attribute, the format ENTI- TY.Attribute (e.g., SAMPLE.Field Code) is used. 1 SAMPLE A collection of fossils, rocks, or sediment taken from a single SITE (see definition of SITE be- low). It can include both loose fossils, fossils in matrix, and drilled cores for paleomagnetic anal- ysis. The entire SAMPLE must be collected on a single trip and only one person is identified as the collector. Primary key: Project sample number Foreign key: Site Visit Number Alternate key: Field Code + Collector ID 1.1 Project sample number: The number identi- fying each SAMPLE collected by a PPP partic- ipant. Numbers are assigned by the data manager in the order in which they are inventoried at the Smithsonian Tropical Research Institute (STRI). Domain: PSNO00001-PSN999999. Not null. 1.2 Site Visit Number: Tells on which SITE VIS- IT the SAMPLE was collected. Domain: Same as SITE VISIT.Site Visit Number. Not null. 1.3 Field Code: The private sample code used by each collector and recorded in his field notes. 1.4 1.5 1.6 1.7 The codes are unique for each collector and in many cases have been used for many years pre- ceding this project. The combination of Field Code and Collector ID is unique and serves as an alternate key to SAMPLE. This key is not used within the database, but will be useful for investigators needing to combine PPP data with their own records. An assistant taking SAMPLEs assigns his supervisor’s code. Domain: Numbers, characters, spaces and **-” up to 12 characters long. Not null. Collector ID: The initials of the person col- lecting the SAMPLE. The initials are unique for each person in the project and in case of conflict are assigned by the data manager. SAMPLEs tak- en by an assistant are assigned the supervisor’s initials. Together with Field Code, uniquely iden- tifies the SAMPLE collected. Domain: Same as SUBSAMPLE.Curator ID. Not null. Type of Sample: lected. Types are: The type of SAMPLE col- micro Small bag of sediment used for mi- crofossils strib Large bulk SAMPLE kept at STRI for processing basb Large bulk SAMPLE sent to the Naturhistorisches Museum, Basel, for processing spec Specimens collected individually from sediments in place float Specimens collected individually from sediments not in place litho Lithified rock sample, e.g., basalt, coquina pmag Drilled core for paleomagnetic analysis unknw Type unknown to the data manager Domain: The above codes, all lower case. Not null. Bags: SAMPLEs are usually put into one or more cloth bags in the field. Domain: Integers 1 to 99. Default is 1. Sample Collection Date: The date the SAM- PLE was collected. Sometimes, collections are made at a SITE over a period of several days, so different SAMPLEs may be collected on differ- ent dates from the same SITE VISIT. Hence Sample Collection Date is an attribute of SAM- PLE, not SITE VISIT. 1.8 1.9 1.10 PPP DaTA MODEL: KAUFMANN Domain: All calendar days since the beginning of the project. Not null. Wash Date: The date that either: 1) a specimen or float SAMPLE was washed, glued, sorted, and boxed or 2) the date that a micro, basb, or strib SAMPLE was cooked and washed. (The date of a second processing step called Preparation Date appears as an attribute to SUBSAMPLE.) The date can be approximate. If null, SAMPLE has not been processed or SAMPLE is not of a type that needs processing. Domain: True or false. Can be Null. Default is false. Lithology: A description of the lithology of the SAMPLE by the person who collected it. Domain: Up to 80 characters. Default is blank. True if SAMPLE is lost. Domain: True or false. Not null. Default is false. Lost Sample?: Sample Comment: Includes information on preservation, lost SAMPLEs, whether a SAM- PLE was barren for calcareous nannoplankton, bryozoans, etc. Domain: Up to 80 characters. Default is blank. 2 SUBSAMPLE (Supertype) 2.1 2.2 2.3 A group of fossils, such as foraminifera, calcar- eous nannoplankton, gastropods, or corals from a SAMPLE, or a collection for a paleomagnetic analysis. These are sent to members of the pro- ject with the corresponding specialization. Ma- terial remaining after one or more Subsamples are extracted is also considered a Subsample. Primary key: Project Sample Number + Sub- sample Number Foreign key: Project Sample Number Project Sample Number: Tells from which SAMPLE the Subsample came. Domain: Same as SAMPLE.Project Sample Number. Not null. Subsample Number: A unique number start- ing from 1 for each Project Sample Number as- signed by the data manager when the Subsample is prepared. In reports, the Subsample Number is connected to the Project Sample Number with a dash, e.g., PSNO00234-2. Domain: Integers from 1 to 99. Not null. Taxon Type: The code for a general group of taxa that are included in the Subsample. Differ- ent instances of SAMPLE.Type of Sample have 2.4 2.5 2.6 2.7 277 different sets of Types of Subsample that are al- lowable. For SAMPLE.Type of Sample = micro ben benthic foraminifera faunal slide pla plankic foraminifera slide nan slide prepared for nannofossils ost ostracode slide For SAMPLE.Type of Sample = strib, basb, spec or float acor ahermatypic corals barn barnacles bry bryozoans cor corals clam clams crab crabs ech echinoderms hcor hermatypic corals mol mollusks sna snails For SAMPLE.Type of Sample = micro, strib or basb orig wood For SAMPLE.Type of Sample = pmag original sample material pieces of wood pmag paleomagnetic sample Domain: The above codes, all lower case. Not null. Repository ID: The code for the name of the institution where the Subsample is currently kept. Domain: Five letters, all lower case. Not null. Curator ID: The initials of the person respon- sible for the Subsample. If a participant’s initials are not unique, the data manager will assign a code. Domain: Four letters, all lower case. Not null. Transfer Date: The date that the SUBSAM- PLE was sent to the curator. Domain: Same as SAMPLE.Sample Collection Date. Not null. Preparation Date: The date a particular group was picked from washed residue of a micro, basb, or strib SAMPLE; was picked for a partic- ular group; or the date a specimen or float SAM- PLE was washed, glued, sorted, and/or put in boxes. If null, SAMPLE was not prepared. Domain: Same as SAMPLE.Sample Collection Date. Default is null. 278 2.8 BULLETIN 357 Barren Subsample?: True if a PROCESSED SAMPLE is found to have no fossils relevant to its Type of Subsample. Domain: True or False. Default is False. 3 AGE DETERMINATION (Subtype of Subsample) 3. = 3.2 S23 Planktic foraminifera and calcareous nanno- plankton from a SAMPLE can be used to deter- mine an age for a SITE. A paleomagnetic mea- surement of polarity, in conjunction with the lo- cal pattern of polarities from other SAMPLEs in the same stratigraphic sequence, can further re- fine the age. The geologic age returned may of- ten not correspond exactly to that obtained from another Subsample, because of the limits of each dating system, but by combining them, the youn- gest and oldest possible ages may be determined even more precisely than the results of the in- dividual determinations taken alone. There is substantial collaboration among the participants on individual SAMPLEs before they return their results. Primary key: Project Sample Number + Sub- sample Number Foreign key: Project Sample Number + Sub- sample Number Project Sample Number: Tells to which SAMPLE the AGE DETERMINATION applies. Domain: Same as SAMPLE.Project Sample Number. Not null. Subsample Number: Together with Project Sample Number, tells which Subsample was used for the determination. Domain: Same as Subsample.Subsample Num- ber. Not null. Age Zone Determined: The age determined by examination of a Subsample by a specialist. Usually, this is from calcareous nannoplankton, planktic foraminifera, or from a paleomagnetic sample combined with previous biochronologic results. The format of the returned age is vari- able. It can consist of a calcareous nannoplank- ton zone (e.g., NN17), a planktic foraminifera zone (N12), an absolute age (3.5 Ma) or a range, consisting of any combination of these (NN16— NN17, 3.5 Ma—3.2 Ma, N12-3.5 Ma). Note that even a zone designation can include an absolute age if the proper index fossil is found (or not found) indicating that only a part of a zone is present in a SAMPLE. This format for ages is 3.4 3.5 3.6 S37) well established and easily understood by the us- ers, but does not lend itself to SQL queries. Ap- plication specific functions (Youngest and Old- est) use the appropriate lookup table to find the currently established ages for zones and return either the youngest or the oldest age for any of the formats above. These functions are then used to fill in the derived attributes in the table cor- responding to the AGE OF SITE entity. Domain: 11 characters and digits, the minus sign, blank, and decimal. The letters are NN or N which must precede a two digit integer (with a leading zero if necessary), or Ma, which must precede a decimal number from 0.0 to 9.9. The minus sign is used as a connector when a range is given. Not null. Age Documentation Code: The reference to the document submitted by a participant which contains his findings from a Subsample. All such documents are given a catalog number and stored as a paper record at several institutions. Domain: Catalog number or numbers used for documents from participants. Age Entry Date: The date that a value for Age Zone Determined was entered into the database, or if corrected or updated, the date of the last change. This information is used in conjunction with Date of Age Interpretation in the AGE en- tity to determine whether Youngest Age and Old- est Age need to be re-interpreted. Domain: Same as SAMPLE.Sample Collection Date. Default is current date. Not null. Age Unreliable?: Whether an AGE DETER- MINATION is considered in error and not to be used in interpreting the age of a SITE. This at- tribute is marked as True in those cases. Domain: True or False. Not null. Default is False. Age Comment: Comments about the age de- termination. Domain: 80 characters or numbers 4 ENVIRONMENT DETERMINATION (Subtype of Subsample) Many types of Subsamples are used to determine the depositional environment for the SAMPLE. For some Subsamples, such as benthic forami- nifera, a determination of paleowater depth is possible as well. Primary key: Project Sample Number + Sub- sample Number 4.1 4.2 4.3 4.4 4.5 4.6 4.7 4.8 PPP DaTA MODEL: KAUFMANN Foreign key: Project Sample Number + Sub- sample Number Project Sample Number: Tells to which SAMPLE the ENVIRONMENT DETERMI- NATION applies. Subsample Number: Together with Project Sample Number, tells which Subsample was used for the determination. Environment Determined: The text of the ENVIRONMENT Determination returned by a participant. May be blank but only if Depth De- termined is blank. Domain: Up to 40 characters. Default is blank. Depth Determined: The paleowater depth de- termination returned by a participant. Is blank if nothing is returned. Domain: Up to 40 characters. Default is blank. Environment Documentation Code: The ref- erence to the document containing the findings from the SAMPLEs. (See note about these doc- uments for AGE DETERMINATION.Age Doc- umentation Code). Domain: Catalog number, or numbers, for doc- uments from participants. Environment Entry Date: The date that a val- ue for Environment Determined or Depth Deter- mined was entered into the database, or if cor- rected or updated, the date of the last change to either. Both need to be considered together in interpreting the ENVIRONMENT of SITE. Domain: Same as SAMPLE.Sample Collection Date. Default is current date. Not null. Environment Unreliable?: Whether an EN- VIRONMENT DETERMINATION is consid- ered in error and not to be used in interpreting the ENVIRONMENT OF SITE. This attribute is marked as True if either Environment Deter- mined or Depth Determined is not considered valid. Domain: True or False. Not null. Default is False. Environment Comment: Comments about Environment Determination or Depth Determi- nation. Domain: 80 characters or numbers 5 TAXON OCCURRENCE An observation that a particular taxon was found in a Subsample. There is a one-to-many rela- 5.1 5.2 5.3 5.4 Sys) 279 tionship between Subsample and TAXON OC- CURRENCE. All of the TAXON OCCURR- ENCEs taken together for a given Subsample can be considered a species list for that subsam- ple. Primary key: Project Sample Number + Subsam- ple Number + Taxon Type + Original Taxon ID Foreign key: Project Sample Number + Sub- sample Number Foreign key: Taxon Type + Original Taxon ID Project Sample Number: Tells the Project Sample Number in which this TAXON OCCUR- RENCE was found. Domain: Same as SAMPLE.Project Sample Number Subsample Number: Together with Project Sample Number, tells the Subsample in which this TAXON OCCURRENCE was found. Domain: Same as Subsample.Subsample Num- ber Original Taxon ID: Together with Taxon Type, identifies a taxon that was observed to oc- cur in the Subsample, using the taxon name in use at the time of the observation. It is not un- common for the name of a particular taxon to change as more taxonomic information about it is changed, and to avoid confusion, the original name is recorded here. The Original Taxon ID, and that taxon’s full name, is resolved by the many to one relationship to TAXON NAME, and by that entity’s many to one relationship to itself. Domain: Same as TAXON NAME. Original Tax- on ID Taxon Type: Tells the code for the group of taxa being searched for in the Subsample. Re- searchers working on different groups of taxa may inadvertently use the same Original or Cur- rent Taxon ID for different taxa. To avoid con- fusion, Taxon Type is made part of the key so that identifiers for taxa need only be unique with- in a particular group. Domain: Same as Subsample.Taxon Type Abundance Code: A collector specific esti- mate of abundance in the SAMPLE, appropriate for the particular taxon and Type of Subsample. Domain: 12 characters and/or numbers. 6 TAXON NAME Taxa identified in a SAMPLE are given a code which is used to record their occurrence. This 6.1 BULLETIN 357 entity records this code, the Original Taxon ID, and the full name which it represents. The code may represent the name of a family, a genus, species and subspecies or even an informal des- ignation, such as “urchin spine” or “unknown coral fragment’’. Using a full genus and species appellation for recording and entering species observed in a SAMPLE, and for analyzing the data afterwards, is both time consuming and error prone. Here, eight-letter abbreviations are used in the data- base. Experience shows that this is long enough so that a person familiar with a particular group can easily decipher it, yet not so short that it is ambiguous. Often, scientific names differ in only one or two letters out of many. It is easier to check whether one of 8 letters is in error than to see if one of 20 letters is in error. In producing summary reports for internal use and when re- viewing the names on a computer screen, eight- letter abbreviations are much easier to read and to format into tables. A good data entry program should require typing only one to three letters to enter the full eight-letter code automatically. To keep track of changes to the currently used name for a taxon, this entity enters into a recur- sive relationship with itself, using Taxon Type + Current Taxon ID as a foreign key to look up the current full name of any taxon recorded in the database. This method of resolving the currently used name of a taxon is not a substitute for a taxo- nomic database (e.g., Paleobank, Krebs ef al., 1996). Maintenance of this data cannot be done automatically by accessing such taxonomic da- tabases, since most changes to Current Taxon ID will be to record the substitution of a scientific name for an informal designation of a taxon, or the correction of an error in identification. It is the responsibility of each researcher to maintain the names of the taxon with which he is working. Primary key: Taxon Type + Original Taxon ID Foreign key: Taxon Type + Current Taxon ID. This foreign key is used in a one to many re- cursive relationship to resolve the Current Taxon ID given the Original Taxon ID. Because Orig- inal and Current Taxon ID are unique only with- in a given Taxon Type, Taxon Type is a neces- sary part of the key for resolving these relation- ships. Original Taxon ID: An abbreviation repre- senting the taxonomic name used at the time a particular specimen is recorded. This name will 6.2 never be changed so that the name in the data- base will always refer to what is actually written down on the original data sheet. This avoids con- fusion in case the name currently used changes several times. Domain: Eight letters, both upper and lower case, and digits 0 to 9. Spaces and punctuation are not allowed and the first character cannot be a digit. These restrictions allow the abbreviation to be used as legal field (column) names for the most commonly used databases. Current Taxon ID: Tells the currently used abbreviation for a given taxon. Often, a research- er will change the name for a taxon, either to correct an error in identification, to reflect a more precise identification, or because of a change in the taxonomy. By recording the currently used name for an obsolete name, the new name can automatically be used when reports are printed. Merely substituting the new code for the old in the database can cause great confusion and de- stroys information about the history of names used for a particular taxon. The rules for changing the Current Taxon ID are as follows: Whenever Current Taxon ID is changed for an existing Original Taxon ID, a new instance must be added to this entity. This new in- stance must have the same value for both Original Taxon ID and Current Taxon ID. The other information in the original instance, in- cluding Namel, Name2, and Name3, is left unchanged to preserve the history of changes to the Original Taxon ID and its full name. If yet another change is made to Current Taxon ID, then another instance is added and both previous instances must have Current Taxon ID updated. There is a recursive relationship between this entity and itself which allows a reference to the Original Taxon ID to return the Current Taxon ID, along with the current full name. All instances then, where Original Taxon ID and Current Taxon ID are different represent obsolete usages. To prevent ambi- guity in archived species lists, the combina- tion of Taxon Type and Taxon ID, even for obsolete usages, must remain unique. This means that an Original Taxon ID, such as Cor- alsp, cannot be recycled for use with a differ- ent taxon, once a more accurate identification is supplied for the original usage. Domain: Same as Original Taxon ID. 6.3 6.4 6.5 6.6 PPP DATA MODEL: KAUFMANN Taxon Type: The taxon or group of taxa for which a set of unique Taxon IDs is constructed and maintained by a researcher Domain: Same as Subsample.Taxon Type Namel: A genus name, a higher taxonomic name, or a short unstructured description. Domain: Not Null. Up to 20 letters. Can include “> Name2: A species name, the letters ‘sp.’ pos- sibly with a letter or number after, or a contin- uation of a short description from Name 1. Domain: Up to 20 letters. Can include *?’. De- fault is blank. Name3: A subspecies name, the letters ‘ssp.’ possibly with a letter or number after, or a con- tinuation of a short description from Namel. Namel, Name2, and Name3 taken together are referred to in this paper as the full name. Domain: Up to 20 letters. Can include *?’. De- fault is blank. 7 SITE VISIT (Supertype) 7.1 A visit to a single SITE by one or more persons at the same time where one or more SAMPLEs are taken. If two people taking different kinds of SAMPLEs agree that their SAMPLEs are equiv- alent in age because of their close proximity on the outcrop they are considered to be from the same SITE VISIT. Since SAMPLEs are consid- ered equivalent in age if their upper and lower ranges in the section are the same, if one SAM- PLE spans only a small part of the vertical extent of another SAMPLE, such as a microfossil sam- ple taken next to a very large coral head, then the two SAMPLEs are considered to be from different SITE VISITs. It is also possible that two SAMPLEs are taken that are of equivalent age but that fact may not be known to the data manager when the Site Visit Number is assigned, and they will receive different Site Visit Num- bers. The primary key for SITE VISIT, Site Visit Number, is also known as the PPP_Number and is used to identify collections in published work. Primary key: Site Visit Number (PPP_-number) Site Visit Number: The code assigned to the SITE VISIT. Having one number for all SAM- PLEs corresponding to the same time range in the section facilitates reference to the age ulti- mately returned by the collectors of those SAM- PLEs by providing a publishable number at an 7.2 281 early stage. It is possible that two SAMPLEs col- lected at exactly the same time and place could receive different Site Visit Numbers because of the way they were handled during collection and subsequent processing. Domain: PPP-O00001 to PPP-999999 Site Number: Tells to which SITE the SITE VISIT belongs. Domain: The set of existing SITE VISIT num- bers, but with the prefix S instead of PPP. 8 FORMER SITE VISIT NUMBER (Subtype of SITE VISIT) 8.1 8.2 8.3 Early in the project, SAMPLEs were assigned what was called a ‘“‘default CJ number”’ to iden- tify the same entity that the SITE VISIT number now identifies. Older labels on stored SAMPLEs and references in field notes use this code. An even earlier code, not used on sample labels, may be found in Coates’ field notes. Primary key: Site Visit Number Site Visit Number: Tells to which SITE VISIT the FORMER SITE VISIT NUMBER refers. Domain: 12 characters. Not null. Former Site Visit Number: The ‘Default CJ number” on some labels and in some field notes. No longer assigned. Domain: 12 characters Default is blank. Obsolete Site Visit Number: The code found only in Coates’ early field notes. No longer as- signed. Domain: 12 characters. Default is blank. 9 SITE OVERLAP 9.1 A correspondence between a current SITE VIS- IT and an earlier or concurrent instance of a SITE VISIT to that SITE or to an adjacent SITE. Examples of a correspondence are: a current SITE VISIT being considered exactly the same as another; a current SITE VISIT being a subset or superset of another; or a current SITE VISIT being close to but not overlapping another. This entity is used to produce user views of the data with equivalent and closely associated SITE VISITs listed close to each other to aid in grouping the SAMPLEs by time interval. Overlap Documentation Code: An identifier for a written note describing how the current Site Visit is related to another. Normally, the note is from a field notebook and the owner, volume, 9.2 9.3 9.4 BULLETIN 357 page, and line numbers are given. This field must be unique, but is otherwise unstructured. Domain: 20 letters or numbers. Not null. Current Site Visit Number: Tells which cur- rent SITE VISIT has entered into an association. Domain: Same as SITE VISIT.Site Visit Num- bers. Referenced Site Visit Number: Tells which SITE VISIT is referred to in documentation about a previous, or a concurrent, SITE VISIT. Domain: Same as SITE VISIT.Site Visit Codes. Overlap Type: How the two SITE VISITs are associated. The judgement of how the collections are related is made by the person collecting the SAMPLE. Sometimes a collection of macrofos- sils will be made from a SITE visited the pre- vious year and these will be considered “‘equiv- alent” and an Overlap Type will be assigned in- dicating this. The codes are: equiv The current SITE VISIT is consid- ered equivalent to another one. part The current SITE VISIT is a part, or a subset, of another one. cont The current SITE VISIT contains, or is a superset, of another one. near The current SITE VISIT is close to but not overlapping another one. Domain: The above codes, all lower case. Not null. 10 SITE (Supertype) A part of an outcrop from which one or more SAMPLEs have been taken, all having the same upper and lower stratigraphic boundaries. The lateral extent of the SITE can be as large as fea- sible while still maintaining the same upper and lower boundaries. Typically, microfossil sample SITEs are six centimeters or less in extent while some bulk, float, and specimen sample SITEs may extend for tens of meters along a bedding plane and be a meter or so in stratigraphic thick- ness. If a return visit is made to a site after an ex- tended period of time and additional SAMPLEs (bulk, specimen, efc.) are collected, the collector may record that the site was the same as that visited previously. But micro samples are nor- mally too small and too precise in their upper and lower bounds to be considered equivalent if collected at different times, so the collector will generally record that the new SITE is near but not exactly the same as a previous collection. 10.1 10.2 10.3 10.4 10.5 10.6 10.7 The code for identifying the SITE uses the digital part of the Site Visit Code from the first visit, preceded by the letter ‘S’. If the SAMPLE is being collected for the first time this will have the same digital component as the Site Visit Code. (e.g., PPP001234 and S001234). Primary key: Site Number Foreign key: Section Number Foreign key: Stratigraphic Unit ID Foreign key: Locality Number Foreign key: Correlated Site Code Site Number: Tells which SITE was visited. The default indicates that no Site Number has been assigned. Site Numbers are assigned in batches, after the data for new SAMPLES have been entered. Domain: Same as SITE VISIT.Site Visit Number except that the digital part is preceded by ‘S’ instead of ‘PPP’. Default is ‘SOOO000’. Section Number: Tells the STRATIGRAPHIC SECTION in which the SITE was found. Domain: Same as STRATIGRAPHIC SEC- TION.Section Number. Default is 0. Stratigraphic Unit ID: Tells from which STRATIGRAPHIC UNIT a SAMPLE was tak- en. Domain: Same as STRATIGRAPHIC UNIT.- Stratigraphic Unit ID. Not null. Locality Number: Tells the LOCALITY in which a SITE is found. Domain: Same as LOCALITY.Locality Number. Not null. Correlated Site Code: Tells to which COR- RELATED SITE this SITE has been assigned. Domain: Same as CORRELATED SITE.Corre- lated Site Number Position in Section: The position of the SITE in the composite section, measured in meters from the bottom and adjusted for dip and local differences in section thickness. Domain: Numbers from 0.0 to 999.9. Not null. Latitude: The latitude of the SITE, using dec- imal notation, as accurately as can be measured. Currently, location is measured to within 100 m using a map or a GPS receiver. Latitude and lon- gitude can be expressed in degrees, minutes, and seconds where required by using a function to make the conversion on the fly. Domain: Numbers from —4.00000 to 15.00000 10.8 11 12 12.1 12.2 12.3 Stratigraphic PPP DATA MODEL: KAUFMANN and 999. Default is 999, indicating a missing val- ue. Longitude: The longitude of the SITE, using decimal notation, as accurately as can be mea- sured. Domain: Numbers from -—88.00000_ to —60.00000, and 999. Default is 999, indicating a missing value. STRATIGRAPHIC SECTION A diagram showing the stratigraphic relationship among SITEs, to scale. The Chief Stratigrapher will decide which SITEs can be grouped togeth- er into any given STRATIGRAPHIC SECTION. A SITE may belong to only one STRATI- GRAPHIC SECTION, or it may remain un- grouped. Primary key: Section Number Section Number: The number assigned to the STRATIGRAPHIC SECTION. Domain: Integers from 0 to 999. Default is 0. Section Name: A name given to the STRATI- GRAPHIC SECTION Domain: Up to 20 characters. Not null. Depositional Basin: A geographic area with a common river drainage. Domain: Up to 20 characters. Not null. STRATIGRAPHIC UNIT The smallest established subdivision of the rocks in the study area. Primary key: Stratigraphic Unit ID Stratigraphic Unit ID: A code identifying the Stratigraphic Group, Stratigraphic Formation, and Stratigraphic Member. Since formation names are unique in the study area, the code con- sists of an abbreviation of the formation name, and if used, the member name. Includes a code for unknown. Domain: Up to 12 characters. Stratigraphic Group: The established name of the group. Can be ‘unknown’. Blank indicates that a group name is not used. (The simple name “Group” cannot be used because ‘group’ is a reserved word used by SQL, the query language used by all relational databases.) Domain: Stratigraphic Group names. Default is blank. Formation: The _ established 12.4 12.5 13 13.1 13.2 13.3 13.4 name of the formation. Can be ‘unknown’. Can- not be blank. Domain: Stratigraphic Formation names. Not null. Stratigraphic Member: The established name of the member. Can be ‘unknown’. Blank indi- cates that a member name is not used. Domain: Stratigraphic Member names. Default is blank. Stratigraphic Sequence: A decimal number indicating the position of the STRATIGRAPHIC UNIT in the geological column. It is used to dis- play the names in stratigraphic rather than alpha- betic order. The smallest numbers correspond to the youngest rocks. Blank is 0, ‘unknown’ is 9919} Domain: Integers from 1.0 to 99.9. Not null. LOCALITY The local area where the outcrop is found. The location of an outcrop is described by a reference to a place name (Locality Name) that can be found on a topographic map. To further define the location, a Detailed Locality is used which consists of short directions on how to reach a particular outcrop. Together, they generally lo- cate the outcrop to within about 100 meters. More than one SITE can be found in a given LOCALITY. Primary key: Locality Number Locality Number: A unique number assigned to each LOCALITY. Domain: Five digits, 0-9. Not 00000 and not null. Locality Name: A name that can be found on a 1:50000 topographic map (such as Isla Colon or Rio Azul) which identifies a small area. Domain: Any name on a map up to 20 charac- ters. Detailed Locality: Short directions on how to reach the outcrop. Examples are ‘‘300 meters north of the bridge over the Rio Azul” or “50 m downstream from Site PPP000123” Domain: Up to 80 characters. Country ID: An abbreviation that tells the country in which the SITE is found. The abbre- viations currently in use are: 284 13.5 13.6 BULLETIN 357 Pan (Panama), CR (Costa Rica), Nic (Nicaragua), Ecu (Ecuador), Tri (Trinidad & Tobago), Ven (Ven- ezuela). Domain: The above codes. Not null. Region ID: A subdivision of a country. Domain: Established names of subdivisions of countries, or an abbreviation for them. Up to 12 characters. Ocean ID: An abbreviation that tells into which ocean the streams flow. Domain: P (Pacific) or C (Caribbean). Not null. 14 AGE OF SITE (Subtype of SITE) 14.1 14.2 14.3 14.4 The composite geologic age of a specific SITE, as interpreted from the information returned from AGE DETERMINATIONS. Sometimes this information is contradictory so that the AGE OF SITE cannot be calculated automatically from individual AGE DETERMINATIONS. It requires an experienced researcher to make the final interpretation. Available valid ages from nannoplankton, forams, and paleomagnetic ana- lyses are presented to the person making this in- terpretation, as well as the maximum and mini- mum possible ages in millions of years. These are listed as derived attributes in this entity since the information is available elsewhere in the da- tabase. In cases where there is no direct infor- mation about a SITE, an AGE is interpolated from the position of the SITE relative to nearby directly dated SITEs. Primary key: Site Number Foreign key: Site Number Site Number: Tells to which SITE the AGE OF SITE refers. Domain: Same as SITE.Site Number. Not null. Youngest Age: The youngest composite age in Ma interpreted for a SITE by an experienced re- searcher. Domain: Decimals from 0.0 to 9.9. Oldest Age: The oldest composite age in Ma interpreted for a SITE by an experienced re- searcher. Domain: Decimals from 0.0 to 9.9 Age Interpreter: The initials of the person who made the age interpretation. 14.5 14.6 14.7 15 15.1 15.2 15.3 15.4 15.5 Domain: Same as SAMPLE.Collector ID. Not Null Date of Age Interpretation: The date the in- terpretation was made. If this date is earlier than the date that any AGE DETERMINATION was last entered into the database for this SITE, the AGEs will have to be reinterpreted. Domain: Same as SAMPLE.Sample Collection Date. Not null. Interpolated?: Whether the AGE OF SITEs were obtained by interpolation from other nearby SITEs. Domain: True or false. Default is false. Age Comment: process. Comments about the dating Domain: Up to 80 characters. Default is blank. ENVIRONMENT OF SITE (Subtype of SITE) The composite depositional environment for a specific SITE including paleowater depth. Like AGE OF SITE, an experienced researcher is needed to make the final interpretation from all available reports of AGE DETERMINATION. Primary key: Site Number Foreign key: Site Number Site Number: Tells which SITE has this en- vironment. Domain: Same as SITE.Site Number. Not null. Paleodepth: The determination of the depth of the water at the time of deposition for the SITE. This is an uncoded attribute, reflecting concisely the best estimate of the Environment Interpreter. Domain: Up to 20 characters. Default is blank. Paleoenvironment: The depositional environ- ment, excluding depth, for the SITE. This is an uncoded attribute, reflecting concisely the best estimate of the Environment Interpreter. Domain: Up to 80 characters. Default is blank. Environment Interpreter: The experienced participant making the interpretation of both Pa- leodepth and Paleoenvironment. Domain: Same as SAMPLE.Collector ID. Not null. Date of Environment Interpretation: The date the interpretation of the ENVIRONMENT was made. This information will be used in con- junction with ENVIRONMENT DETERMINA- TION.Table Entry Date to determine if a re-in- PPP Data MODEL: KAUFMANN 285 terpretation is necessary. Domain: Same as Sample.Sample Collection Date. Not null. 15.6 Environment Comment: Comments about the interpretation of the ENVIRONMENT. Domain: Up to 80 characters. Default is blank. 16 CORRELATED SITE A group of SITEs with similar AGEs. The Chief Stratigrapher will group SITEs to provide coars- er divisions of time for the use of project partic- ipants. Primary key: Correlated Site Number 16.1 Correlated Site Code: A code identifying a group of similarly aged SITEs. Domain: Integers from | to 999. 16.2 Defined Youngest Age: The youngest age, in Ma, of SITEs assigned to the CORRELATED SITE. Domain: Decimals from 0.0 to 9.9. Not null. 16.3 Defined Oldest Age: The oldest age, in Ma, of SITEs assigned to the CORRELATED SITE. Domain: Decimals from 0.0 to 9.9. Definitions The following words are used consistently in the names of entities and attributes to modify the remain- der of the name. code: A set of letters and numbers associated with an attribute or entity. ID: A code which is a short abbreviation of a longer name and which is designed to be easily identifiable by a user familiar with the domain of entities or attributes that it describes. A code consisting primarily of digits, sometimes with a fixed set of letters add- ed to the beginning, assigned in numer- ical order. type: One of a short set of codes used to de- scribe a particular class of attributes. date: A particular calendar day. Dates refer to events connected with the activities of the PPP participants, not to geological events. age: Time expressed in millions of years be- fore the present. age zone: Time expressed either as a microfossil zone, a geological age, or in millions of years before the present. Ms This symbol at the end of the name of an attribute indicates that the attribute takes on logical values, true or false. number: ACKNOWLEDGMENTS The original design of the PPP Database was by Karl Kaufmann and Laurel Collins. Subsequent mod- ifications included substantial input by Laurel Collins. Helena Fortunata assisted generously in developing the early stages of the model. The database is available at the internet Web site http://www. fiu.edu/~collins]/. REFERENCES CITED Fleming, C.C., and von Halle, B. 1989. Handbook of Relational Database Design. Addison Wes- ley. Reading, Massachusetts. 839 pp. Krebs, J.W., R.L. Kaesler, E.A. Brosius, D.L. Miller, and Chang, Y.M. 1996. Paleobank, A Relational Database for Invertebrate Pale- ontology: The Data Model. The University of Kansas Pa- leontological Contributions, New Series, #8. The Univer- sity of Kansas, Lawrence, Kansas. 7 pp. Teorey, T.J. 1990. Database Modeling and Design: The entity-relationship ap- proach. Morgan Kaufmann, San Mateo, California, 267 pp. 2 ij ) i ou ba ~ } UJ i] AN , } aT Ue | ’ é ae ee Es. Wy ; . BILLA, * , i ol U mow “alin all » any i ' i aiid " i 4 t il ‘QVAAnG & Th) if ) 1@ A Ve | tok ie fd = x 1% THAD Li ‘A jh evi ie ' Ate od ‘ aint BD pikcoe te (intel ag vil ; were ais et a uit assed) ie 1 (Peer eer ’ ora ali) Slarrares AT == 1 ith) 9en dfs aT rip ay! Palin ye we eG == qa an lini al} ann a ir Wh ay ATA aww One tin Sie a iW erent Mi (eZ © ae Orepiy eth cle Powel! <¢ bie alle idee ghee A esi ww) a Ay te S Pea) 4g) Sew PP 75 ie th A) allen fect act tambien hg: : Ce ee CAT Ale her Scinaaianaaderatidns A coe 7! a ; Feit an 4 APPENDIX A MAPS ANTHONY G. COATES Smithsonian Instituition Smithsonian Tropical Research Institute Washington, D.C. 20560-0580, U.S.A. These maps were produced with the computer pro- grams Atlas Geographic Information System, version 2.1, and Coreldraw, version 7.0. They include PPP numbers for all Caribbean sampling localities to date. The maps are listed below by number in the order presented. Map 1. Colon to Gobea, Panama Map 2. Miguel de la Borda to Calzones River, Panama Map 3. Petaquilla River to Boca de Concepcion, Pan- ama Map 4 & Insets. Escudo de Veraguas, Panama, and insets A, B, C and D Map 1 Map 5. Valiente Peninsula, Panama Insets of Map 5S. Insets A, B, C, D, E and F Map 6. Cayo Agua, Panama Map 7. Popa Island, Panama Insets of Map 6. Insets A, B, C, D, E, EK G and H Map 8. Bastimentos Island, Panama Map 9. Colon Island, Panama Map 10. Manzanillo Point to Bonifacio, Costa Rica. Map 11. Bananito River to Limon, Costa Rica Inset A of Map 11. Portete to Chocolate Creek Inset B of Map 11. Cangrejos to Route 32 Inset C of Map 11. Banano River to Vizcaya River 1307/1308=1503) 1077=1079/1081 1035=1078, 1501/1503, 1094/1099 Caribbean Sea ee V4. : J f Indio River 79" SS la 1656/1657 1659 AN) 1076=1088/1093 Rte. 82\?Marga io Chagres \. River “A S$ \ Galeta Point a Naranjitos Point Toro Point 22/23 Colon 7172/1174 uN =2508 > lal 42/43 | aN Vee a MN rs 37/4128 +s nse VA Gobea Palmas ff 1646 Dam, 44/45 tes $. Bellas = ° ‘ ONG TY x ria ae G 1639/1643 ae : ean, ca 2q T | Se AR tee ea Boca del ee, 1660/1661 Y ia Me io Indio ml 1645 e 1 ¢ 4 = 1644 Lake Gatun ~ aie 25 ~~26 enone a] ahr" ra * sae fe 288 BULLETIN 357 Map 2 ; Caribbean Sea 158/161 =1973 9° 10’ Miguel de la Borda Miguel de la Borda River nee 162,164 celts 163 Cocle del Norte Calzones River River 80° 15’ Map 3 80° 45' 80° 55’ = XG Caribbean Sea Petaquilla River 8° 55’ = Boca de Concepcion Veraguas River 165/166 Maps: COATES Map 4 & Insets te) 0.5 A 2267 2266 4-7 2268 2269 176 Long Bay Point 2265 1234 81° 33’ 48.17" I 81° 33’ 6.78" Caribbean Sea 1232 Escudo de Veraguas Island Km ee yo wh 171,1243,1974 2480 ©60-: 1235, C 1241 Pee 37 1240 VA 1242 1236 368=2253 LZ =2169 te 367 365° QYy, 2271=2483 2259 2258 = 2254 2262=2481 2255 ee S~\\2260 364 2963 — 2261 oa 362 2264,2482 oe 2270 ule 2272 2277 2274 > 2273 Km 2276 2275 04 9° 6’ 11.46" 169=2180 2181 a Booby 2182 Cays 481=2187 B 179=358= 2178 123022171 2177 \ 2176 360=2170 Ww 2179 D Cotton Bay 2280] 2281 2282 2284 iJ 2286 Cl 2287 2288 +2183 2296 2288 ewe 2294 Km =a) 0 0.05 BULLETIN 357 290 Map 5 2145 Cusapin 2139/2142 Bluefield Point 2146 5 2147 o536 s Caribbean Sea 190=1523 182 Grape Point 192 181 g° 10°44 A Tete Sirain a0 Toro 1% Point Poi Bluefield Bay 905 . Cay Caracol Point 130026 6 y hi . “| 2504 r~ Na 254 2522 Wanchi Point 1 - oS 1509 2524/ 2520=1301 3 B _% Plantain Cays 1507 ae 1291 ee 1508 : Tobabe Point 1514 1510 185 Bugori Tobabe Warrie 1299 186 Point 184 ae ce 1297 Sane ooint 1512 4513 ¥ Ne hae 183 Valiente ' D. Chong or Nispero Point 9° a g Shark Hole Patterson Cay 2838 ala : F Point 2837 2096 Senin a ree Old Bess oft : Beach y Cea, ‘am. 1 Point Chiriqui Lagoon 10752008 B é t runo Peninsula Bluff 2217=2486/2487 eceiaro 379=2227 = N 4 1968,2218 376=2226 ee 2488/2489 2219 Km 378 380 fiance ess 381 389 0 2 81° 55’ 81° 50’ | | z a1 Maps: COATES 291 Insets of Map 5 A B 2215 Toro Cay 2216 1292=2149 2156 SE 2298/ Sac \ 2313 AOD w Z i. }) 213/214, 82° 10 82° 05’ 2243/2244 ral AM 35 Zl Popa Island Map 9 ial 20 1782-1835,1175-1181, a 15 * 2221,2246,2531 Swan Cay- Caribbean Sea 1995,2002 Mimitimbi Bluff 1260/1271,1275,1425 2252 Hill Point | 51—-+ 2251 i a Mimitimbi Creek> 9° 25 e Drago Knapp Hole 1285 1286=2248,1287 1423 1424 Bastimentos Island N Carenero Cay = 9° 20 Solarte “a Maps: COATES 295 Map 10 ON | 9° 50’ Bonifaciof> \ 82° 50' 82° 45) k 1 (a. Caribbean Sea IPA (VW Limonal i 1327 1325 = 1326 WE ICS 1328 oO . 2 4767 3 eS, Cahuita Point 1329S Suarez 1770 Catuita SNRiver 726/731 1768 ENS 1765 Carbon Dos 1766 (Dindiri)™ 732/733 ah : Manzanillo Point N Puerto 9° 40’ Km 453/454, 21774 Viejo _, 1318=1319 ) cP? =a 735/737, S Nee 1736,1737 Sk Aan 1989 4323 y — a BS ” Catarata yy pres el Sandbo Costa Rica nae River Manzanillo NH Sixaola Rive’ a 4 to Sixaola and Guabito ¢ a SS 969/977 Peje River <= Caribbean Sea 10° 00 4 724 720/723 Limoncito Rive Bey SP | izcaya River _ J > Dondonia*, . x ‘ Yar Bananito Rive a) Km ea ee BS 0 5 eset ae oe 1990\ . =| 83° 05 = 83° 15) oy | 296 BULLETIN 357 Inset A of Map 11 A 92-00-53/54 1352= N 21126 1355 iss Sa: i 10° 0’ 58.41" e 1333/1339= 1390=1417/1421 1485/1487 Empalme Moin Moin River @ 2061/2062 1346=1363 =2016 9° 59’ 47.27" 1364/1365 =2015 RECOPE Oil Buenos Renner) hice 1342/1344=1366 1435=1440/1442 =1498=2003 1370 119 e 759_ be Ne ae , 1124=1348 978/979 ey =1436 1125 2047 ‘ sias Farm = 1371 667 1316=1359/1362 715/716 a5 =1387/1388= 717/718 a 2056 1392/1409 é =1473/1478 =2011/2014 84 cn 22580 ae oO 21772/1773 e oe —_ 2055 Berta Farm Rte. 32 2057/2058 2065, 1347 eS 699=768 2019/2020 662 2021/2023 2029 663/665 =1377/1380 =1382/1383 660 =1443/1455 83° 5’ 52.87" Maps: COATES 297 Inset B of Map 11 | B 83° 3' 0.17" Ran oes N Cangrejos i See ~~ 738=1771 Km Tae 658 Ue EEE 771=1349 0 1 ~ 657 } Sag, 309=138 Piuta 655.2126 wee: 71389 : 2125 Lee ms pt 638/646=757 123 — ted () =2005/2010, 7 =1982,1088, 2122 oe OW72 NS 2121 Ce? mei 5) Sh \ > sy 2119 ABD ae 2025-2037 | - LD Zi gee: 1358=2004 =2135 WAT) SOF | 9° 58 39.71" + 2060 ital a ae Wee } 2024=2037. 2=770=2135 SS 1351= 1369 4438/1439 1499/1500 Costa Rica Railroad Sy S9N18°35e 1992 Lomas del Mar (East) 943= 942= 1109 4408 ! | \2027=2030/ ‘| 2032=2134 952 >.> 298 BULLETIN 357 Inset C of Map 11 Cc 452,455/459 Bomba ee 668/677,758 ae 696/697 1729/1733,1983/1984 aoa 2066/2109,2162 ‘ 695 698 lai Agua ate: OLS) g' 460/462,685/691 1726/1728,1986 Banano River 1422, x 1488/1497 ~s 451,678,1764, 449/450,679/684, 2110/2116 1391,1456/1472, 1734/1735,1985 692 693/694 2128/2133 = RRR are 9752755:875 925/929 . gop? 83" 59.84" 934 83° 3' 43.74" 5 | APPENDIX B STRATIGRAPHIC SECTIONS ANTHONY G. COATES Smithsonian Institution Smithsonian Tropical Research Institute Washington, D.C. 20560-0580, U.S.A. The stratigraphic sections listed below were drawn with the computer program Logger version 5.0 (Rock- works, 1991), and include PPP numbers for all Carib- bean sampling localities to date. AR WN 0 9 ID 10. ale 12 a. 13: 14. 15), 16. Ie 18. iI@), 20. 21. 22. 23: 24. 25. 26. PANAMA CANAL BASIN . Sabanita to Payardi Margarita to Gatun Toro Point Pina Rio Indio Miguel de la Borda, 1 km to the East Boca de Concepcion, 0.5 km to the East . Concepcion, 0.8 km to the West at Zapato Bluffs . Calzones River, Eastern and Western Sections BOCAS DEL TORO BASIN Escudo de Veraguas, Northern Coast Escudo de Veraguas, Southeastern Coast Valiente Peninsula, Bruno Bluff to Plantain Cays Valiente Peninsula, Toro Point Valiente Peninsula, Toro Cays Valiente Peninsula, Southern Coast Cayo Agua, North Point, Western Side Cayo Agua, Piedra Roja Point, Western Sequence Cayo Agua, Piedra Roja Point, Eastern Sequence Cayo Agua, North Point to Tiburon Point Cayo Agua, South Nispero Point Bastimentos Island, Short Cut Bastimentos Island, Fish Hole, Eastern Sequence Bastimentos Island, Fish Hole, Western Sequence Solarte Cay, Western Tip Swan Cay, North of Colon Island Colon Island, Hill Point * When PPP numbers are given as a range, e.g., 2471/2476, the first number is the highest and the second the lowest stratigraphically. LIMON BASIN 27. Sandbox River 28. Carbon Dos (Dindiri) 29. Banano River 30. Peje River 31. Bananito River 32. Santa Rita 33. Chocolate to Buenos Aires 34. Empalme 35. Pueblo Nuevo, Cemetery 36. Lomas del Mar, Eastern Sequence 37. Lomas del Mar, Western Reef Flank Sequence 38. Lomas del Mar, Western Reef Tract Sequence 39. Vizcaya River Key TO SYMBOLS 2471 PPP numbers, which are assigned to PPP col- lecting sites*. They are referenced in the PPP Database, which can be accessed at the internet site http://www. fiu.edu/~collins|/ x Sample collected within measured section @ Sample not collected within measured section but correlated to its approximate stratigraphic level from nearby exposures [ All samples within bracket were collected from the same horizon I Sample was collected from various stratigraph- ic horizons within this range A Paleomagnetic sample was collected within measured section A Paleomagenetic sample not collected within measured section but correlated to its approx- imate stratigraphic level from nearby exposure R Paleomagnetic sample with reversed polarity N Paleomagnetic sample with normal polarity 9 Paleomagnetic sample with indeterminate po- larity BRE ed Ue) Cd BA) ed a AR GE |e soil alluvium conglomerate conglomeratic stringers conglomeratic sandstone conglomeratic interbeds limestone shaly limestone silty limestone sandy limestone coral thicket calcarenite recrystallize limestone BULLETIN 357 Key to Lithologies and Sedimentary Structures sandstone bioclastic sandstone silty sandstone c. sandstone sandstone & siltstone interbeds sandstone w siltstone interbeds pebbly sandstone sandstone stringers sandstone & conglomerate interbeds sandstone & siltstone tuff basalt bentonite ash beds basement volcanic breccia columnar basalt fault unconformity siltstone pebbly siltstone sandy siltstone & claystone sandy siltstone siltstone w sandstone interbeds siltstone stringers conglomeratic siltstone corals serpulorbis shells bioturbated burrows arthropod burrows coral heads pinna bed logs aed shale mudstone/claystone sandy shale sandy mudstone siltstone & claystone claystone w sandstone interbeds shale w limestone interbeds claystone or mudstone stringers cross beds boulders spherical concretions concretions wood fragments distal turbidites sandstone channel lense with shell turbidite and conglomerate interbeds Ss LL] [LLL Lay (LS a LL PANAMA CANAL BASIN Sheila. Or Rovere 217 0 210 0 203 0 151 0 132 1 113.2 944 nh PPP number 989= 1308= 1503 FOS, OO, OT. TOG 21077, 1079/108 =1086, 1087-1307 =2163/2165 Li thology Change of Scale 1035 1078= 1305 = 1306-485 1030, 1033 1501=229=230 984° 2166/2168 1310 1075 1038 1039+ 92 221, 220- an SS 6 7 11 219, 490 A 218 ae 1/4, 231/233 8/10, 222, 1040) +}e-.o -e- o- , ( n¢ AG CNG Ae N¢ a fel 1 9 Cr C9 ¢ CN ¢ | GG ay dAGNd AAG AGN MEM CNC NC AC a Ac alc AD WN A CA JAG Ae A Gy ( “. Ile I ol ’ ’ qi" SECTIONS: COATES 301 Section 1 Description NEAR GATE IG IRAMmARDIE RERENERY TOP OF GATUN FORMATION SANDSTONE & SILTSTONE Fine, ton weathering with huge abundance of diverse whole mollusks, extensive bioturbotion, A eral, burrows and Thalassinoides systems, turritellids dominant Scattered large concretions NO EXPOSURE: 49m SILTSTONE Tuffaceous, dark grey-green, rich in mollusks SANDSTONE & SILTSTONE Concretions in upper port, burrowed in !ower port | NO EXPOSURE 45 4m ] SANDSTONE & SILTSTONE Silty ond sandy, snails) with Block Factory NO EXPOSURE 34 9m rich_in whole mollusks (bivalves ond scattered 1-15-cm concretions Section 1s opposite the Cativa SILTSTONE: Dork grey-green, richly Fossiliferous. Many whole orticuloted mollusks, (snails, ivalves) and much comminuted shel! hash, of ten packed in large burrow systems, pervasive bioturbation SILTSTONE: With vague, large concretions, densely evenly packed, with large, mostly whole shells SANDSTONE: Grey-green, tuffaceous, Fine sondstone with feldspar and hornblende, moderately indurated, with calcareous cement, leaching densely | packed whole and Fragmented mollusks. Abundant concretions SILTSTONE & SANDSTONE Grey-green, } ond shel! hash packed in tufFaceous, with intense burrow mottling large arthropod burrows SILTSTONE & SANDSTONE Semi-concretionary to almost hard beds, tuf Faceous. with Finely comminuted shel! hash and scattered mo! lusks SILTSTONE. TufFaceous, grey-green, coarse, with large, anastomosing arthropod gallerys, pervasive burrowing shel! hash dominant, bivalves Filling burrows in the lower port SILTSTONE. Highly orthropod-burrowed, tufFaceous, with shel! hash concentroted in burrows ond large mollusks inside abundant concretions Extensive pi! low-mound-shaped ene Re mOnoh y zone With tendency to Form hard beds No trace of bedding pervasive rework ing SILTSTONE: Rubbly, clayey, tufFaceous, with mollusks of very high diversity SILTSTONE. Extensive burrow systems, many whole mollusks, pervasive bioturbation, abundant shel! hash SILTSTONE & MUDSTONE One dense shel! bed hash SILTSTONE Muddy ond si'ty, 5-10-cm concretions many whole shells and dense she! | scattered shells and hash, with occasional —— BULLETIN 357 Change of Scale Seen fe aacontce ee) NO EXPOSURE 68m ie el - 55 [ - 4% - 3 im 27 [ SILTSTONE & SANDSTONE: Deeply weathered, orange-ochre, laminated, with more omer. or_less porallel bedding CROSS BEDDED SILTSTONE: Laminated, tuffaceous, with sublenticular bedding, L } arthropod burrows Common unidirectional Foreset cross bedding CONGLOMERATE: Volcanic conglomerate with 1-2-cm -in-diameter clasts The rc } matrix is Formed by highly weathered tuffaceous arkose eae EAT: BASE OF GATUN FORMATION PANAMA GmiINAls IBiAiSaN SeeGlome ce hcircciminte lomo im m PPP number Li thology Description 671 0 Sa P = | ee a ee 2mm (a) A) RMA ON SILTSTONE & SANDSTONE: With mollusks, scattered hash, whole mollusks, arthropod burrows packed with shells [ 7 le ; NO EXPOSURE 30m ANDSTONE Grey-blue, silty, with abundant, diverse mollusks + 6100 1: t 599.0 lim 988 0 33228 obs “ 5 = } vid cor BENDSTONE uFFaceous, Fonming a Headecrcane with eS GS a Pecten, alectryonid oysters, gastropods NO EXPOSURE: 95m ls, cones, olives, archetectonid 166 5 1295 24.0 190 170 16.0 10 140 130 20 uo 10.0 g0 80 70 60 50 4.0 30 20 10 SSP SE SSS Change of Scale SECTIONS: COATES 303 contd Section 2, SILTSTONE: Biocalcarenitic, mollusks Fine grained, clayey, burrowed, rich in scattered SILTSTONE & SANDSTONE: TufFaceous, with pervasive bioturbotion, abundant mollusks, turritellids, noticids ( ¢ i ( ( ¢ ¢ ( ( ( ¢ ( SILTSTONE: Massive, tufFoceous, clayey, with abundont, diverse mollusks Lorge calcoreous concretion horizons near the top SILTSTONE: Induroted SANDSTONE: Dork brown, weathered, clayey, with muscovite, quortz, Feldspor, abundant shel! hash and micromollusks, sporse whole mollusks aces el Scale NO EXPOSURE: 209m Change of Scale 37 ——- SILTSTONE NO EXPOSURE SANDSTONE & SILTSTONE NO EXPOSURE Abundont, diverse mollusks 114m Silty. Fine grained with scottered sporse mo! lusks 73m CLAYSTONE: White-weathering, Fine grained, bentonitic, interbedded with deeply weathered, crystalline, tuff-like material SANDSTONE & CONGLOMERATE INTERBEOS: Greenish-grey weathering arkosic, tufFaceous sandstone ond Fine-grained conglomerate with extensively leached shel !s NO EXPOSURE: Unknown thickness For covered interval SILTSTONE: Grey-green weathering ashy, tuffaceous, with abundant Feldspor, hornblende, occasional colcoreous and Frequent |ignitic Fragments Some ports may be shal lon-waterlain crystal tuff. No mollusks NO EXPOSURE: 4 Sm SILTY CLAYSTONE: Pole grey weathering massive, bentonitic, with scattered smal! porphyroclasts SILTSTONE Brown weathering, tuffaceous, indurated, lower port is rich in turritellids, cones, etc with abundant scattered mollusks The CONGLOMERATE: Rubbly conglomeratic layer with o chloritic, volconiclastic, sondy or silty motr ix SANDSTONE & SILTSTONE INTERBEDS: Tuffaceous sandstone ond coarse si|tstone with scottered mollusks, reworked by bioturbotion Upper port With densely scattered turritellids, cones SILTY SANDSTONE: Silty, tufFoceous SILTSTONE: TufFaceous With scattered shel! hash CLAYSTONE- Grey-green weathering bentonitic claystone SANDSTONE & SILTSTONE INTERBEOS: Highly burrowed, mottled, tuFFaceous, with strongly comminuted shel! hash, coorse black volcanic grains, and abundont turritellids Slightly indurated SILTY SANDSTONE SILTSTONE: Dork brown weathering tuffaceous, burrow-mottled, with pervasive rework ing Scattered large bivalves and shel! hash NO EXPOSURE: Unknown thickness For covered interval BASE OF GATUN FORMATION 304 BULLETIN 357 172 129 86 15.0 14.0 13.0 120 10 1076= 1088/1093 —| 1656, 1657 —————_—__» 1639 PANAMA CANAL BASIN Section 3 lero” Rei SECTION AT_AMBUSH ROAD TOP OF CHAGRES FORMATION SANDSTONE Ton-weothering massive, very thick bedded (not closely inspected- high in clifF) SANDSTONE: Ton-weathering, massive, very thick bedded (not closely inspected- high in cliff) Dense Thalassinoides burrow systems SILTY SANDSTONE Coorse, poorly sorted, with scattered, disoriented, disarticulated bivalves and rare Strombina NO EXPOSURE: 20m SANDY LIMESTONE Grades from unit below into alternating coquina and shelly, coarse sandstone TOP OF TORO MEMBER Ha LIMESTONE: Shelly, barnacle and echinoid coquina with coorse sandstone to grit-sized clasts ond plates, white-cream ledging unit CROSS BEDS Thin to laminated bedding has steep, prograding foresets abruptly truncated Cross beds ore alternating clayey, 31!ty sandstone, ond blue-grey, 22, 23 1659 1172/11742508 ———s shelly, barnacle coquina Beds ore 2-50cm thick ai LIMESTONE Shelly, barnacle and echinoid coquina with coarse sandstone to 2b IE ar: 12 i 1 grit-sized clasts and plates, white-cream ledging unit BASE OF TORO MEMBER BASE OF CHAGRES FORMATION NO EXPOSURE- 10m SILTSTONE: Blue-grey, with scattered large mollusks TOP OF GATUN FORMATION . NO EXPOSURE: 87 25m SANDY SILTSTONE: Conglomeratic, massive, unbedded, tufFaceous, with abundant shel |_hash and_ oysters; il itel Vv SECTION BELOW GATUN DAM” HYDROELECTRIC PLANT. SANDSTONE: Highly arkosic, tufFaceous. Pervaded with col!ianassid burrows which contain volcanic pebbles CalciFied, scattered large Flot mounds (> 1m in diameter) of siderasteroid corals SANDSTONE Green-brown weathering, tuffoceous, with scattered volcanic pebbles ond large, widely spaced, calcified arthropod burrows containing volcanic pebbles C. SANDSTONE: Massive, coarse, arkosic, quartzose, tuffaceous Occasional stringers of shells Tan-brown weathering mostly unfossi|iferous SANDSTONE Medium-grained, arkosic, ond a conglomerotic base tuFFaceous, with scattered wood Fragments SANDSTONE: Many armoured mud or sand balls with volconic pebbles ina fine to medium-grained, tufFaceous sandstone matrix Upper port has pervasive, small, callianassid-type burrows forming galleries Patchy, densely conglomeratic ockets im the sandstone SANDSTONE Tuffoceous with numerous 1-cm-diameter burrows BASE OF GATUN FORMATION NO EXPOSURE. Unknown thickness For covered interval SECTIONS: COATES 305 PANAMA CANAL BASIN section 4 Pina a PPP number L thology Description 390 IOPSOE TCHAGRES bORMAmLON SANDSTONE: Coarse, leached Section about 2km west of Chagres River at First a \ head! and NO EXPOSURE: 10m b [ 168-108/108 ——_. ne SILTY SANDSTONE Volconic, quartzose, with lithic and Feldepar grains, 364 1690 ae LS scattered thin disarticulated bivalves and pervasive bioturbation, 5-10-cm- [ 1651-1652 ————# SES diameter arthropod burrows SECTION AT & SW OF PINA. in $1 a a 5 5 = SANDSTONE: Grey-green, silty, coarse, with scattered mollusks, articulated l 1b of ae thin-shelled bivalves, Pecten abundant. Dense gallery systems of arthropod burrows NO EXPOSURE: 10m C. SANDSTONE. Volcanic and quartzose, with leached mollusks common SECTEGON At SANTA MARTA CREEK NO EXPOSURE: 160m [ 5 Change oF Scale SANDSTONE: Indurated, greenish-grey greywacke, poorly sorted, subangulor with lithic, Feldspathic and chloritic grains 1653, 1654 SECTION ON RTE SI10 & S1 NEAR SPILLWAY CONGLOMERATIC SANDSTONE: Pebbly, coarse, volcanic, packed with Fine shel | hash and mo! lusks vel Cie welts) nSstlell SANDY LIMESTONE Grades From unit below into alternating coquina and shel ly coarse sandstone LIMESTONE: Shelly, barnacle and echinoid coquina with coarse sandstone to grit-sized clasts and plates, white-cream ledging unit [ 130 CROSS BEDS: Thin to laminated bedding has steep, prograding Foresets abruptly truncated Cross beds are alternating clayey, silty, volcanic sandstone, and blue-grey and shelly barnacle coquina Units 2-S0cm thick LIMESTONE: Barnacle and echinoid coquina with coarse sandstone to grit-sized clasts and plates. White-cream, ledging shelly BASE OF TORO MEMBER BASE OF CHAGRES FORMATION come Lom SANDSTONE: Volcanic, Fine-to medium-grained, blue-green TOP OF GATUN FORMATION SANDSTONE: Volcanic. Upper part coarser (grit-sized), with mollusk hash, bentonitic mudstone LL Pee MUDSTONE Indurated, very Fine grained, with bentoni tes i | NO EXPOSURE. Sm r gS SILTY SANDSTONE: Fine grained, with scattered smal! mollusks and shel! hash BASE OF GATUN FORMATION i Boo NO EXPOSURE: 60m 306 PANAMA CANAL BASIN Rio. Lmao a PFP number p if IsW/166 5 L159 Lite L 10 24 Ae [ © | 5 ———{! eereee Ls Se | | 153/163 —| L 1638 BULLETIN 357 Section 5 Desorption SECTION NEAR JIMENEZ CREEK TOP OF CHAGRES FORMATION RIO INOIO FACIES SILTY SANDSTONE Medium-grained, white-cream weathering, with abundant 1-2-cm-diometer burrows oxidized and weathering out Plant Fragments common NO EXPOSURE: 90m SILTSTONE W SANDSTONE INTERBEDS Grey-green, volcaniclastic, burrow mottled, with scattered macromollusks Thin, calcareous, hard beds SILTSTONE W SANDSTONE INTERBEDS Grey-green, volcaniclastic, burrow mottled, with scattered macromo!lusks Thin, calcareous, hoard beds SECREON TAT REO ENDO NO EXPOSURE SOm SANDSTONE W SILTSTONE INTERBEDS: Massive with scattered, whole mollusks SE CrieceOoNeNeAR GOB EAl ne OF rel Or ENE CLAYSTONE & SILTSTONE: Blue-grey, burrowed, with abundant mollusks NO EXPOSURE: 10 Sm SILTSTONE: Blue-grey, clayey siltstone and silty claystone BASE OF CHAGRES FORMATION REOPENS TOREAaAGCEES m ta i ol L 2.0 10 T IP NORTH COAST PANAMA Section 6 inquei—de-la-Borda al Kkm= tothe bast PPP number SECTIONS: COATES 307 Li thology Description UNNAMED FORMATION CONGLOMERATIC SANDSTONE: Coarse, volcanic, with extensive arthropod burrows ond scattered molls Contains 6" shel! bed NORTH COAST PANAMA Section 7 Eocagee Concepeion, 0.5 Km tothe Hast PPP number <—SENS e CLAYSTONE & SILTSTONE: Silty and clayey, greenish grey with pervasive bioturbotion and scattered Frogmentory Fossils, dominantly mollusks SILTSTONE- Greenish, grey, clayey, with calcareous concretions Concretionary il zone tends to become ao continuous bed rich in whole mollusks CLAYEY SILTSTONE: Intensely burrowed, with Frequent anastamosing arthropod burrows 1-1 5"-diameter Many longitudinal and straight burrows (>20 cm) Li thology Description UNNAMED FORMATION C SANDSTONE: Massive, coarse, with crystal, tufF-like textures Neor base molds and casts of mollusks. Evenly scattered throughout ore irregular to spherical calcareous concretions, ranging 5-15cm in diameter C. CONGLOMERATE: Coarse, volcanic, of semi-rounded, subangulor mostly basalt boulders (up to 30cm) ASH BEDS: Laminated and Fine pebbly, with increasingly Frequent Fine conglomerate layers near top SILTY SANDSTONE Lominated, with coorse burrows and smal | Neptunian dykes mixed with occasional volcanic sandstone and conglomerate Some Finely cross bedded but most wel! laminated and not bioturbated 308 BULLETIN 357 NORTE COAST ‘PANAMA Section 8 Esncepeion, OFS kn to) the West ot Zapato siltias m PPP) nlinber Li thology Description 150 - + ' E UNNAMED FORMATION L 44 =| r | SILTY SANDSTONE Section goes approx 20-30m higher in clifF than [re 23 Dawa] included here Coarse point-bor channels = ae =| F 4 SILTY SANDSTONE TufFaceous, Fine grained, containing abundant large ic ee a logs and trunks, some with giant encrusting barnacles, 1-2cm in (= da) diameter Be F | SILTY SANDSTONE: Tuffaceous, Fine grained, 6-18" large pockets densely Ise 200 al packed with mollusks Large shark’s tooth found - 70 r AG C. CONGLOMERATE Coarse volcanic Ie = SILTSTONE» Greenish-grey, volcanic, with thin clay units which might L 20 | 167 eee Se ei eee be bentoni tes - 40 4 == = = L Sa sj = ee C. CONGLOMERATE: Coarse, volcanic, boulders up to 20cm 1 ADAM AAT ALAM ALA SILTSTONE: Greenish-grey, becoming lominated near top, with rore costs Ge eb sy 2S SS of mollusks, lenses of lignite and small wood Fragments, ond scattered ic Lia) ee pebbles ond sandstone clasts jm 4 = == Nominee CORSH SPaANahiA Sect Ome Calzones River, Eastern and Western Sides m PPP) number Li thology Description |= 16 0 Ee ae 5585259 gene ee UNNAMED FORMATION IE 4 a - CONGLOMERATIC SANDSTONE: Massive, coarse, volcanic, poorly sorted 2 we 4 Zones of |imonitized mollusks, echinoids and one whole crab IRQ lm 4 CONGLOMERATIC SANDSTONE: Coarse, massive, poorly sorted, volcanic, Ip eu | tufFaceous, with silty sandstone motrix Scattered molds and casts of fr Ho 4 thick-shelled mo! lusks - io = IP so ai] = lon ee Ely tien rel |= 60 |= ee pees ml - 40 = Ic al CONGLOMERATIC SANOSTONE: Weothered, tufFaceous, bioturbated volcanic jE IN| conglomerate with mollusks and corals, and clustered in pockets LC CONGLOMERATIC SANDSTONE Densely packed with wel! preserved, |ow- L ay | i diversity bivalve Fauna and shel! hash SECTIONS: COATES 309 Escudo de Veraguas, m PPP number 1283 (w), 171 ots 1249 z 1244 B0 1974 ire 86 5 a 13 86.0 124 R 14 85 5 rel H 1s 85 0 845 840 83 5 83.0 82.5 82.0 7269 815 a 81.0 2268 Ds Z261 2266 80.0 ae a 93.5 1232/1234 Im) = R 1, | 73.0 179, 358/33, ae 1222/1231, 2170/2179, 2279 78.0 75 710 76.5 2213 A 76.0 755 218 A 75.0 749.5 an A 740 735 zn as BUChS BEL TORO BASIN Northern Coast Li thology LSet SS 2F 25 S ree een Beate G2 = a Ole a ee ae Sa exer aaa Sd IS RNS Section 10 Description SECTION ON W. COAST INMEDIATELY SOUTH OF LONG BAY POINT CLAYEY SILTSTONE: Blue-grey, pale, tan-weathering silty clay Intensely ond distinctly burrowed. Mostly callianassid-type Fauna similar to, but much sparser than, PPP 168 Mollusks ore diverse, but Fragments ore o greater percentage relative to whole than in PPP 168 Thin-shelled, Fragile, spatangoid? echinoids are very common, but corals ore mostly absent Logs are rare or few TOP OF ESCUDO DE VERAGUAS FORMATION SILTSTONE: Marker horizons are distinctively weathering slightly more massive units, blocky. Appear to be indurated relative to burrowed zones, possibly minor disconFormities CLAYEY SILTSTONE: Grey-bluish, with diverse mollusk Fauna ond a very common cornute coral. Also common 1s a large pteropod, Cavolina. Lines of irregular calcareous concretions and indurated arthropod burrows are typical No echinoids or logs | NO EXPOSURE: Small but unknown thickness of section not exposed Somple numbers come From exposures along western port of North coast and ore estimoted to fall within this zone SILTSTONE: Bioclastic, clayey, sondy, with pronounced 10-cm-thick burrow zones at top (ie IGRIGOLE MOM AE TENE II CECHEC TCC £ (eA ENE (CIE SE Xe (GANG IGS IG Ik 8 COs Coe Coe ashok Gin G (eee Ge Gal Go nk Ga Ga Gaek Co CeO CORA ICES VYIVVVU UU ; v VYYVYUYUYUYUYUY F SANDSTONE: Silty, bioclastic, with diverse mollusks, scattered and disoriented, ond Cupuladrians SECTION BELOW IS 1.3 KM TO EAST ALONG NORTH COAST, ON EAST SIDE OF BREACH IN REEF AND IS EXTRAPOLATED TO LIE BELOW WEST COAST SEQUENCE . CORAL THICKET: Coral biostrome with slender branching corals, Madracis?, tylophora?, mussid, sand dollars, mollusks 310 BULLETIN 357 Change of Scale Section JG) contd Sens SILTSTONE & SANDSTONE: Blue-grey, clayey, with Fine shel! hash, pervasive 72.0 bioturbation eo MUDSTONE Silty, bioclastic, grey, with scattered mollusks, pervasive aly bioturbation, Fine hash - 690 E 68 0 b ore 24983, 2271 CLAYEY SILTSTONE. Massive, grey, with scottered mollusks, black, larger 66 0 2270 basalt grains, pervasive bioturbation 65 0 640 eeb4 2882 SILTY CLAYSTONE grey-blue, massive, with pervasive bioturbation, abundant, Pata small, thin-shelled mollusks, tellinids, pteropods EF é20 Fr go NO EXPOSURE 16 5m E 60 0 Change of Scale (Saat 2 eeoeeB an F SANDSTONE: Silty, clayey, with scattered diverse mollusks, pervasive GD al gaa a bioturbation, regular horizons of large, irregular concretions eee salecay A CLAYEY SILTSTONE Blue-grey, with occasional shel! Fragments, regular 20 > 2959 - horizons of smal! concretions : “iy 4 CLAYEY SILTSTONE Grey-blue, with regular, subcontinuos concretion horizons, OM Sb SS Few shells and very sparse shel! hash 33:0 os} NO EXPOSURE : #0 4 70 4 Change of Scale 23.0 E CLAYEY SILTSTONE massive, khaki weathering blue-grey when Fresh, iG ev pervasively bioturbated Rich in very small mollusks, often densely = eo aggregated in arthropod burrows Common strombids, rather larger gastropods, 20 0 and thin-shelled, delicate bivalves 19.0 CLAYEY SILTSTONE: grey-blue, with regular horizons of branching arthropod aa galleries, massive, bioturbated with Few whole mollusks, scattered Fine hash F SILTY CLAYSTONE: grey-blue, with irregular concretions, scattered smal |, ie eee solitary corals and rorer larger mollusks, rich in shelly hash and smal | - 160 mollusks - 150 rt 140 K fe isto SILTSTONE: Clayey, grey-blue, burrow mottled, with scattered mollusks, fe yarn pervasive bioturbation lz % SILTY SANDSTONE. ton weathering, Fine grained, massive, with pervasive Eee \ bioturbation, all Fossils randomly oriented, abundont horn corals, diverse t 100 mollusks F 0 NO EXPOSURE. 2-3m. Ie Se SILTY SANDSTONE: light grey, and clayey siltstone, with concretions rare or = 7.0 a absent F 60 z SILTY SANDSTONE: Light grey, Fine grained, and clayey siltstone, with J = = = gnt grey g y F Boy | Sere tess LAS ee EA spherical concretions and occasional whole mollusks. Upper zone of irregular ee ze NAS | ane 5 concretions and thalassinoid-type burrons r 1235, Rog. [Pee Ee a SANDSTONE & SILTSTONE khaki weathering, massive, shelly and Fine grained Ee ee with large logs and rore, whole mollusks Pervasive Thalassinoides burrows ie 20 Note’ This Forms N. point of central north coast of Escudo F at BASE OF ESCUDO DE VERAGUAS FORMATION m 20.0 190 180 170 16.0 15.0 14.0 13.0 12.0 11.0 100 30 80 70 60 5.0 4.0 30 20 10 enim Tin lie is oe Cea LL ee a po er SE ee ee nS Ee ee neler en Sete 141 IBOCAS DEL TORO BASIN Ssetios ai Bselide) Ge Veraglas, Soutbeasterm) Foust PPP number 2294 2293 2292 2291 2290 2189/2187 168/170- 478 431, 2180/2181) 479/483 2289 2288 2287 2286 2183-2182 | 2285 2284 2283 2282 2281 2280 SECTIONS: COATES 311 Li thology Description TOP OF ESCUDO DE VERAGUAS FORMATION ¢ a (' eVse VS CLAYSTONE & SILTSTONE Grey-blue, muddy, with sandy pervasive SS SI bioturbation, scattered mollusks NO EXPOSURE: Sm CLAYEY SILTSTONE: Rich in mollusks, ond horn corals, bioclastic, mottled with pervasive bioturbation BASE OF ESCUDO DE VERAGUAS FORMATION 312 m 1957 0 1956 0 1965 0 1959 0 1953 0 1952 0 1951 0 1950 0 149 0 1348 0 147 0 1346 0 145 0 1944.0 143 0 12.0 11 0 1990 0 1939 0 1938 0 1937 0 1936 0 1935 0 1334 0 1933 0 1932.0 1931 0 1930 0 Tiemann Un De cae TTT a A La a 1929.0 r 1780.0 1762 2 1744.4 fF 17266 f- 1708.8 f- 1691 0 1673.2 eeooong BULLETIN 357 8 3 8 8s = BUCAS DEO TRG BrsiNn section 12 Valiente Peninsula Brune Buri stosR lomtcaime tays Description SECTION AT BRUNO BLUFF TOP OF BRUNO BLUFF MEMBER AND SHARK HOLE POINT FORMATION CLAYEY SILTSTONE Massive, dark grey, with scattered mollusks High in cliff a ae Not examined directly SSS CLAYSTONE Dark, tight, with 25-cm sandy layer of which bottom 5 cm crammed S= 25 sa with mollusks and bryozoans, many whole ond in stable positions 7Storm SS SS deposit Bie ty ae SILTSTONE Massive, bedded, in 15-20-cm units, well sorted, dork grey, with = = tight clay matrix Occasional scattered mo! lusks : a SILTSTONE W SANDSTONE INTERBEDS: Medium-grained, bedded ond extensively bioturbated with small vertical burrows Thin sandy layers are often pulled apart and form load casts SANDSTONE Introformational slump with siltstone clasts and ae Folds Shelly matrix, mica common and Finely distributed organic debris SILTSTONE W SANDSTONE INTERBEDS: Pole grey, with tight clay Fraction Coorse silty sandstone layers. Burrows abundant with scattered smal! mollusks NO EXPOSURE 2m SILTSTONE Massive, grey CLAYEY SILTSTONE Irregulorly lensing, alternating well sorted siltstone ond silty clay, slumped channels, highly bioturbated ond ripped up, irregular, dork and light grey clasts SILTY CLAYSTONE: Grey, alternating with siltstone Irregular, knobby concretion horizons at base BASE OF BRUNO BLUFF MEMBER NO EXPOSURE 157m Chongenom seailic ~—j [es -o— SEG TONRAlm ORDEBESSErOm NT SHARK HOLE POINT FORMATION SILTY CLAYSTONE Paras) weathering, with Fine silt ond irregular concretion horizons. Precise stratigraphic level = PPP385 not noted Te laitny ALG With conchoida! fracturing and with very sparse, scattered mol lusks NO EXPOSURE: 130m 16% 1599 1518 wr 1452 1430 Change of Scale —it | 8 2331/2322 PPP Ph Ph bh be he ee Change of Scale N| Scattered mollusks, including SECTIONS: COATES 313 Section 12, contd SILTSTONE: Massive, Fine, dark grey, with tight clay matrix, sporadic large Dentalium ond other snails, uncommon bivalves SECREONSALASHARK HOLE POINT NO EXPOSURE — 4m CLAYEY SILTSTONE: Dork grey, massive, Finely micaceous, tight, with scottered mollusks Prominent concretionary horizon near the base BASE OF SHARK HOL POINT FORMATION NO EXPOSURE 8 33m TOP OF NANCY POINT FORMATION SANDSTONE: Possibly lensing rich shel! bed NO EXPOSURE 1 66m CROSS BEDDED SILTSTONE: Micaceous, green, lominated, scoured ond cross bedded CLAYEY SILTSTONE Massive, ene green-weathering, grey-blue, tight NO EXPOSURE 4 66m SANDSTONE & SILTSTONE Lominated, greenish CLAYEY SILTSTONE Sparsely FossiliFerous, massive, micaceous Dental ium commo ond other mollusks sporadic NO EXPOSURE: 3 33m SILTSTONE: Fine grained, massive, brown weathering micaceous, and dark blue claystone Lignitized log Fragments 3’ long to 6" moximum diameter CLAYEY SILTSTONE Mossive, micaceous, with lenses and scattered mollusks CLAYEY SILTSTONE Massive, micaceous, with scattered mollusks NO EXPOSURE: 57m SANDSTONE & SILTSTONE INTERBEDS Laminated, silty clay with scottered rare mollusks. Coarse volcanic sandstone channe! up to 1m thick CLAYEY SILTSTONE: Micaceous, with occasional mollusks CLAYEY SILTSTONE: Grey-blue, laminated Channels with volcanic cong|omeratic sondstone containing large clasts of laminated siltstone and claystone NO EXPOSURE 35m SANDSTONE: Muddy, coorse with bimodal, grain size, volcanic and calcareous grains, wood logs and fragments, pervasive bioturbation, occasional mollusks SANDSTONE: Massive, muddy, coorse, with muscovite, Fine shel! hash, volcanic grains, pervasive bioturbation, burrow mottling bimodal grain size 314 r 146 me JE r Lek | 1333 fe ees 1230 - len 1260 f 1245 1230 flak 1200 BULLETIN 357 Section Loc omGel SANDSTONE: Massive, muddy, coarse, with muscovite, Fine shell hash, volcanic grains, pervasive bioturbation, burrow mottling bimodal grain size SANDSTONE: Muddy, bioclastic, volcanic, with bimodal grain size, muscovite, pervasive bioturbation, occasional small mollusks, lignite, burrow mott! ing SILTY MUDSTONE: Packed with lignite Fragments, micro mollusks SANDSTONE Muddy, volcanic, packed with lignite Fragments, micro mollusks NO EXPOSURE 16m SANDSTONE Muddy, greywacke SANDSTONE Thin bedded, muddy, coarse, with bimodal grain size, basalt granules, lignite, sparse mollusks, thin silty mudstone units, burrow mottling Bioclastic SILTY MUDSTONE: Medium grained, evenly bedded, muddy sandstone ond si|tstone that ore turbiditic?, lignitic, burrow mottled CLAYSTONE: Tight, blue, silty, with burrow Fills, occasional massive coarse NO EXPOSURE 20m SILTSTONE: Muddy, |ignitic NO EXPOSURE 18 Sm Change of Scoaille ALN AR ile ries z ea SS y TS 4 Ry Ger Mee tase, pee ee eee 2405/2419 AES a ae ee ae a w ww Vy a | Sy ea ees 2193-401 . ee ASSES — ~ ¢| A, ee PA porercccacceccecereuncore a a | [es a 2404 yO ee: ies 244-2515 /2501 | 2436=2502/2503 cy 4 2496/2499= 2501 ~ 4 sandstone units | exe 9 240 Change of Scale | 3, 2443 2444/2451 | rs 1 2492/2460 — | 4, S 2461/04 | e————#= 4093——___, 410 — a a i Fi SANDSTONE & MUDSTONE: Sandstone 1s pebbly, muddy, burrow mottled, with tubular burrows, mudstone is laminated sublenticularly, bioclastic, lignitic and micaceous NO EXPOSURE 19m >> PPh PP Phe SANDSTONE Muddy, lignitic, micaceous, thick bedded with pervasive broturbat|on | 4L1= 12531714 : 2189 >> >> bpp SANDSTONE & MUDSTONE: Clean, well bedded, burrow mottled sondstone alternating with slightly silty mudstone SANDSTONE Muddy, burrow mottled, with volcanic granules, fine lignite, and sparse, whole mollusks MUDSTONE & SILTSTONE Muddy, with pervasive bioturbation, common whole mo! lusks 724.0 651 6 615 4 SS ee 2324-2314 th <<) 2325-2331 2238-2312 —1288= 1969-1713 2150=2188=2484~q Scale SECTIONS: COATES 315 Section 12, contd CLAYEY SILTSTONE: Micaceous, dork blue Occosional coorse, volcanic sandstone channels near base. Abundant shel! hash and scattered mollusks Large Dentalium. Slabby concretion horizons SANDSTONE: Coorse grained, thick bedded, volconic, vaguely laminated 2313 PP PPP rrr hr hr hb bb>pprrbrrrrrrprerr Change of Scale See ees Ue Oa Soi oe NESSES AX A A Sea me me ee 1 Q queen testutiies Sear a ieee air ose ie Gane q gq as SANDSTONE & SILTSTONE Medium bedded, blue, coarse, quartoze, volcanic, alternating with clayey, silty sandstone ond clayey siltstone, with scattered mollusks BASE OF NANCY POINT FORMATION NO EXPOSURE 398m SANDSTONE: Indurated, quartoze, chippy, pervosively bioturbated, with shelly hash and scattered mollusks, that ore hard to extract TOP OF TOBABE SANDSTONE SANDSTONE: Indurated, quartoze, chippy, pervasively bioturboted, with shelly hosh and scottered mollusks, thot ore hord to extract. Discrete tholossinoid burrows SANDSTONE Induroted, quortoze, chippy, with uncommon mollusks and scattered, vertical, circular-diameter burrows. Scottered to abundant spatangoid echinoids CONGLOMERATIC SANDSTONE Polymict, poorly sorted and vaguely and streak: ly bedded, pebbly mudstone interbedded with conglomerate Clasts rounded, 10-12 cm in diameter BASE OF TOBABE SANDSTONE ee MUDSTONE: Blue-grey, infilling hummocky surface of the basalt No borings TOP OF UNNAMED VOLCANICS BASALT Blue-black, massive, usually columnor NO EXPOSURE 10m SANDSTONE Thick-bedded alternations of coorse, volcanic boulder spreads ] BASALT. Columnar i CONGLOMERATE. Boulder supported, with interbedded coorse sandstone lenses and stringers of boulders. Clasts 6-20 cm in diameter, roughly sorted in each VOLCANIC BRECCIA: Basalt Flow breccia CONGLOMERATE: Coarsely stratified, boulder supported, volcanic NO EXPOSURE 30m C. SANDSTONE Channeled and high-angle cross beds of coarse sandstone alternating with Fine pebble conglomerate Spherical concretions, occosional long tubular, vertical burrows 91.0 0 170 181 Change oF Scale e ad = 2 ’ e 4 ee 2 BULLETIN 357 Section 12, contd. C. SANDSTONE: Channeled and high-angle cross beds of coorse sandstone alternating with Fine pebble conglomerate long, tubular, occasional Spherical concretions, vertical burrows C. SANDSTONE Often wel! !ominated with channels and lenses, 10-15-cm-thick pebble beds Angular, sond supported, basalt grains 1-2 cm, sporadic large basalt boulders in 20-30-cm-thick breccias alternating with Pebbles SJ DS [SFa Gad BEER PEE rey QV BY AT Y CORILOEZORS (So KS) “ se] y > De A IBS 2 DS Zl Se 134A o. AVLAv> Av> NO EXPOSURE: 30m VOLCANIC BRECCIA: Bosalt Flow CONGLOMERATIC SANDSTONE Pebble, volcanic sandstone matrix supported, alternating with coorse NO EXPOSURE 10m VOLCANIC BRECCIA Basalt Flow BASE OF UNNAMED VOLCANICS Valiente Peninsula, m PPP number 166 BOCAS DEL TORQ BASIN SECTIONS: COATES 317; Section 13 Description TOP OF UNNAMED VOLCANIC FORMATION LIMESTONE: Rubbly, bioclastic, silty, polls blue weathering to cream-colored micrite, with scattered, large, coral heads LIMESTONE Massive, vuggy, recrystallized, medium to thick bedded, rubbly, bioclastic, with scattered, large, coral heads, rubble zones CONGLOMERATIC SANDSTONE Grades into volcanic conglomeratic sandstone Grain supported volcanic conglomerate, with pebbles and large cobbles Dork blue conglomeratic, lithic greywocke dl LIMESTONE Rubbly, bioclastic, micritic Similar to upper unit 20-cm heads of diplorids and plocoid Favids, with mixed volcanic grains Strongly bioclastic in part with muddy motrix NO EXPOSURE 12m j 125 [ 104 =| VOLCANIC BRECCIA: Pebble supported CONGLOMERATIC SANDSTONE Basalt cobble, conglomerate with poorly sorted volcanic sandstone matrix SANDSTONE Coorse, Fine pebbly, volcanic CONGLOMERATIC SANDSTONE: Massive, very poorly sorted, with rounded bosalt boulders up to 2m in diameter. Matrix has laminated, ashy sandstone CONGLOMERATE Basalt clasts, subrounded to angular clasts up to cobble sized poorly sorted, channelled base CONGLOMERATIC SANDSTONE: Pebbles with tuffoceous matrix, and scoured undulating upper surface CONGLOMERATE: Basalt cobble supported SOE OIEROTE Volcanic. Cobbles ond pebbles are both angular and rounded, polymic BOULDERS: Volcanic, poorly sorted, large, (up to 40cm) rounded basalt blocks LIMESTONE Massive, bioclastic reef, with abundant, large mound and domed colonies of reeF corals up to 60-B80cm in diameter NO EXPOSURE 20m RECRYSTALLIZED LIMESTONE: Very hard, blue hearted, recrystallized, micritic biocalcarenite Shelly, with occasional Foraminifera No corals 66 0 55 0 iF 99:0 Ih, j= es}ife] f= 22.0 f= 200 2730/2731 + F BRECCIA: Flow breccia basalt BASALT: Mixture of lava Flows and Flow breccias BASALT Mixture of lava Flows and Flow breccias VOLCANIC BRECCIA- Basalt Flow breccia SANDSTONE: Blue-grey, |ithic greywacke SILTSTONE Dork blue~grey, SANDSTONE: Fine, lominaoted, sublenticulor in port with convolute beds, minor burrows, pull aparts, casts at base, stringers of wood muddy, rich in Foraminifera massive, laminated aminated CLAYEY SILTSTONE: Clayey si!tstone and si!ty claystone SANDSTONE Laminated Y MUDSTO Blue-grey ANDSTONE & LTSTON aminated, with bimodal grain size and abundant, coorse, volcanic grains CLAYEY SILTSTONE: Lominoted, dork grey 0 aminated to very_thin bedded rhythmic alternations of whi te- creom-neathering siltstone, 3-dcem thick, alternating with coarse, volcanic sonde.one Lomination of ten SUbilenmicullog to starved lenses, scoured iltstone pull aports Very Few burrows, smal! channels, requent wood Fragments SILTSTONE & CLAYSTONE Dork grey BASE OF UNNAMED VOLCANIC FORMATION 318 BULLETIN 357 Beeceas PEE. TeROmeaSzan Section £4 Valiente Peninsula, Toro Cays m PPP number Lithology Description 55.0 er ein cS FINGER ISLAND ok wl tsar NANCY POINT FORMATION Ss & LO wa) = 2S SY ° ° ire} ~ ‘6 BULLETIN 357 Section 31 Description TOP OF RIO BANANO FORMATION CLAYSTONE Uiepenced shells, more abundant layers of snails, whole clams No original bedding, pervasive bioturbation SANDY CLAYSTONE: Alternating silty SILTSTONE & CLAYSTONE: Clayey ond SU burrow mottled and sandy at top Dispersed Fine shel! hash Larger snails and bivalves BASE OF RIO BANANO FORMATION (do oo ° iClo¢ o o Se eo a co} © 2) ° ° oy or he - oa CIO COE Secs | Omeec Description TOP OF RIO BANANO FORMATION PEBBLY SANDSTONE. with abundont shel! hash (Fine) Glycimeris and other molls ond Amphistegina PEBBLY SILTSTONE: blue-grey, sandy, rich in shel! hash with Amphistegina and molls SILTSTONE rey, weathered, rich in shel! hash, bryozoa especially Cupuladria, and mol BASE GF REG BANANG BRORMATEON SECTIONS: COATES 339 LIMON BASIN - 704 L m [ 696 + ons-200 62 Bas | Ete. 4 4 UT i an a PPP number 668 1342/1344=1366 r “1440/1442=1435 L664 1498-2003 660 768-699 1348-1436 1124 Tala Slee Seale ee an DOD OD (Jo BS T ral a a T g 8 foe ere Eee) Er T T eal ee Ss 1 gy a a oat eriae toad 12a) a n 1396= 1363-1125 ci | =2016 T 2021, 2022, 2029 208 be 7 1345=1364/1365 3 72015 ee r ii [sr if. eee a SS ee Chocolate to Buenos Aires m Li thology Description Bs Sas er 4 } 41 | fl YVYOVVUVVVUVUVYU VYVVUVYUVYUVYUVYU VYVVYUVYUVUVYUVUL UV UYU U 7] VY¥VUYV one | VYYYUVYUVYUVUYYUY VYGVYUVUYUVYUVUS OMEN NE VYUYUYUYUVYUV YU! YWYVVVVVUVUVYUVVUY UY YYUVYUVYUVYUVUY VYGVYUVYUVYUVYYUVU! VYUVYUVYUOYUVUVYU YVYOVUVMUVU VV VS VYQVYUVUYUYUYUY a Uh AOA hae hack seh seh YVYGVYUGVYUVYUVYYVUS VYUVYUVYUVYUVYUVUS VYUVYUVVUVUV UV US UYUVUVYUYUVYUYYU IYOVOVUVOVUV US VYYVYOVYYVYUVYUVU! UYyVYUVUVYUVYUYUY VYVVYUVYUVYUYUY YS VYVVVVUVYUVYUVUY VY QVYyVYUVYUVYUVYU VYUVUVUVYUVYUVYU UVYYVVYGVYUVUYUVYUY VYUYUYUY YUU UVVYIV OV UVOVUY VWYUDVYVUVVUVYUVUVYU UY DVYUVYUYUVYUVYUY UVYVVYUY UYU UYU VYVDVUVYUYUYUVYUY BYOB UVOVUV VY VYBVY OV OVYUV UV UY VY QYUVUVYUVUYUY UYOVUVUVYUVYUVY VYUVYUYUYUYUYUY VYDVUVYUVYUVYUVY VYVVOV OV UVOVU BYU UVUVUVUV US VY UVYOVUVUVY wa vy Y uu L Section 33 TOP OF BUENOS AIRES REEF MEMBER TOP OF QUEBRADA CHOCOLATE FORMATION SILTSTONE aris green, M. annularis, Diploria, agariciids, A cervicornis, Thysanus, Caulastraea SANDSTONE Ton, Fine to medium grained, whole mollusks ond echinoids, M cavernosa, Diploria, A cervicornis, agariciids = CORAL THICKET Claystone matrix, blue, silty, with Porites overlain by A cervicornis CORAL THICKET: Green-blue claystone, with Porites, Colpophy!!io, Monicina, Dichocoenia, Montastraea, Stephanocoenia, Thysanus, Caulastraea, agariciids CORAL THICKET in silty claystone F. SANDSTONE: With mollusk hash, mollusks, Acropora, Stylophora, M annularis, M cavernosa 340 BULLETIN 357 Change of Scale Section 33, contd. 568 0 BASE OF BUENOS AIRES REEF MEMBER ORAL THICKET Silty claystone motrix, with A. cervicornis, Caoulastroeg mussids, agaricias, poritids 1405/1404- 1362. 1403 1402 QUEBRADA CHOCOLATE FORMATION CLAYEY SILTSTONE: Grey-blue, bioturboted, with abundant scattered mollusks, densely packed ; 14001360 SOR Oue Cueue CORAL THICKET Claystone matrix, blue-grey, with Porites, hash Seon) | 1399/1394 My P WU Ty 9 We OWN Y Gy 1393=1359 1392 = 1347 1316*1388 2012/2019 = 1387=2530 556 0 552 0 548 0 544.0 590 0 536 0 532 0 528 0 524.0 520 0 2057, 2058 516 0 $12.0 978, 973, L083 508 0 CS USL LS] SSL pL La LSU (SS ay nt tO | & BY ° oF Scale po Bee CALCARENITE: Coral-reef Facies with diverse corals, Montastraea, Sty lophora [ [ 62 0 440 0 T 418 0 ip T 396 0 3740 NO EXPOSURE: 21m 7 CALCARENITE: Reef rubble, with faa een Stylophora with, branching corals = 220 Acropora, Ey ees Manic ing, ostraea ? r LAYSTON Blue-grey, with scattered mollusks t- 330.0 SANDSTONE: Shelly, volcanic L AYSTONE: Grey-black, with scattered mollusks in upper part ae a SE Reef rubble, with Porites, Coulastraea, ficroporo, Diploria, | Fs = - - — - ~ — ophnorg L 4 NO EXPOSURE: 64m - 2860 4 L a0 4 243.0 242.0 241.0 240.0 233.0 238.0 237 0 236.0 235.0 234.0 233 0 232.0 231.0 230.0 223.0 228.0 216.0 204.0 192.0 180.0 168.0 156.0 144.0 132.0 120.0 108.0 36.0 84.0 72.0 60.0 98.0 36.0 24.0 12.0 SECTIONS: COATES 341 Change of Scale Section 33, contd. SILTSTONE: Blue-grey, grading up to Fine sandstone Mollusks and hosh CLAYSTONE: Porites thicket with S_ siderastrea CONGLOMERATIC SANDSTONE: Rounded cobble conglomerate Pebbles range From 4-Scm to most common size of 1-2cm CLAYEY SILTSTONE Porites coral thicket (colonies of M annuloris ot top o thicket) a 1 CLAYSTONE- Blue-grey, burrowed, with shel!-hash burrow Fil! 1378 ale aX eS Eee A EAN 4 SANDSTONE: Shelly, volcanic, tufFaceous, with arthropod burrows, pocked witH ~ ~~ SS hash BRE ERE ER 4 ae, ww ww — 1772/1773=663 A A Ry Z | ER EN NS Ae NR, A Z SANDSTONE: Shelly, volcanic, burrows, packed with hash tufFaceous, with orthropod CALCARENITE: With reef rubble, A. cervicornis, P_ porites, Cou!lastroeo CALCARENITE W/SANDSTONE: With reef rubble, AL cervicornis, Pp orites, Coulostroea BA SE OF QUEBRADA CHOCOLATE FORMATION SANDSTONE: Blue-grey, mollusk rich clayey, silty, no corals TOP OF RIO BANANO FORMATION 342 100 96 92 88 84 80 76 72 68 64 60 56 52 48 Fata ges aaa aaa Galan Lada SL ee Tr Come in Kamas al Use m f eaes U s eAs Ua Ue a Otte Epo a Nolita ir ete oh a eae i re a er See See eeeeneeee| LIMON BASIN Empalme n PPP number 108 1428 . 104 2064 a 19427 . Li thology 770 2061/2063, 1340 . id. 71/713 1368) . 2h 36+ 1587 ss 715/716 Ag ——. 369 1367 el 1351+ 1438/1433——, 1372 . 1499, 1! 1373 . 1371 . 2017/ 7n8 ofS BULLETIN 357 poritids, Stephanocoenia Section 34 Description 1O2 Of EMPAEMEREER VEMBER TOP OF MOIN FORMATION CALCARENITE W/SANDSTONE: Carbonate sandstone with Caulastraea Montastraea Monicing, agoriciids MUDSTONE & SILTSTONE With Madracis, Montastraea, Stephanocoenia, agariciids F SANDSTONE: Loteritic, with Porites, M. annuloris, Meandring ?Coulastraea SANDSTONE Coral rich with D_ strigosa, D. clivoso, S_ siderastrea, A cervicornis BA 0 MPALM MBER EEE M SILTY CLAYSTONE Grey-blue, mottled, with wood Fragments, Corbulo, snails, shel! hash, bryozoans, oysters, bryozoans encrusting cobbles EMPALME MOLLUSK LOCALITY CLAYSTONE: With A. polmota, M. annuloris, Coulastroeo, Diploria Dichoenia MUDSTONE With wood Fragments M. annularis, M cavernosa, Agaricia, mussids, CLAYEY SILTSTONE. Mottled, with Montastraeo ond other corals MUDSTONE & SILTSTONE: With corals, A. palmota, A. cervicornis, C isis, F. porites, CALCARENITE: Reef rubble S_ Siderastrea, agariciids, Madracis, P_ porites SILTSTONE: Blue, not well exposed CLAYSTONE: Dork grey-blue, with oysters, mollusks, scaphopods = SEQUENCE OVERLAYS QUEBRADA CHOCOLATE FORMATION WITH SMALL BUT UNKNOWN GAP. BASE OF MOIN FORMATION m iz 94 - 4 eee Us al [LIMON BASIN Pueblo Nuevo, PPP number Cemetery Li thology \\ TOP OF MOIN FORMATION iL Micaceous grains and occasional echinoids CLAYSTONE Blocky, pyritic, ripple-lominated, alternating with rhythmic SECTIONS: COATES 343 Section 35 Description WEST CEMETERY SECTION CLAYSTONE: Tuffaceous, pale weathering, light tan, with occasional Fine horizons of massive, blocky mudstone CLAYSTONE Blue-grey, with obundont arthropod burrows crammed with echinoids mo! lusks and che! |ostomes NO EXPOSURE 166m SILTSTONE & CLAYSTONE: Blue-grey, Fine grained, volcaniclastic, rich in organic material and plants BASE OF WEST CEMETERY SECTION NO EXPOSURE Approximately 77m SANDSTONE: Variably laminated, coarse-to-medium goalies volcanic lithorenites with lenticular beds and channels. Exposure 200-300m E-NE From cemetery near Pueblo Nuevo Fae orepn position is tentative extrapolated From strikes at cemetery, E-W ot this exposure PROBABLY EQUALS PUEBLO NUEVO SANDS OF TAYLOR as }(AS)) BASE OF MOIN FORMATION 344 LIMON BASIN BULLETIN 357 Section 36 Lonas=del= Nor bastern sequence m PPP number Li thol ogy Description tr 700 L + 1484 Q 69.0 SANDSTONE: Massive, weathered, lateritic, covers the top of the Lomas de! Mor is | plateau ond is transgressive on the reef Facies - 680 4 L NO EXPOSURE Sm rf 870 | TOP OF LOMAS DEL MAR REEF MEMBER oe eS IF | 627 e 7 Wy So Fr 640 WoT owe IL | Se °° = a REEF LENS: lens with yellow-brown weathering sandstone and siltstone matrix aed ah epee: Madracis, platy Montastraeo, agariciids fe bss0 O92 _,2,2 L 183 ' 2.9.9.9.9 fr 620 9,9 ,9_.o L 1492 O29, 9,9, 9,9, oO. Seine = i SSeS G2 Se _ Sod L | LOLA niios F SANDSTONE: Massive with abundant claystone lenses, rich in mollusks both g S Seeaiiey whole and hash Pervasively bioturboted with Frequent bryozoans and horn i 0 5 Sé SERN corals [ i SI SLT ce SAF v7 or HN A a 59.0 4 1309, 1385, 1389, eucues r + *1410/1416, Danae pelea CLAYSTONE: Highly Fractured and weathered, yellow-orange No macrofossi|s, - ogg — 7205/2011 Sa Se See se |imonite on Fractures SPOR NA, gq 7 ee are Be es NO_EXPOSURE Im 1481 . SNe REEF LENS: 10-15-m diometer (site PPP 946), containing diverse, smal! coral [ 1 ew dir fnhw etext heads. Flanked by bioclastic, bioturboted, massive siltstone, packed with 56.0 1 1480 . QMS diverse mollusks, horn corals, serpulids, bryozoans FLERE Qe. L 55.0 SES SS }/ CLAYSTONE Grey-blue, sometimes interFingers with the siltstone Flank beds L = ] REEF LENS 10-15-m diometer (site PPP 942), containing diverse, smal! coral 1479 1 ae heads. Flanked by bioclastic bioturbated, massive si!tstone, packed with me oO diverse mollusks, horn corals, serpulids, bryozoans L 530 944 CLAYSTONE: Grey-blue, blocky, massive k Co ; SILTSTONE: Bioclastic, Flanking beds to reef lense Packed with diverse mollusks, horn corals, serpulids and bryozoans pee 9942 1 REEF LENS: 10-15-m diameter (site PPP 947), containing diverse, smal! coral [ 945 . heads. Flonked by bioclastic, bioturbated, massive siltstone, packed with el aq? diverse mollusks, horn corals, serpulids, bryozoans [ e? UNS : CLAYSTONE= Blue, with serpul id reef ] REEF LENS: 10-15-m diometer (site PPP 948), containing diverse, smal! coral [ 999 1 heads. Flonked by bioclastic, bioturboted, massive siltstone, packed with iT 80 > diverse mollusks, horn corals, serpulids, bryozoans li 1 965 . CLAYEY SILTSTONE: Bioclostic, rich in mollusks 8 0 964 ' CLAYEY SILTSTONE: Alternating lamination of sticky grey clay and laminated ir 950-963 ' silt. No Fossils, probably intertidal WL L a LIMESTONE: Biocalcarenite which Flanks ond overlaps the reef lens For (as PPP 992, 946), densely packed with mollusks, bryozoans L ak 7 One A REEF LENS: With bioclastic siltstone matrix rich im many genera of large and 952 ‘ smal! coral heads. Underlying PPP 950 but also lapping on reef is a poorly im sol 7031 A sorted calcorenite, densely packed with horn corals, bryozoans, ond mo! |usks, b 2134 = PPP 952 l= nil adie > a SILTSTONE: Rich, bioclastic, with mollusks, horn corals r 2032-2030 REEF LENS: ReeF lense rich in many genera of large and smal! coral heads, fr 0 ¥ with bioclastic siltstone motrix ii SILTSTONE: Rich, bioclostic, with mollusks, bryozoans, horn corals Fr 0 L BASE OF LOMAS DEL MAR REEF MEMBER - 41.0 NO EXPOSURE: 3 Sm (PURSES PS al iy uy Taal Dipl ip es Gs Vis oe Vet oT Tear java | ieee |e | ee Ta EVE Vali Loe ea SECTIONS: COATES 345 Change of Scale Section 36, contd 40.0 eo [1m NO EXPOSURE 1 Sm + 690 BO ony all CLAYSTONE: Dork grey, tight, intensely burrowed Shel! hash in anastamosing 70 eee = Callianasso burrows Abundant horn corals, mollusks % 0 ) SILTY CLAYSTONE: Motrix of grey fine biocalcarenite Lensing patch reef an Domed colonies of M onnularis, Scolymia, M cavernosa, Modracis, several den Agaricia, abundant mollusks 340 644 764 h\ SANDSTONE: Slightly clayey, shelly, rich in Amphistegina and other benthic SSS alc N| Foraminifera, diverse mollusks 20 757= 1988 —— \ {6a | NO EXPOSURE 1 Sn 0 4 me 4 645 } NO LITHOLOGY Lithology undescribed a 953 I CLAYEY SILTSTONE: Mollusk hash 280 \ CLAYSTONE. Grey, silty, shelly 270 167 , | CLAYSTONE: Blocky, blue-grey, with conchoidal Fracturing, smal! mollusks, 1 miliolids 26 0 2033 ate ee === SILTSTONE Light brown, tan, shelly, sandy, with shel! hash ma 954637 anatase CLAYSTONE Sticky, light grey, weathered, blocky, with no structures ao 4% Ve SS CLAYSTONE: Weathered, with dense branching Thalassinoides burrow system 2.0 = CLAYSTONE: Sticky, dork grey, burrowed at base into unit below ae SANDSTONE Rich with molds of mollusks ad CLAYSTONE Blue-black, sticky, with pods of Fine, silty hash ond 130 Foraminifera =e | CLAYSTONE. Tight, blue-grey 16.0 as MUDSTONE: Tight, blue-grey, sticky. Rich in micromollusks, Foraminifera, eee pervasive bioturbation ae 4 SILTSTONE: Blue-grey, bioturbated Occasional shelly and lithic fragments and zo | a horn corals el SANDSTONE: Pebbly, volcanic Approximate stratigraphic position of bryozoan 100 4 thicket. Pebbles encrusted, also abundant mo! |usks 30 s| MUDSTONE ao | W- - --- SANDSTONE & SILTSTONE INTERBEDS: Sparry, cemented, coral-rich calcarenite Wu an : interbedded with unit of clayey siltstone ond mudstone BO 765 . Sis | SANDSTONE: Coarse, tan-weathering gritty to Fine pebbly, volcanic eal Pervasively burrowed, occasional mollusks and pieces of wood 30 20 io BASE OF MOIN FORMATION 346 r 67 L 59 = 52 = 4 + i 7 f= 30 l= 22 LIMON BASIN Lomas de! Mar, n PPP number Li thology - 74 1350-143 BULLETIN 357 Section 37 Western Reef F lank Sequence Description TOP OF MOIN FORMATION MUDSTONE: Massive, blue-grey. Sequence 1s oriented from regional all Stream traverse meanders and dip information absent Stratigraphic order thus uncertain CLAYSTONE: Massive, puesane with shel! hash and abundant smal! mollusks, basalt grains concentrated in burrows CLAYSTONE: Massive, blue-grey, silty, common visible Foraminifera SANDSTONE: Volcaniclastic and calcorenitic with a blue-grey, clayey matrix, packed with small mollusks and visible Foraminifera SILTY CLAYSTONE: Blue-grey SILTY SANDSTONE Blocky, shelly, calcarenitic, volcaniclastic, with clayey matrix. Rich in micromollusks and visible Foraminifera SILTY MUDSTONE: Grey to greenish-blue Rich in visible Foraminifera volcanic, with scattered basalt grains (2mm) he upper surface has arthropod burrows Packed with micromo! lusks | SANDSTONE: Blue-grey, ee SILTY CLAYSTONE: Massive, grey-green-blue, with occasional scaphopods BASE OF MOIN FORMATION iT T T LIMON BASIN SECTIONS: COATES 347 Section 38 this this Bones ee) Mer Neste neem lire tj sequence m PPP number Li thology Description 150 a —— 20%, 1376 PEED TOP OF MOIN FORMATION 14.0 . . so -s eo SILTSTONE: Blue-grey, clayey, massive, blocky Scottered within this ean SEO LSIS EL SLSLSLS lithology ore diverse coral potch reefs, 10-20m in diometer, with =1331+ aaa ae hp eg bioclastic siltstone Flank beds Rich in mollusks, bryozoans, horn 230 = 1433/1434 9.9.0.0. 9-999. corals, echinoids puladria very abundant 2055 == == ‘or ‘awe REEF LENS: with bioclastic silt motrix 120 ex TN TOP OF LOMAS DEL MAR REEF MEMBER ens Sse SILTSTONE- Blue-grey, cloyey, massive ond blocky Scattered within uo 2025 Ch | eos PTI lithology ore diverse coral patch reefs, 10-20m in diameter, with son oe Fes bioclastic siltstone Flank beds. Rich in mollusks, bryozoans, horn 10.0 2044 ae =" corals, echinoids Cupuladria very abundant 13521127 SUS ey SILTSTONE: Blue-grey, clayey, mossive and blocky Scattered within 30 is bd oe lithology ore diverse coral patch reefs, 10-20m in diameter, with Fuad Bay Soa bioclastic siltstone Flank beds Rich in mollusks, bryozoans, horn 80 2043 21> = =o corals, echinoids. Cupuladria very abundant Periodic thalassinoid Guan SLOSES-OLGL burrow systems common 10 @ Ge Se GS G@ Gi REEF LENS: with bioclostic silt matrix 2042 aloe > OY o~ SILTSTONE: Blue-grey, clayey, massive and blocky Scottered within 2024 Pw p tail lithology ore diverse coral patch reefs, 10-20m in diometer, with 60 BOE) cls bioclastic siltstone Flank beds. Rich in mollusks, bryozoans, horn COU tanec ea corals, echinoids. Cupuladria very abundant Periodic thalassinoid a ee Orgies burrow systems common 5.0 2038-1375 SR ey 2041 ae ya 40 Hat fl QI GN S| REEF LENS: with bioclastic silt motrix =1485/1 oe EAS LX =< BASE OF LOMAS DEL MAR REEF MEMBER 2026 "|FX~<. SY SILTSTONE: Blue-grey, clayey, massive ond blocky Scattered within 30 TR Bil Re Cale Ria lithology are diverse coral potch reefs, 10-20m in diometer, with 2060, 2040 @ 00898, 9,8, O_,8 A} bioclastic siltstone Flank beds) Rich in mollusks, bryozoans, horn 20 “ Lo GEOEOES: @ Se @ GI corals, echinoids Cupulodria very abundont Periodic thalassinoid Aare burrow systems common. These units ore transgressed by deep brown : ow Sie fit 2028 . Sa weathering laminated, lateritic, coarse sandstone Filling in and PV ee ae 2S] abutting reefs 2039 NY via eR BASE OF MOIN FORMATION this this 348 BULLETIN 357 LIMON BASIN Section 39 Vizcaya River n PPP number Lithol ogy Description i A A=) | TOP_OF RIO BANANO FORMATION SEN ES SANDSTONE: Fine to medium grained volcanic, with complex Thalassinoides s a y , , burrows | = SANDSTONE: Laminated, bioturbated, medium grained Ww \ “NV SANDSTONE Lominoted, with low-angle, shore-Foce cross beds, coarse channels co . lorge burrows os ~ SANDSTONE: Volcanic, Thalassinoides burrowed Seg SANDSTONE Medium grained volcanic, slightly concretionory, bioturbated ae Opes Seas eee CLAYEY SILTSTONE Massive, with Finely comminuted shel! hash Sivercuits CONGLOMERATIC SANDSTONE Dispersed shells, local densely packed burrous, = bine, with phosphatic pebbles and granules and Cupuladria bryozoans No Bn Saas edding TI SILTY SANDSTONE Lominoted, with pees ond granules Well defined, with = = ; concave-down, large, thin-nested bivalves NO EXPOSURE 15m [ | Beesley laws ee Soe | SILTY SANDSTONE Massive, dispersed shel! bed with densely packed shel! hash Eee ene ee PPP 925 through 930 ond 1082 ore stratigraphically equal to 931/935 rp 2a = NO EXPOSURE: 21m [ | BASE OF RIO BANANO FORMATION INDEX Ahermatypic corals—see Azooxanthellate corals AzoOoxanthellateicorals ryeeaea ie cee eerie 29582, 1109; 112 DIVELSIY Eri ins cco cas cuniee eeees ores ieee cy) eueloiceusne ci 112 AlEGECOLO RY Wry arielcrs cic carci scene ieee eunso tel ageienonemene cee 113 Aral ySISZOMmVallanCew. nie) a-Rasioiene catch ees oes atcha eet trea o2 PMU IES ME OLMAOM! feiiie it cverencs ue etee: ehece ob ale) see neyeeeie a = 74 BananitovRIVer 5 op cieis tig asses gery cise east aioe eyelashes 338 IBANANOWRIVEDSs, =) sys ers os) sss see eyenies. sissy os] Sues euenoutensus 334, 338 | DASHA TESUGENG! (3 wide letote Gloee culos eas o om.e 26, 294 SUC: CH a pitis anne Seay c mene ce oke Meee eRCeee ieee ame hans 329 ENSOMELOl@ mee neuewevisteileencyic sy chiciepe te (ster esc. eckisus assists ust = (eyes 330 Benthic foraminifera .... 82, 91, 92, 93, 94, 100, 101, 106, 212 IDIVERSILY A Vict) ceieuein tee eis s0s-t Shsurt chs isi 91, 92, 93, 100, 101 INDDTSTCA CHAKKES oe oeune Soe boobs omogdoDue eae 4c 93 PACTS soonoaneanuoagandaoeocor 82, 94, 212 IRECentesamMplesy-earwenes-t succes cic cron eeuiedsrs easter heeyensnens 92 Sample processing] G42. fe so ccme nec ee case hese aie 93 TEES tal pio tov coe eacREnERaIOns pier dsehaseprole Hata-o.8 cio Oarpore S 106 Biostrati praphy) aia suaitets eae cesta shee coche: axes Sedans 38, 41, 61 BocamdesConcepciontey wien ioe sc es eis ele 288, 307, 308 Bocasideli Toro Group), ....-.--- 17, 25, 34, 39, 41, 50, 71, 77 BOmbawewie Sees SS mio hot ese es Sac 31, 35, 40, 69, 86, 298 BS AZORSCCORP ALC Mya a7 shee cs etapsuoy ciel eit reshore Sa cneite 9, 119, 124 IBUNOMB UGE Rec tapein =~ << eyetecs Silas, Sar bieg ae Ml, 855) 935 2905312 Bryozoa—see Cheilostome Bryozoa IBUCHOSWAUTES Wey a3 bf. 3 citer ee ici wees 2 tt Ae L2ON296 BuenoseAires: Member eee sae 2 siecle sl oie ete ee OmS2 UT Camb OnMatlOns sees ee stadene: nee ete sche ts vette ss areaine a ate 74 BumCasbeninsulay pyc rs coe nee Mh resi Suepcusue mene tseehe ee Us tle Calcareous nannofossils—see Nannofossils Wal ZOneSWRIV erie PPP mumbersinas see heen ere ei OR eR 9, 275, 281 MOIETIES. 64.5.5 omfone nad abogoonorraooao oe 271, 276 PPP Web site ... 5, 20, 41, 64, 91, 112, 152, 185, 195, 234, 262, 285 Ropavisland! Steere iacienowncseear-trreeu hate ee hee moo 292, 294 POrtetes ccces-. chs eee abe aie lie ee ere lene. ips icles one Roe ne eet 296 Pueblo INUeviow, reenctest se tenee ene nanoreeacine ene creme nei 297 INDEX 351 PueblowNulevoSandstonGss.