Cainozoic 2002 Research, 1(1-2)(2001), pp. 111-120,November Granulina (Gastropoda, Marginellidae) 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 genus 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 Granulina 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), Marginella Only in recent years have these extremely fossil-rich de- (Eratoidea) 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).