<<

Contr. Tert. Quatern. Geol. 35(1-4) 47-61 6 figs, 4 tabs, 2 pis. Leiden, April 1998

Zooxanthellate scleractinian of

the Bowden shell bed,

southeast Jamaica

Ann F. Budd

UNIVERSITY OF IOWA

IOWA CITY, U.S.A.

AND

DonaldF. Mcneill

UNIVERSITY OF MIAMI

MIAMI, U.S.A.

Budd, Ann F. & Donald F. McNeill. Zooxanthellate scleractinian corals of the Bowden shell bed, southeast Jamaica. In: Donovan, S.K.

(ed.). The Pliocene Bowden shell bed, southeast Jamaica. —Contr. Tert. Quatern. Geol., 35(1-4): 47-61, 6 figs, 4 tabs, 2 pis., Leiden, April

1998.

The zooxanthellate fauna of the Bowden shell bed in southeast Jamaica consists of 38 , and is dominatedby extinct free-living

the and Antillia. of the species belonging to genera Manicina, Thysanus,Trachyphyllia,Placocyathus Eighteen species are free-living,

of the distributed the eleven are massive and nine are branching. Only nine 38 species are extant, and all but one species are throughout

3.8-2.7 Ma for the Bowden shell ofthe 38 have last in Caribbean region. Assuming a geologic age of bed, eight species global occurrences

Formation in units of elsewhere in the Bowden Formation, and eight have global first occurrences either in the Bowden or equivalent age

the Caribbean.

Recent collections made intervals the Pera reveal shift between ~3.3-1.8 Ma from extinct at closely spaced through Bowden-Old sequence a

contain mixture that free-living species to extant branching and massive species. The Old Pera collections a of Caribbean coral species

accelerated faunal between 4-1 Ma, and thus a transitional fauna. dominated Caribbean reefs before or afteran episode of turnover represent

Most the Old Pera is the co-occurrenceof now extinct and extant cervicornis. New notable among species Stylophora granulata

beds in units of stratigraphic data reveal that 18 of the 37 Old Pera species have global last occurrences in the Old Pera or equivalent age

elsewhere in the that it have for Caribbean, suggesting may provided a refuge pre-turnover species.

Comparisons with collections of two similar-aged faunas on the north coast of Jamaica, the ManchionealFormation (2-1.4 Ma) and Hope

differences that related environmental north faunas Gate Formation (1.8-1 Ma), show striking appear to conditions. The coast contain high

with indicative of reef environments. associated percentages of extant species massive colony shapes, exposed, open Community change

with turnover have more in reef environments, such as the Old Pera beds. Plio-Pleistocene may proceeded slowly protected, margin

Key words — Pliocene, Jamaica, Bowden shell bed, Old Pera beds, Manchioneal Formation, Hope Gate Formation, scleractinians, reefs, extinctions, chronostratigraphy.

Marine and A.F. Budd, Department of Geology, University of Iowa, Iowa City, Iowa 52242, U.S.A.; D.F. McNeill, Geology Geophysics,

Rosenstiel School of Marine and Atmospheric Science, University ofMiami, 4600 Rickenbacker Causeway, Miami, 33149,U.S.A.

54 Contents References p.

Introduction p. 47 Introduction The Bowden-Old Pera sequence along the coast of Port Morant Bay p. 48 shell bed has been known for the abun- Comparisons with the Manchionealand Hope Gate The Bowden long

52 dance and exceptional preservation of extinct, free-living, Formations on the Jamaican north coast p. flabelloidreef corals that associated with The Bowden fauna and its evolutionary significance p. 53 are typically sea

Conclusions 54 flats (Vaughan, 1919; Robinson, 1969; Budd et al., p. grass 1996; J.W. Wells, It is also the Acknowledgements p. 54 pers. comm., 1988). type 48

at ten is also revised locality for least named Mio-Pliocene reef coral spe- the Bowden-Old Pera sequence newly using

for five data. the taxa identified in cies. Duncan (1863) designated holotypes species palaeomagnetic We compare

from specimens collected in the Bowden shell bed such as these collections with other Jamaican collections that we

barretti, Placotrochus alveolus Placo- made in 1994 coastal of similar Placocyathus (= through sequences geo-

Siderastrea within the Manchioneal Formation Is- cyathus), Thysanus excentricus, grandis (= S. logic age on Navy

siderea) andCyphastrea costata (=Solenastrea bournoni). land and Folly Point near Port Antonio (northeastern Ja-

In Duncan & Wall (1865), he designated holotypes for four maica), and within the Hope Gate Formation near Rio

Bueno and more species, viz. Placocyathus moorei (= P. barretti), Discovery Bay (north-central Jamaica) (Fig. 1).

= T. We then use museum collections of the Bowden shell bed Thysanus elegans (? excentricus), Stylophora granu-

lata and Antillia walli (=Trachyphyllia bilobata). Vaughan to assemble a complete faunal list and re-evaluate the

of the Bowden reef coral fauna. (1901) also based the new species, Antillia gregorii, on a evolutionary significance

specimen from the Bowden shell bed which Duncan

' (1863) originally identified as ‘Montlivaltia ponderosa

Edwards & Haime'.

Despite its importance to , the Bowden fauna

is poorly known. The last complete published description

was that of Vaughan (1919), who listed 13 zooxanthellate

coral species and commented on similarities with what is

now known as the Mio-Pliocene Gurabo Formation of the

northern DominicanRepublic and the early Pliocene Pine-

crest Beds of south Florida. However, Aubry (1993) dated

the Bowden shell bed as early late Pliocene or 3.8-2.7 Ma

and thus than the Gurabo Forma- (Aubry, 1993), younger

tion and possibly also younger than the Pinecrest reef coral

horizons. Therefore, it may represent one of the youngest

predominantly 'pre-turnover' faunas in the Caribbean, be-

fore accelerated origination and extinction transformed the

species composition of Caribbean reefs during late Plio-

cene and early Pleistocene time (Budd et al., 1994; John-

son et al., 1995; Budd et al., 1996; Budd & Johnson,

1997).

1. the Fig. Outline map of Jamaica showing general locations of

the three reef coral faunas considered herein; A — the Fig. 2. Map showing the locations of stratigraphy (DM series) Bowden-Old Pera sequence along the coast of Port Mo- and coral (AB series) sample sites in the Bowden-Old rant Bay, parish of St Thomas; B — the Manchioneal Pera the of Port Morant sequence along coast Bay. Formation at Folly Point and Navy Island near Port An-

tonio, parish of Portland; and C — the Hope Gate For-

in Bueno mation the Rio area, parish ofTrelawny.

THE BOWDEN-OLD PERA SEQUENCE ALONG THE

COAST OF PORT MORANTBAY

The purpose of this contribution is to present a new,

up-to-date description of the fauna based on our 1994 col- Existing stratigraphy of the Bowden shell bed — Several

lections of reef corals from the Bowden shell and bed, recent stratigraphic studies on the Bowden Formation have

through the Bowden-Old Pera the east information the the classical sequence along provided new on age of Bow-

coast of Port Morant Bay (Figs 1, 2). The stratigraphy of den shell bed and adjacent lithologic units. 49

3. and strontium data collected in the Bowden-Old Pera Fig. Biostratigraphic, palaeomagnetic isotope sequence.

Banner & Blow (1965), in their construction of a plank- Bowden Formation, and a zone NN15 assignment for its

tonic foraminiferal biostratigraphy of the Neogene, exam- lower beds (Aubry, 1993). The Bowden Formation type

ined from the Bowden shell section and samples bed and, based on nearby road to Arcadia section was assigned to

their zonation that them N19 at time, assigned to zone zone NN16. Planktonic foraminifera examined by

Robinson cited communica- the (Pliocene). (1969) a personal Berggren (1993) at same correlative sections were as-

tion with W.H. Blow as a revision of the Bowden shellbed signed to zones PL2 to PL3, which spans the early/late

to at least the boundary between zone N19 and the newly Pliocene boundary between the Gilbert (C2Ar) chron and established zone N20, if not entirely in zone N20. Robin- the Gauss (C2An.3n and C2An.2) chron. These foraminif-

son (1969) also indicated that the upper part of the type eral zones are consistent with the age range that comprises

the coast the of section, along near town Bowden, may the above mentioned nannofossil zonal assignment of extend into younger zones. NN16 and part of NN15.

A combination of planktonic foraminiferal biostra-

tigraphy and palaeomagnetic data on the Bowden type New stratigraphic data and provisional interpretation —

section Robinson & Lamb Two was presented by (1970). We developed a provisional stratigraphy for the upper part

sample sites, one near the Bowden shell bed (ER1146 = of the Bowden Formation type section and the overlying

AB94-10 and the Bowden Old Pera beds based combination of fora- to 14) one upsection near type on a planktonic

section (ER1123 = DM94-1 to 3), had normal and transi- miniferal biostratigraphy, strontium-isotope stratigraphy

tional polarity, respectively (Fig. 2). Combined with the and magnetostratigraphy (Figs 3, 4). A total of 19 sites

biostratigraphy, these palaeomagnetic data led to an upper were sampled along the coast south of the town of Bowden

Gauss chron (C2An.ln) correlation, that is, late Pliocene. (Fig. 2). Palaeomagnetic samples were collected at 17

More recent biostratigraphic re-examination of cor- stratigraphic levels within the approximately 20 m section relative sections of the Bowden Formation (Aubry, 1993; and multiple samples were analysed at each level. Micro-

Berggren, 1993) have confirmed a Pliocene age. Calcare- fossil samples were collected at all 19 sites, and strontium-

nannofossil of the sections has shown ous stratigraphy a isotope samples were collected at DM94-4, DM94-16 and

zone NN16 assignment for most of the exposed part of the in the Bowden shellbed. 50

4. for the Bowden-Old Pera Fig. Chronostratigraphic summary sequence.

field of Florida and the standard Palaeomagnetic samples were collected using a drill versity ages assigned using and oriented reference of Farrell al. using a Brunton compass. They were ana- curve el (1995). Sample mineralogy lysed in a shielded room using a 2G-755 superconducting was determined by X-ray diffraction, and stable-isotope

the of and values measured and magnetometer at University Miami, demagnet- were to assure original mineralogy ised using alternating field and thermal demagnetisation detect potential secondary, meteoric alteration.

Unweathered total of four techniques. silty/clay sands from the Bowden A geomagnetic polarity zones were recog-

Formation had stable of normal and The a very component magnetic rema- nised; two two reverse polarity (Fig. 3).

with soft viscous which of the BowdenFormation had which nence only a very overprint was top a reverse polarity removed in the first few alternating field demagnetisation continued into the lower Old Pera beds. A 4 m normal steps. polarity section occurs over the Bowden Formation which

Selected rock-magnetic tests and thermal demagnetisa- in turn is overlain by about 9 m interval of reverse polarity. tion character suggested a magnetite mineralogy as a car- This reverse polarity zone is sampled at only four levels rier of magnetic remanence. Limestones from the Old Pera due to poor outcrop conditions which prevented in situ beds weaker than the Bowden Formation The where were samples, sampling. upper 5-6 m of section, the more but also stable Similar massive limestone were very during demagnetisation. beds of the upper Old Pera beds occur,

the Old Pera became had normal to many Tertiary limestones, samples a polarity (Fig. 3). weak and unstable by thermal steps at about 400° to 450°. Diagnostic biostratigraphic markers and several stron-

The unaltered nature of the Bowden Formation samples, tium-isotope ages allow a correlation of the polarity rever-

time the characteristic response of the limestones to demagneti- sal sequence to the geomagnetic polarity scale. The

of the Bowden Formation in sation, and the stable, linear decay of the remanence are uppermost portion sampled interpreted here to represent a primary component of mag- our section has several age diagnostic planktonic fora-

minifera which constrain Pliocene netisation. The palaeomagnetic results were interpreted a late age (Fig. 3). For

the of and using the geomagnetic polarity timescale of Berggren et al. example, occurrence Globorotalia pertenuis

(1995a). Sphaeroidinella dehiscens sensu stricto have first appear-

Strontium-isotope samples were measured at the Uni- ances at 3.45 Ma and 3.25 Ma, respectively (Berggren et 51

ofthe section we examined, al., 1995a). These first appearances suggest a mid-Gauss directly tied to the stratigraphy

the Bowden shell chron (C2An.2) or younger age for the upper Bowden three explanations can be proposed. First,

Formation. on bed unit is stratigraphically below the base of our section A strontium-isotope age range(3.1-5.3 Ma) which the of the Bowden Forma- a sample about 1 m above the top of the Bowden Forma- sampled uppermost part

tion. This is since the Bowden shell bed site is tion suggests an age constrained between 3.25 to about 3.0 very likely

to the north of section and and Ma. Reverse polarity within this age range would correlate our dips are southerly

which would the shell beds below to either C2An.lr (3.04-3.11 Ma) or C2An.2r (3.22-3.33 southwesterly put our

lowermost In addition, Robinson & Lamb Ma). We are unable to determinewhether a hiatus is pres- sample. (1970)

measured normal the shell bed site while ent between the top of the Bowden Formation and the polarity at we

the measured in the of the Bowden lower part of the Old Pera beds. If no hiatus is present reverse polarity upper part

the of the Formation. The late Miocene strontium-isotope reverse polarity could be correlated to younger age ranges

it could be sub- would the shells to the of the San San two choices (C2An.lr), and if a gap occurs put equivalent age

chronozone C2An.2r. Clay (Aubry, 1993). The two other explanations for the

is correlated the would be that the The overlying normal polarity zone to apparently older-than-expected ages

have upper part of the Gauss chron (C2An.ln, 2.58-3.04 Ma). shells are reworked or that they have precipitated or

fluids. The for This correlation is based on the co-occurrence of S. dehis- been altered in less radiogenic potential

does exist since the for the cens s.str. (FAD 3.25 Ma) and Globigerinoides fistulosus reworking depositional setting

Bowden Formation sediments is a water (3.33-1.6 Ma) as well as Globigerinoides extremus (LAD deeper slope and it does contain intervals of al- 1.77 Ma) in this interval and above (sample DM94-10). (>100 m water depth),

lochthonous material introduced from shallow water The overlying reverse polarity is correlated to the Ma- by

mechanisms Since there are tuyama chron C2r, although only one sample yielded good gravity (Robinson, 1969). only

three at this time no definitive biostratigraphy (Fig. 3). This correlation is based on the strontium-isotope analyses

answer can be reached. additional detailed strati- co-occurrence in sample DM94-10 of G. fistulosus (3.33- Clearly,

and geochemical analyses are needed to resolve 166 Ma) and G. extremus which has a last appearance at graphic

this 1.77 Ma. This LAD of G. extremus coupled with reverse uncertainty.

correlation. polarity would suggest a Matuyama chron C2r

the section has Coral — June 1994, we made collec- The overlying normal polarity which caps assemblages During

constraints which would the east several good biostratigraphic sug- tions of zooxanthellate scleractinian corals along

correlation the Olduvai subchron C2n sites in the Bowden gest a to (1.77-1.95 coast of Port Morant Bay at seven

Ma). Sample DM94-14 marked the first occurrence of Formation and 15 sites in the Old Pera Beds of the overly-

Globorotalia truncatulinoides, which in the Atlantic indi- ing Manchioneal Formation (Fig. 2; Table 1). Each site

cates 2.0 Ma or et al., 2-3 m and 10-15 m wide. At each a age younger (Berggren 1995a). was approximately high

This correlation is also consistent with a strontium-isotope site, specimens representative of the species composition

age range measured from an aragonite shell at the level of were individually extracted from the outcrop with a rock

in sample 16. Last, G. extremus (LAD 1.77 Ma) is still pres- hammer. A total of 138 specimens were collected the

ent in the stratigraphically highest sample that yield Bowden Formation and 156 specimens in the Old Pera

collected were to Iowa, planktonic foraminifers. The massive, upper Old Pera beds beds. All the specimens shipped

and entered into database had normal polarity and, although no biostratigraphy is prepared, identified, a specimen

available the World-Wide Web available, are thought to also be part of the Olduvai sub- on at http://porites. geol-

chron. ogy.uiowa.edu. The material will be deposited at the

the of the Bowden United States National Museum of Natural In summary, age uppermost Forma- History

is the Florida Museum of Natural tion at its type section and the overlying Old Pera beds (USNM), History (UF),

interpreted here to be late Pliocene (Fig. 4). This age is and the Paleontology Repository of the University of Iowa

the consistent with age estimates from several existing (SUI); other figured specimens are from Geology Mu-

studies. The of the of the University of the West Indies, Mona(UWIGM). biostratigraphic age upperportion seum, consistent Old Pera beds has the potential to be similar age with the Specimens were identified to species using a

from Manchioneal Formation on the northeast coast (Robinson, set of characters and states developed morphometric

of and Recent collected 1969) based on the occurrence of G. truncatulinoides, al- analyses Neogene samples across

Budd et 1994; Budd & John- though data from Kohl (1992) suggest a slightly younger the Caribbean region (see al.,

illustrated definitionsof these char- age of earliest Pleistocene age. These may also be age son, 1997). Lists and

equivalent (latest Pliocene to earliest Pleistocene) to the acters together with informationon species authorship and

also available the World-Wide Hope Gate Formation on the north-central coast (Land, synonyms are currently on

Web at the above address. A total of 24 1991) based on initial strontium-isotope age constraints. species (6 extant;

It is interesting to note, however, that strontium-isotope 25%) were identifiedin the Bowden Formationcollections,

three shell from the classic Bowden and 37 in the Old Pera beds age ranges on samples species (14 extant; 37.8%)

shell bed location all older than collections 2; Pis 1, (see on Fig. 2) yielded ages (Table 2).

5 Ma (Fig. 4). Since this Bowden shell bed location is not Of the 24 species identified in the Bowden Formation 52

cies is S. domi- collections, 16 (66.7%) are free-living, and most of these especially significant, because granulata

to free-living species belong the generaManicina (PI. lg-j) nated Caribbean mid-forereef zones during the Neogene

and Thysanus (PI. lk-n) in the family Faviidae (see Budd prior to turnover and A. cervicornis has dominated them

& Johnson, 1998; Johnson, 1998), and to Placocyathus (? since the Plio-Pleistocene (Budd & Johnson, 1997). The

-) in the family (PI. 2c-f). Pla- Old Pera beds thus contain a transitional fauna in which

communities lived cocyathus alveolus (Duncan, 1863) and Thysanus excentri- members of pre- and post-turnover

1863 far the abundant in In this resemble late Plio- cus Duncan, are by most species side-by-side. respect, they early

theTrachyphylliacollections. The free-living coral bilo- cene faunas in the Limon Group of Costa Rica (Buenos

Aires trend) (Budd et al., 1998), but not Plio-Pleistocene bata (Duncan, 1863) (PI. 2b) is also very common. Five

of the Bahamas & species are branching, the most abundant of which are portions Drilling Project cores (Budd

Stylophora monticulosa Vaughn in Vaughn & Hoffmeister, Kievman, 1993), where members of the old and new fauna

do not 1925 (PI. lc) and a new, unnamed species of Acropora, co-occur.

' herein called Z' Massive corals To examine patterns of faunalchange upsection, the 22 ‘Acropora sp. (PI. Id). are sites into coral less common, and consist of two environmentally tolerant collecting were grouped eight assemblages and trends in of and extant species, Solenastrea bournoni Milne Edwards & (Table 1), percentages extant species

with and Haime, 1849 (PI. 2a) and Porites astreoides Lamarck, species branching, free-living, massive, platy

were examined upsection (Fig. 1816 (PI. le). The exceptionally high number of free-living colony shapes qualitatively The shift from free- corals 5). a predominantly in the Bowden shell bed suggests that they were patterns suggest

in to at the Bowden-Old Pera bound- living a protected, muddy environment, perhaps associ- living branching shapes (that is, between assemblages C and D), although beds ated with sea grass flats (Budd et al., 1996). ary with abundant free-living corals occasionally occur within

the Old Pera beds (such as assemblage G). Non-parametric

statistical tests (Mann-Whitney U) indicate that the Bow-

den Formation contains significantly more free-living spe-

cies (Z = -2.236, p = .0253) and fewer branching (Z = -

2.249, p = .0245) and massive or platy (Z = -2.236, p =

.0253) species than the Old Pera beds. Despite this shift,

the lowest Old Pera assemblage (assemblage D) contains a

low percentageof extant coral species, similar to the Bow-

den Formation assemblages. This percentage appears to

increase significantly upsection within the lower Old Pera

beds. Mann-Whitney U-tests indicate that significantly

higher percentages of extant species occur in assemblages

= = A to D than in assemblages E to H (Z -2.309, p .0209).

and The increase can be attributed to a greater abundance

diversity of branching Stylophora lower in the Old Pera

beds section, and to a greater abundance of branching A.

cervicornis higher in the section.

COMPARISONS WITH THE MANCHIONEALAND HOPE

GATE FORMATIONS ON THE JAMAICAN NORTH

COAST

During June 1994, we also made collections of zooxan-

thellate scleractinians in the Manchioneal Formation near Fig. 5. Percentages of extant species, and species with Port and in the Gate Formation Rio branching, free-living, and other (massive and platy) Antonio, Hope near Bueno and In the Port Antonio colony shapes in eight coral assemblages in the Bow- Discovery Bay (Fig. 1).

den-Old Pera Port Morant The locali- collections at sites Point in sequence at Bay. area, we made 12 on Folly (one

ties included are listed in Table 1. the Navy Island Member, 11 in the overlying rubbly Man-

chioneal Formation limestones) and three sites on Navy

In the Old Pera beds collections contain 12 contrast, Island (one in the Navy Island Member, two in the Man- 13 and 11 massive free-living (32.4%), branching (35.1%), chioneal Formation). In the Rio Bueno area, we made col-

The most abundant taxa the Recent species (29.7%). are lections at four sites within 5 m of sea level on the eastern

species Acropora cervicornis and the Neogene species side of Rio Bueno Harbour, at six sites having elevations

The of these two Stylophora granulata. co-occurrence spe- of 5-15 m within the cliff at the edge of the second or 30 m 53

terrace described by Land (1973), and at four sites having All of these species, except M. cylindrica, oc-

the elevations of 30-50 m along the terrace surface south of the cur in the Bowden-Old Pera sequence, indicating that

of the extinct cliff. Study of microfossils in the Manchioneal Formation extinct Hope Gate species represent a subset of the Old of the Port Antonio area has suggested that the Navy Island Bowden-Old Pera fauna. The transitional nature

the Member is Plio-Pleistocene in age or 1.9-1.6 Ma (Robin- Pera and Hope Gate faunas between predominantly

Manchioneal fauna is son & Lamb, 1970; Aubry, 1993). In addition, Robinson & extinct Bowden fauna and extant

to detrended Lamb (1970) used palaeomagnetic data propose a cor- further supported by correspondence analysis relation of the Navy Island Member to the Olduvai sub- (Fig. 6).

is confirmed with chron (C2N, 1.95-1.77 Ma). This age a

correlation to the Calcidiscus macintyrei Subzone of Gart-

Manchioneal ner (1977) by Aubry (1993). The overlying

Formation is earliest Pleistocene (~2.0-1.4 Ma) based on

the data of Kohl (1992) adjusted to the Berggren et al.

has (1995b) timescale. Study of strontium isotopes sug-

gested that the Hope Gate Formation is latest Pliocene to earliest Pleistocene (Land, 1991). Our collections in the

Port Antonio and Rio Bueno areas were made in a similar

those in the Bowden-Old Pera and manner as sequence,

sites were defined using the same criteria. A total of 258

specimens (34 species) were identifiedin the Port Antonio

area collections, and 197 specimens (29 species) in the Rio

Bueno area collections (Table 2). Fig. 6. Scatterplot of the first two axes determined by detrended

In contrast to the Bowden-Old Pera sequence (Fig. 3), correspondence analysis (DCA) of coral mega-

Antonio collections and 21 all of species in the Port area assemblages in the Bowden shell bed (B) [3.8-2.7 Ma],

Old Pera beds Gate Formation (H) out of the 28 species (75%) in the Hope Gate Formation (O) [3-1.8 Ma], Hope [1.8-1 Ma], Navy Island Member of the Manchioneal collections are extant (Table 2). Species with massive col- Formation (N) [1.9-1.6 Ma], Manchioneal Formation dominateboth the Port Antonio faunas ony shapes (52.9% limestones (M) [2-1.4 Ma], and Falmouth Formation (F) in the Navy Island Member; 63.6% in the Manchioneal [-125 Ka]. Each mega-assemblage consisted of all Formation) and the Hope Gate fauna (50%), with free- specimens collected within a stratigraphic unit that was 3- living species being relatively unimportant. Unlike the 5 m high and 300-500 m wide. The analysis was run us- Bowden-Old Pera sequence, species ofthe generaDiploria ing relative abundance codes as defined in Budd et al. andMontastraea prevail, and platy species (in particular of (1998) using PC-ORD, version 2.0 (McCune & Mefford,

and make a significantly Agaricia Mycetophyllia) up 1995). Visual examination of the plot suggests that axis 1

of the fauna. in is related extinct the in higher proportion Although slightly higher to % species. Despite overlap ages

the Navy Island Member, percentages of branching species of the Old Pera beds, Hope Gate Formation, and Man-

and north col- chioneal Formation, the three faunas at different are similar in the Bowden, Old Pera coast appear

lections. stages of faunal transition.

in The similarities in geologic ages and differences in the the variation in taxonomic In general, the high percentage of extinct corals colony shapes suggest that

the five faunas listed in Table 2 Bowden-Old Pera sequence is due to the high numbers of composition among may which extinc- corals, were more susceptible to be environmental. The relatively high number of massive free-living

tion Plio-Pleistocene time (Johnson et al., 1995; coral species in the Port Antonio collections implies ex- during Budd et al., 1996). posed shallow to intermediate depth (< 20 m) forereef

conditions. However, studies of echinoderms and brachio-

pods in the Manchioneal Formation (Harper et al., 1995;

THE BOWDEN FAUNA AND ITS EVOLUTIONARY Donovan & Embden, 1996) indicate water depths of > 100

SIGNIFICANCE m, suggesting that the corals may have been transported

from shallower reef depths. The Hope Gate assemblages

have similarly high proportions of massive species, and Study of museum collections from the Bowden shell bed

also contain slightly higher proportions of platy and free- increases the total number of species known from 24 to 38

living species, again indicating deeper (15-25 m) forereef species (Table 3). This number is approximately one-third

conditions. of the 111 species estimated to have lived in the Caribbean

in the Gate Formation whole between 4-1.5 Ma. With this the The seven extinct species Hope as a addition, pro-

include three branching species (Stylophora granulata, S. portions of extant species and of species with different

in the Bowden Formation remain minor, Caulastraea portoricensis), one massive species colony shapes approxi-

(Montastraea cylindrica), and three free-living species mately the same: nine of the 38 species (23.7%) are extant,

18 are 11 are massive, and (Manicina sp. C, Trachyphyllia bilobata, Placocyathus (47.4%) free-living, (28.9%) variabilis). 54

nine (23.7%) are branching. at the of Miami is magnetics Laboratory University sup- of the 82 A survey Neogene to Recent Caribbean stra- ported by the National Science Foundation and the W.M.

tigraphic units in the 1996 Cenozoic Coral Database Keck Foundation.

(CCD) compiled by Budd & Johnson (1997), reveals un-

usually high numbers of global last occurrences in the

Bowden-Old Pera sequence (Table 4). Eight of the 38 REFERENCES

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occurrences in the Bowden Formation or in units of Berggren, W.A., F.J. Hilgen, C.G. Langereis, D.V. Kent, J.D. elsewhere in the Caribbean equivalent age (most impor- Obradovich, I. Raffi, M.E. Raymo & N.J. Shackleton, 1995a.

the Quebrada Chocolate and Buenos Aires reef Late tantly, Neogene chronology: new prespectives in high-

trends near Limon, Costa Rica). Only one first occurrence resolution stratigraphy. — Geological Society of America

involves extension of than million Bulletin, 107:1272-1287. a range more a years; lacera. With the of Berggren, W.A., D.V. Kent, C.C. Swisher & M.-P. Aubry, exception Acropora sp. Z, 1995b. A revised Cenozoic and chro- most in the to have had wide geochronology species sequence appear geo- nostratigraphy. In: W.A. Berggren, D.V. Kent & M.-P. Au- graphic distributions. bry (eds). Geochronology, Time Scales and Global Strati-

graphic Correlation. Society ofEconomic Paleontologists and

Mineralogists, Special Publication, 54:129-212. CONCLUSIONS Budd, A.F. & K.G. Johnson, 1997. community dy-

namics and over 8 million years of evolutionary time: stasis The large numbers of last occurrences in the Bowden-Old turnover. — Proceedings of the 8th International Coral Reef

Pera and the contrast in taxonomic sequence, composition Symposium, Panama City, Panama, 1: 423-428. with the reef coral fauna of the north of coast Jamaica, Budd, A.F. & K.G. Johnson, 1998. Neogene paleontology in the

suggest that the Bowden-Old Pera sequence may have northern Dominican Republic. The Family Faviidae (Antho-

served — as one of the last refuges for Neogene Caribbean zoa: ), part II. Bulletins of American Paleon- tology (in press). reef corals, especially free-living forms, before their ex- Budd, A.F., K.G. Johnson & T.A. tinction during Plio-Pleistocene time. The differences with Stemann, 1996. Plio-

Pleistocene turnover the the north and extinctions in Caribbean reef coast fauna, in particular, suggest that reef com- coral fauna. In: J.B.C. Jackson, A.F. Budd & A.G. Coates munities in exposed settings established a modern aspect (eds). Evolution and Environment in Tropical America. Chi- faster than their in counterparts protected environments. cago (University of Chicago Press): 168-204. Faunal change associated with Plio-Pleistocene Caribbean Budd, A.F., K.G. Johnson, T.A. Stemann & B.H. Tompkins, turnover occurred at different times and rates, and fol- 1998. Pliocene to Pleistocene reef coral assemblages in the lowed different pathways at different Jamaicanlocations. Limon Group of Costa Rica. In: L.S. Collins & A.G. Coates

(eds). The Neogene of the Isthmus of Panama: A Paleobiotic

Survey of the Caribbean Coast. — Bulletins of American ACKNOWLEDGEMENTS Paleontology, (in press).

Budd, A.F. & C.M. Kievman, 1993. Coral assemblages and reef

We thank A.G. Coates and K..G. Johnson for reviewing the environments in the Bahamas Drilling Project Cores. — Final draft of the Bahamas manuscript; S.K. Donovan, E. Robinson, L.S. Land, L. report Drilling Project, Comparative Sedimentology Rosenstiel School of Marine and Tjalsma and T. Gary for locality information; R.W. Portell, Laboratory, Atmospheric Science, University of Miami. K. Schindler, K. Ketcher, T.A. Stemann and L.A. Guertin Budd, A.F., T.A. Stemann & K.G. Johnson, 1994. Stratigraphic for assistance with field work; B.H. Tompkins, R.A. Peter- distributions of and genera species of Neogene to Recent sen and T.A. Stemann for assistance with specimen prepa- Caribbean reef corals. — Journal of Paleontology, 68: 951- identifications ration, and data entry; and R.W. Portell 977. (UF), S.J. Wood (UWIGM), T. Coffer (USNM) and J. Donovan, S.K. & B.J. Embden, 1996. Early Pleistocene echinoids Golden (SUI) for assistance with museum collections. This of the Manchioneal Formation, Jamaica. — Journal of Pale- work was supported by a grant from the U.S. National ontology, 70: 485-493.

Science Foundation (EAR-9219138) to AFB. The Paleo- Duncan, P.M., 1863. On the fossil corals of the West Indian Is- 55

reefs north Jamaica. — Bulle- lands. Part 1. — Quarterly Journal of the Geological Society Pleistocene by meteoric water,

of London, 19: 406-458. tin of Marine Science, 23: 64-92.

1991. Some ofthe late Cenozoic evolution of Duncan, P.M., 1864. On the fossil corals of the West Indian Is- Land, L.S., aspects

as stratigraphy. lands. Part 2. — Quarterly Journal of the Geological Society north Jamaica revealed by strontium isotope

— of 28: 45-48. of London,20: 20-44. Journal of the Geological Society Jamaica,

B. M.J. 1995. PC-ORD, Multivariate Duncan, P.M. & G.P. Wall, 1865. A notice of the geology of McCune, & Mefford, Version 2.0. Gleneden Beach, Jamaica, especially with reference to the District of Claren- Analysis of Ecological Data,

126 don; with descriptions of the Cretaceous, Eocene, and Mio- Oregon (MJM Design), pp.

Milne H. J. 1849. Recherches sur les polypi- — Edwards, & Haime, cene corals of the island. Quarterly Journal of the Geo-

des Astreides. — An- logical Society of London, 21: 1-14. ers. Quatriemememoire. Monographic

Farrell, J.W., S.C. Clemens & L.P. Gromet, 1995. Improved nates des Sciences naturelles, Paris, (3)12: 95-197.

seawater Robinson, E., 1969. field guide to Neogenesections in chronostratigraphic reference curve of late Neogene Geological

87 6 West Indies. — Journal of the Geological Society of Sr/* Sr. — Geology, 23: 403-406. Jamaica

Gartner, S. Jr., 1977. Calcareous nannofossil biostratigraphy and Jamaica, 10: 1-24.

revised zonation of the Pleistocene. — Marine Micropaleon- Robinson, E. & J.L. Lamb, 1970. Preliminary palaeomagnetic

Plio-Pleistocene of Jamaica. — 227: tology, 2: 1-25. data from the Nature,

Harper, D.A.T., E.N. Doyle & S.K. Donovan, 1995. Palaeoecol- 1236,1237.

of Pleistocene from T.W., 1901. Some fossil corals from the elevated reefs ogy and palaeobathymetry brachiopods Vaughan,

and — the Manchioneal Formation of Jamaica. — Proceedings of of Curasao, Arube, Bonaire. Rijks Geologisch-

1-91. the Geologists' Association, 106: 219-227. mineralogisch Museum Sammlungen,Leiden, 2:

1919. from Johnson, K.G., 1998. A phylogenetic test of accelerated turnover Vaughan, T.W., Fossil corals Central America, Cuba,

Rico with an account of the American in Neogene Caribbean brain corals (Scleractinia: Faviidae). and Porto Tertiary,

Recent coral — National — Palaeontology (in press). Pleistocene, and reefs. United States

Johnson, K.G., A.F. Budd & T.A. Stemann, 1995. Extinction Museum Bulletin, 103:189-524.

of fossil selectivity and ecology of Neogene Caribbean reef corals. — Vaughan, T.W. & J.E. Hoffmeister, 1925. New species

— Bulletin of the Mu- Paleobiology, 21: 52-73. corals from the Dominican Republic.

of 67: 315-326. Kohl, B., 1992. Pleistocene benthic Foraminifera from Jamaica, seum Comparative Zoology, Harvard,

West Indies. — Benthos '90, Studies in Benthic Foraminif-

277-288. era, Sendai, Japan. Tokai (Tokai University Press):

Lamarck, J.B.P.A. de M. de, 1816. Historie naturelle des animaux Manuscript received 25 October 1996, revised version accepted 6

568 1997. sansvertebres, 1(2). Paris, pp. August

Land, L.S., 1973. Contemporaneous dolomitization of middle 56

Coral Equivalent

Assem Stratigraphic AB94 ER Number of Number of

blage Unit localities localities Species Specimens

10-15m ne

A Bowden Fm. 12-14 of ER140 14 28

B Bowden Fm. 10,11 140 9 39

C Bowden Fm. 1,2 180 15 71

D Old Pera beds 48,49 523, 1125 12 27

E Old Pera beds 50,51 530, 1126 10 19

524,1127,

F Old Pera beds 7-9,52 1128 16 40

531,1130,

G Old Pera beds 4-6,53,54 1131 19 53

H Old Pera beds 45,55 532 11 17

Table 1. List of 1994 coral sites in the Bowden Pera beds Ann Edward our collecting Fm. — Old sequence. AB, Budd; ER, Robinson (see Robinson, 1969).

Navy Is Hope

CCD Bowden Mem, Manchioneal Gate Fm,

species Colony Survivor shell Old Pera Port Fm, Port Discovery

Family Species ID no. shape ship bed beds Antonio Antonio Bay

Astrocoeniidae Stephanocoenia interseptaintersepta 002 massive extantextant Xx Xx Xx

Astrocoeniidae Stephanocoenia duncani 003 massive extinctextinct Xx

affinis 005 extinct Pocilloporidae Stylophora branching Xx_

007 extinct Pocilloporidae Stylophora granulatagranulate 007 branching extinct Xx xX

Pocilloporidae Stylophora minor 009 branching extinct Xx xX

Pocilloporidae Stylophora monticulosa 010 branching extinct Xx Xx

Pocilloporidae Pocillopora crassoramosacrassoramosa 015 branching extinct Xx

Pocilloporidae Madracis mirabilis 020 branching extant Xx Xx Xx

Acroporidae AcroporaAcropora cervicorniscervicomis 022 branching extant Xx Xx Xx Xx Xx

Acroporidae Acropora palmatapalmate 023 branching extant Xx Xx Xx Xx

Acroporidae Acropora sp.Z 026.5 branching extinct Xx

Agariciidae Agaricia lamarckilamarcki 030 platy extant Xx Xx Xx

Agariciidae Agaricia undata 032 platy extant Xx Xx

Agariciidae Undaria agaricites 033 platy extant Xx Xx xX Xx

Agariciidae Undaria crassa 034 massive extant Xx xX

Agariciidae Helioseris cucullatacucullata 043 platy extant Xx

Siderastreidae SiderastreaSiderastrea radiansradians 056 massive extant xX

Siderastreidae Siderastrea siderea 058 massive 058 massive extant Xx Xx Xx xX

Poritidae Porites astreoidesastreoides 063063 massive extant xX Xx Xx Xx Xx

Poritidae Porites portoricensis 065 branching extinct Xx

Poritidae Porites baracoaensis 069069 branching extinct Xx Xx

Poritidae Porites branneribranneri 070070 massive extant Xx

Poritidae Porites divaricatadivaricate 075075 branching extant Xx

Poritidae Porites furcatafurcata 076076 branching extant Xx Xx Xx

Poritidae Porites porites 077077 branching extantextant Xx Xx

Poritidae Goniopora imperatorisimperatoris 080080 massive extinct Xx

Faviidae CaulastraeaCaulastraea portoricensis 083083 branching extinct Xx Xx Xx

Faviidae DiploriaDiploha clivosaclivosa 094094 massive extant Xx

Faviidae DiplohaDiploria iabyrinthiformislabyrinthifonnis 095 massive extant Xx Xx

FaviidaeFaviidae 097 massive Diploria strigosa 097 extant Xx Xx

FaviidaeFaviidae Manicina aff.areolata Manicina aff.areolata 100.5 free-living extinct xX xX

FaviidaeFaviidae Manicina massive mayori 101 massive extant Xx Xx xX

Faviidae Manicina Manicina puntagordensis 102 free-living extinct xX xX

Faviidae Manicina Faviidae Manicina grandis 103 free-living extinct Xx

Faviidae Thysanus navicula 104 free-living extinct xX

Table 2. List of zooxanthellate scleractinian coral species identified in our 1994 collections. 57

FaviidaeFaviidae Manicina sp.A 105 free-living extinctextinct xX

FaviidaeFaviidae Manicina sp.B 106 free-living extinctextinct Xx

FaviidaeFaviidae Manicina sp.C 107 free-living extinctextinct Xx Xx Xx

FaviidaeFaviidae Thysanus excentricusexcenthcus 110 free-living extinctextinct Xx Xx

Faviidae Thysanus corbicula 111 free-living extinctextinct Xx

FaviidaeFaviidae ColpophylliaColpophyllia natans 114 massive extantextant Xx Xx Xx xX

FaviidaeFaviidae Montastraea annularis 116 massive extantextant Xx Xx

FaviidaeFaviidae Montastraea faveolata 117 massive extantextant Xx

FaviidaeFaviidae Montastraea franksifranksi 118 massive extantextant Xx Xx

FaviidaeFaviidae Montastraea limbata-1 121.1 massive extinctextinct Xx

FaviidaeFaviidae Montastraea cavernosa-2cavemosa-2 126 massive extantextant Xx Xx xX

FaviidaeFaviidae Montastraea cylindricacylindrica 128 massive extinctextinct Xx

FaviidaeFaviidae Solenastrea bournoniboumoni 131 massive extantextant Xx

Trachyphyllidae TrachyphylliaTrachyphyllia bilobata 135 free-living extinctextinct Xx xX

Meandrinidae Meandrina braziliensisbraziliensis 138 free-living extantextant Xx xX

X Meandrinidae 139 massive extantextant Xx Xx x

Meandrinidae Meandrina sp.A 139.5 massive extant Xx

Meandrinidae Placocyathus alveolus 140 free-living extinctextinct Xx xX

Meandrinidae Placocyathus barrettibarretti 141 free-living extinctextinct Xx Xx

Meandrinidae Placocyathus costatus 142 free-living extinctextinct Xx Xx

Meandrinidae Placocyathus variabilis 144 free-living extinctextinct Xx Xx xX

Meandrinidae stokesistokesi 148 massive extantextant Xx Xx xX

Meandrinidae Dichocoenia stellaris 149 massive extantextant Xx Xx

Meandrinidae DichocoeniaDichocoenia tuberosa 150 massive extinctextinct xX

Meandrinidae DendrogyraDendrogyra cylindricuscylindricus 151 massive extantextant Xx

Mussidae Antillia dentata 153 free-living extinctextinct Xx

Mussidae Antillia gregorii 154 free-living extinctextinct Xx

Mussidae 155 massive extantextant Xx Xx

Mussidae ScolymiaScolymia laceralacera 157 massive extantextant Xx xX

Mussidae Mussa angulosa 158 branching extantextant Xx

massive Mussidae IsophyllastreaIsophyllastrea rigidahgida 165 massive extantextant Xx

Mussidae Mycetophyllia danaana 168 massive extantextant Xx

Mussidae MycetophylliaMycetophyilia lamarckiana 170170 platy extantextant Xx Xx

Mussidae Mycetophyllia reesireesi 171 platy extantextant Xx

Caryophylliidae fastigiata 173 branching extantextant Xx Xx Xx

total # localitieslocalities 77 15 2 13 14

total # specimens 138 156 30 228 197

totaltotal ## species 24 3737 1717 33 2828

32 ## extant species 66 14 1717 32 2121^ # free-livingfree-living species 16 12 0 0 3

# branching species 55 13 66 7 6

# massive species 33 11 9 2121_ 1414

# platy species 0 11 22 5 5

Table 2. List of zooxanthellate scleractinian coral species identified in our 1994 collections (continued), 58

Vokes

CCD Duncan Vaughan Wells coll.coll. UF coll. speciesspecies Present 1863,1864 1901,1919 coll. (USNM) UWI (loc.

ID no. Genus Species Study (BMNH) (USNM)(USNM) (SUI) loc. T705 coll. XJ002)

003 Stephanocoenia duncani X?x? Xx Xx

007 Stylophora granulata H Xx Xx Xx

009 Stylophora minor Xx

010 Stylophora monticulosa Xx Xx

020 Madracis mirabilis (*) Xx Xx

022 Acropora cervicorniscervicomis (*) Xx

023 Acropora palmata (*) Xx

026.5 Acropora sp.Z xX xX xX Xx

058 Siderastrea siderea (*) x(H)x (H) Xx

063 PontesPorites astreoides (*) Xx

069 Pontes baracoaensis Xx X?x? *X X*

080 Goniopora imperatoris X?x? Xx Xx

083 Caulastraea portoricensis Xx

099 Diploria sp.A Xx

100.5 Manicina affaff.areo/ate.areolata Xx

102 Manicina puntagordensispuntagordensis Xx

103 Manicina grandis Xx Xx Xx

104 ThysanusJhysanus navicula Xx Xx

105 Manicina sp.A Xx Xx

106 Manicina sp.B Xx Xx

107 Manicina sp.C Xx Xx

109 Thysanus sp.A Xx

110 Thysanus excentricus Xx H(H)H (H) Xx Xx Xx Xx Xx

111 Thysanus corbicula Xx Xx Xx Xx Xx

121.1 Montastraea limbata-1 Xx Xx

124 Montastraea canalis Xx

131 Solenastrea Solenastrea bournoniboumoni (*) Xx x(H)x (H) Xx

135 TrachyphylliaTrachyphyllia bilobata Xx x(H)x (H) Xx xX

138 MeandrinaMeandrina braziliensisbraziliensis (*) Xx Xx

140 Placocyathus alveolus xX H xX Xx xX xX xX

141 Placocyathus barrettibarretti Xx H(H)H (H) Xx xX xX xX

142 Placocyathus costatuscostatus Xx Xx Xx Xx

143 Placocyathus ?trinitatis? trinitatis xX

144 PlacocyathusPlacocyathus variabilis Xx Xx Xx Xx Xx

150 Dichocoenia tuberosatuberosa Xx

154 AntilliaAntillia gregorii Xx Xx H* Xx Xx

155 ScolymiaScolymia cubensis (*)Q Xx Xx Xx Xx

lacera 157 (*) Xx

Total 24 13 13 7 11 21 14

list of from the Bowden bed. Table 3. Complete zooxanthellate scleractinian species known shell H=holotype, (H)= holotype of a syn-

onymized species, H*=holotype of another museum, (*)=extant species. 59

Global First GlobalLast

OccurrenceOccurrence inIn Occurrence inIn

Bowden Fm (3.8- Bowden Fm (3.8-

Bowden.OldBowden, Old 2.7 Ma) or Old 2.7 Ma) or Old Pera CCD species PenPera GlobalFirstFirst Occurrence excluding Bowden- Global Last Occurrence excluding Bowden- Pera beds (3-1.8 Pera beds (3-1.8

ID no. Genus Species Occurrences OldPenPera sequence OldOkf Pera sequence Ma)Ma) Ma]Ma)

00002 StephanocoeniaStaphanocoania interseptaintarsepta OP PlnecrestPinecrest Ss, Florida(3.5-3(3.5-3 Ma)Ma) Recent

00003 StephanocoeniaStaphanocoania duncani B,OPB.OP AnguHtaAnguMa Fm,Fm. AnqulllaAngullla (22-16.2 Ma) Lomas del Mar, Cos.Rica (1.9-1.5 Ma)Ml) ■OP=OP

"OP 005 StylophoraStytophora affinisafUnis OP GuraboFm. Pom.Dom. Rep. (8.3-7.5 Ma)Ma) La Cruz Fm,Fm, Cuba (3.5-1.6 Ma) -OP

Panama 007 StylophoraStytophora granulata B,OP Emperador Lm, Panama (22-17.6 Ma) Unda-2, Bahamas (1.8-1.7 Ma) ■OP=OP 009 StylophoraStytophora minor B,OP TampaTampa Fm.Fm, FL (23.7-22 Ma) Unda-3, Bahamas (1.9-1.8 Ma)

010 StylophoraStytophora mormctMosamonticulosa B.OP BaitoaBartoa Fm, Dom.Rep. (17.3-13.1 Ma) Unda-2, Bahamas (1.8-1.7 Ma) OP 015 PocÊoporaPocillopora crassoramosacrassonmosa OP BaitoaBartoa Fm, Dom.Rep. (17.3-13.1 Ma) Mao Fm,Fm, Dom.Rep. (3.7-3.4(3.7-3.4 Ma)

020 MadracisMadrads mirabilismtrabfs B,OPB.OP TamanaTamana Fm, Trinidad (15-11.2 Ma) Recent

=B■a 02202 Acropora cervicorniscarvicomis B,OPB.OP Q. Chocolate.Chocolate,Cos.RicaCos.Rica(3.5-3.2 Ma) Recent

=B 02023 Acropora palmatapalmata B Buenos Aires,Alras, Cos.RicaCosRica (3.2-2.9 Ma) Recent =B

B 026.5 Acropora «P^OspZ (*) B none none BB B

033 Undaria agaricitasagaricites OP CercadoFm,Fm. Pom.Dom. Rep. (8.3-7.5 Ma) Recent

034 UndariaUndaha crassacrassa OP CercadoFm,Fm. Dom.Rep. (7.5-5.6 Ma) Recent

058 SiderastreaSidarastraa sidereasjdema B,OPB.OP BaitoaBartoa Fm, Dom.Rep. (17.3-13.1 Ma) Recent

-B 063 PoritesPorttas astreoides B,OPB.OP Buenos Aires, Cos.Rica (3.2-2.9 Ma) Recent =B

065 PoritesPortias portoricensisportoncansis OP Tampa Fm, FL (23.7-22 Ma) Lomas deldal Mar, Cos.Rica (1.9-1.5 Ma) ■OP=OP

069 PoritesPortias baracoaensisbaracoaansis B,OPB.OP Emperador Lm, Panama (22-17.6(22-17 6Ma) Lomas del Mar, Cos.RicaCoa.RIca(1.9-1.5 Ma) =OP

07070 PoritesPortias brannaribranneri OP PinecrestPinecres» Ss, FL (3.5-3 Ma) Recent

07075 PoritesPortias dhrancatadivaricata OP CalooaahatcheeCaloosahatchee Fm, FL (1.8-1.6 Ma)Ma) Recent OP

076 PoritesPortias furcata OP Pinecrest Ss, FL (3.5-3 Ma) Recent

Bahamas =OP 080 GonioporaGontopora imperatorisknparatons B.OP Tampa Fm.Fm, FL (23.7-22 Ma) Unda-3, Bahamas (1.9-1.8 Ma)Ma) =OP

083 CaulastraeaCaulastraaa portoricensisportoncansis B.OP LirioUrio Lm, MoraMona (11.2-5.3 Ma) Lomas del Mar, Cos.RicaCoa.RIca(1.9-1.5 Ma) =OP■OP

=B 099 DiploriaDiptona apAsp.A B Gurabo Fm, Dom.Rep. (7.5-5.6 Ma) Mao Fm, Dom.Rep. (4-3.7 Ma) >B

100.5100.5 ManidnaManicina atl.araolataaff.areolata B,OP Gurabo Fm, Dom.Rep. (5.6-4.5 Ma) Mao Fm, Dom.Rep. (4-3.7 Ma) -B=B

102 ManicinaManidna puntagordensispuntagordansês B,OP Buenos Aires, Cos.RicaCos Rica (3.2-2.9 Ma) Lomas del Mar, Cos.Rica (1.9-1.5 Ma) •B=B ■OP=OP

103 ManicinaManidna grandisgrandis B Cercado Fm, Dom.Rep. (8.3-7.5 Ma) Mao Fm, Dom.Rep. (4-3.7 Ma) =B

104 ThysanusThysanus naviculanavKuia B,OP Cercado Fm, Dom.Rep. (7.5-5.6 Ma) Mao Fm, Dom.Rep. (4-3.7 Ma) OP

Lirio Lm, Mona Gurabo Fm, B 10105 ManidnaManicina sp.Asp A B Urio Lm, Mora (11.2-5.3 Ma) Dom.Rep.Dom.Rep. (4.5-4 Ma)Ml)

106 ManicinaManidna sp.B B Gurabo Fm, Dom.Rep. (8.3-7.5 Ma) GuraboFm, Dom.Rep.Dom.Rep. (4.5-4(4 5-4 Ma) B

107 ManicinaManidna •p.Csp.C B,OPB.OP Cercado Fm, Dom.Rep. (7.5-5.6 Ma) Mao Fm, Dom.Rep. (4-3.7 Ma) OP

Cos.Rica Lomas del Cos.Rica 109109 Thysanus sp.A«P-A B Q. Chocolate, Cos.Rica (3.5-3.2 Ma)Ma) Lomas Mar, Cos.Rica (1.9-1.5 Ma) =B■B

110 Thysanus excentricusaxcantricus B,OP Gurabo Fm, Dom.Rep. (7.5-5.6 Ma) Clino-4, Bahamas (1.9-1.8 Ma)Ma) ■OP=OP

111 Thysanus corbiculacorbicuta B,OP [ChipolaChipdaFm,Fm, FL (18-15 Ma)Ma) |Cak»sahatcheeCaloosahatcheeFm, FL (1.8-1(1.8-1.66 Ma) | |«OP■OP

114 ColpophylliaCdpophyÊa natansnalans OP Gurabo Fm, Dom.Rep.Dom Rep. (5.6-4.5 Ma) Recent

121.1 MontastraeaMontastraaa limbata-1knbata-1 B,OP 7Tamana Fm, Trinidad (14.8-11.5 Ma) Lomas del Mar,Mar. Cos.Rica (1.9-1.5 Ma)

124 MontastraeaMontastraaa canaliscanaas B Tampa Fm,Fm, FL (23.7-22 Ma) Lomas del Mar, Cos.Rica (1.9-1.5 Ma)

131 SolenastreaSdanastroa boumoni B Tampa Fm,Fm, FL (23.7-22 Ma)Ma) Recent

135 TrachyphylliaTrachyphyta bilobatabiobata B TampaTampa Fm, FL (23.7-22 Ma) Clino-4, Bahamas (1.8-1.6 Ma)

138 MeandrinaMaandrtna braziliensisbrazMensts B,OP Cercado Fm, Dom.Rep. (8.3-7.5(8.3-7.5 Ma) Recent

139.5 MeandrinaMaandhna sp.Asp.A OP GuraboFm, Dom.Rep. (5.6-4.5 Ma)Mi) Recent

140 PlacocyathusPlacocyathus alvaolusalveolus B,OP Gurabo Fm.Fm, Dom.Rep. (7.5-5.6 Ma) PlnecrestPinecrest Ss, FL (3.5-3 Ma) OP

14141 PlacocyathusPtacocyattius barrettibarrett B,OPB.OP PinecrestPlnecrest Ss, FL (3.5-3 Ma) Pinecrest Ss, FL (3.5-3 Ma) =B«B OP

142 PlacocyathusPlacocyathus costatuscostatus B,OPB.OP GuraboGurabo Fm, Dom.Rep. (8.3-7.5 Ma) Mao Fm, Dom.Rep. (3.7-3.4 Ma) OP

Trinidad Gurabo 143 PlacocyathusPlacocyathus ? trinitatistrinrtabs B ManzanillaManzanMa Fm, Trinidad (11.2-5.3 Ma)Ml) Gurabo Fm, Dom.Rep. (4.5-4 Ma) B Cos.Rica -OP=OP 14414 PlacocyathusPlacocyathus variabtsvariabilis B.OP Baitoa Fm, Dom.Rep. (17.3-13.1 Ma)Ml) Lomas deldelMar, Cos.Rica (1.9-1.5 Ma)

del Cos.Rica 150 DichocoeniaCtchocoanla tuberosetubarosa B,OP GuraboGurabo Fm, Dom.Rep.Dom.Rep. (7.5-5.6 Ma) Lomas delMar, Cos.Rica (1.9-1.5 Ma) =OP■OP

154 AntMaAntillia gregoriigragort B Tampa Fm, FL (23.7-22 Ma) Tampa Fm, FL (23.7-22(23.7-22 Ma) B

15155 ScolymiaScotyma cubansiscubensis B GuraboGurabo Fm, Dom.Rep.Dom Rep. (7.5-5.6 Ma) Recent

157 ScolymiaScotymia laceralacan B Lomas del Mar, Cos.Rica (1.9-1.5 Ma) RecantRecent B

Pinecrest T 173 EusmiliaEusmta fastigiatatasbgiata |ppOP Plnecrest Ss, FL (3.5-3(3 5-3 Ma) [RecentRecent

4. First and last Occurrence in the Bowden - Old Pera Old from the Bowden Table sequence. B, Bowden; OP, Pera; (*) known only —

Pera Old sequence. 60

PLATE 1

Common and abundant zooxanthellate scleractinian species in the Bowden shell bed (in taxonomic order). All specimens were collected from the Bowden shell bed, except Fig. b, which was collected from the Old Pera beds approximately 1 km south ofthe shell bed.

Figs a, b. Stylophora granulata Duncan, 1864; a — USNM 324779, branch fragment, x 0.9; b — SUI 92037 (CCD 4374), CCD locality

AB94-53 (Old Pera beds), branch fragment, x 0.9.

Fig. c. Stylophora monticulosa Vaughan in Vaughan & Hoffmeister, 1925; UWIGM 11512, UWI locality 10, branch fragment, x 1.

d. SUI 92038 branch 0.9. Fig. Acropora sp. Z; (CCD 4391), locality AB94-11, fragment, x

Fig. e. Porites astreoides Lamarck, 1816;SUI 92041 (CCD 4371), CCD locality AB94-13, whole colony, x 0.65.

Fig. f. Goniopora imperatoris Vaughan, 1919; USNM 65473, colony side, x 1.2.

Figs g, h. Manicina grandis (Duncan, 1864); g — USNM 65474, USGS locality 2580, calical surface, x 0.9; h — USNM 65464, same

0.9. specimen as Fig. g, side view, x

Manicina UWIGM UWI — side 0.9. Figs i, j. sp. C; 11519, locality 10; i — calical surface, x 0.9;j view, x

Figs k, l. Thysanus excentricus Duncan, 1863; SUI 90997, CCD locality AB94-11; k — calical surface, x 0.9; l — side view, x 0.9.

Figs m, n. Thysanus elegans Duncan in Duncan & Wall, 1865 (? = T. excentricus); USNM 63325, USGS locality 2580; m — calical sur-

face, x 0.9; n — side view, x 0.9. 61\1

PLATE 1 62\2

PLATE 2

PLATE 2

Common and abundant zooxanthellate scleractinian species in the Bowden shell bed (in taxonomic order). All specimens were collected

and which collected ER 0.75 from the Bowden shell bed, except Figs b g were from the Bowden Formation (locality 180) approximately km south of the shell bed.

Haime, SUI 93101 CCD whole x 0.5. Fig. a. Solenastrea bournoni Milne Edwards & 1849; (CCD 4366), locality AB94-13, colony,

Fig. b. Trachyphyllia bilobata (Duncan, 1863); SUI 92040, Wells Colin, locality ER 180, calical surface, x 0.65.

alveolus Tulane d — side 0.9. Figs c, d. Placocyathus (Duncan, 1863);USNM 95334, locality 705; c — calical surface, x 0.9; view, x

barretti USNM USGS — Figs e, f. Placocyathus Duncan, 1863; 63371, locality 2580; e calical surface, x 0.65; f — side view, x 0.65.

cubensis Edwards & Haime, SUI 92039, Wells ER calical surface, 0.65. Fig. g. Scolymia (Milne 1849); Colln, locality 180, x

Fig. h. Antillia gregorii Vaughan, 1901; USNM 65469, USGS locality 2580, calical surface, x 1.