Cainozoic Research, 5(1-2), 13-49, November 2006 pp.

The gastropod fauna of the Luchtbal Sand Member (Lillo

Formation, Zanclean, Early ) of the Antwerp region

(Belgium)

² Robert Marquet¹ & Bernard Landau

'institut royal des Sciences naturelles, Sciences de la Terre, Vautierstraat 31, 1000 Brussels, Belgium.

2 InternationalHealth Centres, Av. Infante D. Henrique, Areias Sdo Joao, 8200Albufeira, Portugal; e-mail; [email protected]

Received 23 September2004;revised version accepted 30 June 2005

The gastropod fauna ofthe Luchtbal Sand Member (Antwerp region, Belgium) is listed based on the collections ofthe IRSNB, and new

material collectedin Doel (Oost-Vlaanderen, Belgium) and Antwerp. Fourteen are new for the Pliocene of Belgium; onespecies

is described is redescribed and is as new: deurganckensis nov. sp. Xenophora scaldensisGlibert, 1958 a lectotype designated.

The total faunaamounts to 157 gastropod species, which make the Luchtbal Member the most species-rich Pliocene unit ofBelgium.

Only 29% ofthese species are still living today, which is insufficient for actualistic palaeoecologicalreconstructions. 46% ofthe species

are endemic to the Pliocene or to the Neogeneof the Basin.

KEY WORDS:: Pliocene, Luchtbal Sand Member, Belgium, , new taxon.

Introduction thick layer (10 to 20 m) of sediment dredged from the

Scheldt River, and subsequently built over. The last time

The Luchtbal Sand Member was described as the lower- the Luchtbal Sand Member was exposed on the right bank

most part ofthe PlioceneLillo Formationby de Heinzelin of the Scheldt River was during the 1970’s, when a junc-

(1955c) following Leriche (1912). A type section and lo- tion between the Amerikadok and Vijfde Havendok was

cality (Tweede en Derde Havendok in Antwerp, 10 to 8 m created in the Noordkasteel area of Antwerp. The material

below the naturalland surface) were proposed by De Meu- collected was never published. Further material, dredged

& who also of the de- from the Scheldt River and various localities ter Laga (1976), gave a diagnosis deposited at

posit. Vandenberghe etal, (1998) estimatedthe age ofthe in the Antwerp harbour area between 1960 and 1995 is

Luchtbal Sand Memberas late Zanclean (± 3.5 Ma), based available. These sediments are often composed of ex-situ

on benthic foram association B11, benthic mollusc associa- Luchtbal and Oorderen Sand Member sediments and fau-

tionBM22Aand otholith 18 (Vincken, 1988). The Luchtbal sands characterised zone During nas. are by the presence ,h th the 19 and the beginning ofthe 20 century, the Luchtbal of large numbers of Ditrupa sp. and of characteristic bi-

Sand Member was often exposed during extensions of the valve species, listed by Marquet (2005, p. 119).

Antwerp harbouronthe right bank ofthe Scheldt River. De During the works on the left Scheldt bank harbourat Kallo

Heinzelin (1950a, 1950b, 1955a, 1955b) described the (Beveren), no Luchtbal deposits were found untilrecently

stratigraphy and the rich molluscan association at the (Marquet, 1995). However, derived yellow sandstones con-

Boudewijnsluis section in the 1940’s. His findings were taining large numbers of the bivalve Palliolum gerardi

partly published by Gilbert (1957a-c; 1958a-c; 1959a, b; (Nyst, 1835) were oftenencounteredin the basal shell bed

1960). Later taxonomic studies on the Boudewijnsluis ma- of the Oorderen Sand Member, proving that the Luchtbal

terial were performed by Marquet (1996: Triphoridae; Sand Member had been present but had been destroyed by

1997, 1998a: turrid gastropods and 2001: Seilinae). Addi- erosion. Also massive numbers of Ditrupa sp. were ob-

tional Luchtbal material was collected by van der Mark served in these sandstones.

around 1964 at the B1-B2 Havendok section; his material In March 2000, a wedge of Luchtbal Sand Member was

is deposited in the NNM Naturalis, Leiden, The Nether- discovered in-situ in the Deurganckdok exposures at Doel

lands. (municipality Beveren), extending for about 1 km length

Today, only a small part of the type area of the Luchtbal (see Marquet, 2002).

Sand Member remains northeast of Antwerp, covered by a - 14-

Figure 1. - 15 -

Figure 1.

1. adriatica Costa, 1829. Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6949.

2. Emarginulaplioaspersa Sacco, 1897. Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6950.

3. Doel. Luchtbal Sand Coll. BLP. Emarginula sp. Deurganckdok, Member.

The level was about 1 m thick at the northern quay of the Emarginula adriatica Costa, 1829

dock and consisted there of yellow sand at the base, often Fig. 1/la-c

eroding deeply into the Kattendijk Formation, with burrows

filled with Luchtbal Sand extending up to 1 m into this

Formation. This lower mollus- *1829 adriatica 12. part yielded a species-rich Emarginula Costa, p, 122, fig.

1916 Emarginula adriatica Costa - Cerulli-Irelli, 217, can fauna, accompanied by many Ditrupa tubes. It was p. 24, 35-36. topped by slightly consolidated sandstones with abundant pi. figs

1923 Emarginulaelongata Costa - Banner, 778, pi. 62, stacked valves ofPalliolum gerardi (Nyst, 1835),together p. fig. 9. with a large number ofbryozoa, few sea urchins and scarce

1949 Emarginulaelongata Da Costa - Glibert,pi. 1, fig, 9. This found the southern gastropods. upper part was only at

1964 Emarginulaelongata Costa - Brebion,pi. 1, figs 7-8. quay ofthe dock. Bivalvia, Scaphopoda and Amphineura 1984 adriatica Costa - Emarginula Piani, p. 200, figs 13- from this were included in the works ofMarquet exposure 20. (2002, 2004). The Deurganckdok of Luchtbal exposure 1990 adriatica Costa - & Emarginula Poppe Goto, p. 67, Sand Member contained several gastropod species new to pi. 2, fig. 17. fauna. the The aim ofthe is - Belgian present paper to assess 1994 Emarginula adriatica Costa Giannuzzi-Savelli et

the Luchtbal Sand Member found 73-74. gastropods in the expo- al., p. 46, figs.

2003 Costa - Landau sures at Doel (Deurganckdok), Antwerp (Noordkasteel), Emarginulaadriatica etal., p. 19, pi. and the species in dredged material probably originating 3,figs 1-3.

from this Member. For the location ofthe citedexposures

see Marquet (1997a, fig. 1), for the extension of the Locus typicus — Adriatic Sea.

Luchtbal Sand in the Deurganckdok, see fig. 11.

Stratum typicum — Recent.

Material and methods Material — IRSNB 1ST 6949 (leg. RM): 1 specimen,

Doel, Deurganckdok; Luchtbal Sand Member.

This study is based principally on material collected by

Robert furtherindicated to be — Marquet (Antwerp), as RM, Dimensions Height 2.4 mm, aperture 12.3 x 7.4 mm

transferred to the Institut royal des Sciences naturelles, (incomplete).

Brussels (IRSNB) in the near future, from Doel, Noord- kasteel and dredged from the Scheldt River. Further mate- Description — Medium-sizedshell, oval inoutline (width rial from Doel and material is in the collections of of dredged aperture more than halfoflength), posterior partbroader L. Ceulemans coll. from (Brussels, Belgium, LC), Doel in than anterior. Apex strongly pointed, close to posterior

that ofB. Landau(Albufeira, Portugal, coll. BLP), A. Rat- border. Posterior side nearly straight, anterior clearly con-

inckx coll. AR,), B. Roest Neth- (Antwerp, Belgium, (The vex. Anterior side with slit, running nearly to the apex, erlands, BR). Furthermore, material from the B1-B2 Ha- open for about a quarter of its length, remaining part with collected vendok, by van der Mark, housed in the NNM distinct scales. Ornament consisting ofabout 40 sharp pri-

Naturalis The coll. indicated axial with weaker (Leiden, Netherlands, NNM, mary ribs, one secondary rib in between.

with RGM numbers) is included. Also the older collections Concentric ornament weaker, forming weak scales be-

in the IRSNB examined. were tween, and fainttubercles on the axials.

Discussion — (1923) Harmer mentionedthe presence of Systematic palaeontology Emarginula elongata in the Pliocene Coralline Crag For-

mationof the . This name however, according

to Sabellietal., 1990,is a synonym ofEmarginula octavi-

Class Cuvier, 1797 1839 and Gastropoda ana Coen, was preoccupied by Emarginula elon-

Subclass Orthogastropoda Ponder & Lindberg, 1995 gata Defrance, 1819 (= Subemarginula elongata, Eocene

Order Salvini-Plawen& ofFrance). Emarginula octaviana clearly differs from our

Haszprunar, 1987 material by the smaller number of axial ribs (about half),

Fissurelloidea 1822 Superfamily Fleming, while its apex points less forward; it is clearly figured by 1822 Family Fleming, Giannuzzi-Savelli et al. (1994, p. 46, fig. 77). Most ofthe

1834 earlier Subfamily Gray, records from the Mediterraneanof E. elongata are

Genus Emarginula Lamarck, 1801 more similar to E. adriatica than E. octaviana. - 16-

Figure 2. - 17 -

Figure 2.

BLP. 1. Emarginula sp. Deurganckdok, Doel. Luchtbal Sand Member. Coll.

2. italica (Defiance, 1820). Antwerp, junction Vijfde Havendok-Amerikadok. Luchtbal Sand Member, IRSNB 1ST 6951.

Doel. Member. 3. (Linnaeus, 1758). Deurganckdok, Luchtbal Sand IRSNB 1ST 6952.

4. Gibbula (Steromphala) cineroides (Wood, 1842). Antwerp, Vijfde Havendok. Luchtbal Sand Member. IRSNB 1ST 6953.

5. Gibbula (Colliculus) crassistriata (Bell in Wood, 1882). Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6954.

This is also the case for most ofthe Pliocene varietiesof E. Figs. 1/3 a-c, 2/1 elongata ‘Costa’, figured by Sacco (1897, pi. 2) and some of his Emarginula cancellata varieties on the same plate. Material — Coll. BLP: I specimen, Deurganckdok, Doel;

Luchtbal Sand Member.

Occurrence — This is the first record of Emarginula adri-

for the of it is in the Luchtbal — 1.95 2.79 2.20 atica Neogene Belgium; rare Dimensions Height mm, aperture x mm.

Sand Member at Doel, where one specimen was found. It is also known from the British CorallineCrag Formation. For Description — Small shell, with regular oval outline; LAV

its general distribution, see Landau et al. (2003, p. 20). ratio of aperture 1.2. Apex pointed, nearly reaching the

anterior is slit concave posterior margin; margin convex. A

is to the that is for present, reaching nearly aperture open a

of its total Emarginula plioaspera Sacco, 1897 quarter length and smooth. The ornament con-

Fig. 1/2 a-d sists of20 strong axial ribs, with a very weak secondary rib

in between. The concentric ornament is rather thick, espe-

*1897 E. elongata var plioaspera Sacco, pi. 2, figs p. 15, cially distinctbetween the axials and forming square tuber- 22,23. cles on them.

Locus — Zinola, typicus Liguria, Italy. Discussion — This species shows a superficial resem-

blance to Emarginula punctura Wood, 1814, described

Stratum — typicum “Astigiano”, probably Zanclean, Early from the Coralline Crag Formation of East Anglia, but it Pliocene. clearly differs by the absence of pits in the surface. Fur-

rib is between each thermore, only one secondary present

Material — IRSNB 1ST 6950 RM): 1 (leg. specimen, pair of primary ribs, instead of three to four. Emarginula Doel; Luchtbal Sand Member. Coll. RM: 1 Deurganckdok, rosea Bell, 1824, also has a single secondary rib, but the Petalocon- specimen, Vrasenedok, Kallo; Kattendijk Fm., primary ribs lie closer together and the umbo reaches be- chus level. yond the posterior shell margin. In E. rosea the slit shows

clear squamae, instead of being smooth as in E. sp. Emar-

Dimensions — 2.91 5.2 3.63 Height mm, aperture x mm. ginula fissura reticulata Sowerby, 1813 has thirty primary

ribs, its apex lies muchcloser to the centre ofthe shell, the

Description — Small, elongate oval shell. Umbo pointed, slit shows squamae and the concentric ornament is mark- before near border. width Slit pre- The unidentified posterior Largest apex. edly stronger. Emarginula species re- from umbo; sent, open to one quarter remaining part ported here is possibly an undescribed species, however

closed, with about 15 clearly distinct scales. Ornament con- our single specimen from Doel is very small and probably

of about 20 axial ribs, with a sisting single nearly equally not adult, and insufficient for describing a new taxon.

strong secondary rib in between. Between the primary ribs,

concentricribs these form dis- numerous are — present; very Occurrence One specimen was found in the Luchtbal

tinct tubercles when crossing the primary ribs. Sand Member of Doel.

Discussion - Although originally described as a variety of

adriatica consider E. Emarginula Costa, 1829, we plioas- Subfamily Diodorinae Gray, 1821 distinct The latter has fewer ribs pera a species. primary Diodora Gray, 1821 (about half), bearing distinct tubercles, and the umbo lies

closer to the posterior margin. We consider these differ-

ences significant enough forseparation at the species level. Diodora graeca (Linnaeus, 1758) Fig. 2/3a-b

Occurrence — The species is new for the North Sea Basin,

been found in the area. Its dis- *1758 784. having previously only type graeca Linnaeus, p.

tribution is limited to the Zanclean Formation 1803 Patella 10. Kattendijk apertura Montagu, p. 491, pi. 13, fig.

1853 nodosa 16. and Luchtbal Sand Member. Eichwald, p. 138, pi. 6, fig.

1853 Fissurella Graeca Linnaeus - Homes, p, 642, pi. 50,

fig. 27. Emarginula sp. - 18-

1897 Fissurella - 8, tioned the of (Glyphis) graeca (Linnaeus) Sacco, p. presence D. apertura (= D. graeca), 3ut ac-

pi. 1, figs 16, 17. Janssen this is Diodora sub- cording to (1984, p. 120)

1916 Fissurella Linnaeus - graeca Cerulli-Irelli, p. 210, pi. costaria (d’Orbigny, 1820), characterised by the different 24, figs. 8-17. internal shape of the apical aperture. Diodora graeca oc-

1923 Fissurella Linnaeus - graeca Harmer, p. 770, pi. 61, curs in Badenian (Miocene, Paratethys) deposits ofPoland fig. 18. (Baluk, 1975), but thispopulation was already extinct long

1960 Diodora Linnaeus - (Diodora) graeca Kojumdgieva before the arrival ofD. graeca in the North Sea Basin, con- & 10. Strachimirov. p. 84. pi. 28. fig, sequently it could not have migrated from there.It is more

1960 Diodora - (Diodora) graeca (Linne) Malatesta, p. 49, that North Sea derive from the Atlantic pi. L fig. 2. likely immigrants

1964 - somewhere the Pliocene. Diodora apertura Montagu Brebion, pi. 1, fig- 12. during

1966 Fissurella Linne - (Diodora) graeca Strausz, p. 29,

pi. 76, figs 7-14.

1974 Diodora - apertura Montagu Malatesta, p. 161, pi. (Defrance, 1820)

28, 10. fig. Fig. 2/2a-b

1975 Diodora - (Diodora) graeca (Linnaeus) Baluk, p, 26,

pi. 1, figs 12-14. *lB2O Fissurella italica Defrance, 79. p.

1988 Diodora Linne - 17. graeca Graham, p. 70, fig. Linne - 71848 Fissurella graeca Wood, p. 168, pi. 18, fig.

1990 Diodora graeca - & Goto, p. 66, 2, (Linne) Poppe pi, 4, fig. 14. 1872 Fissurella costaria - 29. Wood, p. 90, pi. 7, fig.

1992 Diodora - Cavallo & graeca (Linne) Repetto, p. 34,

- 2. 1878 Fissurella graeca Linne Nyst, pi. 2, fig, fig. 12. Linne - 111. 1881 Fissurella graeca Nyst, p.

- Giannuzzi-Savelli 1994 Diodora graeca (Linnaeus) et

1897 Fissurella - (Glyphis ?) italica Defrance Sacco, p. 8, 56-58. al., p. 42, figs pi. 1, figs 18-23.

2003 Diodora - Landau graeca (Linnaeus) etal., p. 26, pi. 1916 Fissurella italica - Defrance Cerulli-Irelli, p. 209, pi. 4, fig. 4. 23, figs. 19-35, pi. 24, figs 1-7.

1960 Fissurella (Lucapina) italica Defrance - Cossmann &

48, 57-59. Locus typicus — Greece. Peyrot, p. pi. 2, figs

1923 Fissurella italica, Defiance - Harmer, p. 771, pi. 61,

fig. 17. Stratum typicum — Recent.

1949 Diodora italica Defrance - Glibert. p. 22, pi. 1, fig.

15, pi. 2, fig. I. Material— IRSNB 1ST 6952(leg. RM): 1 specimen, Doel

1957 Diodora apertura dorsatabMonterosato [sic] - Glib- Deurganckdok; Luchtbal Sand Member. Coll RM: 4 7. ert, p. specimens, same locality and level.

1960 Diodora italica - (Diodora) (Defrance) Malatesta,p.

50, pi. l.fig. 3.

—

Dimensions 5.7 14.4 8.6 1962 Diodora italica Defrance - Height mm, aperture x mm. Glibert, p. 14,

1965 Diodora - Altena apertura (Montagu) van Regteren

- 7. Description Medium-sized patelliform shell, aperture et al., p. 58, pi. 1, fig.

italica Linne - elongate oval; shell moderately elevated, with straight ante- 1966 Fissurella (Diodora) Strausz, p. 29, pi. 76, figs 15-16. rior slope and convex to slightly concave posterior. Apical

1974 Diodora italica Defrance - third behind the anterior Interior of Malatesta,p. 162, pi. 13, aperture at one margin. fig. 2. oval thickened. Ornament of about 30 aperture consisting

1975 Fissurella italica - (Diodora) Defrance Fekih, p. 87, primary axial ribs, with a single secondary and a single or pi. 28, fig. 15. more tertiary ribs in between. Concentric lamellae are dis- 1985 Diodora (Diodora) italica Defrance - Gonzalez Del- crenate, well visible at tinctly developed. Apertural margin 4-6. gado, p. 52, pi. 1, figs the outside. 1990 Diodora italica - Da (Defrance) Silva, p. 6, pi. I, figs

1-3.

— This documented from the Kat- Discussion 1990 Diodora italica - & species was (Defrance) Poppe Goto, p. 66, pi. tendijk Formation and the Oorderen Sand Member by 2, fig. 14.

1992 Diodora italica - Cavallo & Marquet (1995, 1998b). This was an error; the beforemen- (Defrance) Repetto, p. 34, fig. 13. tioned specimens belong to Diodora italica (Defrance,

1994 Diodoraitalica (Defrance) - Giannuzzi-Savelli et al. 1820) treated below. Both species occur in the Luchtbal , 59-60. p. 42, figs Sand Member, D. graeca at Doel and D. italica in the lo-

1995 Diodora - graeca (Linne) Marquet, p. 60, pi. 2, fig. calities on the right bank ofthe Scheldt River. I.

1998 Diodora b- graeca (Linnaeus) Marquet, p. 24, text-fig.

Occurrence — The species is rare in the Luchtbal Sand

2001 Diodora italica - (Diodora) (Defrance) Da Silva, p. Member of Doel and in the British Red Formation; Crag 74, pi. 2, figs 3-4. al. for its further distribution, see Landau et (2003, 26). p. 2003 Diodora italica - Landau (Defrance) et al, p. 26, pi. Its sudden and extinction in the Pliocene ofthe appearance 4, fig. 5. North Sea Basin is puzzling. From the Miocene of the

North Sea Basin, Glibert (1952, p. 6, pi. 1, fig. 2) men- Locus typicus — Italy. - 19-

Figure 3. -20-

Figure 4. -21 -

Figure 3.

1. Gibbula (Colliculus) crassistriata (Bell in Wood, 1882). Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6955.

2. Gibbula (Colliculus) woodi (Harmer, 1923).Antwerp, Noorderlaan, dredged material, Luchtbal Sand Member? IRSNB 1ST 6956

(3/2 a, b), 6979 (3/2 c, d).

3. Margarites belli (Harmer, 1923). Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6957.

Figure 4.

1. Margarites belli (Harmer, 1923). Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6957,

2. Dikoleps cutleriana (Clark, 1848). Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6958 (4/2 a-d. f), 6959 (4/2 e-g).

3. Cirsonella romettensis (Granata. 1877). a, e, f: Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6960. b, c, d: Deur-

ganckdok, Doel. Oorderen Sand Member, Cultellus level. Coll. BR.

4. indet. Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6961.

Stratum typicum — Recent. France from the Early Miocene (Aquitainian, Aquitaine

region) through the Middle Miocene (“Pontilevian”,

Material— IRSNB 1ST 6951 (leg. RM); 1 specimen, Ant- Langhian, Touraine region) until the Late Miocene (“Re- werp Noordkasteel, Luchtbal Sand Member. donian”,Tortonian, Anjou region). Its isolated Paratethyan

occurrence in the Early Miocene of Hungary (Strausz,

— Dimensions Height 15.0 mm, aperture 43.3 x 30.0 mm. 1966) is somewhat enigmatic and this material should be

reconsidered. Diodora subcostaria (d’Orbigny, 1820),

Description — Rather large but relatively low patelliform from the Miocene of France and the North Sea Basin,

shell, oval in outline, with an apical aperture at one third strongly resembles Diodora italica in ornament, and may behind the anterior shell margin. Anterior margin straight, form part ofthe same evolutionary lineage. posterior only slightly convex. Apical aperture oval,

strongly thickened at the inside. Muscle scars also often

distinct. Ornament consisting of about60 to 80 axial ribs, Superfamily Rafmesque, 1815

which size are unequal in but can not be divided clearly Family Skeneidae Clark, 1851

into and ribs. Intercostal Genus primary secondary areas are nar- Dikoleps Hoisaeter, 1968 row; concentric ornament is relatively weak, never forming

scales. Apertural margin crenate, visibly mostly at the

shell’s interior. Dikoleps cutleriana (Clark, 1848)

Fig. 4/2a-g

Discussion — This species clearly differs from Diodora

cutleriana - Rolan graeca (Linnaeus, 1758) by the larger size, the relatively 1983 Tubiola (Clark) Mosquera, p. 100,

flatter 40. shell (this character is however very variable), the text-fig.

1983 - of axial the Dikoleps cutleriana (Clark) van Aartsen et ah, presence more ribs, poorly distinguishable pri- p. 121, fig. 42. mary and secondary ribs, the weaker concentric ornament

1988 Skenea cutleriana - 45. (Clark) Graham, p. 134, fig. lacking scales and the less distinctly crenate outer edge of

1994 Dikoleps cutleriana (Clark) - Giannuzzi-Savelli et the apertural margin. The internal thickening of the aper- al., p. 106, fig. 353. also in adult shells of ture seems stronger fully D. apertura

than in D. graeca.

Locus typicus — Mediterranean

Occurrence — The species is rare in the Kattendijk Forma-

tion, the Luchtbal Sand Member and the Oorderen Sand Stratum typicum — Recent.

Member from Antwerp. In the North Sea Basin it is also known from Dutch Pliocene beach material and in the Brit- Material — IRSNB 1ST 6958, 6959 (leg. RM), Doel,

ish Coralline Crag Formation. For its further distribution, Deurganckdok; Luchtbal Sand Member.

Landau al. see et (2003, p. 27). Juvenile specimens are relatively higher than adults, such as the specimen figured Dimensions — 4/2a, b, c, d, f: height 0.834 mm, width 1.05

0.90 width 1.14 here (Fig. 2/2). Juvenilesresemble D. graeca in shape, but mm; 4/2e, g: height mm, mm.

their ornament always differs (see above). Miocene speci-

mens from the French Touraine (‘Pontilevian’) are never as Description — Very small turbiniformshell, height usually

less than with rounded whorls. large as Pliocene specimens and their shape consequently width, smooth, remains relatively higher. There are no Miocenerecords of consisting of 1.25 whorls, delimitedby a distinct prosocline

D. italica from the North Sea Basin, suggesting it did not ridge. Ornament ofadultshell starting with about five spi- arrive in the North Sea Basin until the Early Pliocene.This ral ribs, which are narrower than the intercostal spaces. The

possibly correlates with the breakthrough ofthe Channel in number of spiral ribs increases to about 50 on the body

the Pliocene Vliet-Lanoe al. whorl. of and narrower Early as proposed by van et They are equal strength, slightly

than the intercostal into the umbilicus. (2002). The species has a continuous record in Atlantic spaces and continue -22-

Figure 5. -23-

Figure 5.

1. Skeneidae indet. Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6961.

2. Lodderena sphaeroideum (Wood, 1842). a, b, c, e: Deurganckdok, Doel, Luchtbal Sand Member. IRSNB 1ST 6962. d, f: Deur-

ganckdok, Doel. Oorderen Sand Member, Cultellus level. Coll. BR.

3. Alvania whitleyi (Bell, 1898). B1-B2 Kanaaldok, Antwerp. Luchtbal Sand Member. ROM 550 001.

A microsculpture is present, consisting of irregular back- whorl of teleoconch smooth, only with microsculpture in

ward pointing spines at the abapical side of the ribs. Be- the subsutural depression. Thenfirst a narrow abapical spi-

tween the ribs, small tubercles are present. The suture is ral, closely followed by two broader adapical ones, de-

very deep, the sutural ramp covered with microscopic tu- velop. Body whorl with six to nine spirals; adapically bercles. The umbilicus is distinct but narrow. broader than intercostal spaces, abapical on the shell base

These ribs for narrower. spiral are smooth except the three

Discussion — The HAV ratio of this species is variable: abapical ribs that surround the umbilicus, which possess

most specimens are rather flat, but Graham (1988) figures a tubercles. The latter are strongest on the abapical spiral.

distinctly higher specimen than those found in Doel. The Between the spiral ribs, a microsculpture of dense irregular,

concentric and umbilicus axial lines is heavy ornament narrow clearly straight prosocline present. Umbilicus narrow

differentiate this species from all other Skeneidae present and deep.

in the Pliocene of Belgium.

Discussion — This species is attributed to the genus Lod-

Occurrence — Only two specimens were found in the derena of because the presence of three periumbilical scaly Luchtbal Sand Member of the Deurganckdok exposure. It ribs and because ofthe protoconch shape, which is slightly is the first record of the species for the Pliocene of the flattened to one side, although the shell certainly is not North Sea Basin and possibly the first record. The planispiral (see Waren, 1991, p. 135). Waren (1980, pi. 1, now seems to live in rather shallow has an species waters, “ fig. 7-9) figured a specimen, labelled Cyclostrema” Atlantic-Mediterraneandistribution; its northernlimits are sphaeroideum (Wood, 1842) by Jeffreys, collected on the SW , Ireland and the Hebrides (Graham, 1988). Travailleur expedition, noting that it probably did not be-

long to Wood’s species. Landau et al. (2003) figured a

similarEarly Pliocene shell from Estepona, Spain and at- Genus Lodderena Iredale, 1924 tributed its authorship erroneously to Jeffreys. This is a

quite different species, which is much higher instead of

rather flat, with more tumid whorls and axial ribs between Lodderena sphaeroideum (Wood, 1842) the spirals instead of a granular microsculpture; the adult Fig. 5/2a-f ornament starts after the protoconch with axial ribs instead

of spirals. The Recent material figured by Waren and the *1842 533. sphaeroideumWood, p, to the Estepona specimens probably belong genus Parvi-

Wood - 1848 Turbo sphaeroideum Wood, p. 122, pi. 15, turboPilsbry & McGinty, 1945. This species already bears fig. 9. the name Turbo lecointrae Dollfus & Dautzenberg, 1899

1872 Turbo - 86. sphaeroideum Wood Wood, p. 218, 9, after material from the Touraine 71923 - (p. pi. figs 5-6), Cyclostrema sphaeroideum(Wood) Harmer, p. 752, ‘Pontilevian’. It is abundant in localities in pi. 60, fig. 15. (France) some the ‘Redonian’ of and Brittany (France) figured in an un-

— PhD thesis Brebion Locus typicus Sutton, East Anglia, . published by (1964, pi. 3, fig. 17). The

name ofthis species should therefore be Parviturbo lecoin-

trae Stratum typicum — Coralline Crag Formation, Early Plio- (Dollfus & Dautzenberg, 1899) and its range encom-

the Middle and Late Miocene and cene. passes (‘Pontilevian’

‘Redonian’) of Atlantic France, the Early Pliocene (Zan-

Material— IRSNB 1ST 6962 (leg. RM): 1 specimen, Doel, clean) ofEstepona (Spain) and living in the eastern Atlan-

Deurganckdok; Luchtbal Sand Member. Coll. BR: 1 tic Oceanbetween the Bay of Biscay, the Sargasso Sea and

Cultellus specimen same locality, Oorderen Sand Member, Senegal.

level.

Occurrence — Lodderenasphaeroideum (Wood, 1842) is

— from the Coralline Formation of Dimensions 5/2a, b, c, e: height 0.66 mm, width 0.778 a species Crag Gedgrave and Sutton, found for the first time in the Luchtbal mm; 5/2d, f: height 0.90 mm, width 1.14 mm. now

Sand Member in Belgium. Harmer (1923) mentionedthe

Description — Very small, turbiniformshell consisting of species from the Atlantic Pliocene of St. Erth, Cornwall,

but as far as can be from his this about three tumidwhorls with a distinct, impressed suture. judged very poor figure

Protoconch smooth, tumid, consisting of 1 whorl, which is specimen is smooth and it could belong to any other spe-

cies of the Skeneidae. As discussed above, material men- clearly delimited by a slightly prosocline, curved ridge; left

tioned under the from and the side of protoconch less curved than right. First quarter same name France, Spain -24-

Atlantic is not conspecific, so L. sphaeroideum is an en- species occurring in the Kattendijk Formation (see Mar-

demic North Sea basin species. quet, 1995,pi. 6, fig. 2). The main difference, according to

Waren (1992, p. 160) is the size of the protoconch, which

250-290 in Cirsonella measures pm romettensis and 400-

Genus Cirsonella 1877 550 in Skenea basistriata. As be in Angas, pm can seen specimen

in which the limit Fig. 4/3b, protoconch is indicated by an

arrow, it is comparable in size to C. romettensis.

Cirsonella romettensis (Granata, 1877)

Fig. 4/3a-f Occurrence — The species is here mentionedfor the first

time for the North Sea Basin. few Only a specimens were

1874 Turbo romettensis MS - Seguenza Jeffreys, p. 113 found in the Luchtbal and Oorderen Sand Members of

(nomen nudum). Doel. It is a southern, Lusitanian and Mediterraneanspe- *1877 Oxystele romettensis 5, Granata, p. cies and its northern limit was until now the Bay ofBiscay. 1883 Cithna adamsi III. Jeffreys, p. Forits distribution, see Landauetal. (2003, general p. 38). 1889 de 285. Cyclostrema funnazzensis Gregorio, p.

1980 Tharsis romettensis Jeffr - Seguenza ms, eys Waren,

p. 18, pi. 3, figs 1-2. Skeneidae indet.

1992 romettensis - Cirsonella (Granata) Waren, p, 160, Fig. 4/4a-b, 5/la figs 4c, 9e, 1 Ic-e, 12a.

1994 Cirsonella romettensis (Seguenza) - Giannuzzi-

Savelli 350. Material— IRSNB 1ST 6961 1 et al., p. 116, fig. (leg. RM): specimen, Doel,

1996 Cirsonella romettensis (Seguenza) - Palazzi & Vil- Deurganckdok, Luchtbal Sand Member.

107. lari, p. 246, fig.

2003 Cirsonella romettensis - Landau et al. (McGinty) , — p. Dimensions Height 2.82 mm, width 2.44 mm. 38, pi, 8, fig. 8.

Description — Small, naticiformshell, consisting ofthree

Locus — and half which typicus Rometta, Sicily, Italy. a whorls, are tumid, except for the adapical

part on the last whorls; suture deeply incised. Protoconch

Stratum - typicum Unnamed beds, Early Pleistocene. large, about500 pm in cross section, consisting oftwo and

half whorls. These a are smooth and considerably broaden

Material— IRSNB 1ST 6960(leg. RM): 1 specimen, Doel, towards the protoconch-teleoconch boundary, which is

Deurganckdok, Luchtbal Sand Member. CollBR, 2 speci- marked by a sharp ridge. Protoconch and teleoconch mens, Deurganckdok, Oorderen sand Member, Cultellus smooth. Aperture oval, except at the parietal side, which is level. straight, higher than broad, detached, not thickened. Um-

bilicus with very narrow, a ridge running into it.

Dimensions — 4/3 f: 0.66 width a, e, height mm, 0.778 mm; 4/3 c, d: height 0.753 mm, width 1.08 mm; 4/3b: Discussion — None ofthe known Mediterraneanor Atlan- height 0.820 mm, width 1.08 mm (damaged). tic Skeneidae shows the combinationof a large protoconch,

towards the lack of and broadening end, any sculpture a

— small, turbiniformshell narrow umbilicus. Description Very with low spire Even its genus cannot be identifiedwith and two and a halftumidwhorls, protoconch included. Su- certainty. Insufficient material is available to describe this ture deep, but withoutsubsutural incision. Protoconch con- taxon as a new species. There is a superficial resemblance

of sisting two and a quarter tumid whorls, appearing between this species and the Recent Northern Atlantic spe- smooth, but with very small, irregular granules under high cies Skenea trochoides (Friele, 1876)and Skenea peterseni magnification; protoconch/teleoconch boundary marked by (Friele, 1877), both figured by Waren(1991, figs 2e, 2g, 6e 3f, a ridge; cross section about 250 pm. Aperture entire, de- and 4a-b, 6d respectively). Both, however, have a more

circular for the rounded and their tached, nearly except short, straight colu- aperture protoconch does not widen sig- mellar lip. Teleoconch nearly smooth, except for three to nificantly towards the protoconch-teleoconch boundary. four strong spirals around the umbilicus and for very fine

lines. Occurrence — prosocline growth Between the spirals, granular mi- Only a single specimen was found in the crosculpture is present. Umbilicus deep, wide in the speci- Luchtbal Sand Member at Doel, Deurganckdok. mens studied.

Discussion — No adult specimens of this species were Family Rafinesque, 1815 found. These have a very characteristic umbilicus, nearly Subfamily Margaritinae Stolizcka, 1868 closed by an expansion ofthe columellar lip. Furthermore, Genus Margarites Gray, 1847 they are relatively higher and the whorls are less tumid, with a shallow suture. Waren (1992, fig. 11 c-e) figures

Juveniles similar belli growth stages of the species. are very to Margarites (Harmer, 1923) those ofSkenea (Lissospira) hasistriata(Jeffreys, 1877), a Fig. 3/3 a-c, 4/1 -25-

*1923 belli Harmer, 751, — Eumargarita p. pi. 60, fig. 12. Locus typicus Boyton, East Anglia, United Kingdom.

1957b Harmer - Margarites belli Gilbert, p. 10, pi. 1, fig 4.

Stratum typicum — “? Crag”, presumably Red Crag For-

Locus — East United typicus Boyton, Anglia, Kingdom mation, Late Pliocene.

Stratum typicum — Coralline Crag Formation, Plio- Early — Material IRSNB 1ST 6955, 6956 (leg. RM): 2 speci-

cene. mens, Doel, Deurganckdok, Luchtbal Sand Member. Coll

RM, 2 specimens, same locality and unit.

Material— IRSNB 1ST 6957 (leg. RM), Deurganckdok,

Luchtbal Sand Member. Coll RM, 3 same local- specimens, Dimensions — 2/5 3/1 a-c, a-c: height 4.53 mm, width 4.95 ity and unit. mm; 3/ld, e: height 3.90 mm, width 3.73 mm.

Dimensions — Height 1.98 mm, width 3.37 mm. Description — See Marquet (1995, p. 69, pi. 5, fig. 4).

it is Although partially eroded, so that no microsculpture

— Rather small, turbiniform Description shell, broader be the limit of can seen anymore, the protoconch of one than with suture and rounded high deep up to seven whorls. be It is specimen can recognised. slightly larger than one Protoconch consisting ofa smooth bulbous nucleus ofhalf whorl and not clearly delimited.The teleoconch ornament

a whorl and a smooth protoconch II, consisting also of half starts with an abapical spiral close to the suture, followed a

a whorl and clearly delimitedby a strong, prosocline ridge of quarter a whorl later by prosocline axial ribs and later from the teleoconch. After the protoconch/teleoconch again by an adapical spiral. One whorl after the onset ofthe

boundary, two very smooth abapical spirals develop. On teleoconch sculpture, spirals and axials begin to form tu- subsequent whorls, the number of spirals increases, to 12 bercles. Fig. 3/1 d, e shows a juvenile specimen, in which the on body whorl. These spirals partially continue on the the umbilicus is still open, while the axial ornament on the shell base, but are absent around the wide umbilicus. The basal part is more developed than in adult shells.

spiral ribs are very faint and broader than the intercostal

The aperture is ovate. spaces. Occurrence — This species has been foundbefore in Bel-

gium only in the Oorderen Sand Member. It is slightly less

Discussion — This resembles species strongly Margarites rare in the Luchtbal Sand Member ofDoel. trochoides Wood, 1842, figured by Marquet (1995, pi. 3,

but the latterhas rounded fig. 4), a more aperture, a slightly

higher spire and almost lacks the spiral ornament, except Gibbula (Colliculus) woodi(Harmer, 1923) around the umbilicus, which is the smooth area in only Fig. 3/2 a-d

Margarites belli. Both probably form an evolutionary line- age. 1848 Trochus montacuti Wood - Wood, p. 129, pi. 14, fig.

I.

1878 Wood - — Trochus Nyst, Occurrence Margarites belli exclusively occurs in the montagui pi. 7, fig. 3.

1881 Trochus Wood - 109. montagui Nyst, p. Luchtbal Sand Member in Belgium, but very rarely. The *1923 Trochus woodi other localitiesknown (Gibbula) Harmer, p. 736, pi. 69, figs only are in the Coralline Crag For- 14-16. mation of East Anglia. Margarites trochoides is not un-

1957b Gibbula woodi Harmer - (Colliculus) Gilbert, p, 20, common in the Kattendijk Formation. pi. 1, fig. 17.

Locus typicus — Orford, East Anglia, United Kingdom. Subfamily GibbulinaeStolizcka, 1868

Genus Gibbula Risso, 1826

Stratum typicum — Coralline Crag Formation, Early Plio- Subgenus Colliculus Monterosato, 1888 cene.

Material— IRSNB 1ST 6956, 6979 (leg. RM), Antwerp, Gibbula (Colliculus) crassistriata (Bell in Wood, 1882) industrial area near Noorderlaan, dredged, presumably Fig. 2/5a-c, 3/la-e from the Luchtbal Sand Member. Coll RM, 2 specimens,

same locality and unit. Coll. RM, 2 specimens, Antwerp *1882 crassistriata Bell in Margarita Wood, p. 10, pi. 1, Noordkasteel, Luchtbal Sand Member. fig. 15.

1923 crassistriata - Margarites (Bell) Harmer, p. 749, pi,

Dimensions — 3/2 a, b: height 5.26 width 5.08 60, fig. 8. mm, mm; 3/2 1965 c, d: height 5.15 mm, width 4.50 mm. Margarites crassistriata (Bell in Wood) - van Reg-

Altena teren et al, p. 8, pi. 2, fig. 11.

— 1995 Gibbula (Colliculus) crassistriata (Bell in Wood) - Description Medium-sized, turbiniform species, nearly

Marquet, p. 69, pi. 5, fig. 4. triangular in side view, with about five flat whorls. Suture

in 1998b Gibbula (C.) crassistriata (Bell Wood) - Marquet, slightly incised. Protoconch unknown. Ornament solely p. 46, unnumbered text-fig. consisting ofabout 15 spiral ribs, which are nearly twice as -26-

broad the intercostal On the shell 12 with 10 which than those the as areas. base, slightly spiral ribs, are stronger on stronger spiral ribs occur. Aperture nearly round, with whorls. Aperture ovate, with an indistinct columellartuber- straight columellar lip. Umbilicus closed by thickenedand cle. Umbilicus narrow, but surrounded by a broad depres- expanded columellar lip. sion.

Discussion — Species ofthe subgenus Colliculus are com- Discussion — The shells figured and described by Wood mon in the Pliocene of Belgium, including Gibbula (C.) (1848) as Trochus cineroides differmarkedly from those beetsi van Regteren Altena, 1954 (Luchtbal to Kruisschans figured by Harmer (1923), Gilbert (1957b) and in this pa-

Member), G. (C.) obconica nehalenniae van Regteren Al- per, in having strong tubercles on the spirals and rather tena, 1954 (Luchtbal and Oorderen Member), G. (C.) ob- tumid whorls. It could be that Wood’s (1848) figures and conicaobconica (Wood, 1842) (Kattendijk Formation), G. description are based on specimens of Gibbula (Sterom-

(C.) petala Marquet, 1998 (same) and G. (C.) crassistriata phala) octosulcata (Nyst, 1835), which differs from the

(Bell in Wood, 1852) (Luchtbal and Oorderen Member) present species in these characters. The umbilicus of G.

(see Marquet. 1995). Gibbula (C.) woodi (Harmer, 1923) cineroidesreported here is nearly completely closed, which clearly differs from all these species in having flat whorls, is not the case in Wood’s (1848) material, while the tuber- with a shallow suture and a triangular cross section. Fur- cles figured by Wood are much stronger than those of

all other have tubercles their thermore, species on spiral Nyst’s species. Wood (1848) did not mention the occur- ornament, which can become very heavy, lacking in G. rence of G. octosulcata in the British Pliocene, although woodi. Harmer (1923) does. None of the Trochidae present in the

Pliocene of Great Britain or Belgium really fit Wood’s

Occurrence — This species is limited to the Luchtbal Sand (1848) figure and description. Here, the subsequent de-

Member in Belgium and the Coralline Crag Formation in scription of Harmer is followed, but the possibility exists

Britain. No specimens have been found in the Luchtbal that the species will have to be renamed.

Member in Doel.

Occurrence — In Belgium, the species is present in the

Luchtbal Sand Member, but only from the right Scheldt

Subgenus Steromphala Gray, 1848 bank. In Britain, the species seems to be found solely in the

Red Crag Formation, which is surprising since the faunaof

the Luchtbal Sand Member is closest to that of the Coral-

Gibbula (Steromphala) cineroides (Wood, 1842) line Crag Formation.

Fig. 2/4a-b

*1842 Trochus cineroides Wood, p. 531. Subclass Orthogastropoda Ponder & Lindberg, 1995

71848 Trochus cineroides Wood - Wood. 131, 14, p. pi. Superorder Cox, 1959

8b. fig. Order SorbeoconchaPonder & Lindberg, 1997

1923 Trochus (Gibbula) cineroides (Wood) - Harmer, p. Suborder Ponder & Lindberg, 1997 730. pi. 58. fig. 24. Superfamily Rissoidea Gray, 1847

cineroides - 1957b Gibbula (Steromphala) Wood Gilbert, p. Family Gray, 1847 17, pi. I, fig. 12. Subfamily Rissoinae Gray, 1847

Genus Alvania Risso, 1826 Locus typicus — Walton-on-the-Naze, East Anglia, United

Kingdom.

Alvania zetlandica (Montagu, 1815) Stratum typicum — Red Crag Formation, Piacenzian, Late Fig. 6/1 a-d Pliocene.

Material— IRSNB 1ST 6953 1 *1815 Turbo (leg. RM): specimen, Vijf- zetlandica Montagu, p. 94, pi. 13, fig. 3.

1848 Rissoa zetlandica, - de Havendok, Antwerp, Luchtbal Sand Member. CollRM: Montagu Wood, p. 101, pi. 11, fig. 7. 1 specimen, same locality and Member.

1895 Rissoa - (Flemingia) zetlandica(Montagu) Sacco, p.

30. Dimensions — Height 14.1 mm, width 18.2 mm.

1914 Rissoia (Flemingia) zetlandica Montagu - Cerulli-

49. Irelli. p. 194. pi. 15, figs 48, Description — Medium-sized, thick-shelled, nearly recti-

Manzonia - 1920 zetlandica (Montagu) Harmer, p. 622, pi. linear conical shell, with apical angle of90°. Whorlsnearly 50, figs 51, 52. become suture flat-sided, tending to imbricate, shallow; - 1946 Alvania (Taramellia) zetlandica (Montagu) Beets, body whorl rounded abapically. Ornament consisting of p. 36. about 15 broad spiral ribs on the body whorl, which are 1965 Alvania (Alvania) zetlandica (Montagu)- van Regte- The intercostal form inci- unequally strong. spaces narrow ren Altena et al., p. 13, pi. 4, fig. 37.

1978 Alvania - Fretter & sions between the spirals. They contain very faint proso- zetlandica (Montagu) Graham, p. cline lines. Tubercles lacking on the spiral ribs. Shellbase 180, figs 154, 155. -27-

1987 - Manzonia zetlandica 1925 Alvania - (Flemellia) (Montagu) whitleyi (Bell) Harmer, p. 606, pi. 50, fig.

Moolenbeek & 49-51. Faber, p. 166, figs 45. 40.

1988 - Manzonia zetlandica Graham, 252, 1997b Alvania - (Montagu) p. (Alvania) whitleyi (Bell) Marquet,p. 13, pi.

98. fig. 3, fig. 1.

1990 Alvania zetlandica - & Goto, (Montagu) Poppe p.

102, pi. 12, fig. 31. Locus typicus — St. Erth, Cornwall, UnitedKingdom.

Alvania - 1996 zetlandica (Montagu) Waren, p. 225, fig.

17a.

Stratum typicum — St. Erth Beds, Pliocene.

1997 Alvania zetlandica (Montagu) - Di Geronitno & La

Pema, pi. 2, fig. 2.

Material— RGM 550 001 (leg. van der Mark): 1 sp, B1- 1997 Ivania zetlandica (Montagu) - Giannuzzi-Savelli et B2 Kanaaldok, Antwerp (1964), Luchtbal Sand Member. al..p. 112, figs. 476-479, 480d. IRSNB 1ST 5890; 1 specimen, Kruisschans Zeesluis, Oor- 2004 - Landau Alvania zetlandica(Montagu) et al, p. 45, deren, OorderenSand Member? pi. 9, fig. 4. Antwerp;

Dimensions — 3.22 width Locus typicus — “Isle ofNoss, Zetland” (Shetland), United Height mm, 1.66 mm.

Kingdom.

— See (1997b, Description Marquet p. 13).

Stratum typicum — Recent.

Discussion — This is the second specimen ofthis rare spe-

cies from the is Material— IRSNB 1ST 6963 (leg. RM): 1 specimen, Doel, Antwerp region. It strongly eroded, making it but Deurganckdok, Luchtbal Sand Member. Coll BLP (leg. nearly unrecognisable, the typical shape ofthe shell, the round and the MG), 1 specimen, same locality and Member. small, aperture spiral ornament of the

protoconch allow idenfication.

Dimensions — 6/1 b 3.45 width 2.19 a, height mm, mm;

Occurrence — The first of Alvania 6/1 c, d: height 2.63 mm, width 1.79 mm. specimen whitleyi

(Bell, 1898) collected in Antwerp originated fromthe basal

ofthe “Scaldisien”, which — attributed Description Small, rissoiform shell with an elevated crag Marquet (1997b)

to the basal of the Oorderen Sand Member. spire and deep, angular suture. Protoconchnot preserved in crag It could

the material. Teleoconchwith about five whorls. Ornament however have derived from the Luchtbal Sand Member.

The second described is from the consisting of equally strong axial and spiral elements, specimen, here, certainly Luchtbal Sand Member. forming rather weak tubercles at the intersections. On the

axials body whorl, 14 and 5 spirals are present in the most

and axials form complete specimen. Spirals rectangles or Superfamily 1822 hexagons, which are covered by a very dense spiral mi- Capuloidea Fleming, Family 1822 crosculpture, which however could be lost due to wear. Fleming,

Aperture elongate oval, entire and with strongly thickened Subfamily Capulinae Wenz, 1840

margin. Umbilicus closed. Genus Capulus Montfort, 1810

Discussion — the Rissoidae Among present in the Doel and Kallo sections, only Alvaniabeani (Hanley in Thorpe, Capulus deurganckensis nov. sp. 6/4 a-f 1844) shows some resemblance (see Marquet, 1997b,pi. 2, Fig.

fig. 6). The latter clearly differs, however, by its shallow

1872 and Pileopsis var. sinuosus Brocchi - Wood, suture more numerous axial and spiral ribs. ungaricus p. 88, pi. 7, fig. 26.

1878 Pileopsis ungaricus var. sinuosus Brocchi - Nyst, pi. Occurrence — Only two heavily damaged and worn speci- 7, fig. 26. mens were collectedin the Luchtbal Sand Memberof Doel; 1881 sinuosus Brocchi - Pileopsis ungaricus var. Nyst, p. the first they are specimens recorded in the Pliocene of 118. Belgium. In Britain, the species has been found in the

1923 Brocchia sinuosa - (Brocchi) Harmer, p. 768, pi. 61, Coralline and the Red Crag Formations. Material is also fig. 10. known from The Netherlands, collectedfrom boreholesand 1958a Brocchia sinuosa Brocchi - 23. Gilbert, p. from beaches (Beets, 1946; van Regteren Altena et al,

1965). For its distribution, from general ranging — Early Locus typicus Deurganckdok, Doel, municipality Beve-

Pliocene to Recent, see Landau et al. (2004, 45). p. ren, prov. Oost-Vlaanderen, Belgium.

Stratum typicum — Luchtbal Sand Member, LilloForma- Alvania whitleyi (Bell, 1898) tion, late Zanclean, Early Pliocene.

Fig. 5/3 a-c

- Material — Holotype IRSNB 1ST 6964 (leg. RM). Para- *1898 Rissoa whitleyi Bell, 153 p. type: IRSNB 1ST 6965 (leg. RM). -28-

Figure 6. -29-

Figure 6.

1. Alvania zetlandica (Montagu, 1815). a, b, d: Deurganckdok, Doel. Luchtbal Sand Member. Coll. BLP. c: Deurganckdok, Doel.

Luchtbal Sand Member. IRSNB 1ST 6963.

2. Nassarius (Uzita) reticosus (Sowerby, 1815). Antwerp, Noorderlaan, dredged material. Luchtbal Sand Member? Coll. LC.

3. (Pseudomurex)bracteata (Brocchi, 1814). Antwerp,Noorderlaan, dredged material. Luchtbal Sand Member?Coll. LC.

4. Doel. Luchtbal Sand Member. IRSNB 1ST 1 Capulus deurganckensisnov. sp. a, b, d, e: holotype Deurganckdok, 6964. c, e: paratype

Deurganckdok, Doel. Luchtbal Sand Member, IRSNB 1ST 6965. f; paratype 2 Deurganckdok, Doel. Luchtbal Sand Member. Coll.

BLP.

5. Nassarius (Uzita) spectabilis vandewouweri (Glibert, 1959). Antwerp, Noorderlaan, dredged material. Luchtbal Sand Member?

IRSNB 1ST 6966.

Other material: Coll. RM: 5 specimens; IRSNB n.n.: see genus Brocchia by all European taxonomists were a xeno-

Coll. Landau Nyst (1881, pi. 7, fig. 26). BLP (leg. MG); et morphic sculpture associated with a living mode on pecti- al. (2004, pi. 15, fig. 2). nid bivalves; specimens living on smooth pectinids would

lack plicae whereas those living on ribbed pectinid species

Dimensions— Holotype: height 17.6 mm, aperture 39.0 x would develop plicae with sharp digitations at the shell

31.3 mm; Paratypel: height 17.4 mm, aperture 41.1 x 34.5 margin corresponding to the ribs ofthe pectinid. Certainly

mm. the series illustratedfor the species Capulus danieli(Cosse,

1858)from New Zealandis convincing, showing forms that

— Named after the would have been considered within the Derivatio nominis type locality. certainly genus

Brocchia. However, this raises some questions within the

Diagnosis — A Capulus species with a paucispiral proto- Pliocene North SeaBasin and European faunas; specimens

conch of one whorl, with spiral plicae on the whole shell with plicae are lacking in the Oorderen or Kruisschans

that distinct before the umbo. Sand Membersof and are most Kallo Doel, although smooth as well

as ribbed Pectinidae abound in these Members, as well as

Description — Large shell, with a smooth protoconch, de- in the Luchtbal Sand Member. The most common pectinid

limited by a thin ridge and consisting of only one whorl. in the Luchtbal Sand Member is the smooth Pseudamus-

Protoconch diameter770 with nucleus pm, a large (diame- sium gerardi (Nyst, 1835).Also in Recent European popu-

ter 289 shellabout two whorls. lations the pm). Complete Shape very comparable specimens are lacking, despite con-

variable, oval in outline, aperture broader than wide. Aper- tinued presence of both Capulus and pectinids in the Re-

tural margin very irregular, probably following the outline cent fauna. We provisionally accept the synonymy ofBroc- of the substrate. Umbo very close to anterior margin, at chia with Capulus, noting that the observations made by about of total width. Anterior whilst for danieli one eighth apertural margin Beu (2004), convincing Capulus are not rather straight. Seen in side view, to the right of the umbo, as obvious in the European faunas. Beu (2004) suggested

the shell more or less strongly bulges and lowers gently to that Capulus hungaricus and Brocchia sinuosa might be

abruptly at the posterior end. Ornamentofthree to six deep conspecific. We do not agree with this synonymy for our

incisions and which of plicae, run nearly over the complete material. There are differences in the size their proto-

shell, but are especially distinctbefore the umbo.Further- conch (Landau et al., 2004) and in the thickness of the shell

more, a large numberof fineaxial ribs is present. Concen- and surface ornament.

tric ribs are mostly fine, some becoming stronger than the Marquet (1997c, 1998b) figured another type of shell from axial sculpture. Doel and Kallo (Oorderen Sand Member) under the name

of Brocchia sinuosa (Brocchi, 1814). It differs from Capu-

Discussion — As mentionedin Landau et al. lus (2004, p. 69), deurganckensis nov. sp. in having a triangular shape, this species clearly differs fromthe MediterraneanPliocene with the umbo distinctly surpassing the apertural margin.

Capulus sinuosus (Brocchi, 1814)in having a paucispiral The axial ribs are finer. The plicae are distinct only to the

protoconch with one instead oftwo and a halfwhorls. Fur- right of the umbo when seen from above and they run ir-

in the Mediterranean thermore, species the plicae nearly regularly, not in a spiral. This species occurs in the Oor- run straight, while in the new species they form a clear spi- deren Sand Member. It has been figured under different ral. Axial ornament is nearly absent and the umbo nearly or names by Woodand Harmerand its correct name is Capu-

completely overhangs the anterior apertural margin. Capu- lus partimsinuosa (Wood, 1848). Synonyms are: Capulus

lus sinuosus (Brocchi, 1814) is with militaris var. partim-sinuosaWood (1848, 156, 17, probably conspecific ...... p. pi.

Capulus laevis (Bronn, 1831). Capulus ungaricus (Lin- fig. 3 a-b, non 3c-f), Brocchia plicata Harmer (1923, ,, r p.

naeus, 1758) also shows some resemblance to the new 769,--. pi. 61, fig. 11), Brocchia partim-sinuosa (Wood) in - - — spe- r . r

cies, but its protoconch consists of two and a quarter Harmer(1923, p. 769, pi. 61, fig. 12) and Capulus ? incer-

whorls, with a very tiny nucleus, its shell shape is much tus Bell1(1870, p. 216), Brocchia incerta (Bell) in Harmer

more regular and spiral plicae are lacking. (1923, p. 770, pi. 61, figs 13-14).

Similar species have been considered as a separate genus,

Brocchia 1827. Beu that the Occurrence — Bronn, (2004) argued ‘plicae’ Capulus deurganckensis nov. sp. is an en-

seen on the posterior slope of specimens included in the demic North Sea Basin species. -30-

Figure 7. -31 -

Figure 8. -32-

Figure 7.

1. Xenophora scaldensis Glibert, 1958. d, e, f: Antwerp,junction Vijfde Havendok-Amerikadok. Luchtbal Sand Member. IRSNB 1ST

Havendok-Amerikadok. Formation. IRSNB 1ST 6968. h; 6967. a, b, c: Antwerp,junction Vijfde Kattendijk g, Antwerp,Ring High-

exit Derived Pliocene. IRSNB 1ST 6969. way Borgerhout.

2. truncatula (Bruguiere, 1792). Deurganckdok. Doel. Luchtbal Sand Member. IRSNB 1ST 6970.

3. Polygyreulima glabella (Wood, 1848).B1-B2 Kanaaldok, Antwerp. Luchtbal Sand Member. RGM 550 003.

Figure 8.

1. Xenophorascaldensis Glibert, 1958. Antwerp. Ring Highway exit Borgerhout. Derived Pliocene.IRSNB 1ST 6969.

2. Acirsa (Hemiacirsa) penepolaris (Wood, 1872). Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6971.

3. Amaea cancellata (Brocchi, 1814). Deurganckdok, Doel. Oorderen Sand Member, Cultellus level. Coll. SM.

4. Amaea woodi (Deshayes, 1861). Deurganckdok, Doel. Luchtbal Sand Member, coll. IRSNB 1ST 6972.

5. () obtusicostata (Wood, 1848)Deurganckdok, Doel. Luchtbal Sand Member. Coll. BLP.

6. Epitonium frondosum (Sowerby, 1829). Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6973.

In Belgium it is limited to the Luchtbal Sand Member, Amerikadok, Antwerp; Kattendijk Formation, Ditrupa- while the figured British records are from the Coralline Glycymeris bed. IRSNB 1ST 6969 (leg RM), Ring High-

Crag Formation. Harmer (1923, p. 768) also mentions re- way near exit Borgerhout, Antwerp; derived mixed Plio- cords from the Red Crag Formation, but his identification cene beds. Coll RM: 3 fragments, Junction Vijfde Haven- has to be checked. dok-Amerikadok, Antwerp; Luchtbal Sand Member. Coll

RM: 4juvenile specimens, Deurganckdok, Doel; Luchtbal

Sand Member. Coll. RM: 2 specimens. Junction Vijfde

Superfamily Troschel, 1852 Havendok-Amerikadok, Antwerp; Kattendijk Formation,

Family Xenophoridae Philippi, 1853 Ditrupa-Glycymeris bed. Coll. RM: 1 specimen, Ring

exit derived mixed Genus Xenophora Fischer von Waldheim, 1807 Highway near Borgerhout, Antwerp;

Pliocene beds. Coll RM: 1 fragment, Kallo, Oorderen Sand

Member, basal crag (probably derived). Coll NNM: 4

Xenophora scaldensis Gilbert, 1958 specimens, Antwerp dock works, level unknown, probably

Fig. 7/1 a-h, 8/1 Luchtbal Sand Member. Coll IIRSNB: 20 specimens from

the Kattendijk Formation and 18 specimens from the

Luchtbal Sand Member of different localities. 1946 Xenophom (Xenophora) cf. deshayesi (Michelotti) -

Beets, p. 46.

1958a Xenophora deshayesi Michelotti subsp. scaldensis Dimensions — 7/1 d-f: height 42.6 mm, width 58.6 mm;

24. Glibert, p. 7/1 a-c: height 39.9 mm, width 63.5 mm; Ig, h, 8/1: height

1997 scaldensis Glibert - Xenophora Marquet, p. 72, pi. 1, 35.6 mm (incomplete), width 58.5 mm.

fig. 7.

1998b Xenophora scaldensis Glibert - Marquet. p. 80, text- Diagnosis — Medium-sizedXenophora with an apical an- fig- gle ofabout 80°, few attachments near the suture, ornament

axial ribs and shell base only consisting of very irregular

Locus typicus — Province Antwerp, Belgium. with curved growth lines, which become sinuous near the

aperture.

Stratum typicum — Luchtbal Sand Member, Early Plio- cene. Description — Shell triangular in cross section. Apical

angle about80°. Shell broader than high, height 62-72%of

material— Glibert includedin his Type (1958a) originally width. At least seven whorls are present, but the protoconch

Xenophora deshayesi Michelotti, sp. 1847 subsp. scalden- is lacking in all material studied. Suture very shallow, with

from the Formation sis only Pliocene material, Kattendijk short flange on last whorl. Whorls straight. Few attach-

and the Luchtbal Sand without Member, designating a type. ments are present, mainly near the abapical suture. Sculp-

Glibert (1963, p. 24) tried to rectify this, but unfortunately ture only consisting of very strong, irregular prosocline

the chose a Miocene specimen as “holotype”: specimen plicae, which are more or less well developed and termi-

Glibert This action is figured by (1952, pi. 5, figs 3a, b). nate occasionally before reaching a suture. Aperture small,

not conforming to the International Code of Zoological ovate. Umbilicus wide, oval, bordered by the glossy colu-

Nomenclature (ICZN). Any subsequent designated type mellar lip of the aperture. Basal sculpture consisting of

would be A be which rounded specimen a lectotype. lectotype can only dense irregular growth lines, are to slightly

selected from a syntypical series and the Miocenematerial angular halfway and end in a sinuous curve near the aper-

does not belong to this series. Therefore, a lectotype was ture.

chosen and its designation will be proposed to the ICZN.

Material— IRSNB 1ST 6967 (leg. RM) Junction Vijfde Discussion Glibert (1963) supposed his “holotype” from

Havendok-Amerikadok, Antwerp;Luchtbal Sand Member. Borgerhout (Antwerp) could originate from the Late Mio-

Junction Havendok- This IRSNB 1ST 6968 (leg. RM), Vijfde cene Deume Sand Member of the Diest Formation. -33-

Member yields only few gastropod species in Borgerhout, Family Aclididae Sars, 1878

The molluscan mostly Epitoniidae. remaining faunais es- Genus Adis Loven, 1846

pecially rich in calcitic Pectinidae and Ostreidae, so it is

very improbable that a complete aragonitic Xenophora

wouldbe found. Xenophoridae are only found as steinkems Adis ascaris (Turton, 1822)

atthat locality in the “Sables aHeterocetes” sandstone, also Fig. 9/la-b

belonging to the Diest Formation. The specimen figured by

Glibert still contains 1842 Alvania (1952) black, glauconitic sediment, supra-nitida Wood, p. 534.

1848 Alvania ascaris Turton - excluding the “Sables a Heterocetes” sandstone and the Wood, p. 99, pi. 12, fig. 11.

derived 1914 Adis supranitida Wood - Cerulli-Irelli, 243, pi, Pliocene crag above it as origin. It probably was p. 37-41. found in 21, figs the Antwerp Sand Member, as Glibert (1952)

1922 Adis - supranitida (Wood) Harmer, p. 868, pi. 65, originally supposed. This Member contains two species of fig. 24. Xenophoridae, which clearly differfrom X. scaldensis and

1965 Adis minor - van Altena (Brown) Regteren etal., p. were well-figured by Janssen (1984) from the MiddleMio- 21, pi. 8, fig. 77. cene from Miste (The Netherlands). The first, X. deshayesi

1983 Adis ascaris - Rolan (Turton) Mosquera, p. 189, (Michelotti, 1847), becomes much than X. scalden- larger text-fig.

sis. but remains relatively flatter and the area around the 1988 Adis ascaris - (Turton) Graham, p, 505, fig. 213.

narrow or closed umbilicus is convex. The second 1997c Adis ascaris - 2. very (Turton) Marquet, p. 84, pi. 4, fig.

X. 1998b Adis ascaris - species, burdigalensis (de Grateloup, 1847) comes (Turton) Marquet, p. 113, unnum-

closer in size and shape to X scaldensis, but its umbilicus bered text-fig.

1999 Adis ascaris (Turton) - et is always closed by a callus. The growth lines on the base Giannuzzi-Savelli al, p. 58, figs 102-105. of the shell make an angle of nearly 90° at three quarters

distance from the umbilicus; they gently curve in the Plio-

Locus typicus —- Great Britain. cene species. The “holotype” of X scaldensis designated

by Glibert (1963) probably belongs to this taxon.

Stratum — Recent. Several other Pliocene typicum European Xenophora species are known. The species occurring in the ItalianPliocene(Siena

Material— IRSNB 1ST 6974(leg. RM): 1 Doel, Basin) were reviewed by Manganelli et al. (2004). Xeno- specimen, Deurganckdok, Luchtbal Sand Member. Coll. BR: 1 phora crispa (Konig, 1825) from the Pliocene to Recent of specimen, Kallo, Verrebroekdok, Kattendijk Formation, the Mediterranean and adjacent Atlantic remains smaller, has material attached bed. more on its shell, and has very dis-

tinct tubercles on the shell base and angular basal growth

Dimensions — Height 1.57 mm, width 0.715 mm lines. Xenophora infundibulum (Brocchi, 1814) (Mediter- ranean Pliocene) has a completely closed umbilicus cov-

— See and ered callus, few attachments, Description Marquet (1997c, p. 84) Marquet by very strong basal ornamen- To these (1998b, 113). descriptions the following can be tation and very distinct interrupted axial sculpture above p. added. The protoconch/teleoconch is delimited the sharply protruding carina. Xenophora plioitalica boundary

a curved (Sacco, 1896) (MediterraneanPliocene) is broad and flat sharply by strongly sinusoid, slightly prosocline,

line, after which the teleoconch ornament develops instan- and resembles Xenophora deshayesi (Michelotti, 1847)

much in it couldbe taneously. The adult spirals are very narrow, one-third very ornament; a Pliocene descendant only the width of the intercostal but delimited. of the latter species. Xenophora davoliiManganelli, Spad- spaces, sharply The is aperture ovate rather thanrounded as in ini & Cianfanelli, 2004 is more than three times larger, it elongately

has the fragmentary juvenile specimen figured strongly angular whorls with a sharply keeled periph- by Marquet eral flange and its umbilicus is closed. (1998b).

Ponder (1983, p. 18) gave a diagnosis ofXenophora. Shell

Occurrence — Until now the had been found in characters include: the narrow peripheral flange, which is species Belgium only in the Oorderen Sand Member. Its is simple and non porcellaneous ventrally, the umbilicus is range

extended to include the Luchtbal Sand Member and the moderate to closed and foreign objects are attached to all Kattendijk Formation (Petaloconchus bed). whorls and usually occupy more than one third ofthe dor-

sal surface. The Belgian Pliocene species concurs with all

characters mentioned; only the extent of the attachmentof Family Epitoniidae Berry, 1910

foreign objects is rather variable, some of the specimens Subfamily Epitoniinae Berry, 1910

are covered for less than one third. This, however, is also Genus Opalia H. & A. Adams, 1853

the case in Xenophora infundibulum (Brocchi, 1814), Subgenus Nodiscala de Boury, 1889

which Ponder includes in the (1983) same genus.

Opalia (Nodiscala) obtusicostata (Wood, 1848)

Suborder Gray, 1853 Fig. 8/5a-d

Janthinoidea 1810 *1848 Scalaria obtusicostata Superfamily Lamarck, Wood, p. 95, pi. 8, fig. 21 -34-

Scala obtusicostata - — 1920 (Pliciscala) (Wood) Harmer, p. Material IRSNB 1ST 6972 (leg. RM): 1 specimen,

554, pi. 68, figs 24-25. Deurganckdok, Luchtbal Sand Member. Coll. RM: 1 fr.,

1958a Opalia (Pliciscala) obtusicostata Wood - Gilbert, p. same locality and Member. 11, pi. 2. fig. 7.

1965 Opalia (Pliciscala) obtusicostata (Wood) - van Reg- Dimensions — Height 4.19 mm, width 1.78 mm. Altena 68. teren et al, p. 19, pi. 7, fig.

Description — Medium-sized species, rather broad (H/W Locus typicus — Sutton, East Anglia, UnitedKingdom. ratio 2.35), with tumidwhorls and deep suture. Protoconch

incomplete in our material, multispiral, with at least four Stratum typicum — Coralline Crag Formation, Early Plio- tumid smooth whorls, which have a microsculpture of ir- cene. regular pits. Their width increases slower than that ofthe

teleoconchwhorls. Ornament starting with closely packed Material — Coll. BLP: 1 specimen, Deurganckdok, axial ribs thatbecome separated on subsequent whorls. On Luchtbal Sand Member. the second teleoconchwhorl, spiral ribs appearbetween the

axials. The axials remain distinct, about as broad as the

Dimensions — Height 3.61 mm, width 1.41 mm. intercostal spaces. They are nearly opisthocline, but turn

prosocline near the abapical suture. On the body whorl of Description — Elongate shell, consisting of more than 10 the figured specimen, 20 axials and about 20 much less whorls. Suture deeply incised, whorls tumid. Height of distinct spirals are present. The shell base is not delimited multispiral protoconch 1.37 mm, consisting of about seven by a keel. The shell is smoothexcept for growth lines. Ap- tumid, smooth whorls. The teleoconch ornament starts with erture entire, nearly circular, but pointed adapically. an axial rib. On the body whorl, 16 strong, nearly ortho- cline ribs axial are They are about as broad as the present. “ ” Discussion — This species clearly differs from Opalia intercostal spaces. Between the axial ribs, circa 15 very obtusicostata (Wood, 1848) by the presence ofmore axial shallow spiral ribs are present, slightly broader than the ribs, the absence of a basal keel and from lines ofpits on intercostal spaces. The aperture is entire, nearly circular. the base. The specimen figured is juvenile, a fragment col- Base ofbody whorl delimited by a carina, abapical of the lected attains a height of 13.7 mm. Amaea cancellata axial and spiral ornament. Belowthis carina, 10 spiral rows (Brocchi, 1814) also occurs in the North Sea Basin. It was of pits are present; above, no pits occur. mentionedby Harmer (1920) from the CorallineCrag For-

mation, but one almost perfect specimen has been found Occurrence — This species is extremely rare: only one recently at Doel (Deurganckdok), Oorderen Sand Member, specimen was foundin the Luchtbal Sand MemberofDoel. Cultellus level (coll. SM). This species differs from A. Its also limited to to range seems this Member and the Brit- woodi(Deshayes, 1864) by its ornament: the axial ribs are ish Coralline Crag Formation. The material, mentionedby finer and equal in strength to the spiral ribs, while the shell van Regteren et al. (1965) from The Netherlands was col- base is not smooth but covered by dense spiral ribs (Fig. lected ex-situ on beaches. 8/3a-d). Amaea cancellata is new for the Pliocene ofAnt-

werp.

Genus Amaea H. & A. Adams, 1853 Occurrence —Amaea woodi (Deshayes, 1861) is very rare

in the Luchtbal Sand Member of Doel and other localities

in Belgium. One specimen was found by the authors (RM) Amaea woodi (Deshayes, 1861) in the Oorderen Sand Member ofDoel. In Britain, it occurs Fig. 8/4a-f in the Coralline Crag Formation, in The Netherlandsit has

been found in beach material.

1848 Scalaria cancellata Brocchi - Wood, p, 95, pi. 8, fig.

22.

*1861 Scalaria woodi 339. Deshayes, p. Genus Acirsa Morch, 1857

1920 Scala (Clathroscala) woodi - Deshayes Harmer, p. Subgenus Hemiacirsa de Boury, 1890 552, pi. 48, figs 10-11.

1952 Scala (Clathroscala) cancellata Brocchi - van Der

2. Burg, p. 51, pi. 3, fig. Acirsa (Hemiacirsa) penepolaris (Wood, 1872)

1958a Amaea woodi - (Clathroscala) Deshayes Glibert, p. Fig. 8/2a, b 12.

1965 Amaea (Clathroscala) woodi (Deshayes) - van Regte- *1872 Turritella ? penepolaris Wood, p. 53, pi. 4, 20, Altena 71. fig. ren et al., p. 19. pi. 7, fig.

1879 Turritella Wood - (Mesalia) penepolaris Wood, p.

26, pi. 2, fig. 14.

Locus typicus — Sutton, East Anglia, United Kingdom. 1920 Scala chatwini 48, (Plesioacirsa) Harmer, p. 557, pi.

figs 38-39.

Stratum typicum — CorallineCrag Formation, Early Plio- 1958b Acirsa Wood - (Plesioacirsa) penepolaris Glibeit, p. cene. 10. -35-

- 1997c Acirsa aff. Mar- — (Hemiacirsa) penepolaris (Wood) Dimensions Height 15.3 mm, width 6.7 mm.

quet, p. 85, pi. 4, fig. 3.

1998b Acirsa (Hemiacirsa) cf. penepolaris (Wood) - Mar- Description — Shell with at least eight tumid whorls (no unnumbered quet, p. 116, text fig. complete specimens available). Suture deep, shellrelatively

broad and rapidly increasing in width. Body whorl with 12

Locus typicus — Orford, East Anglia, UnitedKingdom, axial ribs. These are very sharp, nearly orthocline to

slightly prosocline andthey cover the whorls from suture to

Stratum typicum — Corraline Crag Formation, Early Plio- suture. The axials are thinand much narrower than the in- cene. tercostal than areas (less a quarter). A sharp, detached

is about fifth of spine present at one a whorl below the

Material — IRSNB 1ST 6971 (leg. RM): 1 specimen, Doel, adapical suture. Aperture circular. Deurganckdok, Luchtbal Sand Member. Coll. RM: 15 fr. and specimens, same locality and unit. Coll. AR: 1 speci- Discussion — The nature of this species has been poorly men, Kallo, Vrasenedok, ex-situ. understood. Wood (1848) mentionedthe presence of9 to

10 axial ribs, but said also that too many (about 15) were

Dimensions— Height 19.3 mm, width 4.8 mm. drawn on his figure (pi. 8, fig. 15). Scala (Lamelliscala)

frondosa Sowerby as figured in Harmer (1920) is much

— slender (H/W ratio > 4), Description Elongate, very broader than the real Epitonium frondosum the ribs are , with about 12 nearly flat-sidedwhorls separated by a shal- strongly prosocline and it is clear that the specimens fig- low suture. Protoconch indistinguishable from teleoconch ured belong to Epitonium foliaceum instead. Harmer’s by wear in the material examined. Teleoconch ornament (1920) Scala (Nobiliscala) belgica (a synonym of Epito- consisting of about 20 spiral ribs of unequal strength, but nium foliaceum), figured in pi. 48, fig. 26, is too much much broader than the always very narrow intercostal inci- worn to allow unambiguous identification. It originates sions. Shell base smooth. Aperture oval, pointed above, from Selsey, a locality yielding only Late Pleistocene fos- forming an angle between columellarand parietal lip. sils. Also the ribs of the specimen figured are too thick to

belong to Epitonium foliaceum. So probably this specimen

Discussion — This species differs clearly fromboth afore- is wrongly identified, but its preservation is too poor to mentionedAmaea its species by narrow shell, low whorls, allow certain identification. Epitonium f. frondiculum and especially by the absence of axial ornament. The differs from (Wood, 1842) E. frondosum in having more specimen figured by Marquet (1997c, pi. 4, fig. 3) lacks ribs (about 15), which are much broader (halfof the inter-

these ribs due to wear. costal spaces). The spines are less clearly detachedand lie

closer to the adapical suture. The whorls increase in width

Occurrence — The specimen mentionedin Marquet (1997) less rapidly than in E. frondosum. Epitonium foliaceum

was collected ex-situ. Acirsa penepolaris is not rare in very has (Sowerby, 1825) about eight ribs on the last whorl. the Luchtbal Sand Member of Doel; it also occurs in the These are markedly prosogyrate. The shell is relatively Kattendijk Formation (Gilbert, 1958a). In Britain it is broader in relation to height than E. frondosum. found in the Coralline and Red Crag Formations.

Occurrence — Epitonium frondosum is known with cer-

tainty only from the British Coralline Crag Formationand Genus Epitonium Roding, 1798 from the Luchtbal Sand Member ofBelgium. Very similar

specimens were figured by Cerulli-Irelli (1914) from the

Early Pleistocene of Monte Mario, Rome, Italy, but no Epitonium frondosum (Sowerby, 1829) Pliocene material is known from the Mediterranean. Fig. 8/6a-c

*1829 Scalaria frondosa Sowerby, p. 149, pi. DLXXVII, Superfamily EulimoideaH. & A. Adams, 1853 fig. 1. Family H. & A. Adams, 1853

1848 Scalaria - frondosaSowerby Wood, p. 92, pi. 8, fig. Genus Sacco, 1892 15. Polygireulima

1914 Scala (Spiniscala) frondosa Sowerby - Cerulli-Irelli,

41. p. 226, pi. 20, fig. Polygireulima glabella (Wood, 1848)

Fig. 7/3a-b Locus typicus — Sutton, East Anglia, United Kingdom.

*1848 glabella Wood, 95, 19, 2. , p. pi. fig. Stratum typicum — Coralline Crag Formation, Early Plio-

1872 Eulima Wood - glabella Wood, p. 27, pi. 7, fig. 4. cene. 1923 Eulima - glabella Wood Harmer, p. 844, pi. 64, fig.

27.

Material — IRSNB 1ST 6973 (leg. RM): 1 specimen, 1969 Melanella (Polygyreulima) glabella (Wood) - Jans- Deurganckdok, Luchtbal Sand Member. Coll. RM: 3 fr., sen, p. 156, pi. 2, fig. 5, pi. 8, fig. 2.

same locality and Member. 1984 Balds - (Polygyreulima)glabella(Wood) Janssen, p. -36-

178, 51, 8. pi. fig. Coralliophila (Pseudomurex) bracteata (Brocchi, 1814)

2001 Melanella (Polygyreulima) glabella (Wood, 1848)- Fig. 6/3a-c

Wienrich. p. 465, pi. 75, fig. 8, pi. 76, fig, 1.

*lBl4 Murex bracteatus 3. Brocchi. p. 409, pi. 9, fig.

Murex 1904 (Pseudomurex) bracteatus Brocchi - Locus typicus — Sutton, East Anglia, United Kingdom. Sacco, p. 23, pi. 6, fig. 34.

1904 M. bracteatus var. miopericarinata Sacco - Sacco, Stratum typicum — Coralline Crag Formation, Early Plio- p. 24, pi. 6, fig. 35. cene. 1944 Coralliophila (Pseudomurex) cf. bracteatus (Broc-

chi) var. miopericarinata Sacco - van Voorthuyzen,

Material — RGM 550 002 (leg. VDM): 2 specimens, BI- p. 75, pi. 6, figs 17,24-25. B2 Kanaaldok, Luchtbal Sand Member. RGM Antwerp, 1955 Coralliophila (Pseudomurex) bracteata (Brocchi) - 550 1 and unit. 003, specimen, same locality Rossi Ronchetti. p. 192, fig. 101.

1964 Coralliophila (Pseudomurex) bracteata (Brocchi) -

Dimensions — 7/3a: 3.97 width 1.20 179. Fig. height mm, mm; Anderson, p. 245, pi. 21, fig.

Murex Brocchi - Fig. 7/3b: height 3.63 mm, width 1.36 mm. 1978 bracteatus Pinna& Spezia, p. 145,

pi. 30, fig. 1.

1984 Coralliophila - — bracteata (Brocchi) Description Small, elongate shell, consisting of about (Pseudomurex)

Janssen, p. 226, pi. 61. fig. 2. eight smooth whorls. Width about 35% of height, shell

1992 Coralliophila bracteata (Brocchi) - Cavallo & Re- towards the last whorl. Whorls broadening very slightly petto, p. 98, fig. 219. tumid, especially abapically. The first three whorls are

1995 Coralliophila (Pseudomurex) bracteata (Brocchi) - slightly obliquely oriented in respect with the shell’s axis, Baluk, p. 233, pi. 25, figs 6-7. whorls The suture is indistinct.The subsequent are straight. 2000 bracteata - Coralliophila (Brocchi) Chirli, p. 48, pi. ofthe whorl is about third the of the height body one height 19, figs. 5-8. entire shell. Apex blunt, protoconch indistinguishable. 2001 Ap- Coralliophila (Pseudomurex) bracteata (Brocchi) - erture with anal sinus and ovate, pointed above, strong pe- Wienrich. p. 473, pi. 77, fig. 8, pi. 94, figs. 4-5. ripheral bulge. Columellar side markedly thickened.

Locus typicus — “Piacentino”, Italy.

— least Discussion At fourother eulimid species occur in

Stratum typicum — Unnamed beds. Neogene. the Pliocene ofAntwerp. Eulima glabra (Da Costa, 1778) is much narrower, larger, with a straight outer lip.

Material— Coll. EC: 1 specimen, Antwerp, industrialarea Melanellaalba (Da Costa, 1778) also is much larger, rela- near Noorderlaan, dredged, presumably from the Luchtbal tively broader, slightly carinate, with a less curved outer Sand Member. lip. Vitreulimaphilippii (Rayneval & Ponzi, 1854)is ofthe same size, but always very clearly curved and narrower Dimensions— Height 29.0 mm, width 15.5 mm. (see variation as shown in Giannuzzi-Savelli et al„ 1999, figs 312-319). Crinophteiros collinsi (Sykes, 1903) is the Description — Medium-sized, muriciform, thick-shelled closest resembling species, but the body whorl is relatively Suture whorls with species. deep, tumid, a subsutural plat- higher, the shell is broader (width about40% ofheight) and form. Protoconch not present in the studied specimen. Or- the whorls are flatter.The Recent Atlantic-Mediterranean of 25 ribs nament consisting spiral ribs, up to primary on Polygireulima sinuosa (Scacchi, 1836) has more whorls, a with body whorl, a weaker secondary rib in between. Inter- higher aperture and less tumid whorls (see Graham, 1988, costal about spaces as wide as primary ribs. On the spirals, fig. 227). numerous rectangular tubercles are present, which can de-

into small scales. canal velop Siphonal short, open, with a

Occurrence — (Wood, 1848) is Polygireulima glabella narrow pseudumbilicus. found from the Miocene onwards. Janssen (1969, 1984) and Wienrich (2001) mention its occurrence in the Dutch Discussion — This species is very variable in shape and Miste Bed and the German Reinbek Stufe. It also in occurs size, but its ornament is always characteristic. No other the Antwerp Sand Member, Berchem Formation(coll. RM, species in the Luchtbal Sand Member shows any resem- It unpublished). has been found in the Coralline Crag For- blance. mation of East Anglia, U.K. This is the first record from the Pliocene of Belgium. Occurrence — Coralliophila bracteata has hitherto not

beenrecorded from the North Sea Basin Pliocene. It is rare

in the MediterraneanPliocene (Spain, France and Italy). Order Thiele, 1929 The sole specimen at hand was found ex-situ but the , pres- Superfamily Muricoidea Rafinesque, 1815 ence of typical Luchtbal Member like Pseuda- Family Chenu, 1859 Coralliophilidae mussium gerardi (Nyst, 1835) and of large numbers of Genus Coralliophila H. & A. Adams, 1853 Ditrupa sp. make a Luchtbal origin likely. In the North Sea Pseudomurex Monterosato, 1872 Subgenus Basin, the species is known from various Middle Miocene -37-

localities, where it is present in small numbers. In Belgium, Fig. 6/2a-b

it is present in the Zonderschot Sand Member (Berchem

coll. It to lack in the French Atlan- *1815 Buccinum reticosum 2. Formation, RM). seems Sowerby, p. 17, pi. 110, fig.

tic ‘Redonian’ and ‘Pontilevien’, but occurs in the ‘Bade- 1843 Buccinum elongatum Sowerby - Nyst, p. 572, pi. 45,

1-3. nian’ deposits of Poland (Korytnica) and Paratethyan lo- figs

1848 Nassa reticosa Sowerby - Wood, 33, pi. 3, fig. 10. calitiesin Hungaria, Austria and Romania.This distribution p.

1878 Nassa reticosa Sowerby - Nyst, pi. 2, fig. 4. seems to point to faunal exchange between the Paratethys

1881 Nassa reticosa - 22. Sowerby Nyst, p. and the North Sea Basin through Poland.

1914 Nassa reticosa - (Uzita) (Sowerby) Manner,p. 61, pi,

3, figs 4, 14-19.

1920 Nassa reticosa - (Uzita) (Sowerby) Manner, p. 517, Family Nassariidae Iredale, 1916 pi. 46, figs 1-7, 9-11. Genus Nassarius Dumeril, 1806 1946 Nassarius reticosus - 85. (Sowerby) Beets, p.

Uzita H. & A. Adams, 1853 1959a Hinia reticosa - 24. Subgenus (Uzita) Sowerby Gilbert, p.

1965 Nassarius (? Hinia) reticosus (Sowerby) - van Reg-

Altena teren et ai, p. 33, pi. 14, fig. 135 (pars).

1976 Nassarius reticosus - Adam & Nassarius (Uzita) spectabilis vandewouweri (Sowerby) Glibert, p. 56, 2, 9; 6, 6. (Glibert, 1959) pi. fig. pi. fig.

1979 Hinia reticosa - & 74, Fig. 6/5a-b (Sowerby) Geys Marquet, p. pi. 30, fig. 2.

1993 Hinia reticosa - 92. (Sowerby) Marquet, p.

1878 Nassa Lamarck - pygmaea Nyst, pi. 2, fig. 8.

1997c Nassarius (Uzita) reticosus - (Sowerby) Marquet, p.

1881 Nassa Lamarck - 26. pygmaea Nyst, p, 98, pi. 12, fig. 5. *1959a Hinia vandewouweri (Uzita) Glibert, p. 23, pi. 3, fig.

1998b Nassarius reticosus - (Uzita) (Sowerby) Marquet, p. 17. 151, unnumbered text-fig. 1976 Nassarius spectabilis vandewouweri - Adam & Gli-

7-8. bert, p. 43, pi. 2, fig. 5; pi. 6, figs

Locus — 1997 Nassarius (Uzita) spectabilis vandewouweri (Gli- typicus Holywell near Ipswich, East Anglia,

- 98. bert) Marquet, p. United Kingdom.

1998b Nassarius (Uzita) spectabilis vandewouweri (Glibert)

- unnumbered Marquet, p. 149, text-fig. Stratum typicum — Red Crag Formation, Late Pliocene.

Locus typicus — Austruweel, Antwerp, Belgium. Material— Coll.LC: 1 specimen, Antwerp, industrialarea

near Noorderlaan, dredged, presumably from the Luchtbal Stratum typicum — Luchtbal Sand Member, Lillo Forma- Sand Member. Coll. RM; 1 specimen, Doel, Deurganck- tion, Early Pliocene. dok, Luchtbal Sand Member.

Material— IRSNB 1ST 6966(leg. RM): 1 specimen, Ant- Dimensions — Height 29.0 mm, width 15.5 mm. industrial Noorderlaan, werp, area near dredged, presuma-

bly from the Luchtbal Sand Member. Coll. RM: 4 speci-

— See Description Marquet (1997c, p. 98), Marquet mens, same locality and Member. Coll RM: 3 specimens; (1998b, p. 151). The specimen figured here belongs to the Doel, Deurganckdok; Luchtbal Sand Member. Coll IRSNB form deformis, also mentionedby Wood (1848, pi. 3, fig. 1ST 5005 (holotype). 1 Of). It has five regularly coiledwhorls, but the body whorl

is oriented in oblique respect with the earlier whorls and is Dimensions— 20.5 width 11.3 Height mm, mm. furthermore angular and flat-sided.The ornament fades on

the body whorl; only growth lines and irregular plicae re- Description — see Marquet (1997c, p. 98) and Marquet main. The deformis form only represents intraspecific (1998, p. 149). variation; it is not a separate taxon. Two incisions seem to

mark of the after repairs shell, possibly attack by a crab. Discussion — The specimens figured here and in Marquet The specimen from Doel has the normal ornament for the (1998) illustrate the variability of the species. The illus- species. trated specimens have a stronger parietal tooth and the ax-

ial sculpture lies closer together than in the specimen fig-

Occurrence — The deformis form is recorded here for the ured by Marquet (1998). Typical for the species is however first timefrom Belgium. Nassarius (Uzita) reticosus (Sow- the small in always very aperture, relation to shell height. is erby, 1815) very rare in the Luchtbal Sand Member. It is

increasingly common in the Oorderen Sand Member, to Occurrence — The species is relatively more common in become a dominantmember ofthe molluscanfaunain the the Luchtbal Sand Member than in the Oorderen Sand Kruisschans Sand Member. In the Merksem Sand Member Member, in which it has been found previously in Kallo. it is rarer. Specimens from the Kattendijk Formationofthis

species referred to by Glibert (1959a) probably belong to a different, Nassarius (Uzita) reticosus (Sowerby, 1815) still unidentified species. -38-

Figure 9.

1. Adis ascaris (Turton, 1819). Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6974.

2. material, Luchtbal Sand Member? IRSNB 1ST 6975. Amydina sp. Antwerp, Noorderlaan, dredged

3. Raphitoma (R.) antonjansei Marquet, 1998. Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6976.

4. Phasianema sulcatum Wood. 1842. Antwerp, Noorderlaan, dredged material. Luchtbal Sand Member? IRSNB 1ST 6977

Wood (1848) mentioned N. reticosus was very common in Amyclina sp.

Red Formation, whilst Harmer (1920, 517) 9/2a-b the Crag p. re- Fig. corded it as rare in the stratigraphically older Coralline

Crag Formation.

Material— IRSNB 1ST 6975 (leg. RM): 1 specimen. Ant

industrial area near Noorderlaan, werp, dredged, presuma

Genus Amyclina Iredale, 1916 bly from the Luchtbal Sand Member. -39-

— Altena 171. Dimensions Height 17.5 mm, width 9.2 mm. et al, p. 42, pi, 18, fig.

* 1998a Raphitoma (Raphitoma) antonjanseni Marquet, p

— 274, pi. 3, fig. 1. Description This is a medium-sized, fusiform, thick-

1998b Raphitoma (Raphitoma) antonjanseni Marquet - shelled nassariid, relatively broad, with slightly tumid unnumbered Marquet, p. 181, text-fig. whorls and shallow suture, consisting of about seven

whorls. H/W ratio circa 0.47. Greatest widthslightly below

Locus typicus — B2 Havendok, Antwerp, Belgium. adapical end of aperture. The ornament is very weak, con-

sisting ofeight spiral ribs, which are much broader than the

Stratum typicum — Luchtbal Sand Member, Early Plio- intercostal rather and distinct spaces. They are irregular cene. the ad- and of the only at abapical parts the body whorl,

area in between is nearly smooth. The most adapical spiral Material— IRSNB 1ST 6976 (leg. RM): 1 specimen, Doel, is more deeply incised, resulting in a slight subsutural de- Deurganckdok, Luchtbal Sand Member. IRSNB 1ST 6313 pression. Axial ornament is only seen on the apertural side (holotype), 1ST 4378 (paratype), Antwerp, “Scaldisien”; of the shell; it consists ofabout seven broad ribs, distinct IRSNB: for further material see Marquet 1998a, p. 275, near the adapical suture, forming tubercles when crossing table 15. the spiral ornament; the axials fade towards the shell base.

The H of the aperture is about halfthe H ofthe shell. The

Dimensions — Height 3.99 mm, width 1.57 mm. distinct outer lip is thickened, and a narrow innerlip callus

varix is each whorl. Fifteen is developed. A present on pli-

Description — See Marquet (1998a, 274) and Marquet in the The ad- and p. cae are present outer lip. abapical plicae The of (1998b, p. 181). protoconch the juvenile specimen are most distinct. The second most abapical plica is figured in Fig. 9/3 is better preserved than the protoconch stronger developed as a blunt tooth, the adapical-most is ofthe holotype. The nucleus consists ofhalf a whorl, with smaller and lies deeper in the shell. Siphonal canal very weak, mainly spiral sculpture. The subsequent two and a short. The protoconch is not preserved in the only speci- whorls quarter protoconch have eleven strong spiral riblets, men at hand. crossed by irregular axials. The remaining protoconch

whorls also show a spiral ornament, but with predominant Discussion This species could be new, but the uncer- adapical axial ribs, while therest of the whorl is covered by tainty of its stratigraphic origin as well as the single speci- irregular prosocline and opisthocline threads. men available preclude a description. A similar species was

figured by Brebion (1964, pi. 10, figs 34-39) as Amyclina Discussion — The name antonjanssenei, which was given cf. lambertiei Peyrot, 1926 from the ‘Redonian’ of Brit- originally to this species, is incorrect, because it is derived tany, France. The original illustrations ofthe latter species from Janse; therefore, it has to be changed into antonjans- (Peyrot, 1926, pi. 3, figs 84-86) from the ‘Sallomacian’ of sei. The protoconch is multispiral, clearly different from the Aquitaine Basin (France) shows a shell with less tumid thatof Raphitoma histrix (de Christofori & Jan, 1832). whorls, shallower suture, stronger, more regular spiral or-

nament and a relatively broader callus. There is also a

Distribution — This seems to be an endemic North Sea of slight resemblance to very worn specimens Nassarius Basin Pliocene species, foundin the Kattendijk Formation (Telasco) lamellilabrus { Nyst, 1835)(see Marquet, 1997c, and in the Luchtbal, Oorderen and Kruisschans Members pi. 7, fig. 10), which has a relatively higher aperture, a nar- of the Lillo Formation. Specimens from the British Crags rower shell and less tumid whorls. with intact protoconch have not been found, so their identi-

ficationremains uncertain.

Family Conidae Fleming, 1822

Subfamily Daphnellinae Deshayes, 1863 Subclass Gray, 1840 Genus Raphitoma Bellardi, 1847 Order Fischer, 1885 Subgenus Raphitoma Bellardi, 1847 Superfamily Gray, 1840

Family AmathinidaePonder, 1988

Genus Phasianema Wood, 1842 Raphitoma (Raphitoma) antonjanssei Marquet,

1998 nom. emmend.

Fig. 9/3a-c Phasianema sulcatum Wood, 1842

Fig. 9/4a-b

1872 Pleurotoma Jan - 3. hystrix Wood, p. 41, pi. 6, fig.

1878 Pleurotoma histrix, Jan - Nyst, pi. 3, fig. 13.

1881 Pleurotoma Jan - 46. 15. histrix, Nyst, p. *1842 Phasianema sulcata Wood, p. 535, pi. 5, fig.

1915 Clathurella - Harmer, 1848 Fossarus sulcatus Wood lineolatus - hystrix (Jan) p. 240, pi. 28, figs var. Wood, p.

24-25. 121, pi. 8, fig. 23a, b, aa.

1946 histrix - 107. 1878 Fossarus sulcatus Wood - 20. Raphitoma (Jan) Beets, p. Nyst, pi. 6, fig.

1960 histrix - 18. 1881 sulcatus Wood lineolatus - Raphitoma (Jan) Gilbert, p. 17, pi. 4, fig. Fossarus var. P Nyst, p.

1965 Raphitomahystrixc (Cristofori & Jan) - van Regteren 91. -40-

1921 Fossarus sulcatus (Wood) - Harmer, 670, 53, *1792 truncatula 377. p. pi. Bruguiere, p.

1836 35. Bulla truncatula - fig. Bruguiere Philippi, p. 122, pi. 7,

1956 Phasianemasulcata Wood - 21. vanRegteren Altena, p. fig.

64. 1844 Bulla truncatula - Bruguiere Philippi, p. 96.

Phasianema - 1958a sulcatum Wood 1848 Bulla Adams - 3. Gilbert, p. 17, pi. 2, truncata Wood, p. 176, pi. 31, fig.

12. 1856 Adams - 5. fig. Bulla truncata Homes, p. 621, pi. 50, fig.

1897 Retusa truncatula - 78-79. Sacco, p. 40, pi. 3, figs

1910 Tomatina - Locus typicus — Sutton, East Anglia, United Kingdom. (Retusa) truncatula (Bruguiere) Cerulli-

Irelli. p. 34, pi. 3, figs 69-72.

1923 Utriculus truncatulus (Bruguiere) - Harmer, 794, Stratum typicum — Coralline Crag Formation, Early Plio- p,

p. 63, fig. I. cene.

1933 Tomatina - (Retusa) truncatulaBruguiere Peyrot, p.

209.pl. 12, figs 52-53. Material— IRSNB 1ST 6977 (leg. RM): 1 specimen, Ant-

1928 Tornatina truncatula - Bruguiere Friedberg, p. 544, industrial werp, area near Noorderlaan, dredged, presuma- pi. 36, figs 1-2. bly from the Luchtbal Sand Member.

1952 Retusa truncatula - (Retusa) Bruguiere Glibert, p.

393.

Dimensions — 11.3 mm, width 9.0 mm. 1960 Retusa truncatula - Height (Retusa) Bruguiere Glibert, p.

393, pi. 15, fig. 3.

— Medium-sized shell 1962 Retusa truncatula - 54. Description of about four whorls. (s.s.) Bruguiere Glibert, p.

1966 Retusa truncatula - Strausz, Protoconch heterostrophic and submerged, tonnoid-shaped, Bruguiere p. 474, pi. 74, figs 27-29. broad, with very low spire. The H of the body whorl is

1988 Retusa truncatula (Bruguiere) - Thompson, 34, about94% of H ofthe shell. Suture deep, whorls strongly p. fig. 8. inflated. Ornament consisting of 16 very broad, irregular

1990 Retusa truncatula - & (Bruguiere) Poppe Goto, p. spiral ribs, which are much broader than the shallow inter- 194, pi. 37, fig. 5. costal spaces. Umbilicus narrow, but open. Aperture high, 1992 Retusa truncatula (Bruguiere) - Cavallo & Repetto, oval, with a distinct columellar callus. Near the adapical 474. p. 166, fig, end of this callus, a blunt parietal toothoccurs. 1993 Retusa truncatula - Da (Retusa) (Bruguiere) Silva, p.

276.

— Discussion The here has - specimen figured a distinctly 2005 Retusa cf. truncatula(Bruguiere) Vandenbergheet lower spire than those figured by Glibert (1958a), Wood ai, p. 30.

and Harmer (1848) (1921). However, the shape and num- ber of axial ribs is the and it is that same, possible previous Locus typicus — illustrations represented juvenile shells. It differs clearly from Phasianema zelandica (Bloklander, 1949), figured by Stratum typicum — Recent.

Marquet (1997c, p. 110, pi. 10, fig. 7), by its fewer and broaderribs and its which is by higher aperture, more regu- Material— IRSNBIST 6970(leg RM): 1 specimen, Doel, larly oval. Phasianemazelandica has been foundin Kallo Deurganckdok, Luchtbal Sand Member. only in the Oorderen Sand Member, but according to Glib- ert it shouldbe in the Formation — (1958a) present Kattendijk Dimensions Height 2.52 mm, width 1.2 mm. and in the Luchtbal Sand Member as well. Phasianema

woodi which has been found — van Regteren Altena, 1956, Description Small, involute shell. The protoconch is only in the BritishCoralline Crag Formation, has even less involute. Teleoconch consisting of about four whorls that

ribs (8 to 10). with central constriction. The of spiral are flat-sided, a slight top

the shell is flat or slightly concave. The outer lip extends

Occurrence — Phasianema sulcatum Wood, 1842, is above the top plane. The aperture is very narrow adapi- known only from the North Sea Basin. It has been found in cally, broadening in the abapical third of the shell. Orna- the British Coralline Formationand in Crag the Kattendijk ment, including growth lines, are absent.

Formation and the Luchtbal Sand Member in Belgium.

Discussion — This differs from the species Cylichnina spe- cies found in the Pliocene of Belgium by its flat-sided,

Subclass Milne-Edwards, 1848 nearly tubular shell and by the outer lip that extends above Order Fischer, 1883 the apex. Furthermore, the shell is smooth. Only a single

Retusoidea 1931 whorl is Superfamily Thiele, visible on adult Cylichna cylindracea (Pennant,

Family Thiele, 1931 1777) specimens. Furthermore, that species is much larger

Genus Retusa Brown, 1827 and shows clear microsculpture.

Occurrence — Retusa truncatula has not been found in

Retusa truncatula (Bruguiere, 1792) Doel or Kallo previously. The species is known at least

Fig. 7/2a-c from the Middle Miocene to Recent in the Atlantic, -41 -

Figure 10.

1. Philine quadrataWood. 1839. Deurganckdok, Doel. Luchtbal Sand Member. IRSNB 1ST 6978.

2. Philine intricata Monterosato, 1884. B1-B2 Kanaaldok, Antwerp. Luchtbal Sand Member. RGM 550 005 -42-

the Mediterraneanand 1994 Philine intricata Monterosato - der the Paratethys, but North Sea Basin van Linden, p.

1-6. records start only in the Pliocene. In Belgium it has been 41, figs

1995 Philine intricata Monterosato - van der Linden, foundin Miocene Campine boring material, considered as p. 73. belonging to the “Breda Formation” (Vandenberghe et al.

2005). The species has been reported from depths between

200 Locus — 60-90 0 to m, between Norway in the north to the Canary typicus Palermo, Sicily, Italy; m depth.

Islands and the Mediterranean in the south and southeast.

Stratum typicum — Recent.

Superfamily PhilinoideaGray, 1850 Material— RGM 550 004 (leg. VDM): 2 specimens, BI-

Family PhilinidaeGray, 1850 B2 Kanaaldok, Antwerp, Luchtbal Sand Member. RGM

Genus Philine Ascanius, 1772 550 005, 1 specimen, same locality and unit.

Dimensions — Fig. 10/2b: H 1.82 mm, W 1.34 mm; fig.

Philine quadrata (Wood. 1839) 10/2a, c-g: H 2.06 mm, W 1.54 mm.

Fig. 10/la-d

Description — Tiny, very fragile, involute rhomboidal

1. *1839 Bullaea quadrataWood, p. 461, pi. 7, fig. shell, slightly rounded above, basal side nearly straight,

1848 Bullaea Wood - Wood, 179, 21, quadrata p. pi. fig. consisting of one whorl. On the umbo, a narrow ridge is

9. from present, separating an oval, smoothpart the remainder

- der 1952 Philinequadrata (Wood) van Burg, p. 52, pi. 3, ofthe shell. the umbo. Aperture very large, narrowing near fig. 11. Ornament consisting of about 50 spiral grooves. These

1960 Philine Wood - 30. quadrata Gilbert, p. 25, pi. 4, fig. consist of elongate hexagonal impressions in the central

1988 Philine - 23. quadrata (Wood) Thompson, p. 64, fig. of part the shell; near the shell base, the incisions become

- der 1995 Philine quadrata (Wood) van Linden, p. 75, and narrower the hexagonal pattern is lost. Near the umbo figs 18,22-23. the becomes ornament much more pronounced, sinuous and 1997c Philine - 115. quadrata(Wood) Marquet, p. the hexagons are only visible on part of the incisions. - 217. 1998b Philine quadrata(Wood) Marquet, p.

Discussion — Four other of Philinidaeoccur in the Locus typicus — Sutton, East Anglia, United Kingdom. species Pliocene of Antwerp. Philine scabra (Muller, 1776) is

— much than P. intricata and has Stratum typicum Coralline Crag Formation, Early Plio- larger an elongate shape. Philine is of but cene. ventrosa (Wood, 1839) comparable size,

has a rugose reticulate ornament. Philine denticulata(Ad-

1800)is smooth, devoid ofsurface Philine Material— IRSNB 1ST 6978 (leg RM): 1 specimen, Doel, ams, sculpture. is its is Deurganckdok, Luchtbal Sand Member. quadrata (Wood, 1839) larger, shape subquadrate and has it about90 rows of circular pits.

Dimensions— Height 8.13 mm, width 6.65 mm.

Occurrence — This has found fossil species been as a only

in the Pliocene Luchtbal Sand Member of as ear- — Antwerp Description See Marquet (1997c, p. 115).

lier mentioned by van der Linden (1995). Today, it has a

southern distribution:the Mediterranean, the Atlantic from Discussion — This is the first nearly complete specimen

France to the Madeira and from Doel, although its ornament is somewhat eroded. Brittany, Cape Verde, Azores, the . Usually, the spiral lines are composed ofchain-likerows of round impressions but in the materialstudied here they are reduced to simple spirals, with occasional remnants ofthe impressions. Discussion and conclusions

The fauna of the Luchtbal Sand Member con- Occurrence — Rare in the Kattendijk Formation and the gastropod

tains at least 157 The of5 Luchtbal Sand Member ofAntwerp; CoralhneCrag Forma- species (Table 1). status species (Capulus hungaricus (Linnaeus, 1758), Capulus tion in Britain. Recent distribution see Marquet (1997c, p. unguis alveolata 115). (Sowerby, 1816), Scalaspira (Sowerby, 1829),

Nucella incrassata (Sowerby, 1825) and Spinucella tetra-

from gona (Sowerby, 1823)) recorded the Luchtbal mem-

ber is uncertain: The Luchtbal Sand Philine intricata Monterosato, 1884 Member, Kattendijk

Formation and Oorderen Sand Member share 49 Fig. 10/2a-g species.

The Luchtbal Sand Member shares a further 44 species

*1884 Philine intricata 47. with the Oorderen Sand Member and 31 with the Kat- Monterosato, p.

1897 Philine desmotis Formation. have been found Watson, p. 237, fig. 5. tendijk Twenty-eight species

1905 Philine intricata Monterosato - Sykes, p. 332, fig. only in the Luchtbal Sand Member. The exact distribution -43-

of4 species is uncertain. Most ofthe 28 species, character- Doel. A. Dhondt, J. Van Goethemand K. Wouters allowed

istic ofthe Luchtbal Sand also in the Brit- Member, occur us to use the resources of the “Institut royal des Sciences ish Coralline Crag Formation. naturelles de Belgique”, especially of its collections and

SEM. F. Wesselingh allowed us to study the NNM Natu-

ralis collection. The photography is the work of J. Cilis

(SEM, IRSNB), M. Wagenaar and M. Grigis. The authors

thank L. Ceulemans, M. Grigis, S. Mermuys and G. Roest

for their help in the field and for use of their collections.

We are especially grateful to F. Wesselingh and R. Janssen

who commented critically on the text.

References

Aartsen, J.J. van, Menkhorst, H.P.M. & Gittenberger, E. 1983.

The marine Mollusca of the Bay of Algeciras, Spain, with

general notes onMitrella, Marginellidae and Turridae. Bas-

teria, Supplement 2: 1-135.

Adam, W. & Gilbert, M. 1976. Observations sur le “groupe” de

Nassarius clathratus (Bom, 1778) (Mollusca, Prosobran-

chia). Bulletin de I’Institutroyal des Sciences naturelles de

Belgique, Biologic, 51: 1-69.

Anderson, H.-J. 1964. Die Miocane Reinbek-Stufe in Nord- und

Westdeutschlandund ihre Mollusken-Fauna. Fortschritte in

der Geologic von Rheinlandund Westfalen, 14: 31-368. 11. the Luchtbal Figure Location of Sand Member on exposure Baluk, W, 1975. Lower Tortonian Gastropods from Korytnica, areal of the Doel. an photograph Deurganckdok, Poland. Part I. Palaeontologia Polonica, 32: 1-186.

Baluk, W. 1995. MiddleMiocene (Badenian) gastropods from

Korytnica, Poland. Part II. Acta Geologica Polonica, 45:

The entire mollusc fauna of the Luchtbal Sand Member 153-255.

C. 1946. The and lower includes at least 277 species: 157 gastropod, 117 bivalve, 1 Beets, Pliocene Pleistocene Gastropods

and 2 in the collections ofthe GeologicalFoundation in the Nether- amphineuran scaphopod species (Table 2). There is a lands (with some remarks on other Dutch Me- difference in the bivalves collections). large species longevity amongst dedeelingenvan de Geologische Stickling, C, IV, 1, 6: 1-66. and gastropods: 51% of bivalve species found in the Bell, A. 1870. On some new or little-known Shells of the Crag Luchtbal Sand Member are extant, whilst only 29% ofthe Formations. Annals and Magazine of Natural History, 4; gastropods occur today. As a result, the bivalve assemblage 213-217.

is more useful for actualistic reconstruction of Pliocene Bell, A. 1898. On the Pliocene shell-beds at St. Erth. Transac- environments including depth. The of the bathymetry tions ofthe Geological Society of Cornwall, 12: 111-166.

Luchtbal Sand Member estimated at 35-45 deposit was m Beu, A.G. 2004. Marine Mollusca of the of oxygen isotope stages

(2004). The numberofextant with the last 2 million in New Zealand. Part 1: by Marquet Gastropoda years Revised ge-

neric and of known bathymetric range is too small to add further infor- positions recognition warm-water and cool-water

mation. migrants. Journal of the Royal Society of New Zealand 34: ,

111-265. The longevity ofthe bivalve species (mean 10.7 Ma: Mar- Brebion, P. 1964. Les Gasteropodes du Redonien et leur signifi- quet,2004), however, makes them less useful stratigraphi- cation. Paris. 775 PhD pp. (unpublished thesis). cally than gastropod species, which survive for an average Brocchi, G. 1814. Conchiologiafossile subapennina con osser- of 1.0 Ma. However, the limited geographic rangeofmost vazione geologiche sugli Apennini & sul suola adiacente. species reduces their usefulness; 46% of the Gastropoda Tomo Secondo. 712 Milano. pp. Stamperia Reale, and 27% ofthe Bivalvia are endemic to the North Sea Ba- J.G. 1792. ordre des Bruguiere, Encyclopedic methodiqueoupar sin. Consequently, they can only be used for intra-basinal matieres. Histoire naturelle des vers, des mollusques. 2. Pan- correlations. The limited duration of species gastropods ckoecke, Paris, pp. 345-758.

to be correlated with a shift towards benthic lar- appears a Burg, W.J., van der 1952. Over enige tot nu toe uit het midden-

val life-cycle as represented in a diminished number of plioceen van Nederland onbekende molluskensoorten. Bas- protoconch whorls. teria, 16: 49-64.

Cavallo, O, & Repetto, G. 1992. Conchiglie Fossili del Roero.

Atlante Iconografico. Memorie dell'AssociazioneNaturalis-

tica Piementose, 2: 1-251. Acknowledgements Cerulli-Irelli, S. 1910. Fauna malacologica mariana. Parte 4.

Paleontographia Italica. Memorie dipaleontologia, 16: 23- We are indebtedto the “Ministerie van de Vlaamse greatly 70. Gemeenschap, Afdeling Zeeschelde” and the staffmembers Cerulli-Irelli, S. 1914. Fauna malacologica mariana. Parte 6. ofthe T.V. Deurganckdok for their to work at permission Paleontographia Italica. Memorie dipaleontologia 18: 141- -44-

170. Glibert, M. 1959b. Pelecypodes du Diestien, du Scaldisien et du

Cerulli-Irelli, S. 1916. Fauna malacologica mariana. Parte 8. Merxemien de la Belgique. Quatriemenote (Fin). Bulletin de

Paleontographia Italica. Memorie di paleontologia, 22: 71- Tlnstitut royal des Sciences naturelles de Belgique, 35: 1-24.

220. Glibert, M. 1960. Gastropodes du Diestien, du Scaldisien et du

Chilli, C. 2000. MalacofaunaPliocenica Toscana; vol. 2; Super- Merxemien de la Belgique. Quatrieme note. Bulletin de

Muricoidea. C. 142 I'Institut Sciences naturelles de 36:1 -44. famiglia Chirli, Firenze, pp. royal des Belgique,

Cossmann, M. & Peyrot, A. 1916. Conchologie neogenique de Glibert, M. 1962. Les et fossiles du Ceno-

1’Aquitaine.Actes de la Societe Linneenne de Bordeaux, 69: zoique etranger des collections de ITnstitut royal des Sci-

157-365. ences naturelles de Belgique. Institut royal des Sciences

Costa, O.G. 1829. Catalogo sistematico e ragionata del testacei naturelles de Belgique. Memoires, 2 serie, 70: 1-140.

delle Due Sicilie. Tipografia della Minverva, Napoli, 8+ 132 Glibert, M. 1963. Les Mesogastropoda fossiles du Cenozoique

collections 1’Institut pp. etranger des de royal des Sciences

Defiance, J.L.M. 1804-1845. Dictionnaire des Sciences naturelles naturelles de Belgique. Deuxieme partie. Fossariidae a Fici-

dans lequel on traite methodiquement les differentes etres de dae (inclus). Institut royal des Sciences naturelles de Bel-

la nature. Paris, 61 volumes. gique. Memoires. 2 serie, 73: 1-154.

Deshayes, G.P. 1861. Description des animaux sans vertebres Gonzalez-Delgado, J.A. 1985. Estudio systematico de los Gas-

bassin de Paris servir de del de de decouverts dans le pour supplement teropodos Pliocene Huelva (SW Espana). I: Ar-

d la description des coquilles fossiles des environs de Paris chaeogastropoda. Studio GeologicaSalamanticensia, 20: 45-

77. comprenant une revue generate de toutes les especes ac-

tuellement Tome Deuxieme. Paris. 1 - A. 1988. Molluscs: and connues. Bailliere, Pp. Graham, prosobranch pyratnidellid gas-

432. tropods: keys and notes for the identification of the species.

Eichwald, E. 1853. Lethaea Rossica ouPaleontologie de la Rus- Synopses of the British Fauna (n.s.), 2e ed. The Linnean So-

sie. Demiere London The periode. Schweizerbart, Stuttgart: 533 pp. ciety of & Estuarine and Brackish-Water Sci-

Fekih, M. 1975.Paleoecologie du Pliocene marin au Nord de la encesAssociation, E.J. Brill/Dr. W. Backhuys, Leiden. 662

Tunisie. Annales des Mines de 1-195. et Geologic, 27: pp.

W. 1928. Mollusca miocenica Poloniae. Pars I J. 1877. de Friedberg, (Gas- Granata, Grillo Description quelques especes nou-

Museum Imienica velles S. 15 tropoda et Scaphopoda). Dzieduszyckich oupeu connues. Marchese, Napoli. pp.

5:441-361. A. de 1889, Esame di Gregorio, taluni Molluschi vivente e

Geys, J. & Marquet, R. 1979. Veldatlas van de Cenozoische fos- terziari del Bacino Mediterraneo. Naturalista Siciliano, 8:

sielen van Belgie. Deel 1. Neogeen. Publicatie van de Belgi- 275-292.

sche Vereniging voor Paleontologie, 2: 1-123. Harmer, F.W. 1914-1918. The Pliocene mollusca of Great Brit-

Giannuzzi-Savelli, R., Pusateri, F., Palmed,A. & Ebreo, C, 1994, ain, being supplementary to S. V. Wood's Monograph of the

Atlante delle conchiglie marine del Mediterraneo. Vol. I (Ar- Crag Mollusca. Volume I. Monographs of the Palaeonto-

La Rome. 125 London. 1-200 201-302 chaeogastropoda). Conchiglia, pp. graphical Society, Pp. (1914), pp.

A. & C. 1999. 303-461 Giannuzzi-Savelli, R., Pusateri. F„ Palmed, Ebreo, (1915). pp. (1918).

Atlante delle conchiglie marine del Mediterraneo. Vol. III. Harmer, F.W. 1920-1925. The Pliocene mollusca ofGreat Brit-

Rome. 128 S.V. Wood’s of Evolver, pp, ain, being supplementary to Monograph the

M, 1949. Miocene Gilbert, Gasteropodes du moyen du Bassin de Crag Mollusca. Volume II. Monographs of the Palaeonto-

la Loire. Premiere Partie. Institut des Sciences London. 653-704 royal graphical Society, Pp, 485-652(1920), pp.

naturelles de Memoires, 2 s6rie, 30: 1-240. 705-856 Belgique. (1921), pp. (1923). pp. 857-900 (1925).

M. 1952. du Miocene du Bassin de J. de 1950a. Gilbert, Gasteropodes moyen Heinzelin, Stratigraphic pliocene et Quaternaire

la Loire. Deuxieme Partie. Institut royal des Sciences observee au Rruisschans. I. - Analyse stratigraphique. Bulle-

naturelles de Belgique. Memoires, 2 serie, 46: 1-450. tin de Tlnstitut royal des Sciences naturelles de Belgique, 25:

Gilbert, M. 1957a.Pelecypodes du Diestien, du Scaldisien et du 1-38.

Merxemien de la Belgique. Premiere note. Bulletin de Heinzelin, J. de 1950b. Stratigraphic pliocene et Quaternaire

I'Institut royal des Sciences naturelles de Belgique, 33: 1 -40, observee auKruisschans. II. - Conclusions. Bulletin de T In-

Gilbert, M. 1957b. Gastropodes du Diestien, du Scaldisien et du stitut royal des Sciences naturelles de Belgique, 25: 1-22.

Merxemien de la Belgique. Premiere note. Bulletin de Heinzelin, J. de 1955a. Deuxieme serie d’observations stratigrap-

Tlnstitut royal des Sciences naturelles de Belgique, 33: 1-27. hiques au Kruisschans. Coupes de Fecluse Baudouin. I. -

Gilbert, M. 1957c. Pelecypodes du Diestien, du Scaldisien et du Analyse stratigraphique. Bulletin de Tlnstitut royal des Sci-

Merxemien de la Belgique. Deuxieme note. Bulletin de encesnaturelles de Belgique, 31: 1-29.

Tlnstitut royal des Sciences naturelles de Belgique, 33: 1-28, Heinzelin, J. de 1955b. Deuxieme sdried’observations stratigrap-

Glibert, M. 1958a. Gastropodes du Diestien, du Scaldisien et du hiques au Kruisschans. Coupes de Fecluse Baudouin. II. -

Merxemien de la Belgique. Deuxieme note. Bulletin de Conclusions. Bulletin de Tlnstitut royal des Sciences naturel-

Tlnstitut royal des Sciences naturelles de Belgique, 34: 1-36. les de Belgique, 31: 1-14.

Glibert, M. 1958b. Tableau stratigraphique des mollusques du Heinzelin, J. de 1955c. Considerations nouvelles sur le Neogene

Neogene de la Belgique. Bulletin de Tlnstitut royal des Sci- de FOuest de FEurope. Bulletin de la Societe beige de Geo-

ences naturelles de Belgique, 34: 1-19. logic, 64: 463-476.

Glibert, M. 1958c. Pelecypodes du Diestien. du Scaldisien et du Homes, M. 1851-1870. Die fossilen Mollusken des Tertiar-

Merxemien de la Belgique. Troisieme note. Bulletin de Beckens vonWien, Abhandlungender kaiserlichen und kd-

Tlnstitut royal des Sciences naturelles de Belgique, 34: 1-27. niglichen Geologischen Reichsanstalt, 3: 209-296 (1853),

Glibert, M. 1959a. Gastropodes du Diestien, du Scaldisien et du 461-736(1856).

Merxemien de la Belgique. Troisieme note. Bulletin de Janssen, A.W. 1969. Beitrage zur Kenntnis des Miozans von

Tlnstitut royal des Sciences naturelles de Belgique, 35: 1-36. Dingden und seiner Molluskenfauna 2. Geologica etPaldon- -45 -

tologica, 3: 153-193. Marquet, R. 1997c. Pliocene Gastropodfaunas from Kallo (Oost-

A.W. uit het Mioceen Janssen, 1984.Mollusken - van Winterswijk- Vlaanderen,Belgium) Part 3. Caenogastropoda: Aporrhai-

Miste. Een inventarisatie met beschrijving en afbeeldingen dae to , and Part 4. to Helicidae. Con-

alle van aangetroffen soorten. Koninklijke Nederlandse Na- tributions to Tertiary and Quaternary Geology, 34; 69-149. tuurhistorische Vereniging, NederlandseGeologische Vere- Marquet, R. 1998a. The Pliocene turrid Gastropods of Belgium.

& niging Rijksmuseum van Geologic en Mineralogie, Am- Part 2: Conidae (genera Asthenotoma, Comarmondia, Cy-

451 sterdam. pp. tharella, Mangelia, Lusitanops, Raphitoma and Philbertia).

Jeffreys, J.G. 1874. Someremarks onthe mollusca ofthe Medi- Bulletin de l’Institutroyal des Sciences naturelles de Belgi-

terranean. the the British As- Sciences de la 68: Report of fortythird meetingof que, Terre, 263-287.

sociation/or f/ieAdvancement 1873: 111-116. o/Science, Marquet, R. 1998b. De Pliocene gastropodenfauna van Kallo

Jeffreys, J.G. 1883. On the mollusca the procured during “Light- (Oost-Vlaanderen, Belgie). Publicatievan de Belgische Ver-

and ning” “Porcupine” Expeditions 1868-1870 (Part VI). eniging voor Paleontologie, 17: 1-246.

the London, Proceedings of ZoologicalSociety of 1883: 88- Marquet, R. 2001. A study of some Neogene European species of

115. Seilinae (, Gastropoda). Bulletin de Tlnstitut

B. 1960. Les de Kojumdgieva, E.M. & Strachimirov, fossiles royal des Sciences naturelles de Belgique, Sciences de la

Bulgarie VII; Tortonien. Academic des Sciences deBulgarie, Terre. 71: 195-208.

Sofia, 317 pp. [in Bulgarian]. Marquet, R. 2002. The NeogeneAmphineuraand Bivalvia (Pro- Landau, B., R. & M. 2003. The Marquet, Grigis, Early Pliocene tobranchia and Pteriomorphia) from Kallo and Doel (Oost-

Gastropoda (Mollusca) ofEstepona, Southern Spain. Part 1: Vlaanderen,Belgium). Palaeontos, 2: 1-99.

1-87. Vetigastropoda. Palaeontos, 3: Marquet, R. 2004. Ecology and evolution of Pliocene bivalves

Landau, B., R. & M. 2004. The Pliocene Marquet, Grigis, Early from the Antwerp Basin. Bulletin de Tlnstitut royal des Sci- (Mollusca) of Southern Part 2: Gastropoda Estepona, Spain. encesnaturelles de Belgique, Sciences de la Terre, 74: 205-

Orthogastropoda, Neotaenioglossa. Palaeontos, 4: 1-108. 212.

Leriche, M. 1912. Le Neogene des environs d’Anvers. Reunion Marquet, R. 2005. The Neogene Bivalvia (Heterodonta and extraordinaire du 27 aout au 6 septembre. Bulletin de la Soci- Anomalodesmata) and Scaphopoda from Kallo and Doel ete' Geologiquede France 4e serie, 12: 725-727. , (Oost-Vlaanderen, Belgium). Palaeontos, 5: 1-142. J. der 1994. Philine intricata Linden, van Monterosato, 1884,an Meuter, F.J. de & P.G, 1976. and bio- overlooked Laga, Lithostratigraphy species from the North-East Atlantic and the stratigraphy based on benthic ofthe Neogene (Gastropoda, Opisthobranchia: Philini- deposits ofnorthernBelgium. Bulletin de la Societe beige de dae). Basteria, 58: 41-48. Geologic, 85: 133-152. Linden, J. van der 1995. Philinidae dredged by the CANCAP G. 1803. Testacea Montagu, Brittanica, or natural history of expeditions (Gastropoda,Opisthobranchia). Basteria, 59: 65- British shells, marine, land and fresh-water, the 84. including

most minute: systematically arranged and embellishedwith Linnaeus, C. 1758. Systema naturae. Editio decima, reformata. figures. Romsey, London, 37+606 pp. Tomus I. direct. Laurentii Impensis Salvii, Holmiae, 376 pp. Monterosato, T. di 1884. Nomenclatura generica e specifica di Malatesta, A. 1960. Malacofaunapleistocenica di Grammichele alcune 152 conchiglie Mediterranee. Palermo. pp, Memorie Service alia Carta (Sicilia). per Geologica d’Italia, Nyst, P.H. 1843. Description des coquilles et des polypiers fos- 12: 1-196. siles des terrains tertiaires de la Belgique. Memoires couron- Malatesta, A. 1974. Umbra. Memorie Malacofaunapliocenica per nes de TAcademic royale de Bruxelles, 17: 1-676. Service alia Carta Geologica d'Italia, 13; 1-498. Nyst, P.H. 1878-1881.Conchyliologie des terrains Tertiaires de V. & Manganelli. G., Spadini. Cianfanelli, S. 2004. The xeno- la Belgique. Annales du Musee royal d’Histoirenaturelle de phorid gastropods of the MediterraneanPliocene: the record Belgique. 3: 1-262 (1881), pis. 1-28 (1878). of the Siena Basin. Bollettino della Societd Paleontologica Palazzi, S. & Villari, A. 1996. Malacofauna batiale plio- Italica, 43: 409-451. pleistoceniche dell Messinese 2. Capo Milazzo. Naturalista Marquet, R. 1993. The molluscan fauna of the Kruisschans Siciliana, IV, 30: 237-279. Member (Lillo Formation,Late Pliocene) in the Antwerp area Peyrot, A. 1926.Conchologie neogeniquede TAquitaine(Suite). (Belgium). Contributions to Tertiary and Quaternary Geol- Actes de la Societe Linneenne de Bordeaux, 78: 199-256. 30: 83-104. ogy, Peyrot, A. 1933. Conchologie neogeniquede TAquitaine. Actes Marquet, R. 1995. Pliocene Gastropod faunas from Kallo (Oost- de la Societe Linneenne de Bordeaux, 84: 129-288. Vlaanderen,Belgium) - Part 1. Introduction and Archaegas- Philippi, R.A. 1836. Enumeratio molluscorum Siciliae cum tropoda. Contributions to Tertiary and Quaternary Geology, viventium in tellure turn tertiari in itinere suo 32: 53-85. fossilum, quae

S. Berolini. 267 observavit. Schroppii, pp. Marquet, R. 1996. The family Triphoridae in the Neogene of R.A. Enumeratio molluscorum Siciliae Philippi, 1844. cum Belgium (Mollusca, Gastropoda). Bulletin de Tlnstitut royal viventium in tertiari in itinere turn tellure fossilum, quae suo des Sciences naturelles de Belgique, Sciences de la Terre, 66: observavit. Vol. Secunda. E. Anton, Halis Saxonum: 303 137-149, pp.

P. 1984. Revisione del Piani, genere EmarginulaLamarck, 1801 Marquet, R. 1997a. The Pliocene turrid Gastropods ofBelgium. in Mediterraneo.Atti Simposio: sistematicadeiProsobranchi Part 1: Drillidae, Turridae,Conidae (genusBela). Bulletin de del 1982. Mediterraneo, Lavori della Societd Italiana di Ma- Tlnstitutroyal des Sciences naturelles deBelgique, Sciences lacologia, 21: 193-238. de la Terre, 67: 119-151. Pinna, G. & Spezia, L. 1978. Catalogo dei tipi del Museo Civico Marquet, R. 1997b.Pliocene Gastropod faunas from Kallo (Oost- di Storia Naturale di Milano; I. I tipi di Gastropodi fossili. Vlaanderen,Belgium) - Part 2. Caenogastropoda: Potamidi- Atti de la Societd Italiana de Scienze naturale e Museo Civ- dae Tomidae. to Contributions to Tertiary and Quaternary ico di Storia Naturale di Milano, 119: 125-180. Geology, 34: 9-29. Ponder, W.F. 1983. Xenophoridae ofthe World. The Australian -46-

Museum Memoir, 17: 1-121. continental record in the Maaseik well (49W0220) in the

Poppe. G.T. & Goto, Y. 1990. European Seashells. Volume I Roer ValleyGraben, NE Belgium, Memoirs of the Geological

(Polyplacophora, Caudofoveata, Solenogastra, Gastropoda}. Survey ofBelgium, 52: 1-39.

Wiesbaden. B, Christa Hemmen Verlag, 352 pp. Vliet-Lanoe, van,Vandenberghe,N., Laurent, M.,Laignel, B.,

CO. 1956. in the Regteren Altena, van The genus Phasianema Lauriat-Rage, A., Louwey, S., Mansy, J.-L., Mercier, D., Hal-

Pliocene ofWestern Europe. Basteria, 20: 62-64. legouet, B., Laga, P., Laquement, F., Meilliez, F., Michel,Y„

Regteren Altena, C.O. van, Bloklander, A. & Pouderoyen, L.P. Moguedet, G. & Vidier, J.-P. 2002. Palaeogeographic evolu-

1965. Defossiele schelpen van de Nederlandse stranden en tion of northwestern Europe during the Upper Cenozoic.

druk. Nederlandse 24: 511-541. zeegaten. Eerste deel, tweede Malacologi- Geodiversitas,

sche Lisse. 55 R. The Nortwest Basin. Vereniging, pp. Vincken, (ed.) 1988. European Tertiary

Rolan Mosquera, E. 1983. Moluscos de la Ria di Vigo. Gastero- Geologisches Jahrbuch, Reihe A, 100: 1-508.

de 383 J.H. 1944. Miozane dem podos. Velograf, Santiago Compostella, pp. Voorthuysen, van Gastropoden aus

Rossi Ronchetti, C. 1955. I tipi della “Conciliologie fossile Peelgebiet (Niederlande) (Rissoidae-Muricidae, nach Zittel’s

subapennina” di G. Brocchi, Rivista Italiana di Paleontolo- Einteilung 1924). Mededeelingenvan de Geologische Sticht-

gia e Stratigrafla, 5: 91-259. ing, C. IV, 1,5: 1-116.

Sabelli, B., Giannuzzi-Savelli, R. & Bedulli,B. 1990. Annotated Waren, A. 1980. Marine Mollusca described by John Gwyn Jef-

Vol. Edizioni with the location of the material. check-listof Mediteranean marine Mollusks. I. freys, type Conchological

Libreria Naturalistica Great Britain Bolognese, Bologna: 348 pp. Society of and Ireland,Special Publication, 1:

Sacco, F. 1895. / Molluschi dei terreni terziari del Piemonte e 1-60.

dellaLiguria. Vol. 18. Memorie della Reale Accademia delle Waren, A, 1991. New and little known mollusca from Iceland

Scienze, Carlo Ctaussen, Torino: 1-51. and Scandinavia. Sarsia, 76: 53-124.

Sacco, F. 1897. I Molluschi dei terreni terziari del Piemonte e Waren, A. 1992. New and little known “Skeneimorph” gastro-

della Liguria. Vol. 22. Memorie della Reale Accademia delle pods from the Mediterranean Sea and the adjacent Atlantic

Scienze, Carlo Claussen, Torino: 1-130. . Bollettino Malacologico, 28; 149-248.

Sacco, F. 1904. I Molluschi dei terreni terziari del Piemonte e Waren, A. 1996. Mollusca from Iceland and Scandinavia. Part 3.

della Liguria. Parte XXX. (Aggiunte e correzioni). Memorie Sarsia, 81; 197-245.

della Reale Accademia delle Scienze, Carlo Claussen, Torino: Watson, R.B. 1897. On the marine mollusca ofMadeira. Journal

1-203. of the Linnean Society ofLondon, Zoology, 26: 233-327.

Silva, Marques, C.A.P.F. da 1990. Moluscos pliocenicos da Wienrich, G. 2001. Die Pauna des marinen Miozdns von Keve-

regiao de Caldas da Rainha, Marinha Grande. Pombal (Por- laer (Niederrhein). Band 3. Gastropoda bis .

386-639. tugal). I. . Fissurellidae. Publicacoes Oca- Backhuys, Leiden, pp.

sionais da Sociedada Portuguesa de Malacologia, 15; 1-10. Wood, S.V. 1839. On the fossil shells from the Crag. The Maga-

Silva, Marques, C.A.P.F. da 1993. Gaste'ropodes Pliocenicos zine of Natural History, New Series, 3: 460-465.

Marinhos de Vale de Lreixo (Pombal, Portugal). Sistemdtica, Wood, S.V. 1842. A Catalogue of Shells from the Crag. The

Tafonomia, Paleoecologia. Unpublished MSc thesis, Magazine of Natural History. New Series, 9: 455-462 and

Faculdade de Ciencias da de 312 Universidade Lisboa. pp. 527-544.

Silva, Marques, C.A.P.F. da 2001. Gasteropodes Pliocenicos Wood, S.V. 1848. A monograph of the Crag mollusca, or, de-

Marinhos dePortugal: Sistemdtica, Paleoecologia, Paleobi- scription of shellsfrom the Middle and Upper Tertiaries of

ologia, Paleogeografia. Unpublished PhD thesis, Faculdade the east ofEngland. Part I. Univalves. The Palaeontographi-

de Ciencias da Universidade de Lisbon. cal 1-12 1-208 Liboa, 747 pp. Society, London. + pp.

Sowerby, J. 1816-1818. The mineral conchology of Great Brit- Wood, S. V. 1851-1860.A monograph of the Crag mollusca, or,

ain; or colouredfigures and descriptions of those remains of description of shellsfrom the Middle and Upper Tertiaries of

2. Bivalves. testaceous or Shells which have been preserved at the eastofEngland. Part The Palaeontographical

1-12 various times and depths in the earth. Part 2. Benjamin Me- Society, London, pp. 1-150, pis. (1851), pp. 151-216,

London. 251 13-20 217-342, 21-31 note 1-2 ridith, pp. pis. (1853), pp. pis. (1857),

Sowerby, J. de C. 1829. The mineral conchology ofGreat Brit- (1861).

ain; or colouredfigures and descriptions ofthose remains of Wood, S.V. 1872. Supplement to the monograph of the Crag

testaceous animals or Shells which have been preserved at mollusca, with description ofshellsfrom the UpperTertiaries

various times and depths in the earth. Part 6. Richard Taylor, of the east of England. Volume 3. Univalves and Bivalves.

The 1-31 1-99 London. 250 pp. Palaeontographical Society, London, + pp.

Strausz, L. 1966. Die miozdn-mediterranen Gastropoden Un- Wood, S.V. 1879. Second supplementto the monograph of the

Akademiai Kiado, 692 mollusca, with shells the gams. Budapest. pp. Crag description of from Upper

Sykes, E.R. 1905. On the Mollusca procured during the Porcu- Tertiaries of the east of England. Volume 4. The Palaeonto-

London. 58 pine expeditions. Supplementary note. Part II.Proceedings of graphical Society, pp.

the Malacological Society ofLondon. 6: 322-332. Wood, S.V. 1882. Third supplement to the monograph of the

Thompson, T.E. 1988. Molluscs: Benthic Opisthobranchs (Mol- Crag mollusca, with description of shells from the Upper

lusca: Gastropoda). Synopses of the British fauna: 1-355. Tertiaries ofthe east ofEngland. Volume 5. The Palaeonto-

J, & London. 24 Vandenberghe,N., Laga,P., Steurbaut, E., Hardenbol, Vail, graphical Society, pp.

1998. the southern P.R. Tertiary sequence stratigraphy at

border of the North Sea Basin in Belgium. SEPM Special

Publication, 60: 119-154.

Vandenberghe, N., Laga, P., Louwye, S., Vanhoome, R., Mar-

quet, R., De Meuter, F., Wouters, K. & Hagemans, H.W.

2005. Stratigraphic interpretation of the Neogene marine- -47-

Species <1960<1960 Doel Noord Dreg KFm OSM Occ.

Emarginulafissura reticulata Sowerby, 1813 xX cc xX xX xX xX R

Emarginula adriatica Costa, 18291 829 vr - - R

Emarginulaplioaspera Sacco, 1897 vr xX -

? - - Emarginula sp. 7 vr

crassalta 1874 - R Emarginula crassa Wood, Xx r xX

Diodora - - R graecagraeca (Linnaeus, 1758) r

Diodora italica Defrance, 1820 Xx xX xX xX R

Lepeta scaldensis van Regteren Altena, 1954 Xx r xX xX - xX N

Dikoleps cutleriana (Clark, 1848) vr - - R

Lodderenasphaeroideum (Wood, 1842) vr - Xx N

Cirsonella romettensis (Granata, 1877) vr - xX R

Skeneidae indet. vr - -

Margarites belli (Manner, 1923) xX r - - N

Solariella maculata Wood, 1842 xX c xX Xx xX - N

Calliostomazizyphium (Linnaeus, 1758) Xx r xX - RR

Calliostoma kickxi (Nyst, 1835) r xX - N

Calliostoma simile (Sowerby, 1818) xX xX Xx - xX N

Calliostomaoccidentale (Mighels & Adams, 1842) xX c xX - R

Gibbula cinecineroideswide s (Wood, 1842) xX xX Xx -- -- N

Gibbulaoctosulcata (Nyst, 1835) xX r Xx Xx Xx N

- Gibbula beetsi van Regteren Altena, 1954 xX vr Xx - Xx N

Gibbula crassistriata (Bell in Wood, 1882) vr - Xx N

Gibbula obconica nehalenniaevanvan Regteren Altena, 19541 x c xX Xx -- Xx N

Gibbula woodi (Manner, 1923) Xx xX Xx -- -- N

Tenagodusobtusus (Schumacher, 1817)1817) r Xx -- R

- - Bittium rubanocinctum Gilbert, 19581958 xX Xx - - N

Turritella incrassata incrassata Sowerby, 1814 Xx VCvc Xx Xx Xx Xx

Cingula inusitata (Beets, 1946) Xx c Xx Xx N

Rissoa obsoleta Wood, 1842 xX Xx Xx Xx N

Alvania beani (Hanley in Thorpe, 1844)1844) xX xX Xx R

Alvania whitleyi (Bell, 1898) X ?7

- - Alvania zetlandicazetlandica(Montagu, 1815) vr - - R

- (Wood, 1857) vrvr - xX

Circulus striatusstriatus (Philippi, 1836) Xx xX - R

Circulus supranitidus (Wood, 1842) XX - X

Caecum mamillatumWood, 1842 Xx VCvc - Xx N

Caecum glabrum (Montagu, 1803) vr - Xx R

Tornus belgicus (Gilbert,(Glibert, 1949) xX xX - xX

Aporrhais scaldensis van Regteren Altena, 1954 Xx r Xx Xx Xx N

Aporrhais pespelecani (Linnaeus, 1758) Xx Xx - Xx R

Trichotropis borealis Broderip & Sowerby, 1823 xX r - - R

Calyptraea chinensis (Linnaeus, 1758) xX cc Xx Xx Xx R

Capulus ungaricus (Linnaeus,(Linnaeus, 1758) ?7 7? 7? Xx

Capulus unguis (Sowerby, 1816)1816) ?7 XxxX

doelensis - - N Capulus nov. sp. xX c

Xenophora scaldensis Gilbert,Glibert, 1958 xX vr xX xX - N

Petaloconchus glomeratus (Linnaeus, 1758) xX VCvc xX xX R

Bivonia triqueter {(Bivona,Bivona, 1832) xX c xX xX R

Velutina virgatavirgata(Wood, 1848) xX - - N

- Neosimnia leathesia scaldisia Schilder, 1933 Xx - Xx N

Trivia retusa (Sowerby, 1832) Xx Xx - - N

Trivia tesludinella (Wood, 1842)1842) Xx Xx xX - - N

Trivia c. coccinelloidescoccinelloides(Sowerby, 1823) Xx c Xx xX - xX N

Erato 18941 894 - pernanapemana Sacco, xX xX

Erato cypraeola brittanica Schilder, 1933 X XXX X

Euspira catena (da Costa, 1778) xX vcVC xX xX xX Xx R

Euspira cirriformis gottschei (Kautsky,(Kautsky, 1925) xX r xX - N

Euspira hemiclausa (Sowerby, 1824) cC XXXX

- Euspira exvarians (Sacco, 1891) X - X

Natica crassa Nyst, 1843 Xx VCvc Xx xX Xx N

Galeodea bicatenata (Sowerby, 1813) xX vr xX xxX X -48-

Ficus conditus subintermedius (d’Orbigny, 1852) Xx vr Xxx-X -

Cerithiopsis barleei (Jeffreys, 1887) Xx r Xx - R

Krachia zelandica ( Beets,Beets, 1946) vr Xx Xx - N

Cerithiopsis subulata (Wood, 1848) Xx XxxX

Seila suttonensis Marquet, 2001 Xx c Xx - N

LaiocochlisLaiocochlis woodi vanvan Regteren Altena, 1954 xX r Xx - - N

Marshallora adversa (Montagu, 1803) xX r xX xX xX RR

Obesula scaldensis Marquet, 1996 xX xX - N

IndiaIndia vandermarkiMarquet, 19961996 xX - - N

Adis ascarisascaris (Turton, 1822) vr xX xX RR

Amaea woodi (Deshayes, 1861) xX vr - -

7 - Acirsa penepolaris (Wood, 1872) xX r ? N

Opalia obtusicostata (Wood, 1848) xX vr - - N

Cirsotrema fimbriosumfimbriosumfimbriosum (Wood, 1842)1842) xX vr xx-X X -

Cirsotrema funiculus (Wood, 1872) Xx vr ?7 7?

Epitoniumf. frondiculum (Wood, 1842) X Cc X - X

EpitoniumEpitoniumfrondosumfrondosum (Sowerby, 1829) r - --

Epitonium subulata (Sowerby, 1823) Xx c Xx --

Epitoniumfoliaceum (Sowerby, 1825) xX c xX - xX

Epitonium clathratulum minuta (Sowerby, 1823) c - xX R

Epitonium hennei (Nyst,(Nyst, 1871)1871) xX xX -- N

Eulimaglabra (Da Costa, 1778)1778) xX r xX Xx R

- BalcisBalds albaalba (Da Costa, 1778) xX r xX xX - R

Polygireulima glabella (Wood, 1848)1848) xX

Pterynotus canhami (Wood, 1872) xX xX - Xx N

Pterynopsis binominatabinominata(Staadt(Staadt in Cossmann, 1919)1919) xX xX Xx N

Trophon muricatus (Montagu, 1803) xX c xX xX xX Xx R

Scalaspira alveolata (Sowerby, 1829) 7? - --

Nucella incrassataincrassata (Sowerby, 1825) 7? - --

Spinucella tetragona (Sowerby, 1823) ?7 - XX

Coralliophila bracteata (Brocchi,(Brocchi, 1814) X - --

Atractodon elegans (Charlesworth, 1837) rr xX xX Xx N

Buccinum undatumundatum(Linnaeus,(Linnaeus, 1758) xX - Xx R

Liomesus dalei (Sowerby, 1825) Xx rr xX Xx N

Coins cordatus (Bell,(Bell, 1871) Xx - Xx

Fusinus cf. lamellosus (Borson, 1821) X X --

Amyclina labiosalabiosa (Sowerby,(Sowerby, 1825) Xx rr - xX

vr - - Amyclina spec.spec. vr

Nassarus consociatus (Wood, 1848) Xx rr xxX X

Nassarius dollfusi (Harmer, 1914) Xx xX - Xx N

Nassarius lamellilabrus(Nyst, 1835) Xx rr - xX N

Nassarius ligustica (Bellardi, 1871) Xx - -

Nassarius scaldisianusscaldisianus Adam & Gilbert, 1976 Xx -- xX N

Nassarius spectabilisspectabilis vandewouweri (Glibert, 1959)1959) XX X - X

Nassarius reticosus (J. Sowerby, 1815)1815) XX X - X

Pterygia sulculatasulculata (Wood, 1878)1878) Xx rr Xx - - N

Mitrella scaldensis (van(van Regteren Altena, 1956) Xx xX xX xX xX N

- Vexillum ebenus (Lamarck, 1811) Xx Xx - R

Scaphella lambertilamberti (Sowerby, 1816) Xx rr xX XxxX

N Cancellaria harmeri Glibert, 1960 Xx vr xX xX

Sveltia jonkaireana(Nyst, 1835) Xx c xX Xx xX N

Babylonella wouweri (Harmer, 1916) xX xX xX - - N

Babylonella venerandaveneranda (Beets, 1946)1946) Xx xX Xx - - N

Babylonella gracilentagracilenta (Wood, 1872)1872) Xx r Xx xX xX xX N

Brocchinia mitraeformis parvula (Beyrich, 1856) X X -

- Gemmula antwerpiensis (Vincent,(Vincent, 1890) xX vr xX xX - xX N

Fusiturris porrecta (Wood, 1848) xX - - N

Acamptogenotia intortaintorta nysti Glibert, 1954 X XXX-X X X -

Cerodrillia nysti (Harmer, 1815) xX r xX Xx xX xX N

Spirotropis c. confusa (Seguenza, 1880) vr xxX X

Asthenotoma ornataomata (Defiance, 1826) xX r xX - xX

Cytharella substriolata (Harmer, 1915) xX r Xx xX xX xX N

Cytharella vandewouweri (Glibert, 1960) xX - xX N -49-

- Bela belgica (van Regteren Altena, 1959) xX r xX xX - Xx N

Bela gliberti Marquet, 1997 xX xX xX xX N

N Bela antwerpiensis Marquet. 1997 Xx r Xx Xx Xx N

Bela tenuistriata (Bell, 1871) xX r Xx Xx Xx Xx N

Bela consimilis (Harmer, 1915) Xx Xx xX - Xx N

- Mangelia gracilior (Bell, 1871) xX - xX N

Comarmondia gracilis (Montagu, 1803) xX r xX - xX R

Philbertia pseudoleufroyi Marquet, 1998 Xx xX xX - xX N

RaphitomaRaphiioma antonjanssei Marquet, 1998 xX r xX xX N

Raphitoma perpulchra (Wood, 1848) Xx c Xx Xx - Xx N

Terebra i. inversa (Nyst, 1845) Xx c Xx - Xx N

Terebra canalis Wood, 1842 r -- xX N

Pyramidella laeviuscula Wood, 1842 rr Xx -

Chrysallida stefanisi (Jeffreys, 1869) Xx r Xx - R

Chrysallida indistincta (Montagu, 1808) vr Xx - Xx R

Eulimella acicula (Philippi, 1836) xX Xx xX R

Odostomia (auct ? Brocchi, 1814) xX xX Xxx?X ?

- Brachystomia eulimoides (Hanley, 1844) vr - xX R

Turbonilla internodulaintemodula(Wood, 1848) xX r xX xX R

Turbonilla filosa Wood, 1842 xX - - N

Turbonilla senistriata (Wood, 1879) Xx r - Xx N

Turbonilla kendalli Bell in Harmer, 1920 Xx Xx -

- - Phasianemasulcata Wood, 1842 Xx Xx - - N

Phasianema zelandica (Bloklander, 1949) Xx - Xx N

Acteon noae Sowerby, 1822 xX xX xX xX N

Acteon tomatilis (Linnaeus, 1758) xX r xX - R

Acteon subulatus Wood, 1842 xX r Xx - N

Acteon levidensis Wood, 1842 Xx r Xx - N

Cylichnina elongata conuloidea (Wood, 1851) Xx r XxxX

Retusa truncatula (Bruguiere, 1792) xX vr xX 7? R

Volvulellaacuminata (Bruguiere, 1792) Xx Xx xX R

Ringicula buccinea (Brocchi, 1814) xX c xX xX xX R

Ringicula ventricosa (Sowerby, 1824) xX - -

Philine scabra (Muller, 1776) xX - Xx R

Philine quadrata(Wood, 1839) r Xx -- R

Philine intricata Monterosato, 1884 xX R

Roxania utriculus (Brocchi, 1814) Xx Xx Xx R

Cylichna cylindracea (Pennant, 1777) Xx c Xx Xx Xx R

Scaphanderlignarius (Linnaeus, 1758) xX c Xx xX xX R

Spiratella atlanta (Linnaeus, 1758) Xx xX xX R

Table 1. List of gastropod species occurring in the Luchtbal Sand Member.<1960: material listed by Glibert (1958b), with additions of

Marquet (1996; 1997 a, b, c; 1998 a, b, c; 2001) and D1-D2 Havendok in NNM; Doel: collected in the Deurganckdok section in

Doel; Noord: collected in Antwerp,junction 5e Havendok-Amerikadok at the Noordkasteel;Dreg: material dredgedfromthe Scheldt

River, possibly originating from the Luchtbal Sand Member; KFm; also occurring in the Kattendijk Formation;OSM: also occurring

in the Oorderen Sand Member;Occ.; occurrence; R: to Recent, N: North Sea Basin endemic species; xpresent; r rare; c common; vr

very rare.

Number ofspecies Percentage Extant Percentage NSB endemic

Bivalvia 117 51 27

Gastropoda 157 29 46

Amphineura 1

Scaphopoda 2

Total 111

Table 2. Number of species, percentage ofstill living and of endemic species in the Luchtbal Sand Member. NSB North Sea Basin.