October 2003 CainozoicResearch, 2(1-2) (2002), pp. 87-107,

Contributions to triviid systematics (, ),

6. Early Pliocene from the western Mediterranean

² Dirk Fehse¹ & Bernard Landau

1 Nippeserslrafle 3, D-12524Berlin, Germany; e-mail: [email protected] 2 International Health Centres, Avenida Infante de Henrique, Areias Sdo Jodo, P-8200-261 Albufeira, Portugal: e-mail: [email protected]

Received 30 October 2001; revised version accepted 1 November 2002

The rich Pliocene triviid faunas in four viz., and from the Early (thirteen species genera, , , ) Estepona

revised and recorded from elsewhere. One C.area (southern Spain) are systematically compared to Neogene assemblages species,

esteponica, is described as new.

KEY WORDS:: Gastropoda, Triviidae, Zanclean, Miocene-Pliocene, western Mediterranean, Italy, systematics, new taxon.

Introduction BS Bellardi & Sacco collns, Torino;

CS F.A. Schilder Colin, currently at Humboldt

In our review of trivioid gastropods from the western Universitat, Museum fur Naturkunde, Berlin;

Mediterranean, we started with the family (Fehse DEB D. Fehse Colin, Berlin;

& Landau, 2002). Like the Eratoidae, the same comments HNC Haus der Natur, Cismar;

triviids. Institut des naturelles de Bel- on difficulty and controversy apply to Many of IRScNB royal Sciences

these the result of problems are poor original descriptions gique, Brussels;

and figures, further confused by Schilder & Schilder MCSNM Museo civico di Storia naturale di Milano;

(1971), who, in our opinion, reviewed much ofthe classifi- NHM The Natural History Museum, London.

cationbased on the figures without having seen the actual

specimens. Even Cate (1979) in his review of the family

limited of the descriptions to copying the original many Systematic palaeontology

data, without amplifying on the older, often incomplete or

inadequate descriptions. Superfamily Trivioidea Troschel, 1863

We have adopted several of Schilder’s methods in our Family Triviidae Troschel, 1863 but describe the descriptions, attempted to conchological Subfamily Triviinae Troschel, 1863

features ofall the species fully. Several specific terms are

used: dorsal rib count, characteristics of the dorsal sulcus

and the formof the and anal canals siphonal (see Schilder, Trivia Broderip, 1837

1933a, p. 288, text-fig. 6).

The work is based on material collected at three present — Type species Cypraea europaea Montagu, 1808, by between 5.5-9 km northeast of The localities, Estepona. monotypy.

deposits consist ofa variety of differentfacies, from coarse

sands (Velerin Sands), representing nearshore or beach

deposits, to fine clayey sands (Velerin Carretera), deposited Trivia coccinelloides (J. de C. Sowerby, 1823) at relatively greater depths and a coarse conglomerate

(Velerin Conglomerates). All these deposits are of Early Figure 1/1 a-c Pliocene (Zanclean) age (Vera-Pelaez et ai, 1996, fig. 1).

* 1823 J. de C. Cypraea coccinelloides Sowerby, p. 107,pi.

Abbreviations— 378, fig. 1.

1843 coccinella Def. — Cypraea Nyst, p. 609, pi. 45, fig.

BLP B. Landau Colin, Albufeira, Portugal; 14. -88-

1848 Mont. — 1932bTrivia coccinelloides coccinelloides Cypraea europaea Wood, p. 17, pi. 2, fig. (Sowerby, 1832)

— 107. 6. Schilder, p.

1872 — 1933bTrivia coccinelloides coccinelloides — Schil- Cypraea europaea Mont. Wood, p. 5, pi. 5, fig. (Sow.)

24. der, p. 9, text-figs 2-4.

— 1881 Mont. — 1941 Trivia Cypraea europaea Nyst, p. 59, pi. 5, fig. (Trivia) coccinelloides Sowerby ‘23 Schil-

2a, b. der, p. 73.

1894 Trivia — 1958 Trivia coccinelloides 1823 cocci- europaea var. coccinelloides(Sow.). Sacco, Sowerby sp. subsp.

27. nelloides — Gilbert, 2, 25a, b. p. 46, pi. 3, fig. p. 27, pi. fig.

— Trivia coccinelloides Sow- 1914 Trivia sphaericulata Lamarck Harmer, p. 49, pi. 1971 (Trivia) coccinelloides

1823 — Schilder & 16. 2, figs 15, 16. erby, Schilder, p.

— 1914 — 1998 Trivia C. Trivia pisolina (Lamarck) Harmer, p. 50, pi. 2, fig. (T.) coccinelloides (J. de Sowerby, 1823)

59. 17. Marquet, p. 84, fig.

— 1920 Trivia — Trivia europaea (Montagu) Harmer, p. 507, pi, 45, 2001 sphaericulata (Lamarck, 1810) da Silva, p.

fig. 11. 242, pi. 10, figs 4-9.

Key to the genera of Triviidaefrom Estepona (Spain)

1. Shellwith dorsal sulcus 2

Shell without dorsal sulcus Trivia

2. Dorsal sulcus very narrow or crossed by ribsribs 3

Dorsal sulcus wide and smooth, the ribs forming tuberclestubercles at thethe edge Pusula

3. Shell very small, inflated; dorsal sulcus weakly developed

and not extending across thethe whole dorsum Cleotrivia

Dorsal sulcussulcus clearly developed, extending the whole length ofthethe dorsum,dorsum,

bisecting, narrowing oror depressing the ribs Niveria

Distribution — Sutton (East Anglia), Coralline Crag 5.4 mm, respectively; LAV ratio 1.21.

(Wood, 1848); Walton-on-the-Naze (East Anglia), Red

Crag; Antwerpen-Doel, Kallo (Belgium), ‘Scaldisien’ Description — Shell small, relatively fragile, and oval.

(Nyst, 1882); Kattendijk and Lillo formations, Belgium Spire covered by callus, but visible beneath three vertical

(Marquet, 1998); ‘Astigiana, Piacentino, Liguria, Piacen- riblets extending upwards from the posterior terminal. ziano ed Astiano’, Italy (Sacco, 1894); Val do Freixo Body whorl globose and rounded, equalling about 90% of

(Pombal, Portugal), Lower/Middle Pliocene (da Silva, total height, with the terminalswell produced. The termi-

2001); Certaldo (Firenze, Italy), Piacenzian(DFB Colin); nals are rounded adapically, straight abapically. Dorsum

Latsia, near Nicosia (Cyprus), Pliocene (BLP Colin); humped in the posterior third, completely covered by 20-26

Lucena (Huelva, Spain), Arenas de Huelva Formation, strong ribs. Base slightly convex, with the terminalsrecur-

Zanclean (BLP Colin); Castellace (Italy), Siciliano Forma- ved. The aperture is narrow, strongly curved adapically and tion, Pleistocene (DFB Colin). slightly widened in the area ofthe fossula. Outer lip broad,

widest in towards convex, mid-portion, becoming narrower

Material studied — From the Velerin Conglomerates, 19 the terminals and tapering slightly inwards. The lip bears specimens (BLP Colin), 8 specimens (DFB Colin); from 19-23 strong, equal teeth. Siphonal and anal canals are

indented. Columella Velerin Carretera, 3 specimens (BLP Colin); from the slightly weakly convex, tapering

Velerin Sands, 6 specimens (BLP Colin); from Badagnano, steeply inwards, bordered internally by a strong carinal

2 specimens and from T. Stirone, 15 specimens (Piacen- ridge, bearing 18-19 ribs, which continue onto the carinal zian, Italy; BLP Colin); from Certaldo (Firenze, Piacen- ridge, where they become slightly stronger. Fossula broadly zian, Italy), 1 specimen (DFB Colin); from Kallo (Oor- concave, not clearly delimited from the rest of the colu- deren Member, Lillo Formation), 12 specimens (BLP mella.

Colin) and 1 specimen (DFB Colin); from Walton-on-the-

Naze (Red Crag, Pliocene), 1 specimen (DFB Colin); and Range ofvariation— This species is characterised by the from Castellace (Pleistocene), 1 specimen (DFB Colin). aperture being well to the right ofthe ventral mid-line, the

fossula not delimited from the columella and the inner

Measurements — Maximum length and width up to 8.2 border of the columella and fossula not covered by the

and 5.5 6.5 and mm mm, respectively, but on average mm outer lip when seen in ventral view. -89-

Figure 1. Triviid gastropods from the Zancleanof Estepona (Spain) (photos B. Landau)

coccinelloides = la-c Trivia (J. de C. Sowerby, 1823),Velerin Sands; original size 7.1 mm (BLP Colin)

2a-c Trivia candidula(Gaskoin, 1836), Velerin Conglomerates; original size 7.8 mm (BLP Colin).

3a-c Trivia acuminata Schilder, 1932a,Velerin Conglomerates, original size = 8.0 mm (BLP Colin).

4a-c Trivia sphaericulata (Lamarck, 1810),Velerin Carretera, original size 17.1 mm (BLP Colin).

5a, b Trivia perobsoleta Sacco, 1894, VelerinConglomerates, original size = 15.6 mm (BLP Colin). -90-

Figure 2. Triviid gastropods from the Zanclean of Estepona (Spain) (photos B. Landau):

la-c, 2a, b Trivia frigida Schilder, 1932b, Velerin Conglomerates and Veledn Sands, respectively, original size = 6.2 mm and 6.1 mm,

respectively (BLP Colin).

3a-c Niveria dimidiata (Bronn, 1831), Velerin Conglomerates, original size = 8.2 mm (BLP Colin).

4a-c Niveria permixta (de Cristofori & Jan, 1832), Velerin Conglomerates,original size = 13.8 mm (BLP Colin).

avellanula Velerin = 5a-c Niveria (Sacco, 1894), Conglomerates, original size 14.1 mm (BLP Colin).

6a-c Niveria dorsolaevigata(Cocconi, 1873),Velerin Conglomerates,original size = 17.2 mm (BLP Colin). -91 -

The shells vary slightly in the development of the dorsal Description — Shell small, relatively solid, and oval. Spire

hump, and in the number of dorsalribs and labial teeth. covered by callus. Body whorl globose and rounded, about

90% of total height, with the terminals depressed and

Discussion — Trivia coccinelloides, best known from the straight. Dorsum humped in the posterior third, completely

Pliocene of the North Sea had extended covered Basin, an geo- by 22-28 strong ribs. Base very slightly convex,

the the with the terminals graphical range during Early Pliocene, reaching slightly recurved. The aperture is narrow

southern Atlantic Iberian coast. and strongly curved adapically. Outer lip very broad,

We have also collected specimens from Zanclean deposits convex, widest in mid-portion, becoming narrower towards

at Huelva, Spain, showing that it migrated through the the terminals and tapering slightly inwards. Outer margin

Straits ofGibraltar into the Mediterranean. is also found It of the lip angularly callused. The lip bears 19-22 strong, occasionally in Pliocene strata ofItaly and Cyprus. equal teeth. Siphonal and anal canals indented. Columella

Schilder described (1933b, pp. 8, 13) a comparable weakly convex, tapering steeply inwards, bordered inter- form from the North Sea Basin under the name Trivia nally by a strong carinal ridge, bearing 16-18ribs, which

coccinelloidesparvula. However, in the Kattendijk Forma- continue onto the carinalridge, where they become slightly tion of T. coccinelloides and T Belgium, c. parvula co- stronger. Fossula broadly concave, not clearly delimited

occur, which means that these shouldbe considered distinct from the rest ofthe columella. The carinal ridge is covered the at species level. The North Sea Basin triviids will be by the outer lip, when seen in ventral view.

discussed in detail in a forthcoming paper.

Range ofvariation — This species is characterised by the

broad outer lip, which has an angular outer edge, the

and the Trivia candidula(Gaskoin, 1836) centrally placed aperture carinal ridge which is not

visible in ventral view. The height of the dorsal hump and

the character ofthe ribs in Figure l/2a-c are very variable, being narrow

some specimens, while very broad with narrow interspaces

G. in others. 1757 Cypraea bitou Adanson, p. 73, pi. 5, xi, fig. 3

(pre-Linnean).

* 1836 Cypraea candidula Gaskoin, p. 201. Discussion — The absence of a dorsal sulcus, the central

1932b Trivia candidula — 108. (Gask.). Schilder, p. position of the aperture as well as other features discussed

1941 Trivia candidula Gaskoin ‘36 — (Trivia) Schilder, p. in Fehse (1999, p. 8), in our opinion favour assignment of 73. this to species the genus Trivia rather than to Cleotrivia as

1979 Niveria (Cleotrivia) candidula (Gaskoin, 1836) — suggested by Cate (1979, p. 53). Cate, p. 53, figs 61, 61a. This species is easily distinguished from T. coccinelloi-

1999 Trivia — candidula (Gaskoin, 1836) Fehse, p. 7, pi. des by the thickened outer lip, which has an angular outer 1, fig. 2; pi. 2, fig. 6, and the margin centrally placed aperture. Despite its wide 1996 Pusula candidula — non (Gaskoin, 1836) Giannuzzi-

et Recent distribution, we are not aware of previous fossil Savelli al., p. 160, fig. 666a-c [= Cleotrivia any record of T. candidula.We have extensive Caribbean werneri Fehse, 1999]. and

MediterraneanNeogene collections (BLP and DFB collns)

at hand, in which this is This would species missing. sug-

— gest a western Mediterranean as T. candidula does Type Lectotype, designated by Cate (1979, p. 53, pi. 15, origin, in Italian figs 61, 61a), isNHM 1874.12.11.55. not occur faunas, having migrated to the western

Atlantic in relatively recent times.

Distribution — Extant in Florida (Fehse, 1999) and the

Caribbean (Redfem, 2001); offthe Canary Islands, western

Africa to Malaga, western Mediterranean. This is the first Trivia acuminata Schilder, 1932a

fossil record ofthe species.

Figures l/3a-c; 6/5a-c

Material studied — From the Velerin Conglomerates, 20

1894 — specimens (BLP Colin), 8 specimens (DFB Colin); from Trivia europaea var. cocinelloides (Sow.) Sacco, 46, pi. 3, fig. 27. Velerin Carretera, 5 specimens (BLP Colin); Recent p. 1911 Cypraea europaea var. pediculoides Cerulli-Irelli specimens (DFB Colin) are from Lauderdale, Florida (1), 16. (partim), p. 273, pi. 26, figs 15, Dry Tortuga Islands, USA (1), Malaga (trawled in 70-80 m * 1932aTrivia acuminata 2. Schilder, p. 257, text-fig. depth (5); Palmeira, Sal Island, Cape Verde Islands (2) and

1932b Trivia acuminata Schil. — Schilder, p. 108. Almadies, Senegal (4). 1941 Trivia (Trivia) acuminata Schilder ‘32 — Schilder,

p. 73.

Measurements — Maximum length and width to 8.1 up 1970 Trivia (Trivia) acuminata Schilder, 1932 — Pavia & and mm 6.2 mm, respectively, but on 6.8 mm and average Demagistris, p. 131, pi. 1, fig. 1. 5.4 LAV ratio mm, respectively; 1.26. 1971 Trivia (Trivia) acuminata acuminata Schilder, 1932f -92-

— Schilder & Schilder, p. 17. Both T. coccinelloides (see above) and T. candidula

1984 Trivia var. cocinelloides 1823) europaea (Sowerby, (see above) have more rounded shells, with a dorsal hump,

— Ferrero Mortara 8a-c. et al., p. 153, pi. 26, fig. fewer ribs, and have the fossula not protruding. In both, the

siphonal and anal canals are indented, whereas in T. acu-

Type — Holotype is CS 3411. minata the canals follow the shell contour. Trivia can-

didula has a similarly callused outer margin ofthe lip, but

Distribution — Miocene of NE Upper Atlantic (DFB be may distinguished by the more centrally placed aperture, Colin), Pliocene (Pavia & Demagistris, 1970; Ferrero with the carinal ridge covered by the apertural lip when Mortara et al., 1984) and Lower Pleistoceneof the western seen in ventral view; T. coccinelloides has a non-callused Mediterranean(Cerulli-Irelli, 1911). lip margin.

Schilder (1941, pp. 73, 114) described T. nana from the

Material studied — From the Velerin Conglomerates, 43 Upper Miocene (‘Tortonian’) of Montegibbio, Italy; the specimens (BLP Colin), 9 specimens (DFB Colin); from holotype (CS 5084) is here illustrated in Figure 6/7a, b. VelerinCarretera, 3 specimens (BLP Colin); from Certaldo This is a smaller species, with a more inflated, globular (Firenze, Lower Piacenzian), 3 specimens (DFB Colin); shell; ribs are fewer and coarser, the dorsum is considera- from 4 Castell’Arquato (Lower Piacenzian), specimens the bly more elevated, aperture very narrow and the ex- (BLP Colin), 8 specimens (DFB Colin); from Brigne tremities are only weakly developed. The differences in (Anjou, France, Lower Pliocene), 1 specimen (DFB Colin). shell morphology are significant enough to consider them

separate at the specific level.

Measurements — Maximum length and width to 7.8 up Trivia acuminata is restricted to the Mediterraneanand mm and 5.7 mm, respectively, but on 7.0 mm and average NE Atlantic. This is the first record from the Lower Plio-

5.1 mm, respectively; LAV ratio 1.37. cene; previously, the earliest record was Middle Pliocene

of Valle Botto (Pavia & Demagistris, 1970, p. 131, pi. 1,

Description — Shell small, solid, and elongated oval. Spire fig. 1). covered by callus. Body whorl somewhat elongated, glo- bose and rounded, about 85% of total height, with the terminals produced and slightly rounded. Dorsum evenly Trivia sphaericulata (Lamarck, 1810) rounded, completely covered by 24-30 strong ribs. Base almost straight, with the terminals weakly recurved. The Figure l/4a-c aperture is relatively narrow, curved adapically and wid- ened in the fossular Outer region. lip broad, convex, regu- * 1810 Cypraeasphaericulata Lamarck, p. 107.

widest in narrower larly rounded, mid-portion, becoming 1894 — Trivia sphaericulata Lk. Sacco, p. 47, pi. 3, fig. towards the terminals. Outer margin of the lip slightly to 29.

callused. The bears 1894 Trivia strongly, angularly lip 21-22 strong, sphaericulata var.parvosphaera Sacco, p. 48, equal teeth. Siphonal and anal canals follow the shell pi. 3, fig. 30.

1894 Trivia sphaericulata var. retusoides 48, profile. Columella convex, tapering steeply inwards, bor- Sacco, p. pi. 3, fig. 31. dered internally by a weak carinal ridge, bearing 18-21

1894 Trivia sphaericulata var, obsoleta (Bon.) — Sacco, ribs, which continue onto the carinal ridge, where they do p. 48, pi. 3, fig. 32a, b. become Fossula not significantly stronger. broadly con- 1894 Trivia sphaericulata var. pseudavellanaSacco, p. 49, cave, not clearly delimitedfrom the rest of the columella pi. 3, fig. 34. and slightly protruding. 1894 Trivia sphaericulata var. propeavellana Sacco, p. 49,

pi. 3, fig. 35.

Range ofvariation — The typical features of this species 1932b Trivia sphaericulata sphaericulata (Lam.) — Schil- are the somewhat rounded dorsum, 107. elongated shape, evenly der, p.

high numberofribs and the slightly protruding fossula. The 1932b Trivia testudinellapseudavellana Sacco — Schilder,

callus the 104. marginal on outer lip varies from slightly devel- p.

1941 Trivia Lamarck ‘10 — Schil- oped to forming a marked angular step at the periphery. In (Trivia) sphaericulata der, 73. a few specimens the shape is more globose. p. 1971 Trivia (Trivia) sphaericulata sphaericulata Lamarck,

1810 — Schilder & Schilder, 17. p. Discussion — Schilder in the (1932a, p. 258), discussing 1971 Trivia (Sulcotrivia) testudinellapseudavellana Sacco, Italian Pliocene Triviidae, considered the absence of a

1894 — Schilder & 17. Schilder, p. dorsal sulcus to be the main characteristic feature of this 1984 Trivia sphaericulata (Lamarck, 1810) — Ferrero In a he 1933b, species. subsequent paper, (Schilder, p. 19) Mortara 153. etal., p. did not the note presence of T. coccinelloides in the Medi- terranean. Therefore, the absence ofthe dorsal sulcus alone

Distribution — Plioceneof the western Mediterranean. cannot be used to distinguish Pliocene triviids from the

it is Mediterranean; simply a typical feature of the genus Materialstudied — From the Velerin Conglomerates, 32 Trivia. -93 -

specimens (BLP Colin), 1 specimen (DFB Colin); from the ribs, but a weak depression in the dorsum as described

Velerin Carretera, 1 specimen (BLP Colin); from Rio by Sacco. Our opinion is that Schilder based his decision

Torsero (Italy), Lower Piacenzian, 7 specimens (DFB on Sacco’s figures, misinterpreting the strength of the so-

Colin); from Bacedasco (Italy), Lower Pliocene. 1 speci- called ‘sulcus’ mentionedand figured. There are no other men (BLP Colin). features that distinguish the forms separated by Schilder

from the nominal with in type. We agree Sacco considering

Measurements — Maximum length and width up to 15.4 all illustrated material of Trivia sphaericulata, with the

and 12.9 but 14.6 mm mm, respectively, on average mm exception of T. sphaericulata var. parvavellana and T. s. and 12.4 L/W ratio mm, respectively; 1.18. var. perobsoleta, as a single species, without specifying

varieties.

— Shell solid, and Schilder & Schilder mentioned Description large, spherical. Spire (1971, p. 17) yet an- obscured callus. by Body whorl globose and rounded, other subspecies of T. sphaericulata, T. s. crassa Cocconi, about 95% of total with the terminals 1873 from the Italian ‘Tortonian’. could height, weakly pro- Unfortunately, we

duced and rounded. Dorsum rounded, without true not evenly a obtain a copy ofCocconi’s description and there is no

dorsal sulcus, although a few specimens show a weak materialofthis taxon in F.A. Schilder’s Collection.

dorsal depression, which does not interrupt the ribs. Dor-

covered which sum completely by 22-36 ribs, vary in thickness and become close more set mid-dorsally. Base Trivia perobsoleta Sacco, 1894

and terminals are evenly convex. Aperture wide and

curved, especially in the adapically portion. Outer lip very Figures l/5a, b; 5/3a-d broad, widest in mid-portion, becoming narrower towards the terminals. In the is elevated *1894 Trivia some specimens lip adapi- sphaericulata var. perobsoleta Sacco, p. 49,

The is keeled in 33. cally. lip roundly cross section, with the pi. 3, fig.

1932bTrivia Sacco — 107. narrower inner portion, sloping steeply inwards and the perobsoleta Schilder, p. 1971 Trivia (Trivia) perobsoleta Sacco 1894 — Schilder wider outer portion, less so, outwards. Outer margin ofthe & Schilder, p. 17. lip slightly to strongly, angularly callused. The lip bears 19-

25 teeth. strong, equal Siphonal and anal canals follow the

shell Columella profile. convex, tapering steeply inwards, Type — Holotype is BS.043.12.007.

bordered internally by a weak carinal ridge, bearing 16-20

which continue onto the carinal where do ribs, ridge, they Distribution — Pliocene ofthe western Mediterranean.

become not significantly stronger. The carinal ridge is

strongly protruding in the fossular region. Fossula broadly Material studied — In addition to the holotype, ten speci-

concave and not clearly delimited from the rest of the mens from the Velerin Conglomerates (BLP Colin). columella.

Measurements — Maximum and width to 16.3 length up

Range of variation — The characteristic features of this mm and 14.2 mm, respectively, but on average 14.0 mm taxon are the large size, and 12.1 LAV globular shape, weakly developed mm, respectively; ratio 1.16. terminals, the curved aperture and the strongly protruding

carinal at the fossular All the other characters ridge region. Description — Shell large, solid, and spherical. Spire

are remarkably variable. The number and of the strength covered by callus. Body whorl globosely inflated and ribs and teeth, the outline in the which is typical form, rounded, about 95% of total height, with the terminals not rounded, is modified in of the with some specimens an produced and rounded. Dorsum strongly elevated and

elevated to the outer adapical part lip. evenly rounded, without a dorsal sulcus. Dorsum bearing

subobsolete ribs in 70% of specimens, 30% smooth and

Discussion — This large and beautiful shell is easily with ribs. glossy no Base and outer lip are evenly convex. distinguished from by its large size and congeners spherical Aperture wide and somewhat curved, especially in the

shape. It is one of the larger members of the with genus, adapically portion. Outer lip very broad in mid-portion,

specimens from even than those Estepona being larger narrows abruptly towards the terminals. The lip is rounded

found in Sacco (1894, 48, 49) indicated in his in but Italy. pp. cross section, anteriorly concave and sloping steeply various ‘varieties’ the wide of variation of this range inwards. Outer margin ofthe lip slightly angularly callused.

species. Schilder (1932b, p. 104) and Schilder & Schilder The bears 18-24 fine which lip denticles, are only barely

referred Sacco’s as a (1971, p. 17) variety pseudavellana continued as ribs on the outer lip. Siphonal and anal canals

subspecies to Niveria testudinella a (Wood, 1842), noting follow the shell profile. The anal canal is indistinct. Colu-

depression on the sulcus. Both at Rio Torsero (Italy) and mella nearly straight, tapering steeply inwards, without an all of Sacco’s varieties and intermediatesbetween Estepona inner carinal ridge, bearing 14-18 weak denticles, which those described, are We note that the ‘sulcus’ seen present. are distinctly continued as fine ribs onto the columellabut

by Schilder is not a real furrow, interrupting or depressing obscurely so onto the base. The inner fossular margin -94-

Fossula shallow and delimited Schilder 1932 — & 17. slightly protrudes. clearly Schilder Schilder, p.

from the rest of the columella.

Types — Holotype is CS 221; paratype is CS 5168.

Range ofvariation — The characteristic features of this

are the Distribution — Pliocene of the western Mediterranean, species large size, very globular shape, very weakly Altavilla developed terminals, the curved aperture and, most impor- Milicia (Sicily), ‘Monte Pellegrino, Ficarazzi, Sciacca tantly, the character of the ribs, always obsolete on the Gravina’; Lower Pleistocene (Schilder, 1932b).

sides ofthe shell and subobsoleteto lacking on the dorsum.

These ribs be variable in Material studied— From the Velerin can strength, sometimes barely Conglomerates, 9

whilst in other 2 from visible, specimens they are reasonably specimens (BLP Colin), specimens (DFB Colin);

strongly developed on the dorsum, but invariably absent on the Velerin Sands, 10 specimens (BLP Colin); from Cutro-

fiano the sides even in well-preserved, non-eroded material. (Lecce, Italy); Calabriano Formation (Lower Pleisto-

cene), 13 specimens (DFB Colin).

Discussion — In discussing this species, it is important to

it from Measurements — Maximum and width to distinguish juvenile specimens of T. sphaericulata length up 6.3

(see above); the latterare also devoid of ribbing, but show mm and 5.1 mm, respectively, but on average 5.6 mm and

4.6 LAV the feature of an immature triviid, namely a lightweight mm, respectively; ratio 1.22.

shell, a spire not covered by callus, a non-callused outer lip

and poorly developed dentition.Adult shells of T. perob- Description — Shell small, relatively fragile, inflated and soleta thick with the covered are shelled, spire by callus, oval. Spire covered by callus, faintly visible beneath 3-4 and have thickened and denti- a outer lip well-developed vertical riblets extending upwards from the posterior tion. The other features similar those of are to T. terminal. Body whorl globose and rounded, about 90% of from the obsolete dorsal T. sphaericulata. Apart ribs, total height, with the terminals slightly produced, the is perobsoleta somewhat more globular, the posterior end anterior straight, posterior rounded. Dorsum evenly ofthe is the callused of parietal lip obsolete, outer margin rounded, completely covered by 26-30 fine ribs. Base the labial is less the base is lip shouldered, more convexly rounded, with the posterior terminal strongly recurved. The

inflatedand the columellaand the fossula are less devel- is curved aperture fairly wide, adapically, further widened without oped, an inner adaxial carinal In view of in the fossular Outer ridge. region. lip relatively broad, convex, these constant differences, we keep these two forms dis- regularly rounded, widest in mid-portion, becoming nar- tinct. rower towards the terminals. Outer margin of the lip Some of the from have specimens Estepona a com- slightly angularly callused, becoming subobsolete in mid-

smooth but ribs the ventral pletely dorsum, present on side, portion. The lip bears 18-20 fine, equal teeth. Siphonal and

similar to the Recent South African Triviella Jous- anal canals indented. Columella genus slightly convex, tapering 1884 and the Trivia seaume, Oligocene francisca Schilder, steeply inwards, bordered internally by a weak carinal 1925 and Trivia calva 1928 from Schilder, Germany. A ridge, bearing 16-17 ribs, which continue onto the carinal

similar occurs in the Atlantic Miocene where species Upper ridge, they do not become significantly stronger.

of Sceaux which will be Fossula not delimited (Lower ‘Redonian’) (NW France), broadly concave, clearly from the discussed in a forthcoming paper. rest ofthe columellaand slightly protruding. Schilder and Schilder Schilder (1932b, p. 107) &

(1971, 17) considered T. perobsoleta to be p. synonymous Range of variation — The characteristic features of this with Niveria permixta, but did not discuss their point of species are the small size, inflated shell, large number of view. The also in is different latter, present Estepona, quite fine in relation to the small size and the ribs, weakly pro- (see below). duced terminals. The other shell features are relatively

constant.

Triviafrigida Schilder, 1932b Discussion — Philippi (1836, p. 237) misidentified his

material as Cypraea coccinella Lamarck, 1810. Schilder Figures 2/la-c, 2a, b; 4/2a-c renamed (1932b, p. 109) Philippi’s taxon, but failed to

provide notes or illustrations. Therefore, it is questionable

1836 coccinella — Cypraea Lam. Philippi, p. 237 (non if Schilder actually saw Philippi’s originals, since he Lamarck) designated two specimens in Philippi’s Collectionas types.

1844 Lam. — 200. Cypraea coccinella Philippi, p. The holotype is here illustrated for the first time (Figure 1913 Trivia — 532 frigida(Monterosato, MS) Gignoux, p.

(nomen nudum). 4/2a-c). Ours is the first record of the from * 109. species the Lower 1932b Trivia mediterraneafrigida Schilder, p.

1933b Trivia Schil. — 14. Pliocene. Trivia differs from T. acuminata frigida Schilder, p. frigida (see

in 1941 Trivia Schilder ‘32 — 73. less with (Trivia) frigida Schilder, p. above) being elongated, more inflated, a larger

1971 Trivia (Trivia) multilirata frigida (Gignoux 1913) numberofribs in relation to size, which are finer and more -95 -

close set. Trivia coccinelloides and T. candidulaboth have into the aperture. All marginal portions ofthe base and lip

dorsal and fewer ribs and broader elevated. The ribs of and a hump, stronger a outer are are coarse, equal strength lip. relatively widely spaced. None ofthe species discussed in and with such Yet another species of Trivia occurs in the Italian the present paper are squat, a depressed

central base. candidula is but lacks the Pliocene, T. subdilatata Schilder, 1941, the type (CS 5063) Trivia similar,

above-mentioned features. subdilatata is of which was not figured in the original paper. Here (Figure Trivia easily

subdilatata from T. candidula the different of 6/6), we figure the type for the first time. Trivia distinguished by shape

shell with curved and the the wider columellaand the wider has a very squat a narrow, aperture, aperture, fossula, the columella, fossula and labral lip are very wide. The apertural lip and somewhat finerribs, which are closer set base and labral lip are concave centrally where they slope on the dorsum.

Trivia Key to the identification of the genus from Estepona, Spain

1. Adult shell <10<10 mm 2

Adult shell >10 mm 5

2. Adult shell with callused outer lip 3

non-callused callused 4 Adult shell with non-callused or weakly callused lip margin 4

Shell with and covered carinal T. candidula 3. Shell with centrally placed aperture covered ridge T.

Shell more elongated, thinner outer lip T.T. acuminata

4. Shell with dorsal hump, strong ribs and no lip margin T.T. coccinelloides

Shell evenly rounded, numerous fine ribs, lip margin weak T.Tfrigida

5. Ribs fully developed T.T. sphaericulata

Ribs dorsally obsolete or absent T. perobsoleta

Genus Niveria Jousseaume, 1884b (=Sulcotrivia Schilder, 46. 1894 dimidiata laevisulcata Sacco, 53, 3, 1933b) Trivia var. p. pi. fig. 48.

1911 Cypraea europaea var. pediculoides Cerulli-Irelli Type species — Cypraea nivea Gray, 1824 (= Trivia nix (partim), p. 273, pi. 26, figs 19-21. Schilder, 1923), by original designation. 1925 Trivia () dimidiata (Bronn, 1831) —

Schilder, p. 63.

Discussion — Schilder (1933b, p. 18) erected the genus — 105. 1932bTrivia dimidiata(Bronn, 1831) Schilder, p. dimidiata Sulcotrivia, with Cypraea Bronn, 1831 as type, 1970 Trivia (Sulcotrivia) dimidiata (Bronn, 1831) — Pavia Sulcotrivia shares features in shell although important & 9. Demagistris, p. 132, pi. 1, fig. with 173- morphology Niveria (see Schilder, 1939, pp. 1971 Trivia (Sulcotrivia) dimidiata dimidiata (Bronn,

In both similar. — & 18. 174). addition, type species are closely 1831) Schilder Schilder, p.

There are not enough differences to separate the members 1984 Trivia dimidiata (Bronn, 1831) — Ferrero Mortara et

al., 155, pi. 27, fig. 6a-c. of both genera even on a subgeneric level, which is why p.

1992 Trivia dimidiata — Cavallo & Re- Fehse (in press) favours synonymising Sulcotrivia with (Bronn, 1831)

subscribed here. petto, p. 66, fig. 118. Niveria, a view to

1998 Trivia subaffinis Bugnone, 1880 — Smriglio et al., Species of Niveria are not dorsally humped, but regu- 35-38. p. 161, text-figs larly rounded and have a clear dorsal furrow, not seen in

Trivia. The dorsal sulcus is a distinct groove, which either

depresses, interrupts or bisects the ribs and which is not Distribution — Pliocene of the western Mediterranean

just a weak depression or dimple in the centre of the dor- (Sacco, 1894).

in of T. sum, as seen some specimens sphaericulata (see

above). Materialstudied — From the Velerin Conglomerates, 25

specimens (BLP Colin), 6 specimens (DFB Colin); from

the Arenas de Huelva Formation of Lucena (Lower Plio-

Niveria dimidiata (Bronn, 1831) cene), 1 specimen (BLP Colin); from San Gimignano

(Italy), LowerPiacenzian, 9 specimens (BLP Colin); from

Figure 2/3a-c Valle Botto and Baldichieri (Italy), Upper Piacenzian, 1

and 5 specimens, respectively (DFB Colin).

*1831 16. Cypraea dimidiataBronn, p.

1894 Trivia — dimidiata(Bronn) Sacco, 53, pi. 3, fig. — p. Measurements Maximum length and width up to 8.6 -96-

* mm and 5.9 mm, respectively, but on average 7.4 mm and 1832 Cypraea permixta de Cristofori & Jan, p. 15.

1932bTrivia Crist. & Jan, 1832 — 5.1 mm, respectively; L/W ratio 1.44. permixta Schilder, p. 105.

1941 Trivia (Sulcotrivia) permixta Cristofori-Jan — Schil- Description — Shell medium sized, relatively solid and

der, p. 74. elongated oval. Spire covered by callus. Body whorl elon- 1970 Trivia (Sulcotrivia) dorsolaevigata Sacco, 1894 — gated-globose and rounded, almost 100% of total height. Pavia & Demagistris, p. 133, pi. 1, fig. 4. The terminals are weakly produced, especially the posterior 1971 Trivia (Sulcotrivia) permixtapermixta Cristofori-Jan, terminal and rounded. Dorsum rounded, completely cov-

1832 — & Schilder Schilder, p. 18. ered by 26-36 strong ribs. A dorsal sulcus, of variable 1978 Cypraea permixta De Cristofori & Jan, 1832 —

strength, extends longitudinally across the centre of the & Pinna Spezia, p. 139, pi. 21, fig. 1, la. dorsum, in the of most ofthe majority specimens bisecting 1992 Trivia dorsolaevigata Cocconi, 1873 — Cavallo &

the latter thickened the furrow. 119. ribs, becoming adjacent to Repetto, p. 66, fig.

Base slightly convex, with the terminals recurved. The

is narrow in the and aperture relatively posterior portion Type — Lectotype, designated by Pinna & Spezia (1978,

curved adapically, wider in the anterior part. Outer lip 139, is i.4314. p. pi. 21, fig. 1, la), MCSNM somewhat widest in broadened, convex, mid-portion,

narrower towards the terminals and becoming regularly Distribution — Pliocene of the western Mediterranean.

rounded. Outer margin weakly angularly callused. The lip

bears 19-24 teeth. and anal canals strong, equal Siphonal Materialstudied — From the Velerin Conglomerates, >50 Columella are slightly protruding. convex, tapering steeply specimens (BLP Colin), 10 specimens (DFB Colin); from

inwards, bordered internally by a weak carinal ridge. the Velerin Sands, 3 specimens (BLP Colin); from Certaldo Columella which continue the bearing 18-21 ribs, onto (Italy), Piacenzian, 1 specimen (DFB Colin).

carinal ridge. Fossula broadly concave and delimitedfrom

the rest of the columella. Measurements — Maximum length and width up to 14.7

and 11.2 but mm mm, respectively, on average 12.6 mm

variation — The characteristic features of this Range of and 9.9 mm, respectively; L/W ratio 1.26.

species are the elongated shape, the low profile and the

dorsal which bisects most of the ribs. The number sulcus, Description — Shell large, solid, somewhat elongated to

of ribs and width are somewhat variable. The strength of oval. covered very inflated, Spire by callus, but faintly the sulcus is also but the variable, always present extending visiblebeneath 3-5 vertical riblets extending upwards from

whole length ofthe dorsum. The outer lip is not very broad the posterior terminal. Body whorl varies from somewhat

in comparison with other congeners and usually only elongated and globose to greatly inflated, about 90% of

callused. A number of weakly specimens are smaller, total height. The terminals are rounded, the anterior and the dorsal sulcus rather slightly more globose depresses strongly produced, the posterior weakly so. Dorsum than bisects ofthe ribs. The other shell characteristics most roundedto strongly humped in the posterior third, covered

are similarto those of the typical form. by 30-34 ribs. On mid-portion ofthe shell, the ribs become

much wider towards the summit, where in most specimens

Discussion — The ofthe Sulco- present taxon, type genus they are obsolete, to a variable degree. The summit has a what understand dorsal trivia, clearly shows we to be a true polished appearance due to callus deposited over the ribs, sulcus. this almost all the ribs and In species are interrupted rather than abrasion. A dorsal sulcus, of variable strength the bisected ends thickened. Schilder & Schilder (1971, p. and width, extends longitudinally across the centre ofthe

mentionedNiveria selmae as a 18) (O. Boettger, 1901) dorsum, bisecting some ofthe ribs, depressing others. Base

Miocene of N. dimidiata. N. selmae subspecies However, convex, with the terminalsrecurved. The aperture is rela-

is from N. dimidiata the (Figure 6/8a-c) distinguished by tively narrow in the posterior portion and curved adapi- inflated shell with fewer the better more finer, ribs, by cally, wider in the anterior part. Outer lip broadened,

columella and and the ribs are not widest in towards developed fossula, convex, mid-portion, becoming narrower

bisected the dorsal sulcus and thick- by are considerably the terminals and regularly rounded, bearing 20-25 strong,

ened the which is consider these taxa along furrow, why we equal teeth. Outer lip, with a strong angular margin. Si- be distinct. to phonal and anal canals are slightly protruding. Columella

Niveria common in the dimidiatoaffnis (Sacco, 1894), convex, tapering steeply inwards, bordered internally by a

Atlantic Middle Miocene of the Loire Basin, (‘Pontilevian’ strong carinal ridge. Columella bearing 18-23 ribs, which

is similar to N. dimidiata; this will be the France), species run onto carinal ridge, where they become thickened. discussed in a forthcoming paper. The carinal ridge is strongly protruding in the fossular

region. Fossula deeply concave and clearly delimitedfrom

the rest of the columella. In most specimens a colour

Niveria & permixta (de Cristofori Jan, 1832) pattern is preserved; dorsalpart brown, base and outer lip

Figure 2/4a-c white. -97-

variation — This is characterised Range of species by the bearing 21-23 strong, equal teeth. Outer lip, with a strong callus the on summit of the dorsum and the only slightly angular margin. Siphonal and anal canals follow the shell

terminal. Other features vari- Columella produced posterior are very profile. convex, tapering steeply inwards, bor- the would able, as name suggest. dered internally by a carinal ridge. Columellabears 17-21

ribs, which run onto the carinal ridge, which is strongly

Discussion — the dorsal in Despite hump, seen many ofthe protruding in the fossular region. Fossula deeply concave

the of dorsal sulcus specimens, presence a strong depress- and delimited from the rest of the columella. A colour

all the ribs favours of this is of the ing or bisecting assignment pattern preserved on one specimens, consisting of

the Niveria rather than Trivia. The formed brown either species to genus to two irregularly spots, one at extremity

curious deposition of thick callus onto the summit of the ofthe dorsal sulcus.

dorsum seen in N. permixta is not unique; N. floridana

(Olsson & Harbison, 1952), from the Pinecrest Beds Range of variation — This species is characterised by the

(Pliocene, Florida), and N. dorsolaevigata (see below) large size, the fine ribs and the narrow dorsal sulcus. There show the feature. is variation in the same no significant specimens found.

Discussion — This species most closely resembles N.

Niveria dorsolaevigata (Cocconi, 1873) permixta (see above), but differs in being larger, with finer

ribs (although number of ribs is similar), which do not

Figures 2/6a-c; 5/la-c become thicker towards the summit. The dorsal sulcus is

narrower, and only occasionally are there (see Figure 5/la-

*1873 Trivia affinis Duj. var. dorsolaevigata Cocconi, p. c) a light callus on the summitand the fossula more devel- 162. oped. Lastly, the coloration is different, uniformly brown

Trivia — 1894 dorsolaevigata (Cocc.) Sacco, 52, pi. 3, with p. a white base and lip in N. permixta, white with two 44. fig. the dorsum in spots on N. dorsolaevigata. From N. avella- 1894 Trivia dorsolaevigatavar. sulconitens Sacco, p. 52, nula the differs in (see below), present species being larger, pi. 3, fig. 45. more globose with much finer and more numerous ribs,

1932b Trivia — dorsolaevigata Cocconi, 1873 Schilder, p. with the dorsal sulcus much narrower, the columella taper- 105. much ing more steeply inwards and the fossula being more 1941 Trivia (Sulcotrivia) dorsolaevigata Sacco ‘94 — concave and more Schilder, p. 74. protruding.

1970 Trivia (Sulcotrivia) dorsolaevigata Sacco, 1894 —

4. Pavia & Demagistris, p. 133, pi. 1, fig.

1984 Trivia dorsolaevigata Cocconi, 1873 — Ferrero Niveria avellanula (Sacco, 1894)

Mortara et ai, p. 155.

Figures 2/5a-c; 6/1a-d

Distribution — Pliocene of the western Mediterranean.

* 1894 Trivia avellanula 43. Sacco, p. 51, pi. 3, fig.

Material studied — From the Velerin Conglomerates, 16 1932bTrivia avellanula 1894 — 104. Sacco, Schilder, p.

‘ specimens (BLP Colin); Colli Astesi (Italy), Upper Piacen- 1 94 1 Trivia (Sulcotrivia) avellanula Sacco 94 — Schilder,

74. zian, 1 specimen (BS.043.12.017). p.

1971 Trivia (Sulcotrivia) excoccinella avellanula Sacco,

1894 — Schilder & Schilder, 17. Measurements — Maximum p. length and width up to 17.4

1984 Trivia avellana avellanula — Fer- and 13.1 but var. Sacco, 1894 mm mm, respectively, on average 14.6 mm rero Mortara et ai, p. 155, pi. 27, fig. la-c. and 11.1 mm, respectively; L/W ratio 1.31.

Description — Shell large, relatively solid, inflated and Type — Lectotype, designated by Ferrero Mortara et al. oval. Spire covered by callus, faintly visible beneath 4-6 (1984), is BS.043.12.015.

vertical riblets extending upwards from the posterior terminal. Body whorl globose and inflated, about90-95% Distribution — Middle Miocene of Italy and Pliocene of of total The height. terminals are rounded, the anterior the western Mediterranean.

strongly produced, the posterior weakly so. Dorsum rounded, bearing 30-36 fine ribs. A narrow dorsal sulcus Material studied — From the Velerin Conglomerates, 3 extends longitudinally across the centre of the dorsum, specimens (BLP Colin); from the Arenas de Huelva For-

of the depressing most ribs, bisecting a few in the anterior mation (Lucena), Lower Pliocene, 1 specimen (BLP portion. Base convex, with the terminals strongly recurved. Colin); from Montegibbio, Upper Miocene (‘Tortonian’),

The aperture is relatively narrow below the posterior 1 specimen (BLP Colin); from San Gimignano (Italy), terminal, becoming progressively wider abapically. Outer Lower Piacenzian, 1 specimen (DFB Colin). lip broadened, convex, widest in mid-portion, becoming

narrower towards the terminals and rounded. Measurements— regularly Maximum length and width up to 14.2 -98-

Trivia mm and 11.2 mm, respectively, but on average 13.2 mm 1894 dimidiatavar. eratopsoides Sacco, p. 53, pi. 3,

47. and 10.2 mm, respectively; LAV ratio 1.30. fig.

Trivia Sacco 1894 — 1932b parvavellana Schilder, p. 104.

1941 Trivia (Sulcotrivia) parvavellana Sacco ‘94 — Schil- Description — Shell large, solid, somewhat elongated and 74. der, p. inflated, oval. Spire covered by callus, barely visible 1971 Trivia (Sulcotrivia) dimidiataparvavellana Sacco, beneath 2-3 vertical riblets, extending upwards from the 1894 — Schilder & Schilder, p. 18. posterior terminal. Body whorl elongated globose, about 1984 Trivia sphaericulata var. parvavellana Sacco, 1894 90% oftotal height. The terminalsare rounded,the anterior — Ferrero Mortara et al., p. 154. the produced, posterior weakly so. Dorsum rounded,

elevated slightly in mid-portion, covered by 20-22 strong

Type -— Lectotype, designated by Ferrero Mortara et al. ribs. A fairly wide dorsal sulcus, of variable strength, (1984), is BS.68.043.12.010. extends longitudinally across the centre ofthe dorsum. The ribs become thickened at the edge of the sulcus, narrowing

Distribution— Upper Miocene (Lower ‘Redonian’) of the as they cross the depression. Base very weakly and regu- NE Atlantic (DFB Colin) and Pliocene of the western The larly convex. aperture is relatively wide. Outer lip Mediterranean. broadened, convex, widest in mid-portion, becoming narrower towards the terminals and regularly rounded,

Material studied — From the Velerin Conglomerates, 87 bearing 16-20 strong, equal teeth. Outer lip, with a strong specimens (BLP Colin), 9 specimens (DFB Colin); from angular margin. Siphonal and anal canals are slightly the Velerin Sands, 10 specimens Colin); from Columella bordered (BLP Brigne protruding. concave, tapering inwards, (Anjou), Upper Miocene (‘Redonian’), 3 specimens (DFB internally by a weak carinal ridge. Columellabears 16-17 Colin). ribs, which run onto the carinal ridge, where they become thickened. The carinal ridge is slightly protruding in the

— Maximum Measurements length and width up to 8.4 fossular region. Fossula concave and not clearly delimited mm and 6.3 mm, respectively, but on average 7.2 mm and from the rest ofthe columella.

5.5 mm, respectively; L/W ratio 1.31.

Range of variation — The characteristic features of this

Description — Shell small, solid, inflatedand oval. Spire species are the strong ribs and the character ofthe sulcus. covered by callus, barely visible beneath 2 vertical riblets, Of the two adult specimens available, one is slightly more extending upwards from the posterior terminal. Body whorl inflated. The more depressed specimen is almost identical globose, about 95% of total height. The terminals are to the shell illustrated by Ferrero Mortara et al. (1984, pi. produced, the anterior flattened, the posterior rounded. 27, fig. la-c). Dorsum evenly rounded, covered by 16-20 strong, some-

what ribs. irregular A narrow dorsal sulcus, of variable Discussion — Schilder & Schilder (1971, p. 17) referred strength, extends longitudinally across the centre of the this as a subspecies to Trivia (Sulcotrivia) excoccinella dorsum. The ribs become thickened at the edge of the from the Atlantic Middle Miocene (‘Pontilevian’ of the sulcus and either bisected or narrowed as they cross the Loire Basin, France), but this is more inflated, the dorsal depression. Base almost straight, with the anteriorterminal profile being more rounded, or elevated in the posterior recurved andthe The hardly posterior strongly so. aperture portion, whereas N. avellanula is more depressed, or is narrow, strongly curved adapically and widened in the elevated in The former has mid-portion. more numerous, area of the fossula. Outer lip broadened, convex, widest in finer ribs. The subspecific status given by Schilder &

mid-portion, becoming narrower towards the terminalsand Schilder (1971) would suggest a phylogenetic lineage, regularly rounded,bearing 19-22 strong, equal teeth. Outer which we feel is unclear.

lip, with an angular margin. Siphonal and anal canals are The is from present species distinguished N. permixta slightly indented. Columella concave, tapering inwards, (see above) by the better developed dorsal sulcus, wider bordered weak carinal internally by a ridge. Columella aperture, the less developed fossula, the less steeply in- bears 17-19ribs, which run onto the carinal ridge, which is wardly slanting columella and, above all, the absence of slightly protruding in the fossular region. Fossula concave callus the dorsum. Niveria dimidiata is on (see above) and not clearly delimitedfrom the rest ofthe columella. In much smaller, more elongated, with a much narrower of the clear colour most specimens a pattern is visible; it dorsal sulcus thatbisects most ofthe ribs. consists of a small brown spot on the anterior end of the

dorsal sulcus, a patch surrounding the posterior end and

mid-dorsally a larger patch on either side of the sulcus, Niveriaparvavellana (Sacco, 1894) which, in some specimens, are fused.

3/1 6/4a-d Figures a-c, 2a, b; Range ofvariation — This species is characterised by its

small inflated the character ofthe * size, form, irregular ribs, 1894 Trivia sphaericulata var. parvavellana Sacco, p. 49, the slightly indented canals and the colour The pi. 3, fig. 36. pattern. -99-

strength of the sulcus and the character of the ribs as they shell features and, when considering them to be cross are very variable; in some specimens clearly bisected, stratigraphically isolated, we can understand Schilder & in others continuous and depressed, although in the latter Schilder’s conclusion. However, these types co-occur in

narrowed the Zanclean which makes their case they are across depression. strata at Estepona, subspecific

status impossible. Niveria dimidiatadiffers in being more

Discussion — Schilder & Schilder (1971, p. 18) considered elongated, less globose, and in having a lower dorsal

this taxon to be a subspecies of Trivia (S.) dimidiata, profile. The ribs are more numerous (26-36 vs 16-20) and

d. in Middle of assuming T. (S.) parvavellana to occur the regular in character, whereas those N. parvavellana are

Pliocene and T. (S.) d. dimidiata to be restricted to the wavier and frequently interrupted on both dorsal sides.

Upper Pliocene, both in Italy. The two forms share similar

the Key to identification of the genus Niveriafrom Estepona, Spain

> 1. Adult shell >1010 mm 2

Adult shell <10 mm 4

2. Numberof ribs more than26 3

Ribs strong and fewer than 26, sulcus broad, ribs not bisected,

but thickened on thethe edge of the sulcus, narrowed in the depression N. avellanula

Shell ribs fine and 3. Shell globose, ribs fine and numerous,numerous, sulcus narrow N. dorsolaevigatadorsolaevigata

Shell with dorsum and callus the summit N. humped onon N. perpermixtamixta

4. Shell elongated, with more than 22 ribs, which areare regular N. dimidiata

with fewer which Shell globose, with fewer than2222 ribs,ribs, which are irregular N. parvavellana

Ferrero Genus Pusula Jousseaume, 1884b Mortara et al., p. 155.

Distribution — Pliocene of the western Mediterranean. Type species — Cypraea radians Lamarck, 1810, by

original designation.

Materialstudied — From the Velerin Conglomerates, 30

from — specimens Colin), 4 specimens Colin); Discussion Although some members of the genus (BLP (DFB Colli Astesi display all typical features (see Cate, 1979), the position of (Italy), Upper Piacenzian, 1 specimen

the aperture and the depression of the base are variable. In (BS.043.13.001).

our view, the only constant characters that clearly separate

this from other triviids the dorsal which Measurements — Maximum length and width to 13.6 group are sulcus, up

mm and 9.9 respectively, but on 12.0 mm and in most cases is wide and smooth, with the ribs sharply mm, average

8.6 L/W ratio 1.39. bisected and forming tubercles, spinous in some species, as mm, respectively;

well as the fewer, coarser ribs.

Description — Shell large, solid and elongated oval. Spire

covered by callus. Body whorl elongated-globose and

Pusula subpediculus (d’Orbigny, 1852) rounded, almost 95% of total height. Terminals rounded,

anterior produced, posterior weakly so. Dorsum rounded,

Figures 3/6a-c, 7a, b; 5/2a-d completely covered by 20-22 strong, irregular ribs, which

are somewhat wavy and frequently interrupted on both

1847 L. ? 47. dorsal sides. A smooth dorsal sulcus extends Cypraeapediculus Sismonda, p. strong, wide,

* 1852 170. Cypraea subpediculus d’Orbigny, p. longitudinally across the centre ofthe dorsum, bisecting all

1894 Pusula pediculus var. subpediculus (Orb.) — Sacco, of the ribs, the latter forming prominent tubercles adjacent 54, 3, 49. p. pi. fig. to the furrow. Base almost straight, with the posterior

1932b Trivia 1852 — subpediculus Orbigny, Schilder, p. terminal recurved. The aperture is wide, becoming even 105. wider in the anterior portion. Outer lip somewhat broad-

1941 Trivia (Sulcotrivia) subpediculus Orbigny ‘52 — ened, convex, widest in mid-portion, becoming narrower 74. Schilder, p. towards the terminals and regularly rounded. Outer margin 1971 Trivia (Sulcotrivia) permixta subpediculus Orbigny, angularly callused. The lip bears 13-16 strong, equal teeth. 1852 — Schilder & 18. Schilder, p. Siphonal and anal canals followthe shell profile. 1984 Pusula pediculus var. subpediculus Sacco, 1894 — - 100-

Figure 3. Triviid gastropods from the Zanclean of Estepona (Spain) (photos B. Landau)

= and la-c, 2a, b Niveria parvavellana (Sacco, 1894), Velerin Conglomerates, original size 6.9 mm 7.2 mm, respectively (BLP Colin).

5a-c Velerin Velerin Sands and Velerin Carretera size = 5.4 3a-c, 4a-c, Cleotrivia esteponica n. sp., Conglomerates (3), (4) (5), original

mm, 5.5 mm and 5.2 mm, respectively (BLP Colin).

6a-c, 7a, b Pusula subpediculus (d’Orbigny, 1852), Velerin Conglomerates, original size = 14.2 mm and 12.5 mm, respectively (BLP

Colin). - 101 -

Figure 4. Triviid gastropods from the Zanclean of Estepona (Spain) (drawings D. Fehse):

1 HNC HNC 57307 Velerin a-d, 3a-d Cleotriviaesteponica n. sp.; 57306 ( 1, holotype) and (3, paratype), respectively, Conglomerates,

original size = 7.0 mm and 5.7 mm, respectively.

2a-c Triviafrigida Schilder, 1932b, CS 221 (holotype), Altavilla(Italy), UpperPiacenzian, original size = 12.0 mm. - 102-

Figure 5. Triviid gastropods from the PiacenzianofItaly (photos D. Ormezzano):

la-c Niveria dorsolaevigata (Cocconi, 1873),BS.043.12.017, Colli Astesi (Italy), UpperPiacenzian, original size = 18.8 mm.

2a-d Pusula subpediculus (d’Orbigny, 1852), BS.043.13.001, Colli Astesi (Italy), Upper Piacenzian, original size = 14.2 mm.

3a-d Trivia perobsoleta Sacco, 1984, BS.043.12.007 (holotype), Ventimiglia (Italy), Lower Piacenzian, original size = 14.7 mm. - 103 -

Figure 6. Triviid gastropods from the Mioceneand Pliocene ofItaly (photos D. Ormezzano [1-4] and D. Fehse [5-8]):

la-d Niveria avellanula (Sacco, 1894),BS.043.12.015 (lectotype), Zinola(Italy), Lower Piacenzian, original size = 10.4 mm.

2a-c Niveria cf. avellana (J, de C. Sowerby, 1823), BS.043.12.015/01,Zinola(Italy), Lower Piacenzian, original size = 10.4 mm.

3a, c Niveria cf. dimidiata (Bronn, 1831), BS.043.12.017/01(paratype of N. dorsolaevigata), Colli Astesi (Italy), Upper Piacenzian.

original size = 9.0 mm (compare Smriglio et al., 1998, text-figs 35-38).

4a-d Niveria parvavellana (Sacco, 1894), BS.043.12.010 (lectotype), Rio Torsero (Italy), Lower Piacenzian, original size =6.6 mm.

5a-c Trivia acuminata Schilder, 1932a, CS 3411 (holotype), Empoli? (Toscane, Italy), Lower Piacenzian, original size = 7.5 mm.

6a-c Triviasubdilatata Schilder, 1941, CS 5063 (holotype), Castell’Arquato (Italy), Lower Piacenzian, original size = 6.4 mm.

7a-c Trivia nanaSchilder, 1941, CS 5084 (holotype), Montegibbio(Modena, Italy), ‘Tortonian’(Middle Miocene), original size =

6.0 mm.

8a-c Niveria selmae (Boettger, 1901), CS 5076, Montegibbio(Modena, Italy), ‘Tortonian’(Middle Miocene), original size = 4.9 mm. Columellaconvex, tapering inwards, with the carinal assumed that the two for ridge groups were separated a long time

limited to the fossular Columella region. bearing 14-16 and members of the genus emigrated to the Paratethys

ribs. Fossula and delimitedfrom the the weakly concave poorly during Late Oligocene or Early Miocene (Fehse, 1999,

rest of the columella. A colour pattern is preserved on all p. 8).

specimens; it consists ofa beige background on the dorsum

and white base and Darker brown a lip. patches are super-

the of the at either imposed on beige background dorsum, Cleotrivia esteponica n. sp. side of the sulcus. The patches are arranged in three

at either of the groups, extremity dorsal sulcus and mid- Figures 3/3a-c, 4a-c, 5a-c; 4/la-d, 3a-d dorsally.

Types — Flolotype is HNC 57306;paratype is HNC 57307.

Range ofvariation — This species is characterised by the

elongated shape, flattened base, wide aperture and the wide Type locality and horizon — Velerin Conglomerates, dorsal sulcus. There is little variationamongst the popula- Lower Pliocene (Zanclean), Estepona, Spain. tion from Estepona, with some of the shells slightly more

less than the — inflated, elongated norm. Derivation of name Namedafter the city of Estepona.

— This — Discussion species is quite differentto N. permixta Distribution Known only from Velenn (Estepona, which that Schilder Schilder (see above), again suggests & Spain).

(1971) did not see any of the Pliocene triviids in Sacco’s

Collection. It differs from that species in having a much Material studied — from the Velerin Conglomerates, 9 smooth stronger, dorsal sulcus which completely bisects all specimens (BLP Colin), 4 specimens (DFB Colin); from the ribs. There is dorsal callus in no P. subpediculus, the the Velerin Sands, 7 specimens (BLP Colin); from the aperture is much wider and the fossula less developed. Velerin Carretera, 6 specimens (BLP Colin).

The present taxon more closely resembles the Pliocene-

Recent P. which has — pediculus (Linne, 1758), finer, more Measurements Maximum length and width up to 5.4 numerous and irregular ribs, wavier with a number mm and 4.4 but larger mm, respectively, on average 5.2 mm and of The is interruptions. apertural lip broader, aperture 4.3 mm, respectively; LAV ratio 1.20. subcentral and the anal canal indented. Although Sacco

ancestor of — (1894, p. 54) proposed subpediculus as an Description Shell very small, relatively fragile, and this is both pediculus, impossible, as occur in the Lower spherical. Spire covered by callus, faintly visible beneath

Pliocene, subpediculus in the Mediterraneanandpediculus 3 vertical riblets extending upwards from the posterior in the Caribbean (BLP Colin). A possible ancestral species terminal. Body whorl globose and rounded, about90% of to both may be P. parcicosta (Bronn, 1862) from the total height, with the terminals strongly produced and

Middle Miocene of the Islands flattened. Canary (Schilder, 1928, p. Dorsum evenly rounded, without a true dorsal

275, text-figs 7, 8). sulcus, but a weak dorsal depression, which modifies the

ribs. Dorsum completely covered by 20-22 strong, irregular

ribs, with interruptions. Where the ribs cross the dorsal

Genus CleotriviaIredale, 1930 depression, they are usually modified; interrupted, nar-

rowed and occasionally unaltered. Base and terminalsare

Type species — Cypraea pilula Kiener, 1843, by original evenly convex. Aperture curved, slightly wider in the

designation. anterior portion. Outer lip broad, widest in mid-portion,

becoming narrower towards the terminals. The lip is evenly

— The characteristic Discussion features of this genus are rounded and the outer margin weakly callused. The lip the small, globular or subglobular, bulbously inflated shell, bears 15-18 fine, equal teeth. Siphonal and anal canals the of dorsal sulcus of variable which wide and indented. Columella presence a strength, convex, tapering steeply influencesthe dorsal and short blunt termi- inwards, hardly ribbing bordered internally by a very weak to subobsolete nals. There are two distinct groups within the genus. The carinal ridge, bearing 13-15 ribs, which continue onto the

Recent Indo-Pacific taxa, in which the sulcus is deeply carinal ridge, where they do not become significantly grooved, extending the whole length ofthe dorsum, and the stronger. The carinal ridge protrudes in the fossular region. Atlantic-Caribbean forms, in which the sulcus is most Fossula concave, not clearly delimitedfrom the rest of the strongly developed mid-dorsally, but shorter, not extending columella. all the way between the terminals. In the latter group the sulcus only slightly depresses the dorsal ribbing. Further- Range ofvariation — This species is characterised by the the Atlantic-Caribbean less more, extant species are glo- small size, greatly inflated shell, poorly developed dorsal bose than their Indo-Pacific In the sulcus counterparts. European and short, blunt terminals. The shape is fairly con-

the have combination of the shells Neogene, species a features, a stant, varying mainly in details of the ribs, num- shell like the Recent Indo-Pacific but globular taxa, only a ber of interrupted ribs and character as they cross the weak dorsal sulcus as seen in the Caribbean It is species. sulcus, as well as in the strength of the dorsal depression. - 105 -

Discussion — This is the first member ofthe genus Cleo- graphs ofthe type specimens in Sacco’s Collection. John

be described from the fossil trivia to European record. It is Cooper (formerly The Natural History Museum, London)

similar C. 1999 from the of is thanked to werneri Fehse, coast West for his help in chasing rare literature, while

but also in the Alboran Sea Robert Africa, occurring [Giannuzzi- Marquet (Antwerp) commented on an earlier type- Savelli et al., 1996, p. 160, fig. 666a-c, as Pusula can- script.

didula Cleotrivia (Gaskoin, 1836)]. esteponica n. sp.

differs from the extant species in being more inflated, with

a more elevated dorsum and a weaker dorsal sulcus. In References

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