Contr. Tert. Quatern. Geol. 35(1-4) 95-111 2 pis. Leiden, April 1998

Holoplanktonic

(: Heteropoda and Thecosomata) from

the Pliocene Bowden beds, Jamaica

Arie W. Janssen

XEWKJJA (GOZO), MALTA

Janssen, Arie W. Holoplanktonic Mollusca (Gastropoda: Heteropoda and Thecosomata) from the Pliocene Bowden Beds, Jamaica.In:

Donovan, S.K. (ed.). The Pliocene Bowden shell bed, southeast Jamaica. — Contr. Tert. Quatern. Geol., 35(1-4): 95-111, 2 pis. Leiden,

April 1998.

of Jamaica. Two Three Heteropoda and 14 Euthecosomata (‘Pteropoda’) are recorded from the Pliocene (Piacenzian) Bowden,

Heteropoda, four species of Limacinidae and Clio aff. braidensis (Bellardi, 1873) have not been recorded previously from Bowden.

followed of the for Cavolinia ventricosa Systematic descriptions are by a brief discussion on the age assemblage. Lectotypes are designated (Guppy, 1882) and Diacria digitata (Guppy, 1882).

Key words — Bowden, Jamaica, Heteropoda, Thecosomata, Pteropoda, Pliocene, systematics, biostratigraphy.

A.W. Janssen, Nationaal Natuurhistorisch Museum, P.O. Box 9517, NL-2300 RA Leiden,The Netherlands; or: 12, Triq il-Hamrija, Xewkija

VCT 110, Gozo, Malta.

Contents PectinibranchiataHeteropoda

Atlanta (Atlanta) diamesa Woodring

Atlanta lissa Woodring Introduction p. 95 (Atlantidea)

methods localities 96 Material, and p. discussed detail Collins The pteropods were in by Abbreviations p. 96 (1934, 155), who listed the following nine species and Age of the Bowden shell bed p. 97 p. also mentioned Atlanta one heteropod, sp. (Collins, 1934, Systematic palaeontology p. 97 146): Order Heteropoda p. 97 p.

Order Thecosomata p. 99 Cavolinia telemus (Linne) Conclusions 104 p. Cavolinia ventricosa (Guppy) Acknowledgements p. 105 Cavolinia digitata (Guppy) References p. 105 Cavolinia vendryesiana (Guppy)

Diacria bisulcata Gabb

Cleodora bowdenensis Collins Introduction Styliola sulcifera Gabb

Creseis acicula Rang The classic and renowned shell bed of Bowden (Jamaica) Collins Various Bowdenatheca jamaicensis has been the subject of numerous publications. authors have described the molluscan faunas from this In addition to the genus Bowdenatheca and its type deposit, from the nineteenth century onwards, culminating species, B. jamaicensis, Cleodora bowdenensis was erected in W.P. Woodring's monographs (1925, 1928). Woodring by Collins (1934), who distinguished an 'upper' and a mentionedthe of (1928, p. 113ff) following seven species 'lower' the latter the main shell bed from zone, being holoplanktonic gastropods from Bowden: which Woodring's molluscan materialoriginated. The three

last-mentioned species in the above list came from the Pteropoda Thecosomata lower zone, all others from the upper zone. Cavolinia telemus (Linne) The bulk of the material studied for this paper is in shell Cavolinia ventricosa (Guppy) preservation and originates from sediments yielding abun- Cavolinia digitata (Guppy) dant benthic molluscan material, although a part of the Cavolinia vendryesiana (Guppy) samples show distinct signs of aragonite dissolution. It is Diacria bisulcata Gabb assumed that most of the material comes from the main 96

shell bed, or from close it. layers very to Same locality as PJ 464 (NMB locality 10831), PJ 669

The revision of present the Bowden holoplanktonic (NMB locality 11036), PJ 743 (NMB locality 11110). Leg.

Mollusca comprises three species of Heteropoda and E. Robinson, Myrtle Haase, Alfred Rose, Richard de Souza

fourteen pteropod species. and P. Jung, 12 May 1969.

This sample comprises only a single specimen of

Cavolinia digitata.

MATERIAL, METHODS AND LOCALITIES

Bowden shell bed, Sample 1-4. — Material housed in the

Material studied for the Florida Museum ofNatural Gainesville present paper was largely (samples History, (Florida,

1-1 collected the Bowden shell bed Data follows: to 1-6) at type locality. U.S.A.). are as Bowden (XJ(X)2), Jamaica, St

For and Thomas geographic lithostratigraphic data on this locality, parish. Pliocene, Bowden Formation, in situ. Coll. reference is made to Zans et al. (1963), Robinson (1969) Roger Portell, 28 May 1990. The material comprises nine

and Pickerill et al. (1998). Details of the various samples, samples ofEuthecosomata, apparently picked from wash-

the as known to author, are given below. ing residues. Finer fractions from the same sample received

Reference material listed in the loan to systematic part below on appeared to yield no holoplanktonic molluscs.

includes numbers only sample as defined here, specifying the number of specimens, and followed by registration Bowden shell bed, Sample 1-5. — Material collected by number (for example: sample 1-1/12 specimens, RGM 396 S.K. Donovan, donated to the RGM collections in Septem-

006). ber 1995. Label stating: Pliocene Bowden shell bed, SE

Jamaica. The specimens were picked from washing resi-

Bowden shell bed, Sample 1-1. — Material collected in dues, five species occur. Most, if not all, of this material is

1975 by Dr Jan P. Krijnen, housed in the RGM collections. from unit 2 of the Bowden shell bed sensu Pickerill et al.

The is indicated locality as 'Bowden, road cut exposure, (1998). gully fill'. The material includes a large sample of benthic

molluscs. The discussed herein Bowden shell 1-6. — On the author's holoplanktonics were bed, Sample request, recovered from sediment infillof shells. S.K. and larger gastropod Donovan R.W. Portell collected a quantity of

Contrary to most other samples, the present sample yielded larger benthic gastropod shells at the Bowden type locality, molluscan material in excellent preservation, with hardly in order to recover holoplanktonics from their sediment

dissolution. The of the is for 1-1. The collected in any aragonite quality specimens contents, as sample sample was very good and only this sample yielded small specimens, February 1996 and subsequently donated to the RGM. such as limacinids. Sample 1-1 is therefore considered to Unfortunately, almost all specimens appeared to have been

the represent most complete holoplanktonic molluscan collected from a level of advanced decalcification. Most

in assemblage, which the species occur in almost natural shells were partly to completely dissolved, and sediment ratios. infill had become indurated, forming internal moulds. Still,

A 396 single specimen (RGM 032) in this sample, a a careful inspection of this material by breaking up the

shellof Cavolinia large tridentata, is stated to have come moulds and study under the microscope, yielded quite a from 'a higher level.' This specimen has suffered from numberof holoplanktonics, but exclusively larger species. aragonite dissolution, being preserved as an internal mould Ten species are represented in this material. with shell remains only.

Bowden Formation, Sample 2-1. — Sediment sample

Bowden shell bed, Sample 1-2. -— Material housed in the collected by S.K. Donovan, subsequently donated to the

Naturhistorisches Museum Basel (NMB, Switzerland) with RGM collections, September 1995, labelled: Bowden the following label: Jamaica, SE coast; Port Morant, Bow- Formation, from coastal section, stratigraphically above the den Fm, leg. Jooss and Strasser, 1931; don. Trinidad lease- shell bed. holds. The sediment is decalcified to a high degree and the few

C.H. Jooss and Strasser originally sampled five locali- fossils are represented as moulds and casts, stained dark ties (partly indicated B.l to B.5) within the Bowden For- reddish brown. Identification of the material is difficult mation. However, detailed geographical data are lacking. because of poor preservation. Four species could be recog-

All five localities are assumed to be situated east of Port nised.

2 in Morant, an area not larger than 200 m . This is NMB locality 10635.

ABBREVIATIONS

Bowden shell bed, Sample 1 -3. — Material housed in the

The Naturhistorisches Museum Basel (NMB locality 11146), following abbreviations are used: carrying the following label: PJ 778. Jamaica, Bowden

Shop, Bowden shell bed. Bowden Formation. Outcrop NMB Naturhistorisches Museum, Geology Depart- opened by bulldozer. 1:50,000sheet N; 1:12,500 sheet 235. ment, Basel (Switzerland). 97

material of this reference was studied RGM Nationaal Natuurhistorisch Museum (Palaeon- v the original by

tology Department, Cainozoic Mollusca), Lei- the present author.

den, The Netherlands (formerly Rijksmuseum

van Geologie en Mineralogie).

SMF Senckenberg Museum, Frankfurt am Main ORDER HETEROPODA

(Germany).

Ruthe UF Florida Museum of Natural History, University Family Wiegman &

of Florida, Gainesville(U.S.A.). Genus Atlanta Lesueur

USNM National Museum of Natural History, Smith-

sonian Institution, Department of Paleobiology, Type species — Atlanta peroni Lesueur.

Washington D.C. (U.S.A.).

Atlanta diamesa Woodring, 1928

PI. 1, Fig. la-c

AGE OF THE BOWDEN SHELL BED

v* 1928 Atlanta (Atlanta) diamesa, new species, Woodring,

shell bed the of 133, figs 23-25. In the classic literature on the Bowden age p. these deposits is mostly considered to be Miocene. Recent

material — and two research, however, has demonstrated them to be of a Type Holotype (USNM 369335) from Jamaica' poorly preserved paratypes 'Bowden, (Al- younger age. Collection). Aubry (1993) and Berggren (1993) discussed the drich/Johns Hopkins —The shell has distinct lenticular with biostratigraphy of Neogene deposits in eastern Jamaica in Description a shape double-walled that is broken detail, using calcareous nannofossils and planktonic fora- originally a peripheral flange, The for the The ratio is c. 0.29. minifera, respectively. Berggren's results are incorporated greater part. height/width

number of whorls is c. 4Vi. The has no sharp in Aubry's paper. teleoconch and is Lithostratigraphically, the typical Bowden shell bed boundary with the partly damaged or it be that the first belongs to the Lower Coastal Group, Bowden Formation. covered with sediment. Still, can seen

the of the formation is whorls form a gently elevated cone in the centre of At the type locality only upper part together

The famous Bowden shell bed is the the in shallow and therefore present or exposed. at shell, lying a depression initial whorls base of the section. invisible in side view. Apparently, the are

than the ultimate whorls. No surface Samples from the Bowden type section (Aubry, 1993, relatively higher

ornament is visible the whorls. On the last p. 152) yielded limited nannoplankton assemblages, of low on protoconch

of the whorl there is weak indicationof more or diversity and poor preservation, and could not be assigned part body a

less situated of which 4 are visible 1, to zone. Other nearby outcrops, however, yielded better radially knobs, c. (PI.

material and without exception could all be correlatedwith Fig. la).

fora- On the base of the shell the last whorls are nannoplankton zone NN16. In terms of planktonic only two umbilicus. Both the and miniferal biostratigraphy the BowdenFormation belongs to visible. There is a narrow on upper

is absent. lines zones PL3 to PL5, which means 'early late Piacenzian' lower shell wall spiral striation Only growth

which demonstrate wide forward with (Aubry, 1993, p. 158). Reference is made to Ferrero Mor- are visible, a curve, of the of in the half, tara & Pavia (in press) for a revised interpretation the strongest point curvature peripheral and chronostratigraphical indication 'Piacenzian'. touching the upper suture almost under a right angle the becoming almost parallel to the periphery on opposite

side.

not SYSTEMATIC PALAEONTOLOGY The is elongated heart-shaped, hardly or

indented by the penultimate whorl.

Material from the material, just a fragmen- Symbols used in the lists of synonyms in this chapter are —Apart type ROM those of Richter (1948): tary specimen is available: sample 1-1/1 fragment,

396 006 (PI. 1, fig. 2a-b).

Atlanta diamesa to be related 1881 (in Roman) cited reference contributesto the knowl- Discussion —/ seems closely However, in the edge of the taxon; to the Recent A. peroni (Lesueur, 1817).

* which first valid introductionof the taxon; latterthe early whorls are tighter and more elevated,

indicate shell responsibility for the identification is accepted by characteristics no doubt a juvenile distinctly

from that of the the present author; differing present species. Gabb from the (no symbol) responsibility for the identification is not The late Miocene (?) A. cordiformis in differs in several accepted by the present author, but there is no rea- Dominican Republic (Janssen, press) and of the son for doubt; respects, especially in form ornament early

? in the of the author there is whorls. It to be related to the Recent A. opinion present reason seems closely

to doubt the identification; inflata Souleyet (Janssen, in press). 98

Collins referred (1934, p. 146) to a shell ofAtlanta, cartilagenous material has disappeared, but still a juvenile.

collected in 1932 from a roadside exposure, c. 50 feet Other specimens of this species, as known from the Mediter-

above the base of the Bowden hill, accompanied by several ranean Pliocene(RGM collections), are always much smaller

the which is pteropod species. The present whereabouts ofthis specimen and never reach stage in the spiral ornament

is unknown. completed, as in the specimen at hand.

The juvenile and poorly preserved paratypes, as well as Further fossil occurrences of this species are known from

the fragment discussed here, cannot be assigned to this the early Pliocene ofnorthern Italy. In the Recent fauna this

with is distributedin the and species any certainty. species widely tropics subtropics.

Genus Oxygyrus Benson Genus Protatlanta Tesch

(= AtlantideaPilsbry, objective synonym)

Type species — Oxygyrus keraudrenii (Lesueur).

Type species — Atlanta souleyeti Smith, by monotypy.

Oxygyrus keraudrenii (Lesueur, 1817)

PI. 1, Fig. 3a-b Protatlanta lissa (Woodring, 1928)

PI. 1, Figs 4a-b, 5a-c

* 1817 de A. Atlante Keraudren, Keraudrenii, Lesueur, p. 391.

v* 1928 Atlanta (Atlantidea) lissa, new species, Woodring, p.

Additional — Reference is made the exhaustive synonymy to 134, pi. 2, figs 26, 27.

synonymy of this predominantly Recent species by van der

Spoel (1976, p. 137). Type material— Holotype (USNM 369336) and paratype

Description — Shell c. IVi times wider than high, involute. from 'Bowden, Jamaica' (Aldrich/Johns Hopkins Collec-

The is illustrated in The early whorls, inclusive of the protoconch, are invisible. tion). holotype a juvenile specimen,

than the with rather and Only slightly more body whorl is exposed, and the Woodring (1928), a damaged aperture worn

shell is umbilicated both and covered with The is than deeply above below. The surface, partly glue. paratype more

whorls double the size of the but than are very convex. At the aperture the width of the holotype, barely more an

l whorl is c. 2 A times that of the preceding whorl (apical internal mould on which shell remnants are present.

view). The apertural margin is broken. The shape of the Description — Shell dextral, planispiral, slightly more than

aperture is almost circular, slightly higher than wide and twice as wide as high when adult, relatively higher in juve-

slightly indented by the preceding whorl. nile specimens. The protoconch and early teleoconch whorls

The surface of the shell is covered with c. 20-22 spiral are only partly preserved in the holotype. The number of

lines that whorls is difficult to estimate because of incom- are wavy or demonstrate a very clear zigzag struc- protoconch

ture. On the periphery of the body whorl these spirals leave pleteness and sediment adhering to the shell. The complete

a zone free of ornament, on which only growth lines are shell of the holotype at least has 4 whorls, probably c. 5,

faint. Just behindthe the which that the shell has 5 xh whorls. The aperture spiral ornament stops quite means fully grown c.

abruptly on a slightly thickened set of growth lines. In front early whorls form a depressed cone that is excentric and

of this point the shell surface is smooth. positioned somewhat obliquely with respect to the body

The lines distinct the and de- whorl. At least the last whorl has thin growth are near aperture protoconch a very

scribe a backward curve close to the upper suture followed spiral threadrunning over the middle of the whorl.

by a somewhat wider forward curve above the peripheral There is slightly more than one rapidly expanding teleo-

belt, on which they strongly curve backwards again. On the conch whorl. The aperture is distinctly wider than high

base of the shell the shape of the growth lines is similar. It (damaged in the holotype); at the columellar side it is only

appears that the shell must have been deeply sinuated in early slightly indented by the preceding whorl. The base of the

growth stages. shell has a similar, slightly convex shape as the apical side of

In the only available specimen the aperture is irregularly the shell. The umbilicus is narrow, occupying c. 1/6 of total

but at the and lower suture the shell material shell diameter. The base of the last whorl is broken, upper protoconch

is still present over a short distance. From Recent specimens visible in the umbilicus, its surface is devoid of ornament

it is known that the shell material of fully-grown specimens (only visiblein the paratype).

is cartilagenous, and therefore it seems probable that the The surface of the teleoconch in all available specimens

specimen at hand is in fact complete or almost complete. is poorly preserved, but strongly curved forward growth lines

Material— Sample 1-1/1 specimen, RGM 396 005. are visible under slanting light. No spiral ornament is visible,

Discussion — to der the for two distinct lines the that According van Spoel (1976, p. 138), except spiral on periphery

enclose kind ofbelt the which the fully-grown shell is cartilagenous, covering the calcareous together a on periphery on

early whorls that become absorbed in adult specimens. Thus, growth lines curve strongly backward. However, this periph-

is visible it eral belt in visible in the material. as no dissolution in the present specimen, most not distinctly type

which Material — materialas above, and 1- probably is not a fully grown specimen from the Type specified sample 99

6/1 specimen, RGM 396 0()7. which will be discussed in a future paper.

Discussion —The few specimens from the Bowden Beds,

admittedly rather damaged, closely resemble the late Mio-

Domini- Limacina inflata cene Protatlantarotundata (Gabb) known from the (d’Orbigny, 1836)

PI. 1, 12a-b can Republic (Janssen, in press, pi. 1, figs 3, 4), the only Fig.

differences being the slightly flatter shell and the absence of

* 1836 Atlanta — spiral ornament in the Jamaican specimen. The protoconch (Heliconoides) inflata, d'Orb. d'Orbigny, p.

be less 174,pi. 12, figs 16-19. of P. lissa has a similar oblique position, but seems to ornamented.

Additional — Reference is made to van der Spoel Another Miocene specimen from the Langhian of Italy is synonymy

This (1967) for extensive Janssen (1990, in press) illustrated by Janssen (1995, pi. 1, fig. 3). specimen, an synonymy.

the listed additional Cenozoic records, including data on type internal mould, seems to have a higher protoconch than

Caribbean species, but the height/width ratio of the shell is material and type locality.

— The available material is not well similar. Description very pre-

served. Most specimens are defective, crushed or juvenile. A further interpretation of these Protatlanta specimens

The visible characteristics agree in general with the rich has to be deferred until more material is available, especially Miocene material describedfrom Australia (Janssen, 1990, from the Caribbean. Some more material is waiting to be

15, but there are minor differences. described from various Mioceneand Pliocene localities in the p. pis 2,3,10),

The from Bowden are somewhatmore tightly Mediterranean. specimens

coiled, resulting in a narrower body whorl (apical view).

have half Also the Caribbeanspecimens, when fully grown,

a whorl less and thus remain smaller. Finally, in the Bowden ORDER THECOSOMATA shell wall is form (visible in one specimen only), the slightly

at the of the internalreinforcement. With these Suborder Euthecosomata bulging place

differences, the Bowden shells more closely resemble the Family Limacinidae Gray Recent form ofL. inflata. Genus Limacina Bosc (= Spiratella de Blainville)

Material— Sample 1-1/40 specimens, RGM 396 014-015.

— closer between Discussion There might be a relationship Type species — Clio helicina Phipps. the Australian material of L. inflata and the European L.

miorostralis(Kautsky) than previously thought, since in the Limacinabulimoides (d’Orbigny, 1836) Australian specimens (Janssen, 1990, pi. 2, figs 5-7) the PI. 1, Figs 6-8 internal reinforcements near the aperture do not cause an

* visible of the shell wall. However, in the 1836 Atlanta (Heliconoides) bulimoides, d'Orb. — externally bulging is devel- 36-38. the rostrum only rarely d'Orbigny, p. 179, pi. 12, figs European species apertural oped and the height/width ratio is slightly lower.

Additionalsynonymy — See van der Spoel (1967, p. 53, fig.

21). Limacina 1 Description — In the available material from Bowden the sp.

and PI. 1, 11 larger specimens are all more or less broken deformed, Fig.

frequently with their apices depressed, and only some juve-

nile shells and undistorted. in all — four two of them juvenile are complete They agree Description Only specimens,

and the others available. One of the respects with material of the Recent L. bulimoides, both from strongly defective, are

is illustrated here. The shell is wider than but Atlantic and Pacific localities. A juvenile and an adult Recent juveniles high,

the much than in the shell are illustratedhere (PI. 1, Figs 9,10) to demonstratethis apex clearly protrudes, more so speci-

men described below as Limacina ? 2. The whorls are conspecificity. However, none of the Bowden shells has a sp.

and the base is fully grown apertural margin. For a description of Recent relatively convex perforate.

Material — RGM 396 916-017. specimens, the reader is referred to the relevant literature, as Sample 1-1/4 specimens,

specified above. Discussion — The poor material is insufficient for a specific

— identification.The almost have the 'ideal' Limacina Material Sample 1-1/9 more or less defective/juvenile juveniles

specimens, RGM 395 (K18-011. shape, but cannot with any confidence be assigned to any one

— of the described. This concise and illustra- Remarks So far, this species was only occasionally re- taxa description

corded from Pleistocene deposits. This is the first record tion are therefore only given here to record its occurrence.

frombelow the Pliocene/Pleistocene boundary. In the Recent An interpretation is deferred until more and better preserved

available. fauna this species has a wide distribution in tropical and specimens are

subtropical areas. Limacina 2 Very similarmaterial is available from the early Pliocene ? sp.

of southern France and northern Italy (RGM collections), PI. 1, Fig. 13a-b 100

Description — A single, strongly defective specimen might The growth lines are very slightly curved in apertural

undescribed of Limacina. The direction the ventral side and much represent an species shell, or on more strongly so on

rather what is leftof it, has slightly more than two very flat the dorsal side, curved in a clear V-shape on the lateral

whorls. In lateral view, the apex is hardly or not elevated; the margins. It follows that the dorsal apertural margin is higher

last whorl is carinated. The lines than the ventral Close to the the lines faintly growth are very one. aperture growth

faint, but slightly directed backwards. are stronger and rather regularly distributed. The apertural

Material — Sample 1-1/1 defective specimen, RGM 396 margins are sharp, not thickened or internally reinforced.

Material— 018. Sample 1-1/10mainly defective or fragmentary

Remarks — At first glance this specimen resembles Recent specimens, RGM 396 019-021.

Limacina lesueurii but in that Discussion — Collins his (d'Orbigny), species the apex (1934, pp. 221,222) compared new

is higher, the body whorl is not angular and the growth lines taxon with Vaginella, but quite rightly concluded that it does

less It cannot be excluded that the not to that because of differencesin the are oblique. present speci- belong genus, aper-

in of tural and the absence of lateral carinae. men is, fact, a protoconch a pyramidellid species, but in structures by He

other such the shell wall is found closer resemblance undulata protoconchs always considerably a to Vaginella (Gabb) and

thicker. In the before the shell wall in V. caribbeana with his Bowde- specimen me agrees Collins, which, together

thickness with Limacinidae. natheca he jamaicensis, suggested to 'belong to a distinct, but

closely related group.' Vaginella undulata and V. caribbeana

have recently been reassigned to the genusEdithinella by

Cavoliniidac Fischer Family Janssen (1995, p. 124), but B. jamaicensis demonstrates

Creseinae Subfamily Rampal several differencesand is not consideredto belong in Edithi-

Genus Bowdenatheca Collins nella. The main differences are the absence of transverse

undulations on the dorsal side of B. jamaicensis, the absence

Type species — Bowdenatheca Collins, jamaicensis by of more or less distinctly developed lateral grooves, and the

monotypy. fact that the juvenile shell is shed. In addition, species of

Edithinella have a widened and reinforced aperture when

Bowdenatheca jamaicensis Collins, 1934 fully grown. PI. 1, 14a-c, 15a-c Figs However, assignment to the Creseinae, as proposed

herein, and also by Zorn (1997), has not been substantiated

* 1934 Bowdenatheca jamaicensis n. Collins, 221, pi. sp., p. yet either. Especially the dorso-ventral flattening of the shell 13, figs 13-15. is rather unusual in that subfamily, but for the time being it

1959 Bowdenatheca R.L. Collins — jamaicensis Zilch, p. 51, seems to be the best solution. The taxonomic position of the fig. 171. species shouldbe reconsidered when its protoconch is found.

1980 Collins — 64. Bowdenathecajamaicensis Shibata, p. This species is now known to occur also in the Messinian

1982 Bowdenathecajamaicensis (Collins) [sic] — Lozouet and early Pliocene of the Mediterranean (Janssen, 1995; & 184. Maestraeti, p. Zorn, is 1997). It not yet known from deposits younger than 1982 Bowdenatheca jamaicensis Collins — Bernasconi & the Bowden Beds. A Pliocene of 218. Japanese occurrence a Robba, p.

closely related, if not identical, form was recorded Ujihara 1983 Bowdenatheca Collins — 80. by jamaicensis Shibata, p.

v. 1995 Creseinae ? — (1996) (see list). sp. nov. Janssen, p. 30, pi. 2, fig. 3a-d. synonymy

? 1996 Bowdenatheca ? — sp. Ujihara, p. 780, fig. 5/43-49.

1997 1934 — Bowdenathecajamaicensis Collins, Zorn, p.

35, pi. 4, figs 1-4. Genus Creseis Rang

— Description Shell vaginelliform, regularly conical in Type species — Creseis acicula Rang.

ventral view, slightly arched in lateral view. The protoconch

is unknown, having apparently been shed during metamor- Creseis acicula ? (Rang, 1828)

the is closed smooth phosis, as opening by a septum. The

* dorso-ventral diameter of the shell is 1828 juvenile slightly more Cleodora (Creseis) acicula N., Rang, p. 318, pi. 17, fig.

than the shell but towards the the shell be- 6. width, aperture

comes flattened. The width of the aperture is almost twice the

dorso-ventral diameter.The lengthwise curvature of the shell Additionalsynonymy — An extensive synonymy (mainly

fossil is slight and the ventral side is more strongly curved than the occurrences) may be found in Janssen (in press).

dorsal which is The — conical shell one, virtually straight. transverse curva- Description Elongated fragments, very

ture of the dorsal side is stronger than that ofthe ventral side, slowly increasing in shell diameter and with a very thin shell

which towards the aperture demonstrates a centralswelling, wall.

both sides. The Material— 1-1/11 RGM 396 022. accompanied by a very slight depression on Sample fragments,

lateral — shell parts are gradually rounded without carinae all Remarks The available fragments closely resemble the

over their length. well-known Creseis acicula, but lack the final proof of the 101

found. it be Clio aff. protoconchs, which were not Therefore, cannot braidensis (Bellardi, 1873) 17a-d excluded that these fragments belong to organisms other than PI. 1, Figs 16a-d,

The hold for Thecosomata. same appears to true specimens

*v 1873 Balantium (Flabellulum) braidense Bell. — Bellardi, recorded from Bowden by Collins (1934, p. 208). 12. p. 32, pi. 3, fig.

— further reference is Additionalsynonymy For synonymy, Genus Styliola Gray 62, made to Janssen (1995, pp. 63).

Description — Only three specimens are available, one of Type species — Styliola subula (Quoy & Gaimard). shell is which preserves the protoconch. The triangular,

in apical angle initially c. 30°, but rapidly widening apertural Styliola subula (Quoy & Gaimard, 1827) distance from the from there direction at a short apex, on

* of almost 75°. Side lines are therefore 1827 & Gaimard, enclosing an angle Cleodorealene, Cleodorasubula, Quoy p. of the towards the 233, pi. 8D, figs 1-3. concave in the apical part shell, straight

and in section. The aperture distinctly squarish transverse the — a in apical Additionalsynonymy For further synonymy see Janssen shell has lengthwise curvature, especially part. lateral view it is visible that the carinae bend slightly in (1990, 1995, in press). In The Description — The reader is referred to the relevant literature opposite directions, giving the shell an elegant shape.

is with a distal spine, clearly on the Recent occurrences of this well-known species (for protoconch spherical, sharp

from the teleoconch and connected by a slight example, van der Spoel, 1967). Fossil material was described separated

and discussed in detail by Janssen (1990). constriction of the shell. side of the shell is much flatter than the Material — Sample 1-1/65 specimens, RGM 396 023; The ventral with flat central almost third sample 1-2/1 specimen, NMB H 17789; sample 1-6/5 speci- dorsal side, a rib, occupying one The of the shell width. The lateral fields are virtually flat. mens, RGM 396 024; sample 2-1/1 defective specimen,

RGM 396 025. dorsal side is convex with a swelling in the centre, on which

a is visible: two narrow riblets and — Gabb, (very vaguely) tripartition Discussion The synonymy ofStyliola sulcifera

a wider central one. Lateral fields are concave. A narrow 1873, described from the Pliocene of Santo Domingo, was

marginal zone is present. Under slanting light a transverse discussed by Janssen (1990, p. 34; in press, pi. 3, fig. 7). the lateral of of ornament be seen on fields, consisting Janssen (1995, p. 30) gave the stratigraphical range may

middle Miocene but in the and curved undulations. Styliola subula as to Recent, widely separated

Material— 1-1/3 specimens, RGM 396 029-031. meantimethe species has been found to occur already in late Sample

Discussion — The Bowden specimens differ from those Oligocene sediments of the North Sea Basin, and probably in several also of southwest France. from the early Pliocene, North Italian specimens

of the shell has been respects. The lengthwise curvature not

in the Italianmaterial. The central elevationof observed yet

the dorsal side of the Bowden is Subfamily Clioinae van der Spoel specimens slightly stronger,

Genus Clio Linne but its tripartition is only very vaguely indicated. The same

Bowden is true for the transverse ornament, which in the

shells is traceable under slanting light and only Type species — Clio pyramidata Linne. exclusively is wider on the dorsal side. Also, their apical angle slightly

and reaches almost instead of 70° in Italian C. braid- Clio pyramidata Linné, 1767f. lanceolata 75°, as

of the is such (Lesueur, 1813) ensis. Still, the general appearance specimens is in that a close relationship with C. braidensis obvious, and,

* do allow the intro- 1813 Hyale lanceolee (Hyalaea lanceolata), Lesueur, p. 284, my opinion, the observed differences not

limited material pi. 5, fig. 3A, B. duction of a new taxon, especially so since

is available.

Additional synonymy — For an extensive synonymy of When compared to the Recent C. cuspidata (Bosc) (see

Recent der difference in the dorsal predominantly occurrences, see van Spoel (1967, PI. 1, Fig. 18a-d), there is a major

for fossil Janssen in has central p. 68); occurrences, see (1995, press). side, which in C. cuspidata a sharp carina, ac-

— See der line each side. The Description van Spoel (1967, p. 68). companied by a weaker vertical on

Material— 1-1/1 RGM 396 026; sam- in the Recent Sample protoconch, transverse ornament is very well developed

UF in the Pliocene ple 1-4/1 defective specimen, 68907; sample 1-5/2 species, reaching the same strength as early

defective specimens RGM 396 027; sample 1-6/1 specimen, (Zanclean) C. braidensis from Italy. The protoconch of C.

RGM 396 028. cuspidata closely resembles that of the Bowden specimen,

Discussion — Clio bowdenensis (Collins, 1934) was syn- demonstrating a rather close relationship.

with the form Robba closer with onymised present by (1977, p. 600). At first view, one might suspect a relationship

Janssen (in press, pi. 3, fig. 15) illustrated the holotype. the Miocene C. carinata (Audenino), the dorsal side of which 102

C. well. in C. 1883 Diacria — 435. resembles cuspidata very However, carinata Vendryesiana, Guppy Fischer, p.

ventral side has 1894 — the a strongly different arrangement of Hyalaea (Diacria) vendryesiana, Guppy Cockerell,

118. ornament elements. Its protoconch also differs considerably p.

1903 Cavolinia Guppy — Dail, 1582. (Janssen, 1995, pi. 5, figs 7, 8; pi. 6, fig. 1). In this context it Vendryesiana p.

1928 Cavolina vendryesiana (Guppy) — Woodring, 115, seems plausible that the specimen referred to by Janssen (in p. pi. 1, figs 12,13. press, pi. 3, fig. 12), from the MaoFormation of the Domini-

— 1934 Cavolina vendryesiana (Guppy) Collins, p. 190,pi. can Republic, might also belong to this species rather than to 11, figs 16-18. C. cuspidata.

— 306. 1943 Cavolina vendreysiana [sic] (Guppy) Beets, p.

1962 Cavolina 1873 — vendryesiana Guppy, sp. Gilbert, p.

62. Subfamily CavoliniinaeFischer 1971 Cavolinia cf. — 221. vendryesiana(Guppy) Jung, p. Genus Cavolinia Abildgaard (emend. Philippi) 1975 Cavolinia — 228. vendryesiana Grecchi, p.

1982 Cavolinia cf. vendryesiana (in Jung, 1971) — Bernas-

— Cavolinia tridentata & 215. Type species (Niebuhr). coni Robba, p.

1982 Cavolinia vendryesiana (Guppy) — Bernasconi &

Robba, p. 218.

Cavolinia tridentata(Niebuhr, 1775)

PI. 2, Fig. 1

Comparative remarks — This species closely resembles the

* Recent 1775 Anomia tridentata, 124. C. inflexa and differs in minor Niebuhr, p. (Lesueur) points only. The dorsal side of the fossil species has an obvious reinforc-

rim Additional — ing the which is absent in C. synonymy For an extensive synonymy of this along apertural margin,

In addition, C. has some wide radial folds predominantly Recent species the reader is referred to van inflexa. inflexa on

the dorsal which absent indi- der Spoel (1967). side, are or only very vaguely

Description — The mainly fragmentary material available cated in C. vendryesiana.

Material — 1-1/5 defective 3 from the Bowden locality agrees in general with the Recent Sample ± specimens, frag-

material of this species. ments, RGM 3% 037-038; 11 apical spines, RGM 395 036;

Material—Sample 1-1/1 defective specimen, 4 fragments, sample 1-2/6 ± defective specimens, NMB H 17793; sample

RGM 396 032-033; sample 1-2/1 defective specimen, 3 1/4/2 specimens, UF 68905; sample 1-5/1 defective speci-

fragments, NMB H 17790-2; sample 1-4/1 fragment ?, UF men, RGM 3% 039; sample 1 -6/1 defective specimen, RGM

68903, 7 fragments, UF 68901; sample 1-6/1 defective 396 040.

specimen, RGM 396 035. Discussion — A Recent specimen of C. inflexa is illustrated

— here Remarks One fragment in the available material, a dorsal for comparison (PI. 2, Fig. 3). It has its lateral spines

also more and shell plate (sample 1-2, NMB H 17792), shows traces of an developed bent down,which is not necessarily

in the character. the oblique transverse ornament apical shell part (PI. 2, a specific Probably two species are closely

Fig. 1). This is reminiscent of the early Pliocene Cavolinia related, belong to the same lineage, and should be grouped in

the differenceswith other Cavolinia from the Mediterranean a separate taxon, as most grandis (Bellardi) (Janssen, 1995, p. considerable. 97, pi. 8, fig. 8, holotype). However, in that species the species are

is also This is also known from the Pliocene transverse ornament present on adapertural shell parts species (Piacen- and is of the Mediterranean From the distinctly more accentuated. Therefore, the present zian) area. Japanese Pliocene,

is introduced Cavolinia fragment not considered to be a last representative of C. Ujihara (1996, p. 785, fig. 7/18-30)

but is included which I consider ofC. grandis, in C. tridentata, which agrees with vendryesiana hyugaensis, a synonym

the age of the material. vendryesiana.

Fossil representatives of this species are known from the

Pliocene of the Caribbean and the Mediterranean, whereas in Cavolinia ventricosa (Guppy, 1882) the Recent fauna this species has a circumglobal distribution PI. 2, Figs 4-8 in the temperate to tropical zones.

*v 1882 ventricosa 176, 7, 15. Hyalea Guppy, p. pi. fig.

1903 Cavolinia ventricosa — 1582. Guppy Dall, p. Cavolinia vendryesiana (Guppy, 1873) ventricosa — .v 1928 Cavolina (Guppy) Woodring, p. 114, pi. PI. 2a-c 2, Fig. 1, figs 8, 9.

1934 Cavolina ventricosa — .v (Guppy) Collins, p. 184, pi. 7,

* 16-18. 1873 Hyalaea (Diacria) vendryesiana Guppy, p. 74, pi. 2, fig. figs

2. 1960 Covolinia [sic] ventricosa (Guppy) — Pchelitsev & Ko-

1874 — robkov, 710. Hyalaea (Diacria) Vendryesiana Guppy, pp. 405, p. 252, fig.

441, pi. 17, fig. 2. 1970 Cavolina (Cavolina) cf. C. ventricosa (Guppy) —

1882 — 175. Hyalea vendryesiana, Guppy Guppy, p. Woodring, p. 429, pi. 63, figs 11,12. 103

Collins 1982 Cavolinia ventricosa (Guppy) — Bernasconi & Robba, to belong to this species' were recorded by (1934, p.

218. from localities in Haiti. pp. 217, 185) two

Type material— According to Collins (1934, p. 185) the two

National 'are syntypes, housed in the United States Museum, Genus Diacria Gray

partly covered with glue and cleaning is impracticable', for

he did illustrate — which reason not them, nor designate a Type species Diacria trispinosa (de Blainville).

lectotype. I studied these specimens some years ago; with

some care both could be cleaned and they are illustrated Diacria digitata (Guppy, 1882) herein(PI. 2, Figs 4,7). The better preserved specimen of the PI. 2, Figs 9a-d, lOa-d, lla-d, 12, 13a-b, 15a-d two (PI. 2, Fig. 7) is here designated lectotype (USNM

115625). *v 16. 1882 Hyalea digitata Guppy, pp. 175,176, pi. Type locality — 'Jamaica'.

1903 — 1582. Cavolinia digitata Guppy Dall, p.

— the Description Cavolinia ventricosa closely resembles 1928 Cavolina digatata(Guppy) [sic] — Woodring, p. 114, late Miocene species C. gypsorum (Bellardi) (see Janssen, pi. 1, figs 10,11. 1995, 99, pi. 8, figs 9-12). As stated in that paper the dorsal p. — 115. 1928 Cavolina digitata Guppy (pars ) Woodring, p. shell of C. ventricosa is considerably more convex, as a part — Collins, 194, 11, .v 1934 Cavolina digitata (Guppy) p. pi. result of which the in a few long apertural lip (found speci- figs 1-9.

PI. instead of — & mens only: 2, Fig. 8c) projects horizontally, 1982 Diacria digitata (Guppy) Bernasconi Robba, pp.

in C. with in obliquely as gypsorum (compare Janssen, press, 218, 220.

pi. 4, fig. lb). The ornament of the dorsal side in C. ventri-

— of cosa could be described as consisting of five radial ribs, but Type material Guppy's original material, consisting

in be three and survives in the Smith- the two lateral ones are so weak that, fact, they cannot specimens two fragments,

calledribs. Thus, the ornament comprises three ribs, of which sonian Institution (USNM 115623). Two of these specimens

the better the middle one is usually remarkably narrower than the two are illustrated herein(PI. 2, Figs 9, 11); preserved

their the other others. Commonly, these lateral ribs are separated on one is here designated lectotype (PI. 2, Fig. 9), two

marginal sides by a rather deeply incised furrow (PI. 2, Fig. are paralectotypes.

7a, lectotype). The ventral side of C. ventricosa differs Description — A juvenile shell and the protoconch were

illustrated Collins 11, 7, strongly from C. gypsorum by the absence of the two oblique describedand by (1934, pi. figs 8);

for C. this shell is this studied for the 2, Fig. furrows, so typical gypsorum. Also, part specimen was present paper (PI.

short the end of which relatively higher. Fine and regular transverse growth lines are 13a-b). It has a relatively apical spine,

is inflated an bulb with a rounded visible in a number of specimens, but in the bulk of the to elongate tip, resembling

of the D. The material they are hardly or not preserved. that Recent quadridentata (de Blainville).

and ventral sides are still flat, a The apex of C. ventricosa is slightly curved in dorsal dorsal indicating pre- dorsal side. direction, but the protoconch itself is absent in all available metamorphosis stage; three ribs are visible on the is specimens. The apical angle is c. 135°. The apertural margin strongly

Material — Sample 1-1/1 specimen, 2 fragments, RGM 396 curved and lateral spines project sidewards, just above mid-

041; sample 1-2/1 specimen, NMB H 17794; sample 1-4/1 height of the shell. with specimen, separated from UF 68904, 1 fragment, UF 68906; The fully-grown shell is bilaterally symmetrical VA times sample 1-5/11 ± defective specimens, RGM 396 042; sample distinctly separated dorsal and ventral sides, c.

1-6/9 ± defective specimens, 1 fragment, RGM 396 043-044; higher than wide. Both sides are convex, but the ventral side

?2 than the with distinct mid- sample 2-1/1 specimen, specimens (internal and/or exter- more so dorsal, a swelling at and the nal moulds), RGM 396 045-047. height. The juvenile shell is shed during ontogenesis the sides Discussion — In spite of the rather close resemblance be- opening is closed with a septum. From the septum

the end the lateral slits tween the taxon and C. these species are curve gently to almost horizontal. At present gypsorum, the base of the not considered to be closely related. It is assumed that the between dorsal and ventral shell parts meet

furrows the ventral side of C. and there distinct is less occurrence of oblique on shell, a spine present, commonly

taxonomic value. The in apical direction, but commonly not gypsorum must be given fairly high bending slightly

In the these lie at one fifth only other Cavolinia in which such furrows are found is C. sharply pointed. lectotype spines other the described from the late Miocene of the Do- ofthe shell height, as in most specimens. However, aff. gypsorum, less commonly lie higher, to two fifths of the minican Republic (Janssen, in press, pi. 4, fig. 4), a form spines up

in which the are and apparently also present in Pliocene deposits of Japan. Similar height, case spines sharper project

distorted sidewards furrows have been observed in a single, slightly more strongly (PI. 2, Fig. 15a).

in The inflated ventral shell part is smooth, but for some specimen of C. mexicana (Collins, 1934) (Janssen, press,

an vaguely indicated growth lines that are slightly bent in apical pi. 4, fig. 8), but here these furrows may originate from their middle. The ventral margin is early shell injury repair. direction in apertural

'shells that straight (ventral view) and distinctly thickened. In apertural Apart from the occurrence in Bowden appear 104

this view lip is strongly curved. cene. The last-mentioned author considered his material to

The dorsal side of the shell is but far be considerably flatter, higher probably represent a new subspecies, as as can

the than ventral side and reaching beyond the ventral aper- seen from the illustrations all specimens referred to by the

tural rim. Where the ventral margin touches the dorsal one, Japanese authors resemble closely the Mediterraneanform,

the latter shows constriction. a distinct Threerelatively strong especially by the relatively high position ofthe lateral spines.

radial ribs are present on the middle of the dorsal side,

furrows width. separated by of the same Together, these ribs

less than third of the shell Diacria trispinosa (de Blainville, 1821) occupy one width, leaving wide PI. 2, 14 lateral fields on each side. In some specimens these lateral Fig.

fields are somewhat produced and resemble wide ribs (PI. 2,

* The of this shell is also 1821 Hyaloea trispinosa Lesueur — de Blainville, 82. Fig. 10a). margin part thickened, p. *

1873 Diacria bisulcata Gabb n.s. — Gabb, 200. especially so above the point in touch with the ventral shell p.

part. In apertural view this rim is less strongly bent than the

Additional — Lists of further (mainly ventral one. synonymy synonyms

fossil be foundin Janssen in Material— Sample 1-1/14more or less defective specimens, occurrences) may (1995, press).

— See van der (1967, 84, c. 50 fragments, RGM 396 048-049; sample 1-2/17 speci- Description Spoel p. figs 76-78).

3 internal Material — Typical form: sample 1-1/21 fragments, RGM mens, moulds, 32 fragments, NMB H 17795-7; 396 052; sample 1-2/1 specimen, 9 fragments, NBM H sample 1-3/1 specimen, NMB H 17798; sample 1-4/26 17799-800; 1-4/1 UF 68908; 1-5/1 specimens, 12 fragments, UF 68902, UF 68904, UF 68909; sample fragment, sample specimen, 2 fragments, RGM 396 053; sample 1-6/1 defec- sample 1-5/46 specimens, RGM 396 050; sample 1-6/32 ± tive 2 RGM 396 defective specimens, RGM 396 051. specimen, fragments, 054-055; sample 2-1/1 RGM 396 056. Diacria — specimen (external mould), trispi- Discussion An interesting specimen ofD. digitata was

nosa f. bisulcata: sample 1-2/7 specimens, NMB H 17801-3. illustratedby Collins (1934, pi. 11, fig. 9). It originates from

Discussion — Several specimens amongst the available the 'Miocene' of Port-au-Prince (Haiti), which must have a

less the Bowden Beds material show the typical features of f. bisulcata Gabb. For more or comparable age to (Janssen,

in The is rather a discussion on the status of that taxon see Janssen (in press). press). specimen poorly preserved in a slab The Jamaica shells with of sediment also containing Hyalocylis haitensis (Collins), specimens likewise are small a

height of some 4 mm. Even underslanting light they do not which I consider synonymous with H. striata (Rang). It has

show a subdivisionof the wide central rib. A specimen in the the lateral spines well developed, relatively highly situated NMB collection, showing the typical features quite well, is and pointing downwards. Furthermore, this specimen, al- illustratedherein (PI. 2, Fig. 14). though having passed shell metamorphosis, apparently had its

shell apical part preserved. A new illustration of this speci-

men is here presented (PI. 2, Fig. 12).

Typical Diacria digitata is exclusively known from the CONCLUSIONS

Caribbean (Jamaica and Haiti), but related forms were The molluscan fauna ofthe Bowden shell bed recorded from the Mediterranean and from Japan. Grecchi holoplanktonic

found to three and 14 (1982, p. 723, pi. 54, figs 1-5) introduced Diacria digitata was comprise heteropod pteropod

italicafrom the Plioceneof northern Italy. This taxon differs species:

from typical D. digitata by the relatively high position of the

lateral spines and the more accentuated constriction of the

dorsal side above the juncture with the ventralside. Also, the Heteropoda Atlanta diamesa 1928 lateralfields next to the set ofthree central radial ribs in this Woodring,

keraudrenii form more commonly are swollen and could be interpreted Oxygyrus (Lesueur, 1817) Protatlanta lissa as wide additional radial ribs. The general outline of this (Woodring, 1928)

form therefore resembles that of Diacria trispinosa. A single

Thecosomata specimen from Bowden resembles this form (PI. 2, Fig. 15a-

but it lacks the constriction the dorsal Limacina bulimoides d), on apertural margin (d'Orbigny, 1836) Limacina and rather seems to be an abnormal specimen of D. digitata. inflata (d'Orbigny, 1836)

still Limacina 1 Grecchi, considering the Bowden Beds as Miocene, sp.

his Limacina ? 2 interpreted taxon to be a 'sottospecie allocrona e allopa- sp.

trica.' However, the age of the Bowden Beds being Pliocene Bowdenatheca jamaicensis Collins, 1934

would make Creseis acicula? D. digitata italica a real geographic subspecies, (Rang, 1828)

so far restricted to the Mediterranean. Styliola subula (Quoy & Gaimard, 1827)

Shibata Clio Linne, 1767 f. lanceolata From Japan, (1980, pp. 61, 62, pi. 3, fig. 18), pyramidata (Lesueur,

Shibata & Ishigaki (1981, p. 57, figs 5, 6) and Ujihara (1996, 1813) p. 781, fig. 6/1-6) recorded Diacria digitata from the Plio- Clio aff. braidensis (Bellardi, 1873) 105

Cavolinia tridentata(Niebuhr, 1775) Museum of Natural History, Smithsonian Institution, De-

Profes- Cavolinia vendryesiana (Guppy, 1873) partment of Paleobiology, Washington D.C., U.S.A.); of and Cavoliniaventricosa (Guppy, 1882) sor Stephen K. Donovan (Department Geography of the West Jamaica); Diacria digitata (Guppy, 1882) Geology, University Indies, Kingston, Ronald Janssen Frankfurt Diacria trispinosa (de Blainville, 1821) Dr (Senckenberg Museum, am Main, Germany); Dr Peter Jung (Naturhistorisches Museum, Diacria trispinosa (de Blainville, 1821) f. bisul- Basel, Dr Jan P. of cata Gabb, 1873 Switzerland); Krijnen (Department

Mineral Resources, Geological Survey of New South Wales,

Orange, NSW, Australia); and Roger W. Ported (Florida Naturally, taxa identified in open nomenclature yield no Gainesville, of the of the Clio aff. braidensis Museum of Natural History, U.S.A.). indication age assemblage. Bundesanstalt, Vienna, form of the Mag. Irene Zorn (Geologische is here assumed to be a younger evolutionary Austria), Dr Peter Jung and Dr D.T.J. Littlewood (The early Pliocene (Zanclean) C. braidensis. Amongst the re- Natural History Museum, London, U.K.) critically read the maining species two (Atlanta diamesa and Protatlanta lissa) of which John whereas several others manuscript, the English was improved by are exclusively known from Bowden, W.M. Jagt (Venlo, The Netherlands). are long ranging:

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PLATE 1

otherwise. All specimens figured in Plates 1 and 2 are from the Pliocene Bowden shell bed, southeast Jamaica, unless stated

Fig. 1 Atlanta diamesa Woodring, 1928 (holotype), USNM 369335, apical, frontal and umbilical views, x 12.5.

Fig. 2. Atlanta ? diamesa Woodring, 1928, sample 1-1, RGM 396 006; apical view and transverse section, x 25.

Fig. 3. Oxygyrus keraudrenii(Lesueur, 1817), sample 1-1, RGM 396 005; apical and right lateral view, x 25.

Fig. 4 Protatlanta lissa (Woodring, 1928) (holotype), USNM 369336; apical and frontal views, x 12.5.

umbilical Fig. 5. Protatlanta lissa (Woodring, 1928), sample 1-6; RGM 396 007; apical, frontal and view, x 6.

Figs 6-8. Limacina bulimoides (d’Orbigny, 1836), sample 1-1, RGM 396 009-011; frontal views, x 25.

Figs 9, 10. Limacina bulimoides (d’Orbigny, 1836), Recent, Manihiki Plateau Expedition, sample U 336b, co-ordinates 11° 33.3’S 165° 26.7’

W, water depth 981-1297 m, RGM 396 012-013; frontal views, x 25.

11. Limacina RGM Fig. sp. 1, sample 1-1, 396 017; frontal view, x 25.

Fig. 12. Limacina inflata (d’Orbigny, 1836), sample 1-1; RGM 396 015; apical view and oblique frontal view to show rostrum; x 25.

13. Limacina ? RGM 396 view and left lateral 25. Fig. sp. 2, sample 1-1; 018; apical view, x

Figs 14, 15.Bowdenathecajamaicensis Collins, 1934,sample 1-1; RGM 396 020-021; 14- dorsal view, right lateral view and transverse sections

ataperture and apical part, and ventral view, respectively; 15 - ventral view, right lateral view and transverse section at aperture and

dorsal view, x 6.

Figs 16, 17. Clio aff. braidensis (Bellardi, 1873), sample 1-1, RGM 396 029-030, 16: x 6; 17: x 12.

Fig. 18. Clio cuspidata (Bosc), Atlantic Ocean, E of mid-Atlantic ridge, 45° 21.3’N 27° 9.1’W, boxcore T90-10B, JGOFS Leg-IV expedi-

16th from 43 bottom tion, June 1990, sea depth2,162 m; Holocene, upper cm of sediment, with pteropod-rich layers throughout;

RGM 396 034, dorsal, right lateral, ventral views and transverse section at aperture, x 6. 109\1

PLATE 1 110

PLATE 2

Fig. 1. Cavolinia tridentata(Niebuhr, 1775), sample 1-2; NMB H 17792; fragment showing dorsal shell part preserving weak transverse

ornament in apical part, x 3.

Fig. 2. Cavolinia vendryesiana (Guppy, 1873), sample 1-1, RGM 396 038; ventral, left lateral and dorsal view, x 6.

Fig. 3. Cavolinia inflexa (Lesueur, 1813), Atlantic Ocean, off Morocco coast, 34° 54.5’N 7° 34.6’W, Meteor M53, sample 170 (box core),

sea depth 1,478m, 22nd March 1988; Holocene; RGM 396 430; dorsal view, x 6.

Cavolinia view, Fig. 4. ventricosa (Guppy, 1882), paralectotype; USNM 115625;dorsal, left lateral and ventral x 6.

Fig. 5. Cavolinia ventricosa (Guppy, 1882), specimen illustrated in Woodring (1928), USNM 369313; dorsal, left lateral, ventral and

adapical views, x 6.

Fig. 6. Cavolinia ventricosa (Guppy, 1882), specimen exAldrich/Johns Hopkins University Collection, illustrated in Collins (1934), USNM

left 645204; dorsal, lateral, ventral and adapical views, x 6.

Fig. 7. Cavolinia ventricosa (Guppy, 1882) (lectotype), USNM 115625; dorsal, left lateral and adapical views, x 6.

Cavolinia Fig. 8. ventricosa (Guppy, 1882), sample 1-6, RGM 394 044; dorsal, left lateral and ventral views, x 6.

Figs 9, 11. Diacria digitata (Guppy, 1882), USNM 115623; 9 - lectotype, 11 - paralectotype; adapical, dorsal, left lateral and ventral views,

x 6.

Fig. 10. Diacria digitata (Guppy, 1882), specimen illustrated in Collins (1934, pl. 11, figs 4-6), USNM 645203; adapical, dorsal, ventral

and left lateral views, x 6.

Fig. 12. Diacria digitata (Guppy, 1882), Port-au-Prince, near Pétionville, Haiti; illustrated in Collins (1934, pl. 11, fig. 9), USNM 371903,

ex Woodring & Brown Collection;specimen in sediment slab, showing dorsal side, x 6.

Fig. 13. Diacria digitata (Guppy, 1882), juvenile, illustrated in Collins (1934, pl. 11, figs 7, 8), USNM 645207, ex Andrews & Lynn

Collection; dorsal view (x 6) and protoconch (x 25).

Fig. 14. Diacria trispinosa (de Blainville, 1821) f. bisulcata Gabb, 1873, sample 1-2; NMB H 17802; dorsal view, x 6.

Fig. 15. Diacria digitata (Guppy, 1882), aberrant specimen, illustrated in Collins (1934, pl. 11, figs 1-3), USNM 645202, exAldrich/Johns

Hopkins University Collection; adapical, dorsal, left lateral and ventral views, x 6. 111\2

PLATE 2