Bollettino della Società Paleontologica Italiana, 48 (1), 2009, 33-40. Modena, 15 maggio 200933

Two morphologically close new species of (Archeogastropoda: ) from the Pleistocene of Sicily and Peloponnesus

Vittorio GARILLI

V. Garilli, APEMA Research and Educational Service, Via Alla Falconara 34, I-90136 Palermo, Italy; [email protected]

KEY WORDS - Trochidae, Gibbula, New species, Mediterranean, Pleistocene.

ABSTRACT - Two gastropod species of the genus Gibbula Risso, 1826, G. mariaeangelae n. sp. and G. marialuisae n. sp., are described from the Middle-Upper Pleistocene of Kyllini (NW Peloponnesus) and the Lower Pleistocene of Cartiera Mulino (SE Sicily) respectively. These species are morphologically similar to each other and well distinguishable from congeners, being characterized by a widely umbilicated, quite depressed shell with an almost keeled periphery. They mainly differ in having different patterns of sculpture and preserved coloration: G. marialuisae n. sp., which also has a more depressed shell shape, shows a more delicate spiral sculpture and a color pattern consisting of pale reddish lines and spots on a whitish background, whereas G. mariaeangelae is characterized by whitish-cream spiral cords on a reddish background. This further description of two new Gibbula species, after that of G. olympica Garilli, Crisci & Messina, 2005 (from Kyllini), significantly contributes to a more detailed knowledge of the genus Gibbula in the Mediterranean Pleistocene. Both the described species lived in a palaeoenvironment linked to the present biocoenosis.

RIASSUNTO - [Due nuove specie morfologicamente affini di Gibbula (Archeogastropoda: Trochidae) dal Pleistocene della Sicilia e del Peloponneso] - Vengono presentate due nuove specie di Gibbula Risso, 1826, G. mariaeangelae n. sp. e G. marialuisae n. sp., rispettivamente dal Pleistocene medio-superiore di Kyllini (Peloponneso nord-ovest) e dal Pleistocene inferiore di Cartiera Mulino (Vittoria, Sicilia sud-occidentale). La descrizione di tali specie, dopo quella di G. olympica Garilli, Crisci & Messina, 2005 (dal deposito di Kyllini), fornisce un ulteriore contributo alla conoscenza dettagliata del genere Gibbula del Pleistocene Mediterraneo. Le nuove specie qui descritte appaiono conchiologicamente affini e facilmente riconoscibili dalle congeneri, avendo una caratteristica conchiglia particolarmente depressa e provvista di ampio ombelico. Esse si differenziano principalmente per avere colorazione e scultura differenti: G. marialuisae n. sp. presenta una scultura spirale più delicata ed una colorazione di linee rossicce su un fondo biancastro, mentre G. mariaeangelae n. sp. ha cordoncini spirali bianco crema su un fondo rossiccio. Peraltro, G. marialuisae n. sp. possiede una conchiglia più depressa. G. mariaeangelae n. sp. è stata confrontata con alcune congeneri con le quali mostra, in generale, lievi affinità, inadatte a chiarire eventuali relazioni filogenetiche. In modo particolare, sono stati effettuati confronti tra G. mariaeangelae n. sp., G. ardens (Von Salis, 1793) e G. umbilicaris (Linnaeus, 1767) (con specifico riguardo ad alcune varietà), affini esclusivamente per il modello di colorazione, la scultura della teleoconca e, relativamente a G. umbilicaris, per lo sviluppo ombelicale. G. marialuisae n. sp. è invece difficilmente confrontabile con altre congeneri note, ad eccezione di G. joubini Dautzenberg, 1910 (Africa occidentale) con la quale condivide esclusivamente il tipo di colorazione. Vengono inoltre fornite alcune informazioni di carattere paleoecologico sulle specie descritte, le quali vissero in paleoambienti legati a quelli attuali delle praterie di Posidonia oceanica.

INTRODUCTION The main aim of the present article is to discuss and describe two interesting Gibbula species, recovered Within the Mediterranean fossil and Recent from the Pleistocene deposits of Cartiera Mulino archeogastropods, the genus Gibbula Risso, 1826 (Vittoria, SE Sicily, Italy) and Kyllini (NW Peloponnesus, represents a common malacological component, Greece), considering them to be new taxa after an especially in shallow water assemblages from vegetated extensive check of the malacological works illustrating bottoms. It certainly is one of the most diversified group and/or discussing the family Trochidae from various in the family Trochidae, as slightly more than 20 species geographical realms (e.g. Bucquoy et al., 1884; Pilsbry, were recorded from the Mediterranean Sea (Giannuzzi- 1889; Sacco, 1896; Cossmann & Peyrot, 1918; Savelli, 1997), where some endemisms also occur (e.g. Malatesta, 1960; Nordsieck, 1968; Ghisotti & Melone, G. nivosa Adams A., 1851, from Malta, G. spratti (Forbes, 1972; Malatesta, 1974; Caprotti, 1976; Nordsieck & 1844), from the Aegean Sea, and G. tingitana Pallary, García-Talavera, 1979; Spadini, 1986, 1987; Cavallo & 1901, from the Alboran Sea). Almost the same number Repetto, 1992; Beck, 1995; Giannuzzi-Savelli et al., of species occurred in the Upper Neogene shallow water 1997; Borghi & Vecchi, 2001; Delamotte & Vardala- mollusc-rich associations from Tuscany (N Italy, see Theodorou, 2001; Ardovini & Cossignani, 2004; Chirli, Chirli, 2004), whereas fewer species are recorded from 2004). the Mediterranean Pleistocene, probably because of the lack of exhaustive taxonomic revisions. As a matter of fact, due to the wide intraspecific range shown by its METHODS, MATERIALS AND GEOLOGICAL species, this genus is affected by somewhat unresolved SETTING systematic-taxonomic matter. Also the phylogenetic relationships between its fossil and Recent Two shells were picked from bulk samples (about 70 representatives appear to be rather enigmatic. dm3), collected at the top of a regressive marine Lower

ISSN 0375-7633 34 Bollettino della Società Paleontologica Italiana, 48 (1), 2009

the “Kyllini field-work 2004” from a large amount (about 80 dm3) of sediment, sampled from the N2 layer described by Garilli et al. (2005). As reported by these authors, this stratigraphical layer can be referred to the Middle-Upper Pleistocene, lying 50-70 m over a greyish blue turritellid-rich layer with a late Sicilian (sensu Ruggieri et al., 1984) - early Middle Pleistocene nannofossil association, namely Crenalithus asanoi (Sato & Katayama, 1992) and Gephyrocapsa sp. 3 (Castradori, 1993; Sprovieri, 1993; Di Stefano, 1998; De Kaenel, 1999). The N2 layer, containing leaves and rhizomes of Posidonia oceanica and a molluscan association dominated by herbivorous - detritus feeding taxa (mainly trochids, phasianellids, cerithiids and rissoids), was referred to the HP biocoenosis by Garilli et al. (2005; see also this article for a stratigraphical log containing the N2 layer). In order to describe the two new species, particular attention has been given to teleoconch features. Scanning electronic microscope analyses, addressed to an in-depth investigation of protoconch characters, were not performed in order to preserve the original coloration occurring in the investigated material. However, a simple description of larval shells has been provided by stereomicroscope observation. The following abbreviations are used for measurements: Ht = shell height; Hlw = last height; Ha = height; D = shell maximum diameter; Ilw = inclination of whorl profile to shell axis (measured on the last whorl). The type material was measured with precision of 0.05 mm. A schematic drawing of measured characters is given in Fig. 2. Studied material is housed in the Dipartimento di Geologia e Geodesia, University of Palermo, Italy (DGUP, V. Garilli Coll.) and in the Goulandris Natural History Museum, Kifissia, Greece (GNHM). Fig. 1 - General (a) and detailed locations of the Kyllini-Cape Trypiti N2 layer (b) and the Cartiera Mulino deposit (c, asterisk). KY = Kyllini; TR = Cape Trypiti; VI = Vittoria. Figs. 1a and 1b are modified from Garilli et al. (2005). SYSTEMATICS

Class Family TROCHIDAE Rafinesque, 1815

Pleistocene sequence (Conti et al., 1979; Costa, 1989) Genus Gibbula Risso, 1826 ex Leach ms. underlying a lacustrine succession. This deposit crops out at about one hundred meters from the abandoned Type species: magus Linnaeus, 1758 (S.D., paper-mill named Cartiera Mulino (Vittoria, SE Sicily, Herrmannsen, 1847) Figs. 1a, c). In particular, material comes from the “3D1” layer of Costa (1989), of which the very rich mollusc association is related to shallow, sheltered muddy-sandy Gibbula mariaeangelae n. sp. environments with Posidonia oceanica (Linnaeus) Pl. 1, figs. 7-12 Delile, 1813, namely a SVMC-HP (sensu Pérès & Picard, 1964) ecotone (Costa, 1989). This layer consists of a Description - Shell small, depressed, conical, greyish to yellowish clayey fine sand containing a reaching about 4.6 mm and 8.3 mm in height and diameter molluscan assemblage dominated by trochids (mainly respectively. Protoconch depressed and paucispiral, Jujubinus spp.), rissoids (mainly Pusillina spp. and consisting of slightly more than one whorl starting with Alvania spp.), and cerithiids (mainly Bittium spp.) (see roughly triangular nucleus. Whorls apparently smooth, Costa, 1989 for a comprehensive account of the separated from teleoconch by very weak demarcation. molluscan association from the 3D1 layer and the related Teleoconch growing more in width than in height, Ht/D stratigraphic log). ratio from 0.55 (holotype) to 0.62 (paratype 1), Four further shells come from the northern part of consisting of about 4 (holotype) and 3.5 (paratype 1) the Kyllini-Cape Trypiti sequence (Kyllini, Ileia, NW slightly convex whorls with very short, almost Peloponnesus, Figs. 1a-b). They were collected during horizontal subsutural ramp and Ilw of about 50°. Whorls V. Garilli - New Mediterranean Pleistocene taxa of Gibbula 35

Type material and locality - Holotype (GNHM FA ID 19/22), paratypes 1 (DGUP KIGR 005), 2 (DGUP KIGR 006) and 3 (DGUP KIGR 007) are from the Middle-Upper Pleistocene layer N2 of Kyllini, Ileia, NW Peloponnesus, Greece (UTM 34S 511793E 4199216N).

Etymology - This species is dedicated to Maria Angela Stradi, a lovely friend of mine.

Occurrence - Gibbula mariaeangelae n. sp. is known only from type material. Accordingly, it is only known from the Middle-Upper Pleistocene of Kyllini, Ileia, NW Peloponnesus. The paleoenvironmental setting indicates that the species lived in Posidonia oceanica meadows.

Comparisons and remarks - Gibbula mariaeangelae Fig. 2 - Measurements. Ht = shell height; Hlw = last whorl height; n. sp. is somewhat similar to juvenile shells of G. pennanti Ha = aperture height; D = shell maximum diameter; Ilw = inclination of whorl profile to shell axis. (Philippi, 1846) (see Beck, 1995, pl. 15, figs. 9-10) from which it differs mainly by having a more depressed shell with a proportionally larger . Furthermore, G. pennanti has a larger, markedly elongated adult shell separated by deep, almost horizontal . Early (mean Ht/D of 0.9) with early teleoconch whorls showing whorls more convex and rounded, smooth, with mottles flat, wider spiral cords, separated by narrower interspaces variable in size and shape. Rest of shell sculptured by (see Beck, 1995, pl. 102, figs. 1-2). primary and secondary spiral cords. Primary cords Gibbula leucophaea (Philippi, 1836), as illustrated narrow and distinct, numbering 8 on last whorl; two by Beck (1995, pl. 16), may have a depressed shell (Ht/ stronger cords on subsutural part and on periphery of D down to about 0.7, see Beck, 1995, pl. 16), with a last whorl. Secondary cords very narrow, irregular and similar, acute periphery of the last whorl. However, this almost flat, numbering 4 (holotype) to 2 (paratype 1) species has more convex whorls, with a less inclined on last whorl. Primary cords separated by wide profile to the shell axis; it also has more prominent spiral interspaces, where secondary cords occur in middle. cords, appearing markedly elongated (usually with an Ht/ Very fine, narrowly interspaced spiral threads, mainly D of about 0.95) and stepped (see Beck, 1995, pl. 99, observed on adapical portion of mature whorls, in fig. 3). interspaces between primary and secondary sculpture. Gibbula umbilicaris (Linnaeus, 1767), particularly Whorls covered by numerous, marked, prosocline its depressed form “latior” (originally as Trochus latior incremental lines, reaching from suture to suture, Monterosato, 1878; see Giannuzzi-Savelli et al., 1997, obsolete on spiral cords. Base moderately convex with figs. 242a-b and 243) is comparable to G. mariaeangelae sculpture pattern similar to the dorsal one, having 7 n. sp. Color pattern (whitish suprasutural maculae on a (holotype) and 6 (paratype 1) spiral, widely interspaced, reddish background), sculpture (alternating primary and primary cords, and 3 (holotype) and 2 (paratype 1) secondary spiral cords) and umbilicus shape are the most narrower, spiral secondary cords. Umbilicus wide, remarkable shared characters. G. umbilicaris differs by notably deep, bordered by two spiral cords. Cords having a larger shell and more convex and proportionally crossed by well defined incremental scars, well marked higher whorls, with a more elevated (Ht/D 0.7-0.9) and on remaining basal area. Last whorl well developed, stepped profile. Furthermore, in G. umbilicaris the spiral making up 83% (paratype 1) to 86% (holotype) of entire sculpture, starting from the first teleoconch whorls, is shell height, almost keeled at periphery. Aperture wide, finer, more irregular in thickness, and with fewer primary subrhomboidal, making up 59% (paratype 1) to 62% spiral cords on the last whorl. Also the shell base of the (holotype) and 71% (paratype 1) to 73% (holotype) of form “latior” is different, being less convex (see Beck, last whorl and total height respectively. Outer lip 1995 pl. 25, fig. 4). rounded, thin, internally smooth and thickened close Gibbula ardens (Von Salis, 1793), which is similar to edge. Columellar side sinuous in central-upper part to G. umbilicaris (see also Beck, 1995), is comparable and slightly thickened abapically. Original coloration well with G. mariaeangelae n. sp. as well. G. ardens form preserved especially on holotype and paratype 1. Spiral “barbara” (reported by Monterosato, 1884 as G. cords whitish-cream, white on stronger subsutural and barbara), from the Pleistocene (Ruggieri, 1978) of peripheral cords; background bright red, all through Timpone Pelato (W Sicily), has a quite similar color shell surface. pattern, with whitish spots, usually particularly marked on the subsutural spiral cords and close to the base, on a Measurements - Holotype: Ht = 4.6 mm; Hlw = 3.95 reddish background (Figs. 3a-c). The sculpture, with mm; Ha = 2.85 mm; D = 8.3 mm Paratype 1: Ht = 4.15 primary and secondary spiral cords, is very similar to that mm; Hlw = 3.45 mm; Ha = 2.45 mm; D = 6.7 mm. of the new species. However, the shell of G. ardens is Paratypes 2 and 3: not measured because of their bad larger (maximum height 18.5 mm in the examined preservation. specimens), more elevated (Ht/D 0.8-1.05), with a proportionally narrower umbilicus and more rounded 36 Bollettino della Società Paleontologica Italiana, 48 (1), 2009 whorls, and (in the examined form “barbara”) the holotype only) depressed and paucispiral, probably secondary spiral cords lie closer to the primary sculpture. consisting of about 1 apparently smooth whorl, with a Furthermore, the shell base of the form “barbara” bears nearly triangular nucleus. Protoconch/teleoconch flat, more closely spaced spiral cords. It is noteworthy transition not well marked. Teleoconch of 3.7 (holotype) to remark that G. ardens co-occurs with the new species and about 3 (paratype) whorls growing more rapidly in the N2 layer where it shows differences from G. in width than in height, having a Ht/D ratio of 0.48 mariaeangelae n. sp. similar to those described above (paratype) and 0.51 (holotype). Whorls slightly convex for the form barbara (see Fig. 3d). G. ardens var. (almost flat in paratype), smooth near teleoconch onset, succincta Monterosato, 1878 (as Trochus succinctus), then with narrow spiral cords separated by wide see Bucquoy et al. (1884, pl. 45, figs. 13-16) and Pilsbry interspaces; Ilw is about 56° and 62° in holotype and (1889, pl. 33, figs. 82-83), has a rather depressed shell paratype respectively. Spiral cords not always regular and differs from G. mariaeangelae n. sp. mainly by having in width; 10 cords (holotype) and 8 cords (paratype) more marked spiral cords, a very rounded last whorl on last whorl. Cords lacking from early teleoconch profile and a markedly canaliculated suture. whorls, which appear smooth. Rest of shell with Other Gibbula species from the N2 layer, G. magus prosocline growth lines extending from suture to suture. (Linnaeus, 1758), G. fanulum (Gmelin, 1791), G. Last whorl acute at periphery and well developed, adansoni (Payraudeau, 1826) and G. olympica Garilli, making up about 90% total height. Teleoconch whorls Crisci & Messina, 2005, are not particularly similar to separated by deep, almost horizontal suture. Base G. mariaeangelae n. sp. Rather, the new species is moderately convex, sculptured by 11 (holotype) and 8 conchologically close to another new species herein (paratype) spiral cords and growth lines. Umbilicus very described below. wide and deep, with marked incremental scars and well- It is noteworthy that the Gibbula species compared defined, almost keeled periphery. Aperture wide, nearly above have a higher Ht/D ratio ranging from 0.7 to 1.05, rhomboidal, making up 67% (paratype) to 68% a range characterizing most of the congeners from the (holotype) and 77% (paratype) to 78% (holotype) of Mediterranean Upper Neogene to Recent. Very few last whorl and total height respectively. Outer lip thin, exceptions do exist, e.g. the Pliocene G. distefanoi internally smooth; columellar lip gently arched, with Crema, 1903, with an Ht/D (down to 0.56) matching that slight thickening in lower part. Original coloration well of G. mariaeangelae n. sp. The two species are otherwise preserved: reddish lines or spots (well marked on not similar. adapical portion of teleoconch whorls of holotype) on G. mariaeangelae n. sp. also shows a certain amount spiral cords on whitish-cream background. Very early of ontogenetic variation, the smaller shells (paratypes) teleoconch whorls white (in holotype) or translucent having more convex whorls and a slightly less depressed (in paratype). Base with almost identical coloration as general shape than larger ones. dorsum, having reddish lines (near periphery) and spots The main protoconch characters, namely its depressed (covering most of base) on spiral cords. shape, its apparently smooth surface and the roughly triangular nucleus, are frequent within Gibbula as well Measurements - Holotype: Ht = 3.5 mm; Hlw = 3.05 as in other trochid genera (see Beck, 1995). Also the mm; Ha = 2.35 mm; D = 6.85 mm Paratype: Ht = 2.5 onset of teleoconch sculpture, which is rather similar to mm; Hlw = 2.2 mm; Ha = 1.7 mm; D = 5.15 mm. that shown by G. umbilicaris (see Beck, 1995, pl. 100, fig. 8) is a character shared with different trochids (e.g. Type material and locality - Holotype (DGUP CMRG the genus Osilinus Philippi, 1847, see Beck, 1995, pl. 003) and paratype (DGUP CMRG 004), are from the 103, fig. 8 and pl. 104, figs. 1 and 6). Lower Pleistocene 3D1 layer (described by Costa, 1989) of Cartiera Mulino (UTM 33S 461624E 4089386.7N), about 15 km WNW of Ragusa, SE Sicily. Gibbula marialuisae n. sp. Pl. 1, figs. 1-6 Etymology - This species is dedicated to my mother Marialuisa. Description - Shell small, strongly depressed, conical, reaching 3.5 mm and about 6.9 mm in height Occurrence - Gibbula marialuisae n. sp. is known and width respectively. Protoconch (preserved on only from the type locality. The paleoenvironmental

EXPLANATION OF PLATE 1 figs. 1-6 - Shells of Gibbula marialuisae n. sp. Lower Pleistocene, 3D1 layer of Cartiera Mulino (SE Sicily). 1-3 - Apertural, apical and basal views of holotype (Ht = 3.5 mm, D = 6.85 mm, DGUP, CMRG 003). 4-6 - Apertural, apical and basal views of paratype (Ht = 2.5 mm, D = 5.15 mm, DGUP, CMRG 004). figs. 7-12 - Shells of G. mariaeangelae n. sp. Middle-Upper Pleistocene N2 layer of Kyllini (NW Peloponnesus, Greece). 7-9 - Apertural, apical and basal views of paratype 1 (Ht = 4.15 mm, D = 6.7 mm, DGUP, KIGR 005). 10-12- Apertural, apical and basal views of holotype (Ht = 4.6 mm, D = 8.3 mm, GNHN, FA ID 19/22).

All to scale. V. Garilli - New Mediterranean Pleistocene taxa of Gibbula Pl.37 1 38 Bollettino della Società Paleontologica Italiana, 48 (1), 2009 setting inferred for the 3D1 layer at the Cartiera Mulino variation testifies to the wide diversification of this site indicates that the species lived on sheltered shallow- genus. water bottoms with Posidonia oceanica. Remarkable morphological affinities between the new species described and other congeners are lacking, Comparisons and remarks - Gibbula marialuisae n. especially for Gibbula marialuisae n. sp.: while a single sp. shares remarkable similarities only with G. mariaeangelae n. sp.: the two species have a very depressed shell, with a wide, deep umbilicus, bordered by a sort of keel, and a sculptural teleoconch pattern formed by narrow spiral cords. However, G. mariaeangelae n. sp. has a less depressed shell, almost keeled at the periphery of the last whorl, a less inclined whorls profile (50° vs. 56-62° to the shell axis), and bears more distinct spiral sculpture consisting of primary and secondary cords. Furthermore very fine spiral threads occur in the interspaces between primary and secondary cords of G. mariaeangelae n. sp. The coloration of the two species is also different, consisting of a reddish background with whitish spiral lines in G. mariaeangelae n. sp., and of a whitish background with reddish-orange lines or spots in G. marialuisae n. sp. The coloration pattern of the latter species, never found in European congeners, is comparable to that of the West African species G. joubini Dautzenberg, 1910, which may have brownish linear maculae or lines on the spiral sculpture. However, the eastern Atlantic species has a larger, quite elevated shell (Ht/D 0.9-1.06), with a proportionally narrower umbilicus and stronger and flatter spiral cords starting from the first teleoconch whorl (Beck, 1995, pl. 12, figs. 1-11 and pl. 102, figs. 7-8). Probably due to ontogenetic changes, the smallest shell of Gibbula marialuisae n. sp. (paratype) has less convex whorls, a more acute last whorl periphery and a more inclined whorls profile (about 62° vs. 56°) than the respective holotype. The protoconch characters and the early teleoconch sculpture of G. marialuisae n. sp. are very similar to those of G. mariaeangelae n. sp. and, apparently, do not provide distinguishing characters.

CONCLUDING REMARKS

Gibbula mariaeangelae n. sp. and G. marialuisae n. sp. are morphologically close species sharing remarkable characters: teleoconch sculpture, umbilicus shape and, above all, a very depressed shell (with a Ht/D range of about 0.5-0.6). The last character is very rarely shared with other congeners (e.g. the Pliocene G. distefanoi), as the mean Ht/D of Gibbula is about 0.9, and its wide

Fig. 3 - Gibbula ardens (Von Salis, 1793), with morphs showing coloration pattern and sculpture comparable to those of G. mariaeangelae n. sp. a-c - Juvenile shell of the form barbara Monterosato, 1884 with size comparable to that shown by G. mariaeangelae n. sp. (a, apertural; b, basal; c, apical views), Ht = 9.3 mm, D = 10.1 mm, Pleistocene of Timpone Pelato (W Sicily). d - Apertural view of G. ardens from the Middle to Upper Pleistocene N2 layer of Kyllini (NW Peloponnesus, Greece), Ht = 10.6 mm, D = 11.4 mm. V. Garilli - New Mediterranean Pleistocene taxa of Gibbula 39 or a few main characters (such as sculpture, protoconch Caprotti E. (1976). Malacofauna dello stratotipo Piacenziano shape, coloration pattern) pool, other remarkable (Pliocene di Castell’Arquato). Conchiglie, 12: 1-56. characters decidedly diverge. Although a certain degree Castradori D. (1993). Calcareus nannofossil biostratigraphy and biochronology in eastern Mediterranean deep-sea cores. of similarity does exist between G. umbilicaris and G. Rivista Italiana di Paleontologia e Stratigrafia, 99: 107-126. mariaeangelae n. sp., any attempt to better understand Cavallo O. & Repetto G. (1992). Conchiglie fossili del Roero. the systematic-phylogenetic setting of the new species Atlante iconografico. 246 pp. Associazione Naturalistica would be risky on the basis of the resemblances noted Piemontese, Amici del Museo F. Eusebio di Alba. here. Chirli C. (2004). Malacofauna pliocenica toscana. Vol. 4, The description of Gibbula mariaeangelae n. sp. and Archeogastropoda. 113 pp. Arti Grafiche BMB, Firenze. Conti A., Di Geronimo I., Esu D. & Grasso M. (1979). Il Pleistocene G. marialuisae n. sp. from Cartiera Mulino and Kyllini, in facies limnica di Vittoria (Sicilia meridionale). Geologica together with the recently discovered G. olympica, from Romana, 18: 93-104. the same Greek site (Garilli et al., 2005), contributes to Cossmann M. & Peyrot A. (1918). Conchologie Néogénique de a more detailed knowledge of the genus Gibbula in the l’Aquitaine. Actes de la Société Linnéenne de Bordeaux, 70: 1- Mediterranean Pleistocene. Further studies would be 491. useful for better outlining composition and diversity of Costa B. (1989). La malacofauna pleistocenica della Cartiera Mulino this genus through the Neogene-Quaternary, and for better (Vittoria, Ragusa). In Di Geronimo I. (ed.) Atti 3° simposio Ecologia e Paleoecologia delle comunità bentoniche, Catania- understanding the phylogenetic relations between fossil Taormina 12-16 Ottobre 1985: 477-500. and extant species. De Kaenel E., Siesser W.G. & Murat A. (1999). Pleistocene calcareous nannofossil biostratigraphy and the western Mediterranean sapropels, sites 974 to 977 and 979. In Zahn R., ACKNOWLEDGMENTS Comas M.C. & Klaus A. (eds.), Proceedings of the Ocean Drilling Program, Scientific Results, 161: 159-163. Delamotte M. & Vardala-Theodorou E. (2001). Shells from the Greek My special thanks to Stefano Palazzi (Modena, Italy) for Seas. 316 pp. The Goulandris Natural History Museum, Kifissia. his precious and indefatigable assistance during the field work Di Stefano E. (1998). Calcareous nannofossils quantitative “Kyllini 2004”. He also picked paratypes 1 and 2 of Gibbula biostratigraphy of holes 969E and 963B (Eastern Mediterranean). mariaeangelae n. sp. and reviewed an early draft of the In Robertson A.H.F., Reichter K.-C. & Camerrlenghi A. (eds.), manuscript. I also warmly thank Eugenio Di Liberto (Palermo, Proceedings of the Ocean Drilling Program, Scientific Results, Italy), Luca Galletti and Francesco Pollina (APEMA, Palermo), 160: 155-165. who assisted me during field working at Kyllini. Luca Galletti, Garilli V., Crisci M. & Messina R. (2005). A new species of Gibbula with passion, also helped me in sampling the Cartiera Mulino (Gastropoda: Trochidae) from the Pleistocene of Killini (North- site. Thanks also to Giuseppe Buccheri and Antonino Greco western Peloponnesus, Greece). Bollettino della Società (Dipartimento di Geologia e Geodesia, University of Palermo) Paleontologica Italiana, 44 (1): 47-53. for supporting the above mentioned field trip, to Maria Ghisotti F. & Melone G.C. (1972). Catalogo illustrato delle conchiglie Antonietta Rosso (Dipartimento di Scienze della Terra, marine del Mediterraneo. Superfamiglia Trochacea. Conchiglie, University of Catania, Italy), who allowed access to the Cartiera 8, supplemento 4: 79-146. Mulino collection, and to Evi Vardala-Theodorou (Goulandris Giannuzzi-Savelli R., Pusateri F., Palmeri A. & Ebreo C. (1997). Natural History Museum, Kifissia, Greece), who provided the Atlante delle conchiglie marine del Mediterraneo. Vol. 1 , catalogue number of G. mariaeangelae n. sp. (holotype). I am Archeogastropoda. 125 pp. La Conchiglia, Roma. also grateful to Luca Bertolaso (Correggio, Reggio Emilia, Italy) Malatesta A. (1960). Malacofauna pleistocenica di Grammichele for providing useful literature. This article greatly benefited from (Sicilia). Memorie per servire alla descrizione della carta the comments by the referees Rafael La Perna (Dipartimento di geologica d’Italia, 12: 1-256. Geologia e Geofisica, University of Bari, Italy) and Robert Malatesta A. (1974). Malacofauna pliocenica umbra. Memorie per Marquet (Institut Royal des Sciences Naturelles de Belgique, servire alla descrizione della carta geologica d’Italia, 13: 1-498. Département de Paléontologie, Brussels, Belgium). I am Monterosato (Di Maria) T. (1878). Enumerazione e sinonimia delle extremely grateful to Alan G. Beu (Institute of Geological and conchiglie mediterranee. Giornale Scienze Naturali ed Nuclear Sciences, Lower Hutt, New Zealand), who added Economiche, Palermo 13: 61-115. valuable comments to the revised manuscript and improved my Monterosato (Di Maria) T. (1884). Nomenclatura generica e specifica English. di alcune conchiglie mediterranee e sinonimia delle conchiglie mediterranee. 152 pp. Stabilimento Tipografico Virzì, Palermo. Nordsieck F. (1968). Die europäischen Meeres-Gehäuseschnecken (Prosobranchia). Vom Eismer bis Kapverden und Mittelmeer. REFERENCES viii + 273 pp. Fischer, Stuttgart. Nordsieck F. & García-Talavera F. (1979). Moluscos marinos de Ardovini R. & Cossignani T. (2004). West African Seashells. 318 Canarias y Madera (Gastropoda). 208 pp. Aula de Cultura de pp. L’Informatore Piceno, Ancona. Tenerife. Beck L.A. (1995). Europäische Kreiselschnecken (). Zur Pérès J.M. & Picard J. (1964). Nouveau manuel de bionomie Systematik und Evolution europäischer Trochiden benthique de la mer Méditerranée. Recueil des Travaux de la (Kreiselschnecken) unter besonderer Berücksichtigung der Station Marine d’Endoume, Faculté des Sciences de Marseille, Gattungen Gibbula Risso, 1826, Osilinus Philippi, 1847 und 31: 1-137. Jujubinus Monterosato, 1884 (Gastropoda, Prosobranchia). Pilsbry H.A. (1889). Trochidae, pp. 5-519. In Tryon G.W. & Pilsbry Available on internet at http://www.staff.uni-marburg.de/~beck/ H.A. (eds.), Manual of Conchology. Structural and Systematic. diss1995/trochoid.htm#textinhalt. Vol. 11. Conchology Section, Academy of Natural Sciences, Borghi M. & Vecchi G. (2001). La malacofauna plio-pleistocenica Philadelphia. del torrente Stirone (Pr). Trochidae (parte II). Parva Naturalia Ruggieri G. (1978). Una trasgressione del Pleistocene inferiore della 2000-2001, Museo Civico di Storia Naturale di Piacenza: 11-43. Sicilia occidentale. Il Naturalista Siciliano, II (3-4) serie quarta: Bucquoy E., Dautzenberg P. & Dollfus G. (1884). Les Mollusques 159-171. marins du Roussilon. Vol. I, parte 2: 197-386. Baillière & fils, Ruggieri G., Rio D. & Sprovieri R. (1984). Remarks on the Paris. chronostratigraphic classification of Lower Pleistocene. Bollettino della Società Geologica Italiana, 103: 251-259. 40 Bollettino della Società Paleontologica Italiana, 48 (1), 2009

Sacco F. (1896). I molluschi dei terreni terziari del Piemonte e Sprovieri R. (1993). Pliocene-Early Pleistocene astronomically della Liguria. Parte XXI. 65 pp. Carlo Clausen, Torino. forced planktonic Foraminifera abundance fluctuations and Spadini V. (1986). Contributo alla conoscenza dei Trochidae del chronology of the Mediterranean calcareous plankton bio- Senese: specie nuove o poco conosciute. Bollettino events. Rivista Italiana di Paleontologia e Stratigrafia, 99: Malacologico, 22 (1-4): 85-90. 371-414. Spadini V. (1987). Nota ai Trochidae del Senese: Gibbula (Colliculus) turbinoides (Deshayes, 1832) e G. (Tumulus) umbilicaris (L., 1766). Bollettino Malacologico, 23 (1-4): Manuscript received 21 October 2008 92-94. Revised manuscript accepted 14 March 2009