Contr. Tert. Quatern. Geol. 35(1-4) 129-146 1 fig., 1 table, 4 pis. Leiden, April 1998

Echinoderms of the Pliocene Bowden shell bed,

southeast Jamaica

Stephen K. Donovan

UNIVERSITYOF THE WEST INDIES

KINGSTON, JAMAICA

AND

Christopher R.C. Paul

UNIVERSITY OF LIVERPOOL

LIVERPOOL, ENGLAND

Donovan, Stephen K. & Christopher R.C. Paul. of the Pliocene Bowden shell bed, southeast Jamaica. In: Donovan, S.K. (ed).

The Pliocene Bowden shell bed, southeast Jamaica.— Contr. Tert. Quatern. Geol., 35(1-4): 129-146,1 fig., 1 table, 4 pis. Leiden, April

1998.

The the of unit in fauna of late Pliocene Bowden shell bed of southeast Jamaica is more diverse than that any coeval the

echinoderm have been identified from the Bowden shell bed: asteroid Caribbean. The following taxa goniasterid or astropectinid sp. indet.,

indet., and the echinoids Eucidaris diadematid indet.,Arbacia Echinometra ophiuroid sp. madrugensis (Sánchez Roig, 1949), sp. sp., sp.,

cf. indet. and indet. Adult echinoids Tripneustes sp., carrizoensis Kew, 1914, scutelline sp. spatangoid sp. are invariably

tests have because of their low volume them less preserved as fragments; juvenile may survived to surface area ratio, making prone to

mechanical collapse. At the ordinal level, Eucidaris - diadematoid- Echinometra - arbaciid - toxopneustid - Clypeaster - scutelline -

spatangoid echinoid faunas seem to have been the norm in the Caribbean during the Pliocene. Echinoids across the Pliocene-Pleistocene

in the Caribbean follow similar of faunal that ofbenthic molluscs. interval region may a pattern turnover to

Key words — Bowden shell bed, Pliocene, Echinodermata, Asteroidea, Ophiuroidea, Echinoidea.

S.K. Donovan, Department of Geology, University of the West Indies, Mona, Kingston 7, Jamaica; C.R.C. Paul, Department of Earth

Sciences, University of Liverpool, P.O. Box 147, Liverpool, L69 3BX, England.

Contents species of asteroid, an ophiuroid, plus at least eight species

of echinoid, making it the most diverse Pliocene echino-

derm known from the Antillean Introduction p. 129 assemblage region (Dono-

van & Paul, Systematic palaeontology p. 130 1996).

The echinoids of the Pleistocene deposits of eastern Discussion p. 134

Jamaica have been documented in detail et al., Acknowledgements p. 135 (Donovan

References 135 1994b; Donovan & Portell, 1996; Donovan & Embden, p. 1996) and this analysis is extended into the Pliocene herein.

Prior to the account of Donovan & Paul (1996), the only

of fossil echinoid Introduction report a from Bowden was by Weisbord

who noted that, 'In the late Miocene (1969, p. 309), [Echi-

The known echinoid fauna of the Pliocene of Jamaica is nometra] lucunter is reported from Bowden, Jamaica.' (The sparse (Donovan, 1993). Donovan & Paul (1996) have Bowden Formationwas previously dated as Miocene; see

This has been recently augmented this fauna, otherwise based mainly on Donovan, 1998). specimen(s) not located by

it if it reports of more or less complete tests, in a preliminary S.K.D. However, remains uncertain truly came from

the Bowden shell bed the Bowden report ofthe moderately diverse echinoderms from the late or even Formation. For

Pliocene Bowden shell bed of southeast Jamaica, the fauna comparison, Caldwell (1966) recorded a needlefish jaw of which re-identified the claw of is largely disarticulated. The purpose of the (subsequently as a callianassid

W.C. Blow in Clarke & Fitch, present paper is to document fully the fauna of the Bowden crustacean; 1979), purport- shell which is from the Bowden but the of the bed, now known to consist of at least one edly Formation, description 130

shows it have been collected from locality to an overlying data). The first Pliocene asteroids known from the island

unit, either the Old Pera beds (early Pleistocene) or the Port are from the Bowden shell bed.

Morant Formation (late Pleistocene). Prior to Robinson's The precise taxonomic assignment of this small collec-

(1969) definition of the Bowden Formation sensu stricto, tion of marginal ossicles is problematic. The specimen

all of these units in 'Miocene' were 'lumped together' a figured by Donovan & Paul (1996) appears to be a typical

Bowden Series (E. Robinson, pers. comm.). It should be astropectinid superomarginal (compare with specimens

noted that tests ofEchinometra, assigned to E. viridis A. figured by Blake, 1973; J.W.M. Jagt, written comm.). With

and Echinometrai? Agassiz, 1863, sp., occur in the basal one possible exception, all of these specimens could con-

conglomerate of the Port Morant Formation (Donovan et ceivably have been derived from a single species. The

al., 1994b). It is considered at least possible, perhaps figured specimen shows the typical facet morphology of

these ossicles. probable, that this is the horizon which yielded Weisbord's Externally they are all smooth and unsculp-

at E. lucunter (Linne, 1758). tured or, in least one example (EE 5214[1]), perhaps

The is Specimens used in the present study are deposited in the weakly pitted. only exception the biggest specimen

in lot EE which is broad with collections of the Natural History Museum, London 5237, a weak, pustular exter-

(BMNH EE 5209-EE 5230, all SEM stubs; and EE nal sculpture, the largest pustules being concentrated

to the and distal facets. This indi- 5237-EE 5246) and the Florida Museum of Natural His- adjacent proximal may

the of second until tory, Gainesville (UF 67426-67437, 71283). Descriptive cate presence a species, but, more

terminology of the echinoderm test follows Gale (1987a), specimens are available, it is considered conservative to

Melville & Durham (1966), and Durham & Wagner (1966). include all ossicles in one species under open nomencla-

The classification of echinoids is that of Smith (1981, ture.

1984) and Smith & Wright (1993). Synonymy lists include Asteroids are unknown from the Jamaican Pleistocene.

Two Recent known from the island reported occurrences in Jamaica only. Plates 1-4 illustrate astropectinids are

recorded from bottoms. specimens sputter coated with 60% gold-palladium and Astropectensandy Notably, dupli-

1840 favours 'Soft examined using a Leica 440 scanning electron microscope catus Gray, (1-550 m water depth)

All BMNH sediment of sand shell hash' (SEM). specimens were processed from bulk composed or (Hendler et at.,

from unit 2 of the Bowden shell bed substrate for samples (sensu Pick- 1995, p. 73), suggesting a possible preference

the in the Bowden shell erill el al., 1998) unless otherwise stated. All specimens are species preserved bed.

from the Bowden shell bed unless indicated otherwise.

Class Ophiuroidea Gray, 1840

SYSTEMATIC PALAEONTOLOGY Incerti ordinis

indet. Class Asteroidea de Blainville, 1830 Ophiuroid sp.

PI. 8 Subclass Neoasteroidea Gale, 1987b 1, Fig.

Family Astropectinidae Gray, 1840 or

1996 vertebral ossicles. — Donovan & Paul, GoniasteridaeForbes, 1841 Ophiuroid

table 1. p. 166,

Asteroid sp. indet.

Material— A vertebral BMNH EE 5212. PI. 1, Fig. 2 single ossicle,

Remarks — Like asteroids, no complete fossil ophiuroids

— known from the Jamaican rock record. Disarticulated 1996 Astropectinid or goniasterid. Donovan & Paul, are

p. 166, fig. 1G, table 1. vertebral ossicles have been recorded from the Oligocene

and Pleistocene of the island (Donovan et al., 1993; Dixon

Material, localities and horizons — Thirteen from marginal et al., 1994); they are also known the Miocene (R. W. ossicles from the Bowden shell bed: BMNH EE 5213(1-4], Ported, pers. comm.). The Bowden vertebra differs from

EE and BMNH EE EE those 5214(1-3] 5237(1-6], 5213(1] was previously described from Jamaica in having a strep- illustrated by Donovan & Paul (1996). A further ossicle, tospondylous, hourglass-shaped, 'ball-and-socket' articula-

BMNH EE 5238, collected from the Bowden Forma- tion was (G. Hendler, written comm.; compare with, for exam-

tion at section 5 of Pickerill et al. towards the (1998), top ple, Mortensen, 1933, fig. 2). Smith et al. (1995, p. 236) of the formationand 1 below the micritic over m prominent noted this type of articulation in the Ophiocanopidae,

limestone horizon. Ophiobyrsinae, Hemieuryalinae and the euryalid groups. In

Remarks — No fossil asteroids have been found complete the extant Caribbean shallowerwater fauna, these groups

in the Jamaican rock but disarticulated record, marginal are mainly represented by gorgonocephalids and hemieury- ossicles Previous have are locally common. reports men- alids (Hendler et al., 1995).

tioned specimens from the Upper Cretaceous, Eocene and

Oligocene (Donovan et al., 1993; Dixon el al., 1994); they

are also known from one Miocene locality (S.K.D., unpubl. Class Echinoidea Leske, 1778 131

Subclass Cidaroidea Claus, 1880 fig. ID, table 1.

1996 Indeterminate diadematoid.— Donovan, 35, Order Cidaroida Claus, 1880 p. fig.

IB. Family Cidaridae Gray, 1825

in Diadematoid.— Donovan, fig. 6.12D. Genus Eucidaris Pomel, 1883 pr.

Material— 27 spines, UF 67433 (19 spines), BMNH EE

5216[l-7], EE 5229[2]; and one test plate, BMNH EE Eucidaris madrugensis (Sánchez Roig, 1949) 5210[ 1]. PI. 1, Figs 1, 5-7, 9

Remarks — Diadematoids in the Bowden shell bed are

represented by fragments of spines and rare test fragments, 1996 Eucidaris madrugensis (Sanchez Roig). — Donovan but there is little of these further & table 1. prospect classifying Paul, p. 166, fig. 1C, without material Gordon, However, 1996 Eucidaris madrugensis (Sanchez Roig). — Donovan, superior (C.M. 1990).

1A. the nature of the diadematoidtest the p. 35, fig. precludes possibility

in Eucidaris — that will be in the turbidite pr. madrugensis (Sanchez Roig). Donovan, they preserved deposits (Picker-

fig. 6.12B. ill et al., 1998) of the Bowden shell bed (Greenstein, 1991;

Donovan & Gordon, 1993). Although unknown from

Material— 89 spines, UF 67434 (52 spines), BMNH EE articulated specimens in the Jamaican fossil record, the

5211 1 EE EE BMNH EE diadematoids known from the [ -4], 5229[1], 5230[l-4], distinctive spines of are now

5246f 1 -28]; a genital plate (madreporite), BMNH EE early Palaeocene (Donovan & Veltkamp, 1992), late Plio-

1 and 10 test BMNH EE 5218[ ]; plates/test fragments, cene, early Pleistocene (Donovan et al., 1994b; Donovan &

5219[ 1 -3], EE 5245[l-7]. For a more complete synonymy, Embden, 1996) and late Pleistocene of the island (C.M. see Cutress (1980, p. 70). Gordon, 1990). Two species, Diadema antillarum(Philippi,

Remarks — of Eucidaris Pomel Spines are locally a com- 1845) and Astropyga magnifica A.H. Clark, 1934, are

of Jamaican of late environments in the mon component sedimentary deposits common in modern shallow water middle Eocene age and younger (Donovan et al., 1991; Caribbean (Hendler et al., 1995).

Donovan, 1993). Cutress (1980) identified two nominal species from the fossil record of the Caribbean, Eucidaris

Arbacioida 1900 tribuloides (Lamarck, 1816) and E. madrugensis. The Order Gregory,

Arbaciidae 1855 former species is common in the Recent (Hendler et al., Family Gray, Arbacia 1835 1995) and Pleistocene (Donovan & Embden, 1996; Table Genus Gray,

1 herein) of the island, but E. madrugensis has only been identifiedin Jamaica from the Bowden shell bed (Donovan Arbacia sp. & Paul, 1996), where it forms a distinctive and abundant PI. 2, Figs 2, 8; PI. 3, Figs 1-4,6 part of the echinoid fauna. Primary spines of this species have a low, ribbed crown with a central, raised boss and

— & table 1. 1996 Arbacia sp. Donovan Paul, p. 167, longitudinal rows of spinules typical ofEucidaris, but are

1996 Juvenile regular echinoids incerti ordinis. — Donovan distinctive in having some spinules developed as thorns, table 1. & Paul, p. 167, fig. IE, F, often arranged in circlets (PI. 1, Figs 5-7). 1996 Toxopneustid fragments (pars). — Donovan & Paul, The only Caribbean species with broadly similar spines table p. 167, 1. is the approximately coevalPrionocidaris cojimarensis

(Lambert & Sanchez Roig in Sanchez Roig, 1926), of Material— Three tests, UF 67430, BMNH EE 5215 [1, 2]; However, Miocene-Pliocene(?) age (Cutress, 1980). spines and two test fragments, BMNH EE 5226 [1, 2], For com- of the latter taxon lack the typical Eucidaris crown and Arbacia below. ment on possible spines, see Tripneustes sp. show a much broader range of morphologies than is seen in Remarks — UF 67430 is only moderately preserved (un- the Bowden shellbed. Eucidaris madrugensis is otherwise fortunately, it readily absorbed glue in being mounted for limited to the Miocene-Pliocene of Cuba and the Domini- SEM) and small in size (= juvenile?). However, it is appar- can Republic (Cutress, 1980; but see comments by Kier, Arbacia, in the of the ently an a genus common Neogene 1992, p. 14). southeastern USA and Caribbean (Cooke, 1959; de

Buisonje, 1974; Lewis & Donovan, 1991). Other specimens

attributed to Arbacia herein include certain test fragments Subclass Euechinoidea Bronn, 1860 from the adoral region (note buccal notches; PI. 2, Figs 2, Order DiadematoidaDuncan, 1889 8) and two further, small, juvenile tests (PI. 3, Figs 2-4, 6). Family DiadematidaeGray, 1855 is shown the ab- That the latter are, indeed, juveniles by

and the sence of genital pores (PI. 3, Figs 3, 4) gaps be-

tween sutures within plate columns (PI. 3, Fig. 2). The Diadematid sp. indet. plate smaller have PI. 2, Figs 1, 6 specimen (PI. 3, Figs 4, 6) appears to single

1996 indet. — Donovan & not in the ambulacral BMNH EE Diadematoid sp. Paul, p. 166, pores, pore pairs, regions. 132

5215[1, 2] are not dissimilar to someLytechinus species, faunas throughout the region (Hendler et al., 1995). The

such of four Bowden is as Lytechinus williamsi Chesher, 1968, one shellbed specimen (at least superficially) closer

species of extant Western Atlantic Lytechinus (for further to E. viridis, which has its pore pairs arranged in arcs of

comment this five 1993, on genus, see Tripneustes sp.). However, (Donovan, p. 382).

there appears to be a weakly developed, granular sculpture

present interradially, and on the genital and ocular plates,

in the smallest tests (PI. 3, Figs 2-4, 6); also note what Family Toxopneustidae Troschel, 1872

be Genus L. 1841 appears to a similar sculpture on the smaller test frag- Tripneustes Agassiz,

ment (PI. 2, Fig. 8). This is at least reminiscent of sculpture

seen in Pliocene Arbacia improcera (Conrad, 1843) and

morphologically similar, coeval species (see Lewis & Tripneustes sp.

Donovan, 1991). PI. 2, Fig. 3

that the three Assuming juvenile tests were truly de-

rived from the — same species, then they represent a growth 1996 Toxopneustid fragments (pars). Donovan &

table 1. series from BMNH EE 5215[2] (PI. 3, Figs 4, 6) through Paul, p. 167,

BMNH EE 5215[ 1] (PI. 3, Figs 2, 3) to UF 67430 (PI. 3,

The smallest test is Material— 28 ambulacral fragments, BMNH EE 5226[3], Fig. 1). single-pored, but pore pairs have developed in the specimen of intermediate size. Only BMNH EE 5227[1, 2], BMNH EE 5239(1-25]; and 13

the largest test has developed typically arbacioid naked interambulacral fragments, UF 67428, BMNH EE

zones in the interambulacra. 5240(1-12], BMNH EE 5209(1] is a rotula probably de-

This is only the third species of arbaciid recognised rived from a toxopneustid (compare PI. 1, Fig. 3 with C.M.

from the Jamaican fossil record, after the late Cretaceous Gordon, 1990, text-fig. 2-26). 115 spines (PI. 2, Figs 4, 5)

be derived fromArbacia Echinometra Goniopygus supremus Hawkins, 1924, and an indetermi- may sp. and/or sp.

nate species from the early Pleistocene (Donovan & Emb- and/or Tripneustes sp.; UF 67435 (85 spines), BMNH EE

den, 1996). ExtantArbacia punctulata (Lamarck, 1816) has 5218(2], EE 5229(3], EE 5241(1-28]. A keeled tooth,

not been recognised from modern shallow water environ- BMNH EE 5209(2], is probably derived from Tripneustes

Echinometra ments of Jamaica (J.D. Woodley, pers. comm.). sp. or sp.

Remarks — The following comments by Mortensen (1943,

the p. 383) were of particular relevance in identifying

Order Echinoida Claus, 1876 toxopneustid fragments of the Bowden shell bed: 'The

ambulacral is found in the Family Echinometridae Gray, 1825 simplest [toxopneustid] type

the Genus Echinometra Gray, 1825 (genus]... Lytechinus [amongst others] ... plates being

trigeminate, each with a primary tubercle. The pore arcs are

more or less are oblique ... Trigeminate ambulacral plates

Echinometra found also in here sp. ... Tripneustes [amongst others], but a

PI. 2, 7 is found Fig. primary tubercle only on every second or third (or

the adoral which fourth) plate (except plates have a primary

1996 Echinometra — Donovan & table 1. sp. Paul, p. 167, tubercle each). This results in the plates being lower than

where each plate carries a primary tubercle, and the

Material— An ambulacral with of at five plate an arc least pore-arcs being more horizontal, and the pore pairs be-

pore pairs, BMNH EE 5210[2], coming more separated from each other, so as to form three

Remarks — This single ambulacral plate is the evi- vertical series This is carried in only separate ... to an extreme

dence of the presence of Echinometra in the Bowden fauna Tripneustes, where even the original trigeminate plates

(compare PI. 2, Fig. 7 with Gordon & Donovan, 1992, fig. become partly occluded from the midlineof the ambula-

have been derived from and combine form ...'. 2A), although some spines may crum to pseudo-polyporous plates

this genus (see below). C.M. Gordon (1991) discussed why Fragments of toxopneustid test from the Bowden shell bed

Echinometra has fossil record in the an apparently poor (PI. 2, Fig. 3) are typical of Tripneustes. Indeed, the ar-

a robust however, Caribbean, despite having relatively test; rangement of pore pairs is at least close to that seen in the

it is locally common as disarticulatedelements (Gordon & extant CaribbeanTripneustes ventricosus (Lamarck, 1816)

Donovan, 1992). As noted above,Echinometra lucunter, (compare PI. 2, Fig. 3 with Mortensen, 1943, figs 239c,

noted from Bowden Weisbord by (1969), was probably 299a).

from the basal Port Morant Formation. FossilEchinometra The only other Pliocene toxopneustids noted from the

in Jamaica is best known from the but else- Caribbean Pleistocene, region are Lytechinus variegatus (Lamarck,

where in the region it is also recorded from the Oligocene 1816) and Tripneustes cf. ventricosus (Lamarck) from

and Miocene (Sanchez Roig, 1949; W.A. Gordon, 1963; Venezuela. The former record is based on two fragmentary

Poddubiuk & Rose, 1985). Extant E. lucunter and E. viridis tests (Cooke, 1961, pi. 4, figs 1, 2), while the latter is based

are common components of hard substrate, shallow water on four fragments of test (Weisbord, 1969, pi. 15, figs 133

nominal 6-13). It is probable that Pliocene members of this family the Bowden specimen. This adds yet another

is the Jamaican fossil fauna & are more widespread within the Antillean region than Clypeaster to (Donovan

recognised, but the generally fragmentary preservation Portell, 1996).

probably discourages identification. The two common

Caribbean the toxopneustid generain the at present day are

Lytechinus and Tripneustes (Hendler et al., 1995). Suborder Scutellina Haeckel, 1896

Incertae familiae

Cohort Irregularia Latreille, 1825

indet. Order Clypeasteroida A. Agassiz, 1872 Scutelline sp.

Suborder Clypeasterina A. Agassiz, 1872 PI. 3, Figs 5, 8; PI. 4, Fig. 2

Family L. Agassiz, 1835

indet. — Donovan & Genus Clypeaster Lamarck, 1801 1996 Lunulate scutelline gen. et sp. Paul, 167, 1A, B, table 1. p. fig.

1996 — Juvenile scutelline clypeasteroid. Donovan, p.

35, fig. 2. Clypeaster cf. carrizoensis Kew, 1914

Fig. 1

Material — Two juvenile tests, UF 67429and 67431: plus

fragments of tests; BMNH EE 5217, UF 67432 and 67436.

of There are very numerous fragments clypeasteroid,

including many that are obviously scutelline, in BMNH EE

5220[ 1-4], EE 5224[l-4] (PI. 3, Fig. 7), EE 5225[l-4], EE

5243 (numerous, larger fragments of test, mainly scutel-

line) and EE 5244 (numerous smaller fragments of test).

Remarks — The Bowden shell bed scutellineclypeasteroid

is best recognised from low ambital fragments with rarely

with preserved parts of lunules (PI. 4, Fig. 2). Comparison

complete tests of the approximately coeval Encope homala

Arnold & Clark, 1934, from Jamaica (Donovan et al.,

1994a) suggests that they are not conspecific. Rarejuvenile

with clypeasteroids from Bowden, always a poorly pre-

button-like and served apical region, are low, are presumed

to be conspecific with the fragments in the shell bed,

small have lunules although they are too to developed (PI.

3, Figs 5, 8). Nominal, coeval scutellines from elsewhere

in the Caribbean (Donovan & Paul, 1996, table 2) are

limited to Encope spp.; however, the Bowden scutelline

does not appear to belong to this genus.

The majority of test fragments listed above are scutel-

line in origin, particularly those from the ambital region,

some of the smaller, more nondescript fragments - although Fig. 1. Clypeaster cf. carrizoensis Kew, 1914, UF 71283, a be derived from Clypeas ter cf. carrizoensis Kew. It is apical view, b - oral view, c - lateral view (anterior to- may their wards left). Specimen whitenedwith ammonium chloride interesting to note that, despite high preservation

sublimate. potential (Kier, 1977; Donovan, 1991; Greenstein, 1993),

mature clypeasteroid echinoids in the Plio-Pleistocene

Material— A single juvenile test, UF 71283. For discus- succession of the Bowden area are invariably preserved as

scutelline indet., residence time of dead tests in sion of clypeasteroid fragments, see sp. fragments, suggesting long

below. the sea floor rather than catastrophic entombmentof live

Remarks — The juvenile test from the Bowden shell bed is specimens. At least some breakage of specimens is cer-

to ‘cotteaui’ of due to based sedimen- close in gross morphology the morphotype tainly post-mortem transport, on

.Rose & Poddubiuk (1987) (compare Fig. 1 herein with tological evidence, in both the Bowden shell bed (Pickerill

There least nomi- and the Old Pera beds Cooke, 1959, pi. 12, figs 1-3). are at ten et al., 1998) overlying (Donovan et

nal species of Clypeaster known from the Pliocene of the al„ 1994b).

Caribbean region (Donovan & Paul, 1996, table 1). Of

these, only Clypeaster carrizoensis Kew, illustrated by de

has Buisonje (1974, p. 240, pi. 11, figs 5-7) from Curasao, Order Spatangoida Claus, 1876 and in the same broadly open petals gross morphology seen Incertae familiae 134

Spatangoid sp. indet. across plate boundaries (such as PI. 2, Figs 2, 8), although

PI. 1, Fig. 4; PI. 4, Figs 1, 3, 4 some of this breakage certainly represents damage during

post-collection preparation, whilecomplete, disarticulated

1996 indet. — Donovan & Spatangoid sp. Paul, p. 167, plates were probably derived from long dead specimens

table 1. (for example, PI. 2, Fig. 3; PI. 3, Fig. 7) (Smith, 1984, pp.

in pr. Indeterminate spatangoid. — Donovan, fig. 6.14C. Juvenile survived their 17-19). tests may have because of

low volume to surface area ratio when compared with

Material— Numerous test fragments, BMNH EE 5221, adults, making them less prone to mechanical collapse; it EE 5222[l-3], EE 5223[l-3], EE 5228, EE 5242 (numerous is perhaps significant that the only test to retain spines (PI. specimens), UF 67426 (2 fragments), UF 67427 (2 frag- 3, Figs 4, 6) is also the smallest. and six BMNH EE UF 67437. ments); radioles, 5218[3], At the level of echinoid order, the faunaof the Bowden

Remarks — That this spatangoid followed a burrowing shell bed is diverse than that of other Pliocene more any habit is indicated by common test fragments preserving stratigraphic unit in the Caribbean region (Donovan & fascioles 4, ambulacrum III with (PI. Fig. 1); (anterior) Paul, 1996, table 2). The paucity of known Pliocene echi- the of tube feet for pores indicating presence adapted noderms noted for Jamaica (Donovan, 1993) seems to

a funnel 4, constructing respiratory (PI. Fig. 3; compare extend to the rest of the region. However, as noted by with and of sunken ambu- Smith, 1980); rare fragments Donovan & Paul (1996), comparison of the Bowden shell

lacral 4, The so far petals (PI. Fig. 3). only apical system bed echinoids with coeval Caribbean faunas does suggest identifiedincludes three This is genital pores. arrangement some similarities. Clypeasteroids and spatangoids are the unknown from of the nominal Pliocene any spatangoids, most Nominal widespread groups. clypeasteroids are including various brissids and schizasterids, hitherto identi- limited to Clypeaster spp. and the scutelline Encope spp. fied from the & Paul, 1996, table region (Donovan 2). Spatangoids include various brissids and schizasterids; the McNamara (in press) has noted However, intraspecific fragments from Bowden suggest that the Jamaican species,

variation in the number of in an extant Austra- gonopores too, was a deep burrower. Regular echinoids are poorly

lianSchizaster, suggesting that a genetic relationship may known and are mainly Eucidaris spp., a taxon that is easily nevertheless be between the Jamaican material and possible identifiedfrom its large, distinctive spines (Cutress, 1980). coeval from elsewhere in the specimens region. Venezuelan toxopneustids have been assigned to extant

referred to a the of test Although single species, range Caribbean species. Only the Bowden shell bed seems to thicknesses and variations in tuberculation shown the by include diadematoids, but other faunas have not been Bowden shell bed indicate that spatangoid fragments may examined for microscopic fragments. Echinometra is

more than one taxon is Partial tests of spatangoids present. known from complete tests in Jamaica and Curasao. At the have been collected from the overlying BowdenFormation, ordinal level, it therefore seems probable that Eucidaris - and are & R.W. Portell, awaiting preparation (Donovan - diadematoid - Echinometra - arbaciid - toxopneustid research in progress). Clypeaster - scutelline - burrowing spatangoid echinoid

faunas may have been the norm for the Caribbean during

the Pliocene.

DISCUSSION Table 1 illustrates the distribution of known echinoid

taxa in the Pliocene to Pleistocene interval of Jamaica, in

Two distinct preservational styles are shown by echinoid rocks representing a number of depositional environments.

specimens in the Bowden shell bed. Only juveniles occur At the level of echinoid order/family, the fauna of the

as complete tests, begging the question - were live adults Bowden shell bed (although almost certainly including

shows not present in the original fauna or were they all destroyed species from a range of palaeoenvironments) good

by transport in the turbidity currents? It is already known agreementwith that seen in Jamaican Pleistocene deposits

that tests of small echinoids can survive transport and from both shallow and deeper water shelf environments;

similar association of taxonomic is deposition as part of a turbidite(see, for example, Donovan indeed, a higher groups

& Pickerill, 1993), so the occurrence ofjuveniles (at least seen in the modern shallow-water fauna of the Caribbean

three out of the eight species) is perhaps not surprising. (Hendler et al., 1995). Those groups that are noticeable

Although 'fresh' tests ofmature echinoids that were tum- common throughout this range are Eucidaris, diadematids, bled by Kidwell & Baumiller (1990) showed only slow Echinometra, Clypeaster, scuteilines and spatangoids.

mechanical breakdown, this would presumably have been However, the rare species (Eucidaris madrugensis, Cly-

accelerated under the abrasive conditions of greatly more peaster cf. carrizoensis) from the Bowden shell bed that are

turbidite deposition. Therefore, it is at least probable that recognisable to the level of species, however tentatively,

the echinoid debris documented herein is fragmentary a differ from those of the same genus that are common

mixture of remains representing specimens that were both throughout the Pleistocene. Although the data available alive and dead at the time of deposition, the former being from Bowden and elsewhere(Donovan & Paul, 1996, table

derived from 'live' broken up during transport. Fragments 2) are far from perfect, there is no reason to doubt that echinoids be broken may represented by test fragments echinoids across the Pliocene-Pleistocene interval in the 135

H.G., 1860. Klassen und Ordnung des Thier-Reiches, 1. Caribbean region may follow a pattern of faunal turnover Bronn, Amorphozoen. Leipzig/HeidelburgC.F. Winter, 434 [not similar to that shown by the benthic molluscs (Jackson et pp. seen]. ai, 1993; Allmon et al., 1993). Buisonje, P.H. de, 1974. Neogene and Quaternary geology of

Aruba, Curasao and Bonaire. — Natuurwetenschappelijke

Studiekring voor Suriname en de Nederlandse Antillen, Uit- ACKNOWLEDGEMENTS 78: 1-293. gaven,

Caldwell, D.K., 1966. A Miocene needlefish from Bowden,

This research was initiated during the period of a to grant Jamaica. — Quarterly Journal of the Florida Academy of Sci- the authors from the Leverhulme which we 339-344. Trust, grate- ences, 28(1965):

We thank Walter R. Brown and Susann fully acknowledge. Chesher, R.H., 1968. Lytechinus williamsi, a new from

G. Braden of the SEM Laboratory, National Museum of Panama. — Breviora, 305: 1-13.

Natural History, Smithsonian Institution, for their assis- Clark, A.H., 1934. A new sea-urchin from Florida. — Journal of the Academy of Science, 24: 52, 53 seen]. tance in the preparation of SE micrographs for Plates 1-4. Washington [not Clarke, M.R. & J.E.Fitch, 1979. Statoliths of Cenozoic teuthoid These were taken whileS.K.D. was in receipt of a Smith-

cephalopods from North America. — Palaeontology, 22: sonian Institution Senior Research Fellowship, which he 479-511. gratefully acknowledges. Roger W. Portell is thanked for Claus, C.F.W., 1876. Grundziige der Zoologie (3rd edition). arranging the loan of UF specimens. We are particularly 822 Marburg/Leipzig, pp. [not seen]. grateful to Gordon Hendler for his remarks concerning the Claus, C.F.W., 1880. Grundziige der Zoologie (4th edition). affinities of the Bowden Edward Robinson for ophiuroid, vii 821 Marburg/Leipzig, + pp. [not seen]. units his comments on the nomenclatureof the at Bowden 1843. of and of Conrad, T.A., Description a new genus, 1969, and Dirk Nolf for Clarke & Fitch and fossil shells the prior to to bringing twenty-nine new Miocene, one Eocene of

(1979) to our attention. Special thanks to Carla Gordon and United States. — Proceedings of the Academy of Natural Sci-

of 305-311 Hal Dixon for their help in the field, and to Yolande Nor- ences Philadelphia, 1: [not seen]. Cooke, C.W., 1959. Cenozoic echinoids of eastern United States. man for washing samples. David Lewis (BMNH) and John

— United States Geological Survey Professional Paper, 321: W.M. Jagt (Natuurhistorisch Museum Maastricht) are iii 106 review This + pp. thanked for their constructive comments. paper Cooke, C.W., 1961. Cenozoic and Cretaceous echinoids from was completed during the period of National Geographic

Trinidad and Venezuela.— Smithsonian Miscellaneous Col- Society grant 5722-96 to S.K.D., which is gratefully ac- lections, 142(4): 1-35. knowledged. Cutress, B.M., 1980. Cretaceous and Tertiary Cidaroida (Echino-

dermata: Echinoidea) ofthe Caribbean area. — Bulletins of

American Paleontology, 77(309): 1-221.

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PLIO P-P PLEISTOCENE

EARLY LATE

BSB RHB Man OPB PMF FaF

Order CIDAROIDA

StylocidarisStylocidaris afftnisaffinis - - + - -

Cidaris (Tretocidaris)(Tretocidaris) bartlettibartletti - - + + - -

Eucidaris madrugensismadrugensis + - - - -

- Eucidaris tribuloides - + + + + +

Order DIADEMATOIDA

Diadematid sp. indet. + - + + - +

Order PHYMOSOMATOIDA

Arbacia .....- - - Arbacia sp. +

Arbaciid sp. indet. - - + - -

Order TEMNOPLEUROIDA

? LytechinusLytechinus sp. indet. ....- - - . ?

Tripneustes sp. indet. + -- - - - ?

Order ECHINOIDA

Echinometra lucunter - - - +

- - - Echinometra viridis + - + +

Echinometra indet. - - - Echinometra sp.sp. indet. + + +

Order HOLECTYPOIDA

Echinoneus cyclostomuscyclostomus - - + - -

Order CASSIDULOIDA

- Echinolampas depressa -- - + - - -

Order CLYPEASTEROIDA

ClypeasterClypeaster cf. carrizoensis + - - - -

Clypeaster lampruslamprus - - + - -

Clypeaster rosaceus - + - + - -

Clypeaster subdepressuslsubdepressus - - + -

Encope homala - + - .-

Scutelline - - - Scutelline sp. indet. + +

Clypeasteroid sp.sp. indet. - -- - + - +

Order SPATANGOIDA

- Meoma ventricosaventricosa -- - - + -

PaleopneustesPaleopneustes cristatus - - + - -

Schizaster doederleini ....- - - +

indet. - - Spatangoids sp. + + + +

Incerti ordinis

Indeterminateregulars - - + + - +

Table 1. Stratigraphic distributionof the Plio-Pleistoceneechinoids of Jamaica (expanded after

Donovan & Embden, 1996). Key — PLIO = Pliocene; P-P = Plio-Pleistocene; BSB

= Bowden shell bed; RHB = Round Hill beds, August Town Formation (nearshore

siliciclastics/limestones); Man = Manchioneal Formation (including Navy Island

Member) (deeper water limestones); OPB = Old Pera Beds (siliciclastics); PMF =

Port Morant Formation (siliciclastics); FaF = Falmouth Formation(raised reef); + =

present; - = absent. 140

PLATE 1

Asteroids, ophiuroids and echinoids of the Pliocene Bowden shell bed, Jamaica.

Figs 1, 5-7, 9. Eucidaris madrugensis (Sánchez Roig, 1949), 1 - BMNH EE 5218[1], madreporite, x 25; 5 - primary spine, BMNH EE

5230[1], x 18; 6 - primary spine, BMNH EE 5211[1], x 20; 7 - primary spine, BMNH EE 5230[2], x 18; 9 - interambulacral

plate, BMNH EE 5219[1], x 25.

2. EE of 22. Fig. Astropectinid or goniasterid asteroid sp. indet., BMNH 5214[1], profde marginal ossicle, x

3. BMNH EE 20. Fig. Toxopneustid? sp., 5209[1], rotula, x

4. BMNH EE 14. Fig. Spatangoid sp. indet., 5218[3], spine, x

Fig. 8. Ophiuroid sp. indet., BMNH EE 5212, proximal surface, x 24. 141\1

PLATE 1 142

PLATE 2

Echinoids of the Pliocene Bowden shell bed, Jamaica.

Diadematid - - Figs 1,6. sp. indet., 1 BMNH EE 5216[1], spine, x 18; 6 BMNH EE 5210[1], interambulacral (?) plate, x 18.

2 - - Figs 2, 8. Arbacia sp., BMNH EE 5226[1], x 14; 8 BMNH EE 5226[2], x 30.

3. BMNH EE of 12. Fig. Tripneustes sp., 5227[1], fragment ambulacrum, x

Figs 4,5. Indeterminate regular echinoid spines, 4 - BMNH EE 5229[3], x 7; 5 - BMNH EE 5218[2], x 20.

BMNH EE 25. Fig 7. Echinometra sp., 5210[2], ambulacral plate, x 143\2

PLATE 2 144

PLATE 3

Echinoids of the Pliocene Bowden shell bed, Jamaica.

6.Arbacia - - lateral 4 and view of Figs 1-4, sp., 1 UF 67430, apical view, x 19; 2, 3 view (interambulacrum centre) apical BMNH EE

- view 5215[1], x 38 and x 30, respectively; 4, 6 apical and lateral view (interambulacrum centre; note single ambulacral

pores) of BMNH EE 5215[2], x 38 and x 48, respectively.

8. Scutelline 5 - UF oral 8 - UF surface Figs 5, sp. indet., juvenile tests, 67429, surface, x 25; 67431, apical (petaloid region not pre-

served), x 40.

Fig. 7. Scutelline ? sp. indet.,BMNH EE 5224[1], test fragment, x 25. 145\3

PLATE 3 146\4

PLATE 4

PLATE 4

Echinoids of the Pliocene Bowden shell bed, Jamaica.

from - EE 3 - BMNH EE Figs 1, 3, 4.Spatangoid sp. indet., 1 BMNH 5222[1], test fragment with fasciole, x 20; 5222[2], fragment

adjacent to apical system (anterior towards top of page), showing ambulacra III (right) and IV, and interambulacrum 3, x

18; 4 - BMNH EE 5221, indeterminate test fragment, x 12.

BMNH EE of 12. Fig. 2. Scutelline sp. indet., 5217, ambital fragment of test with edge lunule (left), x