October 2003 CainozoicResearch, 2(1-2) (2002), pp. 3-8,

Heteroninella The (, )

from the Lower Pliocene of Estepona, southern Spain

² Bernard Landau¹ & Pierre Lozouet

1 International Health Centres, Avenida Infante de Henrique 7, Areias Sao Joao, P-8200-261 Albufeira, Portugal;

e-mail: [email protected]

2 Museum nationald’Histoirenaturelle, 55 rue de Buffon, F-75005 Paris, France; e-mail: [email protected]

Received 19 September 2001;revised version accepted 25 May 2002

A 1941 of the turbinid genus HeteroninellaMagne, is described from the Velerin Conglomerates (Zanclean, Pliocene) near

the Heteroninella Estepona (southern Spain); this constitutes first record of this genus from the Neogene of Europe. Originally,

bertarellii & in 1826. (Andreoli Marsigli, 1997)was placed the trochid genus Risso,

KEY WORDS: Pliocene, Spain, Gastropoda, Turbinidae, Heteroninella.

Introduction deposited relatively rapidly or in storm conditions, contains

the richest fauna; a curious admixture of large and small,

The Lower Pliocene (Zanclean) deposits exposed at both waterwom and perfectly preserved, shells in between

wealth of and boulders and ofmudstone of sizes. Velerin are beginning to yield a interesting lumps varying

endemic have been described new, species. Although some

recently (Muniz Solis, 1995; Landau & Marquet, 1999,

unusual than the discussed in 2000), none are more species Systematic palaeontology

the considered this to be present paper. Originally, we a

is Abbreviations — the abbreviations new species; however, it has since turned out that it Below, following are

described Andreoli used: undoubtedly the same shell as that by

& Marsigli (1997), from the vicinity of Serre di Rapolano

(Siena, Italy), under the name of Gibbula bertarellii. AA apical angle;

based Andreoli & Marsigli’s description was on a juvenile H height;

shell and an incomplete adult specimen. The species is MNHN Museum national d’Histoire naturelle, Paris;

obviously extremely rare in Italy, and for this reason we W width.

consider it useful to offer a new description here of the

complete adult shell. Superfamily Raflnesque, 1815

The Malaga Basin is located in the western sector ofthe Family TurbinidaeRaflnesque, 1815

Internal Zones of the Betic Cordillera. The sea gateway, Subfamily TurbininaeRaflnesque, 1815

which extended from the Mediterranean to the Atlantic Genus HeteroninellaMagne, 1941

during the Late Miocene (Tortonian) had shrunk to a small

inthe from the — de basin Pliocene, extending present Malaga- Type species parkinsoni Basterot, 1825, by

Torremolinos area inlandin an E-W directionfor about 30 original designation.

km. The Pliocene series consists of conglomerates and

which to bluish marls and sands, give way laterally grey

sands de Galdeano clays, topped by yellowish-white (Sanz Heteroninellabertarellii(Andreoli & Marsigli, 1997) these & Lopez Garrido, 1991). Although deposits comprise Figures la-d, 2/1-3

a variety offacies, ranging from coarse sands, representing

nearshore beach to fine sands & 1. or deposits, clayey deposited *1997 Gibbulabertarellii Andeoli Marsigli, p. 19, pi.

at relatively greater depths, the gastropod species discussed

herein is found only in a coarse conglomerate (the Velerin Material — Three specimens (MNHN-PL 15297A, PL

Conglomerates). This conglomerate, which must have been 15297B, PL 15297C, leg. B. Landau); six, three subadult -4-

and four incomplete specimens (B. Landau Colin, Description — Shell large, trochiform, with convex whorls.

Albufeira); one complete, repaired adult specimen (R. Protoconch and early teleoconch whorls eroded in all

adult available. Five teleoconch whorls Marquet Colin, Antwerp); and a single complete, specimens preserved in specimen (Vera-Pelaez Colin, Malaga). All material is from fiilly-adult specimens, convex, with the periphery just the Velerin Conglomerates, near Velerin (Estepona, above the . Suture deeply impressed and narrowly

canaliculate. whorls ofthree southern Spain). Ornament on consisting

strong, raised, regularly spaced spiral cords, with a fourth

Diagnosis — Large species of Heteroninella, with very appearing just above the abaxial suture on the penultimate strong elevated spiral sculpture, separated by deeply . excavated interspaces and with a wide, deep .

PL = Figure 1. Heteroninella bertarellii (Andreoli & Marsigli, 1997),MNHN 15297A in various aspects; original height 56.3 mm;

Velerin Conglomerates (Lower Pliocene, Zanclean),Velerin (Estepona, Spain). -5- -6-

Figure 2. Heteroninellabertarellii (1-3) and H. parkinsoni (de Basterot, 1825)(4-6) compared:

= - 56.2 Velerin 1 MNHN PL 15297B; original height mm; Conglomerates (Lower Pliocene, Zanclean), Velerin (Estepona, Spain);

= 2 - MNHN PL 15297C; original height 41.2 mm; same stratigraphic level and locality;

= 3 - Landau Colin, specimen 1; original height 59.1 mm; same stratigraphic level and locality;

= 4 - MNHN PL 5932A; original height 34 mm; Lower Oligocene (‘Stampian’), Gaas (Landes, France);

= 5 - MNHN PL 6020A; original height 32 mm; Lower Oligocene(‘Stampian’), Bordeaux (Gironde).

= 6 - MNHN PL width 10 Lower Gaas 5932B; ; original mm; Oligocene (‘Stampian’); (Landes, France).

The cords are convex but flattened, separated by deeply apical whorls eroded, even in juvenile specimens. Despite excavated interspaces, approximately half the widthof the this, one of the specimens (MNHN PL 15297A) shows cords teleoconchwhorls. the of on early In some specimens some trace original colour pattern on the spiral cords, cords remain broader throughout, while in others the consisting of orange-brown, vertical flammules. As interspaces increase in width abaxially to be equal to or Andreoli & Marsigli (1997) correctly noted, there is no wider the cords whorl. There is than on the body a species in the European Miocene to Recent with which H. tendency for the cord placed at the periphery to be slightly bertarellii is likely to be confused. Those authors discussed stronger. Axial ornament absent, except for weak, close-set its superficial similarity to Gibbula ardens (von Salis prosocline growth lines, which cover the whole surface. Marschlins, 1793), but this is much smaller and does not

flattened have the umbilicus characteristic of Base to slightly convex, bearing eight flattened, very deep the present close-set spiral cords, the peripheral cord being broadest, taxon. In our opinion, H. bertarelliidoes not belong to the the remaining cords subequal in strength and width. Strong family , but rather to the Turbinidae.Indeed, the axial folds visible the half relative are on last whorl, corresponding closest seems to be H. parkinsoni (de Basterot, to the position of earlier apertures. The umbilicusis wide, 1825) (Figure 2/4-6), a species characteristic of the French about half the diameterof the base and very deep, almost Atlantic Lower Oligocene (‘Stampian’). This species shows

the The of the umbilicus is the characteristics of cords extending to apex. slope same sculptural strong spiral shallow up to a broad, indistinct cord arising from the mid- and a deep umbilicus, but the cords in the latterare not as columellar margin, beyond which the umbilicus dips strong or elevated, the interspaces shallower and although steeply. The spiral ornament does not extend into the the umbilicus is equally deep, it is much narrower. The umbilicus. The is tangential, large and is more evenly rounded in H. parkinsoni, subquadrate. The outer lip is regularly arched, with a whereas the base ofH. bertarellii is flattened, giving the shallow notch at the junction with the , coinciding body whorl a subtrigonal outline. with the outer border of the umbilicus. Columella Faunal elements co-occurring with H. bertarellii in somewhat thickened, especially at the mid-portion, erect Malaga are similar to those described by Andreoli & and slightly reflected over the umbilicus. Basal callus thin Marsigli (1997, p. 20), with the same species they listed and restricted. found in association. The Italian beds consist of gravels

and sands, also representing a sandy environment, similar

Measurements(in mm) — to that ofthe Velerin Conglomerates. Andreoli& Marsigli

(1997) dated the Italian beds as late Early to Middle

Specimen H W AA remarks Pliocene, now generally assigned to the Zanclean Stage,

which would be similar in age to the deposits in Estepona.

MNHN PL 15297A 56.3 58.2 70° adult

MNHN PL 15297B 56.2 61.5 90° adult Remarks — From an ecological point of view, the larger

MNHN PL 15297C 41.2 48.5 90° subadult turbinine gastropods may be separated into two groups

Landau Colin, no. 1 59.1 58.6 70° adult (Beesley etal., 1998), namely:

Landau Colin, no. 2 49.5 56.8 88° adult

Landau Colin, no. 3 53.4 56 66° adult 1. the genus Risso, 1826 and its allies, which are

Landau Colin, no. 4 c. 58 56.3 65° incomplete adult predominantly offshore;

Landau Colin, no. 5 47.5 49.8 80° subadult 2. the group of Turbo Linne, 1758 and Astralium Link,

1807, which are predominantly intertidal to shallow

Discussion — Heteroninella bertarellii is somewhat tidal. variable in height; although usually tallerthan broad, some specimens are more depressed. The height measurements The earliest European species ofBolmaare from the Upper

given above are an underestimate, as the apical whorls are Oligocene (North Sea and Aquitaine basins). Outside eroded in all the specimens available; nevertheless, the Europe, Bolma has been recorded from the Upper apical angle varies between 65E and 88E. The ornamentis of (Beu & Ponder, 1979; Darragh & Kendrick,

At relatively constant in number ofribs and character, varying 2000). present, the genus is represented in the

in the ofthe ribs. Mediterranean Sea Bolma only width by rugosa (Linne, 1767). The shell is not well preserved and chalky, with the Vera-Pelaez et al. (1996) listed four species ofBolma -7-

occurring in the Estepona outcrops, Bolma fimbriata Borson, S. 1825. Saggio di orittografia piemontese. Memorie

B. B. and dellareale Accademia delle Scienze di Torino 25, 180-229, (Borson, 1825), granosa (Borson, 1825), rugosa, pi. 5. B. muricata (Dujardin, 1837). Cossmann, M. & Peyrot, A. 1909-1935. Conchologie neogenique Although the oldest undoubted species of Turbo in de I’Aquitaine. Actes de la Societe linneenne de Bordeaux Europe is T. munieri Vasseur, 1882 (Middle Eocene, 600 121 63-86, 3, pp., pis. France), the major turbinid radiation occurred during the Darragh, A. & Kendrick, G.W. 2000. Eocene bivalves and Early Oligocene. For instance, six species of Turbo, gastropods from the Pallinup siltstone, Western Australia, inclusive ofHeteroninella are on record from parkinsoni, with new records from the Eocene and Oligocene of the Lower of the Basin Oligocene Aquitaine (Magne & southeastern Australia. Proceedings of the royal Society of

Vergneau-Saubade, 1971) and three from the Upper Victoria 112, 17-58. F. 1837. Dujardin, Memoires sur les couches du sol Touraine Oligocene. Only one species has been described from the en et descriptions des coquilles de la craie et des faluns. Lower Miocene of Europe, Turbo neuvillei Cossmann & Memoires de la Societegeologique de la France (2)9,211- Peyrot, 1917 (Aquitaine Basin, France). The genus 311, pis 15-21. Astralium comprises only A. aquitanicum Benoist, 1874, Landau, B.M. & Marquet, R. 1999. A new species of Cyllene which ranges from the Upper Oligocene to the Lower (Cyllenina) (, Mesogastropoda) from the Arenas de Miocene (Lozouet et al., 2001). HuelvaFormation(Pliocene, southern Spain), Contributions the the Turbo/Astralium Subsequent to Early Miocene, to Tertiary and Quaternary Geology 36, 41-44.

to be in B.M. & R. 2000. The the group appeared no longer represented the Landau, Marquet, genus Cymbium in

Iberian and European fossil record. It is clear that this group must have Neogene. Contributions to Tertiary Quaternary Geology 37, 23-34, 4 suffered species extinctions at the Oligocene/Miocene pis. Link, H.F. 1806-1808. Beschreibungder Naturalien-Sammlung boundary. In this respect, the occurrence of a species of Universitdt 160 der zu Rostock, + 38 pp. Rostock. Heteroninellain the Lower Pliocene ofthe Mediterranean C. 1758. tria Linne, von per regna naturae, is area particularly astonishing. As indicated by Lozouet secundum classes, ordines, genera, species, cum (1992), the newly collected material from Malaga is both characteribus, differentiis, synonymis, locis. Editio decima views of the Mediterranean Pliocene complementing our 824 reformata, 1(1-2), iii+ pp. Holmiae (Laurenti Salvii). fauna and the for detailed C. von 1767. tria aptly demonstrating necessity Linne, Systema naturae per regna naturae, studies of strata the classic Italian occurring beyond secundum classes, ordines, genera, species, cum

deposits. characteribus, differentiis, synonymis, locis. Editio duodecimo reformata, 1(2), 533-1068. Holmiae (Laurenti Salvii).

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