Cainozoic 2002 Research, 1(1-2)(2001), pp. 111-120,November

Granulina (, ) from the Pliocene of

Málaga (southern Spain) with descriptions of four new species

³ Rafael La+Perna¹,Bernard Landau² & Robert Marquet

1 di di E. Orabona 4,1-70125 e-mail: Dipartimento Geologic e Geofisica, Universita Bari, via Bari, Italy: [email protected]

2 International Health Centres, Avenida Infante D. Henrique 7, Areias Sao Joao, P-8200 Albufeira, Portugal; e-mail:

bernie. landau@btinternet. com 3 Institut royal des Sciences naturelles de Belgique, do Constitutiestraat 50, B-2060 Antwerp, Belgium: e-mail:

Robert.Marquet@antwerpen. be

Received 3 July 2001; revised version accepted 25 November 2001

that nine of the Granu- The Lower Pliocene shelfdeposits crop out near Estepona (Málaga) have yielded species marginellid

G. clandestine lina Jousseaume, 1888, six of which [G. iberica n. sp., G. detruncata n. sp., G. malacitana n. sp., G.? longa n. sp.,

1992 (Brocchi, 1814), and sp. sensu Gofas, 1992] are extinct, while G. marginata (Bivona, 1832), G. boucheti Gofas, and G. guttula La Perna, 1999 still occur in the Mediterranean to this day. Differences in species composition in the Pliocene of

Italy are thoughtto be due mainly to the typically restricted distribution and endemicity of Granulina and ofmarginellids in general.

Affinities between Pliocene representatives of Granulina from the Mediterranean and Recent West African species confirm the

‘warm’ of this the ‘warm-temperate’ to character genus. Subsequent to Middle-Late Pliocene and Pleistocene cooling events, the

species with warmer affinity mostly went extinct, the genus evolved a more temperate character and diversity amongst shallow-water

species decreased.

Key words: Pliocene, Gastropoda, Marginellidae, new species, Spain, Mediterranean, systematics, palaeobiogeography

Introduction marked ecological trend to the deep shelf and upper slope has recently been noted (La Pema, 1999). How-

Until a few years ago, two fossil/extant Mediterranean ever, the ‘warm-temperate’ to ‘warm’ biogeographic

species of the genus Granulina were listed in the litera- affinity of this genus is beyond doubt. In the eastern At-

ture, viz. G. occulta (Monterosato, 1869) and G. clan- lantic its northern limit is along the Iberian coasts, which

several recent have been de- destina. However, papers holds true for both the shallow- and deep-water species.

voted to extant species from the Mediterraneanand Gi- The present paper attempts to fill the gap in our

braltar (Gofas, 1992; Smriglio & Mariottini, 1996, 1999; knowledge of Pliocene species of Granulina, on the ba-

Smriglio et al., 1998; La Pema, 1999) and to Pliocene sis of material collected from Lower Pliocene deposits

and Pleistocene representatives from the Mediterranean exposed near Estepona (Malaga Basin, southern Spain).

the These data of interest. do (La Pema, 1999, 2000). Contrary to previous views, new are particular Not only

genus Granulina has turned out to be well diversified in they increase our knowledge of the genus, they also pro-

the Recent Mediterranean (12 species), three of which vide additional data on the geographical position, close

are endemic to the Gibraltar area. Similarly, during the to the Strait of Gibraltar, ofa gateway that has controlled

Pleistocene, it was well represented, with four extinct biotic exchanges between the Atlantic and Mediterranean

and five extant forms having been recorded from Italy. since the Late Cainozoic.

Numerous Pliocene Mediterraneanspecies probably still For stratigraphic, tectonic and palaeogeographical be four known from the Pliocene data situated remain to studied; are on the Malaga Basin, in the western sector

of of which survives the of the of the Betic reference is Italy, one to present day. Com- Internal Zones Cordillera,

parably, many species have recently been recorded from made to Sanz de Galdeano & Lopez Garrido (1991).

off the West African coast and from the Ibero-Moroccan During the Tortonian (Miocene), an extended seaway

Gulf south to the Guinea Gulf, including the Canary and existed from the Mediterranean (Malaga area) to the At-

Cape Verde Islands (Fernandes, 1987; Gofas & Fernan- lantic, through the Guadalquivir and Ronda basins. In

des, 1988; Fernandes & Rolan, 1991; Gofas, 1992; Pin & contrast, the Pliocene basin was much smaller, extending

Boyer, 1995; Rolan & Fernandes, 1997; Boyer & Rolan, inland for c. 30 km from the present-day Malaga- Torre-

1999; Smriglio et al., 2001; Boyer, 2001). molinos area. Pliocene sediments also crop out along the

Although most species of Granulina, or at least the coast, south to Estepona. This series consists of con-

better known from shallow-water ones, are settings, a glomerates and sands, which are laterally replaced by - 112-

bluish marls and in turn overlain emmost Mediterranean and Atlantic. The fol- grey clays, by yellowish adjacent

white sands, to a maximum thickness of c. 400 m in the lowing taxa are found both at Velerin and in coeval At-

easternmost areas. The age ranges from Early-Middle to lantic deposits at Huelva (Spain): Crepidula lucenica

early Late Pliocene. The uppermost Pliocene and Qua- Landau, 1984, Cyllene (Cyllenina) lucenensis Landau &

ternary sediments are terrestrial. Marquet, 1999 and Cymbium ibericum Landau & Mar- Francisco studied the bivalves from well is Lozano (1998) quet, 2000. More interesting, as as unexpected,

several in the associations of relict outcrops Malaga area. Most the occurrence Miocene species, such as Ver- proved to be indicative of infralittoral and circalittoral micularia milleti (Deshayes, 1839), Euthriofusus burdi-

environments with coarse- to fine-grained substrates. galensis (Defrance in de Blainville, 1824),

Only in recent years have these extremely fossil-rich de- () eratoformis Hoemes & Auinger, 1880, and

in the with the recorded posits received attention literature, em- Perrona jouanneti (Desmoulins, 1842), not

phasis on molluscs (e.g., Vera-Pelaez et al., 1995; Lo- previously from other Lower Pliocene deposits of the

zano Francisco, 1998; Muniz-Solis, 1999; Landau & Mediterranean.

Marquet, 1999, 2000). Not only are they exceptionally with 700 of rich, over species gastropod (BL, pers. obs.),

but the unique geographical position of these deposits Material and methods

also is reflected in the faunal composition. As could be

of Five Rio Velerin expected, there are many species typical an Early localities, del Padron, Carretera, Vel-

Pliocene Mediterraneanfauna, as found in Italy. There is erin Antena, ‘Velerin Conglomerates’ and Parque Antena

also a strong Atlantic influence with typically northern (Table 1), between 5.5 and 9 km northeast of Estepona

species, such as Scaphella lamberti (J. Sowerby, 1816), (see map in Vera-Pelaez et al., 1995), have been sampled

as well as a more southerly influence, exemplified by the by two of us (BL, RM). At each of these, bulk samples

recent discovery of the gastropod genus Cymbium at have been taken and sieved on a 1 mm, and from each, at

Velerin (Landau & Marquet, 2000). Amongst a fairly least 5 kg of residue has been sorted out under a binocu-

large number of new species, most of which have yet to lar microscope.

to to west- be described, a few appear be endemic the

SPECIES VCa I VA I PA I RP I VCo

Granulina iberica ?9 n. sp. common present

Granulina detruncata n. sp. present present

Granulina malacitanan. sp. present present present

Granulina? longalonga n.n. sp. present present

clandestina Granulina clandestina present present

Granulina marginata present

Granulina boucheti present present

Granulina guttula present

Granulina sp. common

Table 1. in VCa = VA =Velerin PA = RP Distribution of Granulina the Malaga area; VelerinCarretera, Antena, Parque Antena,

Rio del Padron, VCo = ‘Velerin Conglomerates’.

At Velerin Carretera, greyish sandy clays and clayey exposed, from a clayey sediment with scattered, well-

sands are exposed in a road-cutting about 4 m in height, preserved shells.

with scattered of The Rio del Padron is the left bank of a fauna, consisting mainly small, su- outcrop on a

perbly preserved molluscs. Velerin Antena is a disused river bed, mostly dry in summer, exposing a sequence of

the Velerin Carretera. >15 it quarry, exposing same lithology as m; appears to represent an upward-shallowing The molluscan fauna generally is small sized and is trend with the main body of sediment consisting of

dominatedin number of specimens by bivalves, many of clayey sand, with a sparse fauna, yielding white corals.

which are still articulated. Most species belong to the The molluscan fauna, however, occurs at a level c. 5 m

families Yoldiidae, Nuculanidae and Semelidae, which below the top of the section, containing coarse-grained

indicate a relatively deep-water (deep circalittoral) as- sand with shells and occasional pockets of gravel. Shells

semblage, which has experienced little or no transport. are often abraded and bivalves occur mainly disarticu-

Parque Antena is a temporary building site, lithologi- lated, and are often decalcified.

cally and palaeoecologically closely similar to the se- The ‘Velerin Conglomerates’ are exposed close to

at Velerin and Velerin Velerin and form the bank quence seen Carretera Antena. Antena, a steep valley on right taken below the level of small seasonal river. Samples have been c. 5 m highest a -113 -

= 1-4. Granulina 2 - 1ST H 3.4 3 - Figures iberica n. sp., from Veledn Carretera; 1, holotype (IRScNB 6422), mm; paratype

= = (IRScNB 1ST 6423), H 3.4 mm; 4 - paratype (IRScNB 1ST 6424), H 3.3 mm (SEM).

1ST H = Figure 5. Granulina aff. iberica n. sp. (IRScNB 6439), from ‘Velerin Conglomerates’; 2.8 mm (SEM).

= 6-8. 7 - 1ST H - Figures Granulina detruncata n. sp., from Velerin Carretera; 6, holotype (IRScNB 6427), 2.8 mm; 8 paratype

(IRScNB 1ST 6428), H = 2.5 mm (SEM).

= 9-11. Granulina malacitana n. from ‘Velerin 10 - 1ST - Figures sp., Conglomerates’; 9, holotype (IRScNB 6430), H 2.1 mm; 11

paratype (IRScNB 1ST 6431), H = 2.0 mm (SEM).

1ST H = Figures 12,13. Granulina? longan. sp., from Parque Antena, holotype (IRScNB 6433), 2.9 mm (SEM).

The conglomerates contain coarse-grained, whitish to conglomerates seem to represent a shingle beach deposit, brown sand which may be indurated in parts, containing containing an admixture of faunal elements of varying

mainly eroded boulders up to 1 m in diameter. In some depths and environments.

places rare pockets of yellowish brown sand occur. The Over 160 shells of Granulina have been examined,

molluscan fauna rich and is dominated of which have been identified The is very by gastro- most to species. re-

which attain size. Some bivalves either pods, may a large are maining specimens were too poorly preserved or but show but immature identified. articulated, most are not. Some specimens too to be Descriptions below are few of others affected. based traces transport, are strongly In on fully-grown shells. All types and illustrated

the eastern part of the same hill exposure, conglomerates specimens are housed in the collections of the Institut

contain small amounts of sand and barely eroded schist royal des Sciences naturelles de Belgique (Brussels).

slabs; only very few molluscan species are present. These -114-

Figure 14. Granulina clandestina(Brocchi, 1814) (IRScNB 1ST 6434), from ‘Velerin Conglomerates’; H = 6 mm (SEM).

Figure 15. Granulina sp. (IRScNB 1ST 6438), from Rio del Padron; H = 2.2 mm (SEM).

17. touched 1992 1ST from ‘Velerin H = 2.1 and 2.4 Figures 16, Granulina Gofas, (IRScNB 6436) Conglomerates’; mm, respec-

tively (SEM).

Figure 18. (Bivona, 1832) (IRScNB 1ST 6435), from ‘Velerin Conglomerates’; H = 2.1 mm (SEM).

19. 1ST from H = 1.9 Figure Granulina malacitanan. sp., paratype (IRScNB 6432), Velerin Carretera; mm (SEM).

= Figures 20-22. Granulina guttula La Pema, 1999; 20 - IRScNB 1ST 6437, from Velerin Carretera; H 2.3 mm; 21, 22 - paratypes

84 H = 2.2 2.1 (UPMC collections), southeast Tyrrhenian Sea, depth m; and mm, respectively (SEM).

and of Granu- — For descriptive terminology Type species Marginella pygmaea Issel, 1869, by

lina, we follow Coovert (1988a), Gofas (1992), Coovert monotypy.

& Coovert (1995) and La Pema (1999). As previously

discussed by La Pema (1999), the traditional placement

Granulina iberica n. of the genus Granulina in the family Marginellidae, sp. Figures 1-4 rather than in the as proposed by Coovert &

Coovert (1995), is here maintained.

Type locality — Velerin Carretera, Estepona (province

of Malaga, Spain).

Stratum - Zanclean (Lower Pliocene). Systematic palaeontology typicum

Type material — Holotype is IRScNB 1ST 6422; para-

types are IRScNB 1ST 6423-6426. = Abbreviations — H = shell height; D maximum di-

Material examined — 86 specimens from Velerin Car- ameter; IRScNB = Institut royal des Sciences naturelles retera, including the types; 28 specimens from Velerin de Belgique (Brussels); UPMC = University Palaeon- Antena; 3 specimens from ‘Velerin Conglomerates’; all tological Museum, Catania; RM = R. Marquet Colin; BL BL and RM collections, except for type specimens. - B. Landau Colin.

Etymology — Latin ibericus, in reference to the Iberian

Peninsula.

Description — Shell small, with immersed , Class Gastropoda Cuvier, 1797 ovoid-pyriform in shape, maximum diameter posterior to Order Thiele, 1929 mid-height of shell, H/D 1.18-1.36.Posterior end slightly Family Marginellidae Fleming, 1828 obliquely truncated. Siphonal notch distinct. Lip strongly Genus Granulina Jousseaume, 1888 thickened in fully-grown stage, slightly bevelled posteri- -115-

denticulations but well defined. Four all orly. Lip fine, Conglomerates’; material, except type specimens, in blade-like columellar plications, slightly excavated inside BL and RM collections. aperture, the uppermost notably small. Parietal callus Etymology — Latin detruncatus meaning beheaded, in

sinus. allusion the of wide, distinct, forming a deep narrow Aperture to clear-cut, ‘upper’ part the shell.

wide. Surface — relatively smooth, shiny, except for faint Description Shell minute, with immersed spire,

growth striae. Measurements of are H 3.4 holotype mm, ovoid-rhomboid in shape, maximum diameter slightly D 2.5 and other material: H 2.7-3.4 mm; paratypes mm. posterior to mid-height of shell, H/D 1.35-1.48.Posterior

Distribution — Known from the Lower Plio- end truncated and covered exclusively strongly by a thick, slightly

cene of the of Granulina into Malaga; commonest species at stepped callus, growing a thin parietal ridge almost the localities sampled. inside the aperture and extending to columellar plica-

Remarks — The degree of thickening of the outer lip tions. Siphonal notch distinct. Lip strongly thickened, varies from weak with the denticulations to strong, markedly bevelled posteriorly, forming a wide, flatfish in the former poorly developed stage (Figures 3, 4). This and slightly rough surface with a somewhat sharp outer is related the character clearly to lip thickening in mar- border. Lip denticulations fine but well defined. Four

ginelliform gastropods, occurring late in ontogenetic notably strong columellarplications, obscurely excavated development & Coovert, inside (Coovert 1995; Nehm, 2001). aperture, the upper two notably smaller and con- Live well fossil shells of specimens as as empty extant or nected by a thread-like callus. Parietal callus fairly dis-

Granulina are found with usually a well-developed lip tinct, forming a wide, shallow sinus. Aperture narrow. its , strength remaining fairly constant within a spe- Surface smooth, but crossed by impressed growth striae. cies. Subadult specimens showing initial of Measurements of stages lip holotype are H 2.8 mm, D 1.9 mm; less found than shells. thickening are commonly juvenile paratypes and other material: H 2.3-2.5 mm.

This would suggest a fairly rapid lip thickening in the Distribution — Known exclusively from the Lower Plio- adult whereas in G. iberica the stage, lip thickening was cene ofMalaga.

slow for the Size also is Remarks — The probably unusually genus. strong posterior truncation, rhomboid somewhat variable, yet there is no relationship to lip outline and markedly thickened and wide lip clearly dis- thickness. tinguish this species from all other fossil and Recent

A shell from the single (Figure 5) ‘Velerin Conglom- European Granulina. However, it may be compared with erates’ differs in being more solid, more ovate and in the shallow-water West African G. ghanensis Rolan & thicker having a notably parietal callus, and is here re- Fernandes, 1997. Specimens of G. ghanensis from Mi- ferred G. aff. iberica. amia available to as (Ghana) to us are notably smaller, with a this Although species is clearly distinct from other rounded posterior rostration and small, close-set plica-

fossil and extant to a certain it excavated inside the European species, extent tions, more deeply aperture by a resembles the deep-water Mediterranean G. tenuilabiata shallow sulcus. This makes them appearalmost ‘double’, La 1999 in and from which it differs Pema, shape size, a character also present in other species (Gofas, 1992; La in thicker mainly having a lip (when fully-grown), a Pema, 1999, 2000). These two species have the follow- much sinus the callus and distinct features in the narrower on parietal a ing common: ‘rhomboidal’ shell outline, posterior truncation. Other species reminiscent of G. the wide lip surface, the well-developed posterior callus, iberica and G. the Pleistocene tenuilabiata, e.g., G. ro- notably thick in G. detruncata, thinner in G. ghanensis sarioi La 1999 and G. ovulina Pema, (Monterosato, but forming a shallow pad near the posterior end, and the and the 1881), extant G. gofasi Smriglio & Mariottini, thin parietal callus ridge running from the posterior cal-

1996, all have particularly large (3.0-3.5 and lus and the two to mm) glo- joining upper three plications. Another bose with shells, near-absent to fine lip denticulations species apparently rather similar to G. detruncata and G. which and lack any parietal callus ridge (see below). It is ghanensis is G. africana Gofas, 1992, from Senegal and worth that these have had the mentioning species or notably Ivory Coast. In Granulina canariensis, described

to deep-water (circalittoral epibathyal) ranges (La Pema, recently from the Canary Islands (Boyer, 2001), also 1999, table Granulina iberica 1). probably had a mainly size, shape and apertural features are similar to those circalittoral distribution well. in G. but as seen detruncata, the former has a narrower lip and a rather sharp posterior end.

An anomalous number of plications, ‘about 5 small

denticles’, was recorded in the original description of G.

Granulina detruncata n. sp. ghanensis (Rolan & Fernandes, 1997) and ‘three oblique Figures 6-8 and 13-15 fine folds pleats on the upper part’ in G. oca- rina (Fernandes, 1987), from the Cape Verde Archipel-

Type locality — Velerin Carretera, of Estepona (province ago. Topotypes G. ocarina show a thin callus ridge ofMalaga, Spain). joining the uppermost columellar folds, similar to that in

Stratum — Zanclean typicum (Lower Pliocene). G. ghanensis and G. detruncata. This ridge is a common

Type material — is IRScNB 1ST Holotype 6427; para- feature in Granulina (Coovert, 1988b; Coovert & Coo-

are IRScNB 1ST 6428,6429. types vert, 1995; La Pema, 1999, 2000) and, whenever slightly

Material — examined 6 specimens from Velerin Carre- the of than four tuberculate, may give appearance more including the 2 from ‘Velerin tera, types; specimens columellar-parietal plications. -116-

Granulina malacitana similar in and size G. but data n. sp. shape to malacitana, no

the micro-ornament ofthe former available. Figures 9-11, 19 on are

Type locality — ‘Velerin Conglomerates’, Estepona

(province ofMalaga, Spain). Granulina? longa n. sp.

Stratum typicum — Zanclean (Lower Pliocene). Figures 12, 13

Type material — Holotype is IRScNB 1ST 6430; para-

type are IRScNB 1ST 6431, 6432. Type locality — Parque Antena, Estepona (province of

Material examined — 12 specimens from ‘Velerin Con- Malaga, Spain).

glomerates’, including the types; 3 specimens from Vel- Stratum typicum — Zanclean (Lower Pliocene). erin Carretera; 3 specimens from Rio del Padron; all Type material— Holotype is IRScNB 1ST 6433.

material, except for type specimens, BL and RM collec- Material examined — 1 specimen (holotype) from Par- tions. que Antena; 3 specimens (1 juv.) from Velerin Carretera;

— from the all Etymology Spanish adjective malacitana, material, except for type specimen, in BL and RM

meaning ‘of Malaga’. collections.

Description — Shell minute, with immersed spire, mark- Etymology — Latin longus, meaning elongated. edly ovoid in shape, maximum diameter at mid-height of Description — Shell small, with immersed spire, ellipti- shell to slightly posterior, H/D c. 1.4. Posterior end ob- cal-subcylindrical, with maximum diameter at mid-height

scurely truncated to regularly rounded, with a distinct of shell, H/D c. 1.65. Posterior end somewhat truncated.

callus producing a shallow rounded relief. Siphonal Siphonal notch fairly well developed. Lip moderately notch weakly developed. Lip thickened with fine den- thickened, roughly rectilinear in its median part. Den- ticulations. Four moderately strong columellar plications ticulations very fine, indistinct. Four blade-like columel-

excavated inside the two lar not excavated inside Parietal slightly aperture, upper notably plications, aperture. smaller. Parietal callus callus not particularly wide and poorly indistinct. Aperture moderately narrow. Surface

demarcated, forming a wide, shallow sinus. Thin parietal smooth, shiny, except for very fine growth striae. Meas-

callus ridge inside aperture, with a slightly tuberculate urements of holotype are H 2.95 mm, D 1.75 mm.

appearance. Aperture narrow. Surface smooth, but Distribution — Known exclusively from the Pliocene of

crossed by fine axial striae, mostly appearing as thin, Malaga.

darker and lighter, regularly spaced bands lacking relief. Remarks — The markedly subcylindrical shape and

Measurements of holotype are H 2.1 mm, D 1.6 mm; straight lip make allocation of this species within the paratypes and other material: H 1.8-2.2 mm. genus Granulina rather doubtful. In addition, there are

Distribution — Known only from the Lower Pliocene of no fossil or extant European marginelliform gastropods

Malaga. that may be compared to the present species, except for

Remarks — The main distinctive features of G. malaci- ‘Marginella’ ovulaeformis Seguenza, 1879, a poorly the marked shallow callus known Pleistocene commented tana are egg shape, posterior species briefly on by La relief, small size and the axial ‘ornament’. This last fea- Pema (1999), which is much larger and pyriform in

ture is reminiscent of the axial pattern seen in G. van- shape. Both species are reminiscent of representatives of

hareni van Aartsen et al., 1984, endemic to Gibraltar the genus Ovaginella Laseron, 1957, from New Zealand

(Gofas, 1992), in which the axial lines are slightly raised and Australia(Coovert & Coovert, 1995), but the present

whereas material is limited examine inner shell features. giving a ‘chagrine’ appearance (Gofas, 1992), too to shallow and of chromatic whorls while they are very mostly nature Internal are partially resorbed in Granulina,

(darker and lighter lines, see Figure 19) in G. malaci- they are unmodifiedin other marginellids.

tana. The true nature of this pattern is not understood.

Under high magnification, very fine granulations are seen

to cover the shell, giving the surface a satin appearance. Granulina clandestina (Brocchi, 1814)

Boyer & Rolan (1999) illustrated the same microsculp- Figure 14

tural pattern in G. vanhareni and in a new species, G.

* 1814 Voluta clandestina 11. fernandesi, described from the Cape Verde Islands. As Brocchi, p. 642, pi. 15, fig.

1992 — is Granulina clandestina (Brocchi, 1814) Gofas, p. 5, seen in SEM images, the pattern of granulation some- fig. 3. what different in the two. In addition, G. malacitana

— La 2000 Granulina clandestina (Brocchi, 1814) Pema, p. does not differ markedly in morphology from G. van- 54, figs 8-10. hareni, both species being notably small, egg-shaped and

with a small posterior callus. However, the latter has a Material examined — 1 specimen from ‘Velerin Con- somewhat cylindrical shape and a marked anterior (si- glomerates’ (IRScNB 1ST 6434); 1 specimen from Vel- phonal) slope break. Granulina fernandesi also is par- erin Carretera (BL colln).

ticularly small and somewhat similar in shape to G. van- Distribution — Pliocene, Mediterranean.This species is hareni. also known from the Pliocene of Italy (Gofas, 1992; La Another from G. species West Africa, mauretanica Pema, 2000), from where it was first described. Pleisto- shows remarkable similarities in shell and Gofas, 1992, cene records from Italy are in need of confirmation (La soft parts to G. vanhareni (see Gofas, 1992), and is also Pema, 1999). - 117-

Remarks — Its ‘cordiform’ shape is unique amongst fos- Granulina guttula La Pema, 1999

sil and extant Mediterranean species. However, G. clan- Figures 20-22

destina is closely similar to the West African G. parilis

La Gofas & Fernandes, 1988 from the Guinea Gulf, which *1999 Granulinaguttula Pema, p. 39, figs 13-16.

differs by having a more ovate and slightly subcylindrical

Material examined — from Velerin shell, with a less marked ‘cordiform’ outline. Carretera, 1 speci-

in men (IRScNB 1ST 6437), 5 specimens BL colln; type

material is housed in the UPMC collections.

Distribution — Granulina marginata (Bivona, 1832) Pliocene-Recent, Mediterranean, previ- known Figure 18 ously only from the type locality, the eastern Tyr-

rhenian Sea, at a depth of84 m.

*1832 Volvaria 5. — marginataBivona, p. 24, pi. 3, fig. Remarks The present shells differ from the types in

1992 Granulina 1832) — Gofas, 6, marginata (Bivona, p. being slightly larger (maximum height 2.7 mm v.s- 2.25 figs 5-8, 25. mm in types) and slightly less rostrate. However, in view

1999 — La Granulina marginata (Bivona, 1832) Pema, p, of our comparatively poor knowledge of G. guttula and 38, figs 36-42. the limited fossil material, we prefer to consider them 2000 Granulinamarginata— La Pema, figs 14, 15. conspecific. A certain difficulty in referring some fossil

shells of Granulina to extant species was stressed by La Material examined — from ‘Velerin Conglomerates’, 1 Pema (1999), a fact related to the evolutionary pattern of specimen (IRScNB 1ST 6435), and 2 specimens in BL this genus (see below). colln. In the original description, a noteworthy feature of Distribution — Pliocene-Recent, Mediterranean and this species was not recorded, i.e. the glossy and slightly Gibraltar. This species is also known from the Pliocene iridescent the fact that the appearance, despite type mate- and Pleistocene of Italy (La Pema, 1999, 2000). In post- rial is not particularly fresh. This is in contrast with the Pleistocene times, it is the commonest and most widely more usual opaline in shells of the shallow-water of the in the appearance genus distributed, species genus Granulina. Two recently described species from north- Mediterranean(Gofas, 1992; La Pema, 1999). west Africa, G. cerea and G. crystallina (Smriglio et al., Remarks — The material examined is limited and poorly characterised 2001), are by a similar, particularly trans- preserved. Identificationis based on size and shell shape, shell parent wall, giving a glossy or waxen appearance. as most apertural features are obscured by indurated ma- different In addition, they are not markedly from G. gut- trix. Granulina marginata also has a parietal callus tula in size, shape and apertural features, differing ridge, which is thin, often indistinct and faintly tubercu- mainly in being more inflated and in having the maxi- late. mum diameter located more posteriorly.

Granulina boucheti Gofas, 1992 Granulina sp. Figures 16, 17 Figure 15

26. *1992 Granulina boucheti Gofas, p. 10, figs 9, 10,

1999 — La compare Granulina boucheti (Gofas, 1992) Pema, p. 38,

1992 Granulina — 4. figs 38-40. sp. Gofas, fig.

2000 — La Granulina sp. Pema, p. 38, p. 54, figs 11-13.

Material examined — 3 specimens from Velerin Carre-

Material examined— from Rio del Padron, 1 specimen tera; from ‘Velerin Conglomerates’ 1 specimen (IRScNB (IRScNB 1ST 6438) and 10 specimens in BL colln. 1ST 6436) and 3 specimens in BL colln.

Distribution - Pliocene, Mediterranean. This was proba- Distribution — Pliocene-Recent, Mediterranean. After bly the commonest species of Granulina in the Mediter- G. marginata, this is the second commonest extant spe-

ranean Pliocene. cies, also fairly common in the Pleistocene of Italy (Go-

Remarks — This species was first but not fas, 1992; La Pema, 1999). The present record is the first identified, named Gofas and recorded from from the Pliocene. by (1992), subsequently

the Pliocene of Sicily by La Pema (2000). It is similar to Remarks — The specimens from Estepona closely match G. boucheti, from which it be distinguished by a extant shells. Granulina boucheti is another species with may

tuberculate of sharper and ‘symmetrical’ posterior rostration. However, a thin, faintly parietal ridge. A comparison this species appears more closely related to the Pleisto- the soft parts and shell of this species with those of G. cene G. jhomisiensis La Pema, 1999. It will be formally marginata was provided by Gofas (1992). These are two named by La Pema & Smriglio (in prep.), on the basis of closely similar, often sympatric species, with a markedly material from northern Italy (Tuscany). shallow-water distribution. They may be related to G. pierrepineaui Pin & Boyer, 1995 from Senegal, which is

similar to G. boucheti in shell and soft parts and which

also has a markedly shallow-water distribution. - 118-

Discussion of the Pliocene fauna, the general pattern of distribution

of the extant Mediterranean and West Atlantic species

The differences in be overlooked. The sole with present paper highlights some compo- cannot species a genuinely sition of the genus Granulina between the Pliocene of Mediterranean to Atlantic (Ibero-Moroccan Gulf) distri- and bution is Malaga (westernmost Mediterranean) Italy (central G. occulta; three species are known only from

Mediterranean). Granulina iberica, G. detruncata, G. the Gibraltar-AlboranSea, while G. guancha (d’Orbigny, malacitanaand G.? longa, occurring at Malaga, are not 1840) appears restricted to the Canary Islands, etc. This known from to date. On the other G. be biased Italy hand, elliptica pattern may by an incomplete knowledge of a La Pema, 2000, is known exclusively from the Pliocene number of species, described recently and often known of while G. known Sicily, guttula, formerly only as an only from the type locality; however, a high rate of en-

extant at Granulina within the is doubt. The species, occurs Malaga. marginata, demicity genus beyond same G. G. clandestina and Granulina known is boucheti, sp. are general pattern found amongst marginellids and mar- from a much wider area, which may include the western ginelliform gastropods; for instance, all nine species of and central Mediterranean. In this it is worth respect, the marginellid genus recorded by Moreno & the wide distribution of and noting Recent G. marginata Bumay (1999) from the Cape Verde Islands are endemic

G. boucheti It seems unlikely that these compositional to the archipelago. The exclusive intracapsular larval

differences are related to distinct palaeoecological char- development of marginellids (Bouchet & Waren, 1985; of the sites all Pliocene records of acters studied, as Coovert, 1986; Coovert & Coovert, 1995) strongly re- Granulina from Italy are from shelf deposits similar to duces the dispersal capabilities, thus promoting specia- those occurring in the Malaga area. Although such dif- tion, endemicity and fast turn-over through (geological) ferences may in part be due to an incomplete knowledge time.

SPECIES CLUSTER GEOGRAPHIC RANGE STRATIGRAPHIC RANGE

Granulina marmarginatagin ata Mediterranean Pliocene - Recent

Granulina boucheti Mediterranean Pliocene - Recent

Granulinapierrepineaui northwest Africa Recent

Granulina clandestina Mediterranean Pliocene

GranulinapariparilisIis central West Africa Recent

Granulina malacitana Malaga Lower Pliocene

Granulinavanhareni Gibraltar Recent

Granulinafernandesi Cape Verde Islands Recent

Granulina mauretanica northwest Africa Recent

Granulina detruncata Malaga Lower Pliocene

Granulinaghanensis central West Africa Recent

Granulinaafricana northwest Africa Recent

\ Granulinacanariensis Canary Islands Recent

Granulinaguttula Mediterranean Pliocene - Recent

Granulinacrystallina northwest Africa Recent

Granulinacereacerea northwest Africa Recent

Granulina iberica Malaga Lower Pliocene

Granulina tenuilabiata Mediterranean Pleistocene - Recent

Granulinarosarioi Sicily Pleistocene

Granulina ovulina Sicily Pleistocene

- Granulinagofasi Mediterranean Pleistocene? - Recent

Granulina sp. Mediterranean Pliocene Granulinajhomjhomisiensisisiensis Sicily Pleistocene

Table 2. Clusters of of Granulina discussed in the with the exclusion of G.? with indication of their species present paper, longa,

geographical and stratigraphical distribution. ‘Mediterranean’stands for fossil and extant species with recorded or inferredwide

distributionwithin this basin (see text for additional comments).

As noted above, several cases of morphological genetic relationships and for outlining the evolutionary similarity may be recognised amongst some species, both and biogeographic history of the genus, allowing also a

extant and fossil, as based on a number of shell features certain taxonomic differentiation at species-group level

(size, shape, apertural features, micro-ornament, etc.). to be attempted. It is worth noting that marked similari-

These similarities have importance for inferring phylo- ties in soft parts occur between species with morphologi- - 119-

cally similar shells, as documented by Gofas (1992) for References

G. and G. boucheti. On these marginata grounds, some

J.J. H.P.M.G. & E. groups or ‘lineages’ may be tentatively recognised (Ta- Aartsen, van, Menkhorst, Gittenberger, ble 2). 1984. The marine ofthe Bay of Algeciras, Spain,

with generalnotes on Mitrella, Marginellidae and Turridae. Granulina marginata and G. boucheti represent an an- cient Basteria, Supplement 2, 1-135. stock which may have northwest African affinities, Bivona, A. 1832. Nuovi gened e nuove specie di molluschi. while G. clandestina and G. the most sp., probably Scienze di Sicilia Effemeride e Lettere 2, 1-19. distributed Pliocene widely species, may be referred to Blainville, H.M.D. de 1824. Memoire sur la classification me- distinct the former two lineages, pointing to a central thodique des animaux mollusques, et etablissement d’une West African while the latter include the affinity, may de la Soci- nouvelle considerationpour y parvenir. Bulletin Pleistocene G. whose with jhomisiensis, relationship ete Philomatique de Paris 1824, 175-180. extant species is not yet clear. A similar ‘southerly’ af- Bouchet, P. & Waren, A. 1985. Revision of the Northeast At-

lantic finity may also be proposed for G. detruncata, G. mala- bathyal and abyssal Neogastropodaexcluding Turri- dae (Mollusca, Gastropoda). Bollettino citana and G. guttula, each of them pointing to clearly Malacologico, Supplement 1, 121-296. distinct lineages. No living representatives of the G. de- F. Boyer, 2001. The genus Granulina (Margmellidae) in the truncata lineage occur in the Mediterranean, while the Canary Islands. Bollettino di Malacologia 37, 27-32. lineage of G. malacitana never seems to have extended Boyer, F. & Rolan, E. 1999. Granulina fernandesi (Gastro- into the Mediterraneanbeyond Gibraltar and the Alboran poda: Volutacea), a new species from Cape Verde Islands, Sea. Granulina iberica and ‘allied’ Pleistocene and ex- the and some considerations on genus IberusGranulina. endemic stock of unclear tant species may represent an 17, 1-10.

and with a marked for 1814. affinity preference deep-water Brocchi, G. Conchiologia fossile subappenina con habitats. Granulina has clear Atlantic sul elliptica a affinity, osservazioni geologiche sugli appenini e suolo adi- demonstrated ‘sister’ acente, 712 Milano as by a species recently found in 2, pp. (StamperiaReale). Coovert, G.A. 1986. A review of Upper Pliocene strata of Portugal (La Pema et ai, in marginellid egg capsules. Marginella Marginalia 1, 13-19. prep.). Coovert, G.A. 1988a. Taxonomic characters in the family Mar- Definition of such a model will be refined by further ginellidae: conchological characters. Marginella Margina- research. However, the present data support the lia 4, 43-47. ‘warm-temperate’ to ‘warm’ biogeographical character Coovert, G.A. 1988b. Marginellidae ofFlorida, part 1: Granu- of Granulina and allow a assessment to be made general lina hadria. Marginella Marginalia4, 1-8. the of this in the Mediterra- concerning history genus Coovert, G.A. & Coovert, H.K. 1995. Revision of the su- The Pliocene shelf nean. species were characterised by praspecific classification ofmarginelliform gastropods. The ‘admixture’ of Nautilus 43-110. an ‘warm-temperate’ and ‘warm’ charac- 109,

G.C.F.D, 1797. Tableau elementaire de ters. Subsequent to the Middle-Late Pliocene and Pleis- Cuvier, I’histoire

naturelledes animaux, xvi + 710 pp. Paris (Bauborin). tocene cooling events, the species with thermophilic af- Deshayes, G.P. 1839. Trade elementaire de Conchyliologie, finity (the ‘African lineages’) mostly went extinct, new les avec applications de cede science a la Geologic 1, 272 and earlier with af- species appeared species temperate pp. Paris. such finity, as G. marginata and G. were more boucheti, C. 1842. Revision de Desmoulins, quelques especes de Pleu- successful. The Mediterranean thus became the most rotomes. Actes de la Societe Linneenne de Bordeaux 12,

northerly ‘outpost’ of Granulina in the eastern Atlantic. 109-185.

Of the twelve extant known from the Mediterra- F. 1987. Descrizione di species Fernandes, tre nuove specie di Mar-

nean, only G. marginata, G. boucheti (entire Mediterra- ginellidae (Mollusca: Gastropoda) delle isole di Capo

Verde. 1 nean), G. vanhareni and G. torosa (Gibraltar) have a Argonauta3, 259-267, pi. Fernandes, F. & Rolan, E. 1991. The Marginellidae (Mollusca, typical shallow-water distribution. The first two also oc-

Gastropoda) of Prince Island (Republica de Sao Tome e curred in the Pliocene, where at least four species Principe). Journal ofConchology 34, 85-90, 1 pi. (Granulina sp., G. clandestina, G. malacitana and G. Fleming, J. 1828. An history ofBritish , exhibiting the elliptica) had a similar shallow-water distribution. This descriptive characters and systematical arrangement of the would imply that the diversity of Granulina in the shelf genera and species of quadrupeds, birds, reptiles, fishes,

waters exceeded the modem one. as far as we However, mollusca and radiata of the United Kingdom, xxiii + 565

the overall did not & know, diversity change significantly, pp. Edinburgh/London(Bell Bradfute). and this related to increased colonisa- Gofas, S. 1992. Le was probably an genre Granulina (Marginellidae) en tion of environments shelf Mediterranee et dans 1’Atlantique oriental. Bollettino de deeper-water (outer and upper slope). Malacologia 28, 1-26. Gofas, S. & Fernandes, F. 1988. The marginellids of Sao

Tome, West Africa. JournalofConchology 33, 1-30. Hoemes, R. & Auinger. M. 1880. Die Gasteropoden der Acknowledgements Meeresablagerungen der ersten und zweiten miocanen

Mediterranstufe in der osterreich-ungarischen Monarchic. We are grateful to Emilio Rolan (Vigo) for the loan of Abhandlungen der kaiserlich-koniglichen geologischen material of G. fernandesi, G. ocarina and G. ghanensis, Reichsanstalt 12, 53-113, pis 7-12. to Carlo for useful and to the Smriglio (Rome) pointers, Issel, R. 1869. Malacologia del Mar Rosso, ricerche zo-

editor for constructive comments. xi 387 journal’s managing ologiche epaleontologiche, + pp., pis 1-15. Pisa. - 120-

Jousseaume, F.P. 1888. Descriptions des mollusques recueilles Smriglio, C., Gubbioli, F. & Mariottini, P. 2001. New data

le la Mer 1888 par M. Dr. Faurot dans Rouge et le Golfe de concerning the presence of Granulina Jousseaume,

Aden. Memoires de la Societe zoologique de France 1, (Neogastropoda, Cystiscidae) along the West African coast

165-223. and description of four new species. La Conchiglia 287,

Landau, B.M. 1984. A discussion of the molluscan fauna of 54-59.

two Pliocene localities in the province of Huelva (Spain), Smriglio, C., Mariottini, P. & Rufmi, S. 1998. Description of

including descriptions of six new species. Tertiary Re- Granulina melitensis n. sp. (Neogastropoda, Cystiscidae) search 6, 135-155, 2 pis. from the Mediterranean Sea. La Conchiglia 287, 53-56.

2000. J. 1816. Great Landau, B.M. & Marquet, R. The genus Cymbium (Gas- Sowerby, The Mineral Conchology of Britain,

tropoda, Volutidae) in the Iberian Neogene. Contributions 2, 115-180. London (The author).

to Tertiary and Quaternary Geology 37, 23-34, pis 1-4. Thiele, J. 1929-1931. Handbuch der systematischen Weichtier-

La Pema, R. 1999. Pleistocene and Recent Mediterranean spe- kunde, 1, 778 pp. Jena (Gustav Fischer). cies of Granulina (Gastropoda, Marginellidae), with de- Vera-Pelaez, J.L., Lozano-Francisco, M.C., Muniz-Solls, R.,

scription of four new species. Bollettino de Malacologia Gili, C., Martinell, J., Domenech, R., Palmqvist, P. &

34, 43-52. Guerra-Merchan, A. 1995. Estudio preliminar de la mala-

La R, 2000. Pema, Granulina elliptica n. sp. and comments on cofauna del Plioceneo de Estepona (Malaga, Espana).

the Mediterranean Pliocene species of Granulina (Gastro- Iberus 13,93-117.

poda, Marginellidae), Bollettinode Malacologia 35, 53-55.

Laseron, C.F. 1957. A new classification of the Australian

with of from Marginellidae (Mollusca), a review species

the Solanderian and Dampierian zoogeographical prov-

inces. Australian Journal of marine and freshwater Re-

search 8, 274-311.

Lozano Francisco, M.C. 1998. Los bivalves (Mollusca, Bival-

via) del Pliocene de la Provincia de Malaga. Malakos 6/7,

16-58.

Monterosato, M.T.A. 1869. Testacei nuovi dei mari di Sicilia,

18 pp., pi. 1. Palermo(Ignacio Mirto). Monterosato, M.T.A. 1881. Relazione fra i molluschi del

Quatemario di Monte Pellegrino e Ficarazzi e le specie viventi. Bullettino della Societd di Scienze Naturali ed

Economiche di Palermo 2, 12-15.

Moreno, D. & Bumay, L.P, 1999. The genus Volvarina (Gas-

tropoda: Marginellidae) in the Cape Verde Islands. Journal

ofConchology 36, 83-124.

R. El Muniz-Solis, 1999. genero Conus L., 1758 (Gastropoda, Neogastropoda) del Plioceno de Malaga, Espana). Iberus

17,31-90.

Nehm, R.H. 2001. Neogene paleontology in the Northern Do-

21. minican Republic, The genus (Gastropoda,

Marginellidae). Bulletins ofAmerican Paleontology 359,

7-46, pis 1-3. Orbigny, A.D. D’ 1840. Mollusques, Echinodermes, Fora-

recueillis iles Canaries MM. miniferes etPolypiers aux par

decrits Webb et Berthelot et par Alcide D’Orbigny. Mol-

lusques 1,117 pp., pis 1-8. Paris.

Pin, M. & Boyer, F. 1995. Tre nuove specie di Marginelle della regione di Dakar (Senegal). La Conchiglia 275,

55-61.

Rolan, E. & Fernandes, F. 1997. The small marginelliform

gastropods from Ghana (Neogastropoda, Cystiscidae). Ar-

gonauta 11, 3-12.

Sanz de Galdeano, C. & Lopez Garrido, A.G. 1991. Tectonic

evolution ofthe Malaga Basin (Betic Cordillera) - Regional

implications. GeodinamicaActa 5, 173-186.

Seguenza, G. 1879. Le formazioni terziarie della Provincia di

Reggio (Calabria). Memorie della Reale Accademia dei

Lincei, Classe di Scienze Fisiche. Matematiche e Naturali

(3)6, 1-446.

C. & P. 1996. Smriglio, Mariottini, Description of a new spe- cies of Cystiscidae Stimpson, 1865 from the Mediterra-

Granulina La 54-56. nean: gofasi n. sp. Conchiglia 281,

Smriglio, C. & Mariottini, P. 1999. Description of Granulina

gubbiolii n. sp. (Neogastropoda, Cystiscidae) from the Mediterranean Sea. La Conchiglia 292, 35-40.