The family Marginellidae FLEMING 1828 in the

Arch. Molluskenkunde

|

144 | (2) | 243–258 |

4 figures

| Frankfurt am Main, 21.12.2015

The family Marginellidae Fleming 1828 in the Miocene (Tortonian) of South Piedmont (Italy), with the description of three new species (Gastropoda: Muricoidea) Maurizio Sosso, M. Mauro Brunetti & Bruno Dell’Angelo

Abstract This study is based on specimens of the family Marginellidae Fleming 1828 from the Miocene (Tortonian) of South Piedmont sites in the surroundings of Stazzano (Alessandria): Rio di Bocca d’Asino, S. Agata Fossili and Castellania. Seven species are here reported, three of which described as new (Dentimargo elusiva, Eratoidea antoniae, and “Marginella” giuntellii), three already known [Stazzania marginata (Michelotti 1847), Volvarina oblongata (Sacco 1890), and Marginella deshayesi Michelotti 1847] and one identified only at generic level (Volvarina sp.). The distribution of the genus Dentimargo is extended to the Miocene (Tortonian) of Italy. Dentimargo has a wider distribution, and was already known from the Eocene of France and the Pliocene of Spain. Eratoidea antoniae is compared with the syntype of E. eratoformis (Hoernes & Auinger 1880) and with specimens of E. cf. eratoformis from the Pliocene of Estepona (Spain), here figured. Key words: Marginellidae, Tortonian, South Piedmont, taxonomy, new species, distribution.

eschweizerbartxxx sng-

Introduction The purpose of present work is to define the family Marginellidae Fleming, 1828 from the Miocene (Tortonian) of South Piedmont sites in the surroundings of Stazzano (Alessandria). Several studies were published about this rich molluscan fauna (Bellardi & Michelotti 1840; Michelotti 1847; Bellardi & Sacco 1873-1904; Bongo 1914; Caprotti 1964, 2010; Robba 1968), but the systematic is often related to old works and taxa are correlated to species of different age, furthermore several small species are frequently incorrectly determined. The family Marginellidae is characterized by a shell of variable dimensions, from small to very large, shell size is often very valuable as a species-level character; painted in various colours, with or without spots of dif-

ferent shape; almost always with protoconch paucispiral; thick lip with or without internal denticulations, often with external varix; siphonal notch present or absent; the columellar lip presents two to six plications; there is no operculum (Coovert & Coovert 1995; Moreno & Burnay 1999). The habitat of Marginellids is of wide range, from the lower intertidal zone to more than 1000 m depth, and from rocky reefs to soft bottom, the feeding habits are not well known, but belike they feed in wide range of animal food, with high selectiveness; one species has been observed parasitizing sleeping fishes (Bouchet 1989) and two other living in the sand are drilling the small bivalves and feed on their flesh (Ponder 1998).

Authors’ addresses Maurizio Sosso, Via Bengasi 4/4, I-16153 Genova; E-mail: [email protected] M. Mauro Brunetti, Via 28 Settembre 1944 n. 2, I-40036 Rioveggio (Bologna); E-mail: [email protected] Bruno Dell’Angelo, Museo di Zoologia, Via Selmi 3, I-40126 Bologna; E-mail: [email protected] © E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller), 2015, ISSN 1869–0963 DOI 10.1127/arch.moll/1869-0963/144/243-258

244

Sosso, M. et al.: The Marginellidae in the Miocene (Tortonian) of Italy

Fig. 1. Location map. (A = Rio di Bocca d’Asino; B = Sant’Agata Fossili; C = Castellania).

The family Marginellidae apparently originated in the ancient Tethys Sea, most Marginellidae are found in the Eocene of Paris Basin (Cossmann 1899; Nieulande 1981), in the Middle Eocene of Nantes and Cotentin Basins (Le Renard & Nieulande 1985), in the Oligocene of France (Lozouet 1999) and Italy (Füchs 1870), in the Miocene of France (Lozouet 1998) and Italy (Michelotti 1847; Seguenza 1880; Sacco 1890, 1904; Robba 1968; Ferrero Mortara et al. 1981), and in the Pliocene-Pleistocene of the Mediterranea sea (Landau et al. 2006; Silva et al. 2011). Some species of Marginellidae were also reported from the Eocene of southern Australia (Cossmann 1899; Glibert 1960; Coan 1965), the Oligocene of Washington State (Coan 1965), and the family subsequently had a rapid diffusion in the tropical seas. Actually the family Marginellidae has a wide diffusion in all the seas of the world (Coovert & Coovert 1995; Engl 2002). In the present work the Marginellidae from the Tortonian of South Piedmont are revised. eschweizerbartxxx sng-

Material and methods The studied material comes from the south-eastern end of BTP (“Bacino Terziario Piemontese”), which include the Borbera, Lemme and Scrivia Valleys (Ghibaudo et al. 1985; d’Atri et al. 2002). The shells were found by M. Sosso, P. Giuntelli, and M.M. Brunetti during many years of research in three localities in the surroundings of Stazzano (Alessandria) (Rio di Bocca d’Asino, Sant’Agata Fossili and Castellania, Fig. 1), all belonging to the Sant’Agata Fossili Fm. of Tortonian age (Boni & Casnedi 1970). In the considered area, this formation is made up of sandy facies with intercalating pelitic levels and an upper, mainly clayey facies with intercalating conglomeratic beds. Mollusc and coral associations are well preserved and point to deepening, from the outer shelf-slope transition to bathyal depths, with an increased content of allochthonous fauna (Ghibaudo et al. 1985).

Fig. 2. Morphological terms used for the shell description.

The classical locality of Rio di Bocca d’Asino (44°53'21''N, 8°53'14''E), placed at 2500 m NW from Stazzano (Alessandria), is already known for its rich mollusc fauna (Bellardi & Sacco 1873-1904; Bongo 1914; Caprotti 2011), and is attributed to the lower part of the Marne di S. Agata Fossili of Tortonian age (Boni

Sosso, M. et al.: The Marginellidae in the Miocene (Tortonian) of Italy 245

& Casnedi 1970; Ghibaudo et al. 1985). Sacco (1890) refered to “Stazzano” several localities included in the neighbourhoods of this area, i.e. Rio di Bocca d’Asino, and others; see Bongo 1914: 398. The site of Sant’Agata Fossili (44°47'06''N, 8°55'40''E) is placed at 500 m E from Sant’Agata Fossili village (Alessandria). Its rich fossil malacofauna was recently studied and attributed to the Tortonian age (Bongo 1914; Robba 1968). The outcrops of Castellania (44°47'55.2''N, 8°55'57.6'' E) are placed near the Rio Mazzapiedi-Castellania. The molluscan fauna is not much frequent nowadays, and the works of Gianotti (1953) and Robba (1968) confirm the attribution to the Tortonian age. We follow Marshall (1991) for the dates of publication of Bellardi & Sacco (1873–1904), and Coovert & Coovert (1995) for the systematics. The following abbreviations are used (Fig. 2): AA DAS DN L

apical angle distance of apical fold to siphonal end diameter of nucleus shell length

LA LLW NI NE W

length of aperture length of last whorl number of internal teeth number of external teeth shell width

Repository: BD MB MGGC MGPT MS MSNG MZB PG SMF

Private collection of B. Dell’Angelo, Genova, Italy (will be deposited in MZB) Private collection of M.M. Brunetti, Rioveggio, Bologna, Italy Museo Geologico Giovanni Capellini (Bologna, Italy) Museo di Geologia e Paleontologia, Università di Torino, Italy Private collection of M. Sosso, Genova, Italy Museo di Storia Naturale “Giacomo Doria”, Genova, Italy Museo di Zoologia dell’Università di Bologna, Italy Private collection of P. Giuntelli, Nole Torinese, Torino, Italy Senckenberg Forschungsinstitut und Naturmuseum, Frankfurt a. Main, Germany

Systematic account Class Gastropoda Cuvier 1795 eschweizerbartxxx sng-

Family Marginellidae Fleming 1828

smooth and glossy (Coovert & Coovert 1995; Moreno & Burnay 1999).

Subfamily Marginellinae Fleming 1828

Volvarina oblongata (Sacco 1890)

Tribe Prunini Coovert & Coovert 1995 Genus Volvarina Hinds 1844 T y p e s p e c i e s : Marginella (Volvarina) nitida Hinds 1844 [= Voluta mitrella Risso 1826], by subsequent designation (Redfield 1871). D i s t r i b u t i o n : The genus is quite cosmopolitan, with high diversity of species in the warm and temperate seas, from intertidal to 1,780 m. Fossil shells are present in the French Eocene, Oligocene to Pleistocene of W. Atlantic, early Miocene of W. Pacific, Miocene of Morocco and Turkey, Miocene to Pleistocene of Italy, Pliocene of Spain, Pleistocene of California (Glibert 1960; Coovert & Coovert 1995; Landau et al. 2006). Recent. R e m a r k s : The genus is characterized by a shell from small to moderately large; spire rarely immersed, or low to tall; shape elongate to moderately broadly cylindrical; aperture narrow to moderately broadly; lip thickened, smooth, not denticulate; external varix usually absent, but weak to strong in some species; siphonal notch absent or weak; columellar lip with 3 or 4 plications, some species with weaker 5th or 6th, combined occupying half or less of the aperture. The shell surface is

Figs 3 M–Q Marginella oblongata (Bon) Sism. – Doderlein: 24 (nomen nudum). 1890 Marginella (Volvarina) oblongata Bon. – Sacco: 28, pl. 2, fig. 13. 1899 Marginella oblongata Bon. – Cossmann: 93, pl. 4, fig. 21. 1904 Volvarina oblongata (Bon.) – Sacco: 92. 1960 Hyalina (Volvarina) oblongata – Glibert: 88; 1968 Hyalina (Volvarina) oblongata (Bellardi) – Robba: 465, 472, 568, pl. 43, figs 7 a, b. 1979 Hyalina (Volvarina) oblongata (Bellardi) – Pavia & Robba: 554. 1981 Marginella (Volvarina) oblongata B ellardi , Bonelli m.s. – Ferrero Mortara et al.: 177, pl. 54, fig. 5. 1996 Volvarina oblongata (Bonelli) – Vera-Peláez et al.: 112. Non 2002 Volvarina cf. oblongata (Bonelli) – Chirli: 28, pl. 14, figs 3–4 (= Volvarina sp.). Non 2002 Volvarina oblongata (Bonelli in Bellardi) – Muñiz-Solis: 278, fig. 18 A – B (= Volvarina sp., fide Landau 2006). Non 2005 Volvarina oblongata (B ellardi , 1890, B onelli m.s.) – Brunetti: 12, figs 1, 2 (= Volvarina sp.). 1862

246




2006 2013

Volvarina oblongata Bellardi – Landau et al.: 37. Volvarina oblongata (Sacco, ex Bellardi ms) – Landau et al.: 208, pl. 32, figs 14–15.

T y p e m a t e r i a l : MGPT Syntype BS.021.01.019, figured by Sacco (1890: pl. 2, fig. 13) and Ferrero Mortara et al. (1981: pl. 54, fig. 5). T y p e l o c a l i t y : Stazzano (Alessandria), Piedmont, Italy. T y p e s t a g e : “Marne di Sant’Agata Fossili”; Miocene: Tortonian. M a t e r i a l e x a m i n e d : Rio di Bocca d’Asino: 4 shells (MS, PG); S. Agata Fossili: 2 shells (MB, PG); Castellania: 1 shell (MS).

D e s c r i p t i o n : Shell of medium size, length 11.0 mm and wide 4 mm; protoconch paucispiral with a superficial suture and poorly distinct junction with teleoconch; teleoconch subcylindrical with the anterior portion more developed than the posterior, weak shoulders at the aperture insertions level; surface smooth; suture indistinct; aperture broad and prosocline inclined, from 65 % to 75 % of total height; outer lip internally smooth, with a strong external varix; siphonal notch absent; posterior sinus rather broad; columellar lip with four plications occupying more than half the aperture, right lip internally smooth, external side thickened but without varix. Va r i a b i l i t y : The range of variability of the characters measured is reported in Appendix 1. The examined specimens show only low variability. R e m a r k s : The species is rare in the Rio di Bocca d’Asino and Castellania outcrops, with shells quite well preserved. Chirli (2002) and Brunetti ( 2005) reported the presence of Volvarina oblongata from the italian lower Pliocene, Muñiz-Solis (2002) from the lower Pliocene of Estepona (Spain). We agree to Landau et al. (2006) to consider these reports as misidentifications, probably to be attributed to undescribed species. These reports are quite distinct from the type material present in the Bellardi & Sacco collection, the dimensions are smaller (7.3 mm for the specimen figured by Chirli and Brunetti, and 8.60 mm max for the specimens reported by Muñiz-Solis) than those of the specimens from Miocene (Coovert & Coovert 1995; Moreno & Burnay 1999), and moreover the species found from the Pliocene show plications decidedly more evident than the typical ones, as already evidenced by Muñiz-Solis (2002: “labro interno….con 4 pliegues inclinados bien prominentes…”). eschweizerbartxxx sng-

Volvarina oblongata has some resemblance with Volvarina parvula Sacco 1890 from the Early Miocene of Valle Ceppi (Turin, Piedmont), from which it differs by the more developed aperture and the posterior sinus at the level of the last whorl suture. D i s t r i b u t i o n : Middle Miocene: Proto-Mediterranean Sea (Serravallian): Karaman Basin, Turkey (Landau et al. 2013). Late Miocene: Proto-Mediterranean Sea (Tortonian and Messinian): Po Basin: Rio di Bocca d’Asino, S. Agata Fossili, Castellania, Borelli (Sacco 1890; Robba 1968; Pavia & Robba 1979; this paper). Volvarina sp. Figs 3 K–L M a t e r i a l e x a m i n e d : Rio di Bocca d’Asino: 2 shells (PG).

D e s c r i p t i o n : The two shells have a length of 7.74 and 8.39 mm, respectively; shape subcylindrical; aperture broad and wider anteriorly, about 70% of total length; columellar lip with four angulated plications occupying about 50% of aperture; right lip internally smooth, external side thickened but without varix. R e m a r k s : We do not attribute these shells to Volvarina oblongata because the only two specimens examined are much smaller and the dimensions represent an important character in species of the family Marginellidae and particularly the genus Volvarina (Coovert & Coovert 1995; Moreno & Burnay 1999). The two shells from Rio di Bocca d’Asino have a smaller length vs. the minimum length reported for the shells of Volvarina oblongata (Appendix 1: 9.93 mm); they have a fully developed lip, and can therefore be considered adult specimens. D i s t r i b u t i o n : Late Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin: Rio di Bocca d’Asino (this paper). Tribe Marginellini Fleming 1828 Genus Stazzania Sacco 1890 T y p e s p e c i e s : Marginella (Stazzania) emarginata Bonelli (= nomen nudum; = Marginella marginata Michelotti 1847), by monotypy. D i s t r i b u t i o n : only known from the Italian Tortonian.

Fig. 3. A–E. “Marginella” giuntellii sp. nov., Holotype MGPT-PU 135031, Rio di Bocca d’Asino, shell length 3.34 mm, A–C) entire shell, D) detail of the aperture, E) apical view. — F–J. Stazzania marginata (Michelotti 1847), Rio di Bocca d’Asino. — F, H-J) shell length 12.93 mm, F) apical view, H–J) entire shell (MS). — G) shell length 13.38 mm, entire shell (MS). — K–L. Volvarina sp., Rio di Bocca d’Asino. — K) shell length 8.39 mm, entire shell (PG). — L) shell length 7.74 mm, entire shell (PG). — M–Q. Volvarina oblongata (Sacco 1890). — M) Rio di Bocca d’Asino, shell length 11.39 mm, entire shell (MS). — N–P) Castellania, shell length 10.4 mm, entire shell (MS). — Q) S. Agata Fossili, shell length 10.17 mm, entire shell (PG).

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R e m a r k s : The genus is characterized by the shell of medium size, broadly biconic; spire medium; aperture narrow; lip thickened, smooth, often with single posterior denticle; external varix present, siphonal notch absent; columellar lip with four strong plications occupying more than half the aperture; collabral parietal callus ridge present, connecting with bifurcated outer ends of plications (Coovert & Coovert 1995). We agree with Landau et al. (2006) to consider Stazzania as a distinct monotypic genus with peculiar characteristics and hitherto known only from the Italian Tortonian. Stazzania marginata (Michelotti 1847) Figs 3 F–J 1847 1862 1873 1881 1899 1890 1890 1904 1904 1905 1914 1914 1960 1968 1973 1981 2002 2006 2006 2011

Marginella marginata Michelotti: 321, pl. 13, figs 10, 11. Marginella emarginata Bon. Sism. – Doderlein: 24 (nomen nudum). Marginella marginata Bon. – Cocconi: 104. Marginella marginata Bon. – Coppi: 47. Marginella emarginata Bon. – Cossmann: 88, pl. 4, fig. 5. Marginella (Stazzania) emarginata Bon. – Sacco: 26, pl. 1, fig. 10a. Marginella (Stazzania) emarginata Bon. Varietà A – Sacco: 27, pl. 1, fig. 10b. Stazzania emarginata (Bon.) – Sacco: 92. Stazzania emarginata var. brunneozonata Sacc. – Sacco: 92. Marginella emarginata Bon. – Sacco: 907. Marginella (Stazzania) emarginata Bon. – Bongo: 438. Marginella (Stazzania) emarginata Bon. var. brunneozonata Sacc. – Bongo: 438. Marginella (Stazzania) emarginata Bonelli – Glibert: 91. Marginella (Stazzania) marginata Michelotti – Robba: 570, pl. 43, figs 9 a, b. Marginella (Stazzania) marginata Michelotti – Marasti: 80, 81, 95, pl. 20, fig. 17. Marginella (Stazzania) marginata Michelotti – Ferrero Mortara et al.: 177. Stazzania emarginata (Sismonda) - Muñiz-Solis: 282, fig. 22 A–B. Marginella marginata Michelotti – Landau et al. 2006: 24, pl. 1, figs 2, 3. Stazzania marginata (Michelotti) – Landau et al. 2006: 29. Marginella (Stazzania) marginata Michelotti – Caprotti: 66. eschweizerbartxxx sng-

T y p e m a t e r i a l : Coll. Michelotti nel Museo di Geologia di Roma (fide Sacco 1890). T y p e l o c a l i t y : “Tortona” (Alessandria), Piedmont, Italy. T y p e s t a g e : “Marne di Sant’Agata Fossili”; Miocene: Tortonian. M a t e r i a l e x a m i n e d : Rio di Bocca d’Asino: 20 shells (BD, MS, PG); Sant’Agata Fossili: 4 shells (MS, PG).

D e s c r i p t i o n : The adult shells have usually a length between 12 and 14 mm, and a width from 6.8 to

7.8 mm; aperture from 65% to 75% of total height, shape biconical with the posterior side more developed than the anterior; shoulder sub-rounded; aperture broad and right; columellar lip with four strong plications occupying more than half the aperture; right lip without denticulation and thickened in the central part, the external side is transformed into a varix; siphonal notch absent. Va r i a b i l i t y : The range of variability of the characters measured is reported in Appendix 1. The examined specimens show low variability, except in the apical angle which shows a range between 73° and 92°. R e m a r k s : This species originally was reported in an unpublished catalogue by Bonelli (1824) as Marginella emarginata, but the first valid description was published by Michelotti (1847) as M. marginata, and this is the valid name to attribute to this species (Robba 1968). Sacco (1890) and Robba (1968) considered Marginella benasterensis Seguenza 1880 a synonym of the present species, but the impossibility to find the type of M. benasterensis do not permit to confirm this synonymy; the types of the fossils from Benestare, Reggio Calabria, were destroyed in the earthquake of Messina in the 1908, or subsequently lost (Laghi & Palazzi 1991; Bertolaso & Palazzi 2000). Moreover the study of the dimension of the Tortonian specimens of Piedmont and the personal communication of Angelo Vazzana, support our conviction to consider M. benasterensis a problematic taxon which needs the study of further topotypic material; in fact the specimen illustrated by Seguenza (1880: pl. 11, fig. 2), described and compared on p. 101 by the author with S. marginata, presents the compatible length of 14 mm, but appears not similar in the shape, with a larger aperture, strong columellar lips and more elevated external varix. The species is more common in Rio di Bocca d’Asino site, less in Sant’Agata Fossili, always with well preserved specimens. D i s t r i b u t i o n : Late Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin: Rio di Bocca d’Asino, Sant’Agata Fossili, Rio Mazzapiedi, Montegibbio, Torrente Stirone (Sacco 1890; Cocconi 1873; Robba 1968; this paper); Algeria (Sahelian) (fide Robba 1968). Genus Dentimargo Cossmann 1899 T y p e s p e c i e s : Marginella dentifera Lamarck 1803, by original designation. D i s t r i b u t i o n : The genus is present from Eocene of France, Eocene to Pleistocene of West Atlantic, Oligocene to Pliocene of Australia, Miocene to Pliocene of Western Pacific, Pliocene of Spain. Actually it is present worldwide in the tropical and temperate sea from intertidal to 1300 m (Coomans 1976; Lipe 1991; Coovert & Coovert 1995; Landau et al. 2006). R e m a r k s : The genus is characterized by shells of small to medium dimensions; usually biconical; spire medium to tall; aperture moderately broad to broad; lip


Table 1: Dentimargo elusiva sp. nov., type material, holotype and 3 paratypes. type

repository

type locality

L

figs

Holotype

MGPT-PU 135033

Rio di Bocca d’Asino

9.84

figs 4 A–D, F

Paratype 1

MSNG 57967


9.02

fig. 4 G

Paratype 2

PG


9.74

Paratype 3

MGPT-PU 135034


6.64

figs 4 E, H

thickened, smooth or denticulate, often with more or less strong single posterior denticle; external varix present, siphonal notch absent, columella with four strong plications occupying more than half the aperture (Coovert & Coovert 1995). Dentimargo elusiva sp. nov. Figs 4 A–H O t h e r m a t e r i a l : Rio di Bocca d’Asino: two incomplete shells (MS). T y p e l o c a l i t y : Rio di Bocca d’Asino (Alessandria), Piedmont, Italy. T y p e s t a g e : “Marne di Sant’Agata Fossili”; Miocene: Tortonian. E t y m o l o g y : referring to the difficulty of finding the specimens, notwithstanding their reasonable size.

D e s c r i p t i o n of the holotype: Shell of medium size, length 9.84 mm; protoconch rounded and paucispiral with a superficial suture and poorly distinct junction with teleoconch regularly biconical, the anterior portion developed more than the posterior; surface smooth; suture indistinct; aperture quite compressed, 48% of total length; outher lip internally smooth with a flat but evident denticulation to delimit the anal channel, with a strong flat external varix, more developed in the superior side; siphonal notch absent; posterior sinus broad; columellar lip with four plications occupying more 70% of the aperture, the upper three very strong and horizontal, the lower one more oblique and weaker; siphonal notch absent. Va r i a b i l i t y : The range of variability of the characters measured is reported in Appendix 1. The examined specimens show a low variability. R e m a r k s : This is the first report of the genus Dentimargo from the Italian Miocene, and particularly from the Tortonian of the Po basin. Dentimargo elusiva shows specific and well recognizable characters and a reasonable size, and it is strange that this species has not been identified to date, probably because of the real rarity. Dentimargo elusiva has some resemblance with Dentimargo bifidoplicata (Edwards 1855) from the Eocene of Paris Basin, from which it differs mainly by the stratigraphic age and the increasing size (ratio ca. 2:1). Two species of Dentimargo have been recently described from the Pliocene of Estepona (Spain) and westcentral Portugal: Dentimargo malakosi (Muňiz Solís eschweizerbartxxx sng-

2002; Silva et al. 2011) and “Dentimargo” lozoueti Landau et al. 2006, (nom. nov. pro Marginella minima Muňiz Solís 2002). Dentimargo elusiva differs from D. malakosi by the larger size (length 6.64–9.84 mm vs. 4.5–5.9 mm) and by the more slender and subcylindrical shape, and from “Dentimargo” lozoueti by the larger size (3.5–4.5 mm in D. lozoueti) and by the decidedly marginelliform shape. D i s t r i b u t i o n : Late Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin: Rio di Bocca d’Asino (this paper). Genus Eratoidea Weinkauff 1879 T y p e s p e c i e s : Marginella margarita K iener 1834, by subsequent designation (Cossmann 1899: 87). D i s t r i b u t i o n : Among the extant marginellids, the genus Eratoidea is known only from the tropical West Atlantic (Coovert & Coovert 1995). Fossil records of the genus are reported from the Middle Miocene (Badenian) Paratethys from Niederleis, Pötzleinsdorf, Forchtenau (Austria), Lissitz (Czech Republic) (Hörnes & Auinger 1879), Korytnica, Poland (Bałuk 1997); one species from the Pliocene of Estepona, Spain, is provisionally referred to this genus (Landau et al. 2006). R e m a r k s : The genus is characterized by shells of minute to medium size, broadly biconical; some species are axially costate; spire low to medium, often stepped; lip thickened, denticulate; external varix present; siphonal notch absent; columella with four strong plications occupying half or more of the aperture. Eratoidea antoniae sp. nov. Figs 4 L–P O t h e r m a t e r i a l : Rio di Bocca d’Asino: 8 shells (MS). T y p e l o c a l i t y : Rio di Bocca d’Asino (Alessandria), Piedmont, Italy. T y p e s t a g e : “Marne di Sant’Agata Fossili”; Miocene: Tortonian. E t y m o l o g y : This species is named after the name of the first author’s wife.

D e s c r i p t i o n of the holotype: Shell of small size, length up to 3,30 mm; protoconch rounded and paucispiral with a superficial suture and poorly distinct junction with teleoconch; teleoconch regularly biconical with the anterior portion developed more than the posterior;

250


Table 2. Eratoidea antoniae sp. nov., type material, holotype and 13 paratypes. type

repository

type locality

L

figs

Holotype

MGPT-PU 135035


3.3

figs 4 L–P

Paratype 1

MGPT-PU 135036


3.3

Paratype 2

MZB 32036


4

Paratype 3

MGGC 23532


3.5

Paratype 4

MSNG 57968


3.9

Paratype 5

SMF 347151


3.4

Paratype 6

MSNG 57968


3.46

Paratype 7

PG


3.7

Paratype 8

MB


3.1

Paratype 9

BD 125


3.3

Paratype 10

MSNG 57968


4

Paratype 11

MZB 32037


3.3

Paratype 12

MGGC 23532


4

Paratype 13

SMF 347152


3.3

surface smooth; suture hardly distinguishable; aperture quite broad, 73% of total height; external varix strong, flat, more developed in the central part; siphonal notch absent; posterior sinus broad; columellar lip with four plications occupying more than 56% of the aperture, becoming from horizontal to very oblique abapically, the last plication forming the lower columellar margin. Siphonal notch absent. Va r i a b i l i t y : The range of variability of the characters measured is reported in Appendix 1. The examined specimens show a low variability, except in the apical angle which shows a range between 60° and 88°. R e m a r k s : Eratoidea antoniae differs from the syntype of Eratoidea eratoformis (Hoernes & Auinger 1880) from Lysice (Czech Republic), by the more globose shape of the shell, the larger developing of the teleoconch’s anterior portion and the less convexity of the whorls, the greater dimension of the apex, the presence of a more pronounced external varix, the concavity of the columellar side, the thickness of columellar plications and outer lip and the breadth of anterior channel; we can not document the eventual presence of denticulations on the outer lip inside, on the base of the image of the syntype of E. eratoformis (figs 4 Q–S). eschweizerbartxxx sng-

Eratoidea antoniae differs from “E.” eratoformis (sensu Landau et al. 2006 = Stazzania parva Muňiz Solis 2002) from the Pliocene of Estepona (Spain) (figs 4 T–V) by the smaller size (3.80–4.60 mm lenght in S. parva vs. 2.67–4 mm in E. antoniae), more concave columellar side, more robust outer lip callus, more elevated spire, slight denticulation on the inside of outer lip, more convexity of the apex, the shoulder shape and the sinuosity of the outer side of the anterior channel. D i s t r i b u t i o n : Late Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin: Rio di Bocca d’Asino (this paper). Genus Marginella Lamarck 1799 T y p e s p e c i e s : Voluta glabella Linnaeus 1758, by original designation. D i s t r i b u t i o n : The genus is present from Miocene and Pliocene of Italy and Pliocene of Spain to Recent Indo-Pacific, West Atlantic, West African, South African sea with distribution from intertidal to more 2,000 m (Coovert & Coovert 1995). R e m a r k s : The genus is characterized by shells of small to very large size; biconical to obconical, without

Fig. 4. A–H. Dentimargo elusiva sp. nov. — A–D, F) Holotype MGPT-PU 135033, Rio di Bocca d’Asino, shell length 9.84 mm, A–C) entire shell, D) apical view, F) detail of the aperture. — G) Paratype 1, MSNG 57968, Rio di Bocca d’Asino, shell length 9.02 mm, entire shell. — E, H) Paratype 3, MGPT-PU 135034, Rio di Bocca d’Asino, shell length 6.64 mm, E) apical view, H) entire shell. — I–K. Marginella deshayesi Michelotti 1847, Rio di Bocca d’Asino, shell length 27 mm, entire shell (BD). — L–P. Eratoidea antoniae sp. nov., Holotype MGPT-PU 135035, Rio di Bocca d’Asino, shell length 3.3 mm, L–N) entire shell, O) apical view, P) detail of the aperture. — Q–S. Marginella eratoformis Hörnes & Auinger 1880, Syntype, NHMW 1865/0015/0008, shell length 3.4 mm, Q–R) entire shell, S) original label. — T–V. “Eratoidea” eratoformis, Estepona (Spain), Pliocene, length 3.76 mm, T) apical view, U–V) entire shell (MS).



252


axial costae; spire of low to medium height; aperture broad; lip thickened, smooth to denticulate, with external varix; siphonal notch usually present; columella with four strong plications occupying more than half the aperture (Coovert & Coovert 1995). Marginella deshayesi Michelotti 1847 Figs 4 I–K 1847 1862 1873 1881 1890 1899 1904 1904 1904 1905 1914 1940 1960 1968 1973

Marginella deshayesi Michelotti: 321, pl. 17, fig. 16. Marginella deshayesii (sic!) Micht. – Doderlein: 24. Marginella marginata Bon. – Cocconi: 104. Marginella marginata Bon. – Coppi: 47. Marginella (Glabella) deshayesi Micht. – Sacco: 25, pl. 2, fig. 9. Marginella deshayesi Mich. – Cossmann: 88. Eratoidea ? deshayesi (Micht.) var. subadentata Sacco (= var. A. Bell.): 91, pl. 19, fig. 40. Eratoidea deshayesi var. fuscomaculata Sacco (= var. B. Bell.): 92. Eratoidea deshayesi var. subrectelabiata Sacco (= var. C. Bell.): 92, pl. 29, fig. 41. Marginella deshayesi Micht. – Sacco: 907. Marginella (Eratoidea) deshayesi Micht. – Bongo: 438. Marginella (Glabella) deshayesi Micht. – Chavan: 106. Marginella deshayesi Michelotti, sp. 1847 – Glibert: 92. Marginella (Eratoidea) deshayesi Michelotti – Robba: 466, 471, 569, pl. 43, fig. 8 a, b. Marginella (Eratoidea) cf. deshayesi Michelotti – Marasti: 80, 81, 95. Marginella (Glabella) deshayesi Michelotti – Ferrero Mortara et al.: 176. Marginella deshayesi Michelotti – Landau et al.: 24, pl. 1, fig. 4. Marginella (Eratoidea) deshayesi Michelotti – Caprotti: 65. eschweizerbartxxx sng-

1981 2006 2011

T y p e m a t e r i a l : Coll. Michelotti nel Museo di Geologia di Roma (fide Sacco 1890). T y p e l o c a l i t y : “Tortone”. T y p e s t a g e : “Marne di Sant’Agata Fossili”; Miocene: Tortonian. M a t e r i a l e x a m i n e d : 8 shells (BD, MS).

D e s c r i p t i o n : Shell large, length up to about 30 mm; aperture from 71 % to 80 % of total length; shape biconical with the anterior side more developed than the posterior; body whorl shows a subsutural depression, shoulder rounded; aperture broad and fairly arcuate; columellar lip with four strong plications occupying 65 % of aperture; right lip shows in the internal side 10–12 slightly knotty denticulations, external side with a strong varix. Va r i a b i l i t y : The range of variability of the characters measured is reported in Appendix 1. The examined specimens show a low variability, except in the apical angle which shows a range between 89° and 105°.

R e m a r k s : The species is present in Rio di Bocca d’Asino and Sant’Agata Fossili, with well preserved shells. The attribution of some previous authors to the genus Eratoidea Weinkauff 1869 is inconsistent mainly for the dimension of the shells (not exceeding 11 mm in the adult shells) and for other characters well evidenced by Coomans (1976) and Coovert & Coovert (1995). Marginella stephaniae C osta 1866, known from the Miocene (Tortonian) of Portugal, placed by Sacco (1890) in synonymy of M. deshayesi, is a distinct species, and does not occur in the Miocene of Italy. Marginella deshayesi has quite a different shell, with an higher spire, more rounded shoulder, narrower aperture and denticulate lip (Silva et al. 2011). D i s t r i b u t i o n : Late Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin (Sacco 1890; Bongo 1914; Robba 1968; this paper); Morocco: Dar-bel-Hamri (Chavan 1940; Glibert 1960; Robba 1968); Algeria (fide Robba 1968). Pliocene: Morocco: Oued Arjet (fide Glibert 1960). “Marginella” giuntellii sp. nov. Figs 3 A–E O t h e r m a t e r i a l : Rio di Bocca d’Asino: 10 shells (MS); S. Agata Fossili: 2 shells (MB, MS). T y p e l o c a l i t y : Rio di Bocca d’Asino (Alessandria), Piedmont, Italy. T y p e s t a g e : “Marne di Sant’Agata Fossili”; Miocene: Tortonian. E t y m o l o g y : The specific name honours our friend Piero Giuntelli, who collected part of the material here presented and greatly contributed to the knowledge of the Miocene malacofauna of Piedmont.

D e s c r i p t i o n of the holotype: Shell of minute size, length 3.34 mm; protoconch paucispiral with a superficial suture and poorly distinct junction with teleoconch; teleoconch biconical with the anterior portion more developed than the posterior, shoulders angulate to carinate; surface smooth; suture indistinct; last whorl with axial sculpture consisting of nine strong axial costae, slightly prosocline and arcuate, which disappear at 3/5 of the base; aperture broad and prosocline inclined, about 63% of total height; outer lip internally smooth, with a strong external varix; siphonal notch absent; posterior sinus rather broad; columellar lip with four strong plications occupying more than half the aperture. Va r i a b i l i t y : The range of variability of the characters measured is reported in Appendix 1. The examined specimens show a scarce variability, except in amplitude of the apical angle which shows a range between 82° and 100°. R e m a r k s : It is difficult to assign this species to a particular genus. The size, the presence of axial sculpture consisting of strong costae, the teleoconch biconical with the anterior portion more developed than the poste-


Table 3. “Marginella” giuntellii sp. nov., type material, holotype and 9 paratypes. type

repository

type locality

L

figs

Holotype

MGPT-PU 135031


3.34

figs 3 A–E

Paratype 1

MZB 32035


3.48

Paratype 2

MSNG 57966


3.22

Paratype 3

SMF 347153


3.25

Paratype 4

MB


3.3

Paratype 5

BD 126


3.13

Paratype 6

MGGC 23531


3.41

Paratype 7

MS


3.16

Paratype 8

MGPT-PU 135032


3.39

Paratype 9

PG


3.07

rior, and the presence of four columellar folds could be consistent with the attribution of this species to the genus Hiwia Marwick 1931, only known by two species, H. amplificata (Marwick 1931) from the Oligocene of New Zealand and H. aldingae (Tate 1878) from the Eocene of Australia (Coovert & Coovert 1995). In the European Neozoic there are no records of species attributable to Hiwia, and for this reason we prefer to attribute provisionally the new species to the genus Marginella. The shells belonging to the genus Marginella have sometimes axial sculpture, that always differs from “Marginella” giuntellii by the number of always less strong ribs. The only comparable shells are placed in the genus Faba Fischer 1883 (= Glabella Tryon, ex majore parte, non Glabella Swainson), which presents a very large shell, a eschweizerbartxxx sng-

bulging siphonal fasciole and the outer lip strongly dentate within (Marwick 1931). “Marginella” giuntelli differs from the other species of Marginellidae from the European Miocene by the axial sculpture consisting of strong costae, while in all other European Miocene species there is no trace of sculpture. Only a species from the French Upper Oligocene, reported as Stazzania rhytidobasis Lozouet 1999, has an axial sculpture formed by thin ribs on the base of the last whorl (Lozouet 1999), different from that of “Marginella” giuntelli, present on the last whorl with nine strong axial costae, slightly prosocline and arcuated, which disappear at 3/5 of the base. D i s t r i b u t i o n : Late Miocene: Proto-Mediterranean Sea (Tortonian): Po Basin: Rio di Bocca d’Asino, S. Agata Fossili (this paper).

Discussion This work greatly extends the knowledge of the Marginellidae from the Tortonian of Piedmont, Italy. Seven species are here reported, three of which described as new (Dentimargo elusiva, Eratoidea antoniae, and “Marginella” giuntellii), three already known (Stazzania marginata, Volvarina oblongata, and Marginella deshayesi) and one assigned only at generic level (Volvarina sp.). These findings represent an important extension of the data reported by Sacco (1890, 1904), where the Marginellidae of the Late Miocene were poorly represented. Marginellidae are found at three localities in the surroundings of Stazzano (Alessandria), and are represented by 89 shells (Table 4), mainly from Rio di Bocca d’Asino, which is the more sampled site. In the European Miocene, Marginellidae only occurred in the European Atlantic (Silva et al. 2011), in the Paratethys (Hoernes & Auinger 1880) and in Italy in the Burdigalian/Langhian of the Turin hills (Sacco

1890; Ferrero Mortara et al. 1984) and in the Tortonian of the Po Basin, in the surroundings of Stazzano, Alessandria (Bongo 1914; Sacco 1890, 1904; Robba 1968; this paper). In the European Atlantic with the exception of Marginella stephaniae from southern and west central Portugal, only Volvarina is well represented (Lozouet et al. 2001). In the Paratethys the only species of Marginellidae reported is Eratoidea eratoformis (Hoernes & Auinger 1880) from Lysice (Czech Republic), that differs strongly from E. antoniae by several characters (see above). In the Turin hills Sacco (1890) reported several species of Marginella (M. borsoni, M. longa, M. excavata, M. affinis) and Volvarina (V. elongata, V. parvula). The four species of Marginella present in the Turin hills all differ from both Tortonian species by different shape, with the posterior part more developed. They also differ from Marginella deshayesi by the smaller size (20–25

254


Table 4. Number of shells found by locality/species. Rio di Bocca d’Asino

S. Agata Fossili

Castellania

Volvarina oblongata

4

2

1

Volvarina sp.

2

Stazzania marginata Dentimargo elusiva sp. nov. Eratoidea antoniae sp. nov.

20

4

4

22

Marginella deshayesi 8 “Marginella” giuntellii sp. nov. 20

2

total

8

1

80

mm vs. 25–30 mm), and from “Marginella” giuntellii by the larger size (20–25 mm vs. 3–3.5 mm, and the absence of axial sculpture. Two species of Volvarina are known from the Turin hills, one of them (V. parvula) differs by the less developed aperture and the more elevated posterior sinus, while the syntype of the other species (V. elongata) is represented by one incomplete larger shell (27 mm), with more acute apex. With regard to the rather clear cut differences between the Tortonian species and those of the Early Miocene of the Torino hills it seems that direct phylogenetic relationships are not clearly evident. This is consistent with observations on several other families (Sacco 1890; Robba 1968; Zunino & Pavia 2009). eschweizerbartxxx sng-

A particularly important result of the present revision of the Tortonian species is the extension of the distribution of the genus Dentimargo to the Miocene (Tortonian) of Italy. Dentimargo has a wider distribution, and was already known from the Eocene of France and the Pliocene of Spain. During the Messinian crisis, salinity conditions became prohibitive for the survival of mollusc macrofauna (Sabelli & Taviani 1984). Thus, it appears that the incongruous paleobiogeographic distribution of the Mediterranean Neogene molluscs must derive from a gap in the fossil record of the Neogene of the Atlantic region between Morocco and Portugal. The only reasonable explanation is that a part of the Miocene fauna survived in the Atlantic Ocean (very probably not far from the area of Gibraltar) and re-immigrated in the Mediterranean Sea with the re-establishment of acceptable hydrological conditions (Sabelli & Taviani 1984; Silva et al. 2011). Genera such as Marginella and Volvarina were well represented in the European Atlantic Miocene, so their presence in the Atlantic and Mediterranean Pliocene was predictable where many species of Marginellidae are known (Muñiz-Solis 2002; Landau et al. 2006; Silva et al. 2011). Acknowledgements We thank Piero Giuntelli (Torino, Italy) and Bernie Lan(Albufeira, Portugal) for the loan/gift of material from their own collection; Mathias Harzhauser (Naturhistorisches Museum Wien, Vienna, Austria) for the photos of the syntype of Eratoidea eratoformis; Angelo Vazzana (Reggio Calabria, Italy) for the informations on Marginella benasterensis. Ronald Janssen (SMF) and an anonymous reviewer provided very constructive criticisms of the manuscript. dau

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P onder , W.F. (1998): Family Marginellidae. — In: B ees ley P.L., Ross C.J.B. & Wells A.: Mollusca: The Southern Synthesis. Fauna of Australia, 5: 838–841; CSIRO Publishing. Redfield, J.H. (1871): Catalogue of the known species, Recent and fossil, of the family Marginellidae. — Americal Journal of Conchology, 6: 215–269. Robba, E. (1968): I molluschi del Tortoniano-tipo (Piemonte). — Rivista italiana di Paleontologia e Stratigrafia, 74: 457–646. Sabelli, B. & Taviani, M. (1984): The paleobiogeographic distribution of the Mediterranean benthic mollusk and the messinian salinity crisis or where did the mollusks go?. — Annales Geologiques des Pays Helleniques, 33: 263–269. Sacco, F. (1890): I molluschi dei terreni terziarii del Piemonte e della Liguria, 6. Volutidae, Marginellidae, Columbellidae. — Memorie della Reale Accademia delle Scienze di Torino, 2 (40): 295–368 [reprint Torino (C. Clausen): 1–76, 2 pls; Marshall 1991].

Manuscript submitted: 02.03.2015 Revised manuscript accepted: 07.10.2015


Appendix 1 – Range of variability of the characters measured. Volvarina oblongata (Sacco, 1890)

L

W

LLW

LA

DAS

AA

DN

NI

NE

W/L

min

9.93

4.02

7.90

6.48

3.66

53

0.30

4

0

0.37

LLW /L 0.72

max

11.61

5.01

10.49

8.87

4.35

61

0.52

4

0

0.45

mean

10.89

4.41

9.55

7.72

3.95

57

0.42

4

0

0.40

0.31

0.81

0.72

0.24

3.74

0.07

0

0

0.02

st. dev. 0.59

0.67

DAS / LA 0.44

0.93

0.77

0.58

0.88

0.71

0.52

0.06

0.05

0.05

LA / L

Volvarina sp.

L

W

LLW

LA

DAS

AA

DN

NI

NE

W/L

min

7.74

3.48

7.13

5.22

2.64

51

0.31

4

0

0.44

LLW /L 0.90

0.67

DAS / LA 0.51

max

8.39

3.71

7.54

5.80

3.04

55

0.47

4

0

0.45

mean

8.07

3.60

7.34

5.51

2.84

53

0.39

4

0

0.45

0.92

0.69

0.52

0.91

0.68

0.51

st. dev. 0.46

0.16

0.29

0.41

0.28

2.83

0.11

0

0

0.01

0.02

0.01

0.01

DN 0.21

NI 4

NE 0

W/L

LLW /L

LA / L

DAS / LA

0.58

0.85

0.70

0.60

LA / L

Stazzania marginata (Michelotti, 1847)

L

W

LLW

LA

DAS

min

9.87

6.60

9.01

7.40

4.91

AA 74

max

14.17

8.70

12.30

11.07

6.70

92

0.33

4

0

0.72

0.92

0.81

0.72

5.69 0.85

85

0.30

4

0

0.65

0.90

0.73

0.67

11.67

0.05

0

0

0.06

0.02

0.03

0.03

mean

11.69 st. dev. 2.74

7.55 1.01

10.50 2.16

8.47 1.73

Dentimargo elusiva sp. nov.


L

W

LLW

LA

DAS

AA

DN

NI

NE

W/L

min

6.64

3.61

5.45

3.38

2.53

50

0.70

4

1

0.53

LLW /L 0.77

0.48

DAS / LA 0,73

max

9.84

5.28

7.83

5.17

3.86

51

0.81

4

1

0.54

0.82

0.53

0,75

mean

8.81

4.73

6.98

4.44

3.30

50

0.78

4

1

0.54

0.79

0.50

0.74

st. dev. 1.49

0.77

1.07

0.76

0.56

0.5

0.05

0

0

0.01

0.02

0.02

0.01

LA / L

Eratoidea antoniae sp. nov.

L

W

LLW

LA

DAS

AA

DN

NI

NE

W/L

min

2.67

1.75

2.35

1.80

1.22

60

0.19

4

0

0.58

LLW /L 0.85

0.64

DAS / LA 0.54

max

4.00

2.40

3.50

2.90

1.72

88

0.26

4

8

0.67

0.92

0.73

0.69

mean

3.44

2.14

3.05

2.37

1.44

76

0.23

4

3.53

0.62

0.89

0.69

0.61

st. dev. 0.02

0.09

0.08

0.06

0.13

7

0.02

0

1.41

0.03

0.02

0.01

0.04

LA / L

Marginella deshayesi Michelotti, 1847

L

W

LLW

LA

DAS

AA

DN

NI

NE

W/L

min

21.80

15.60

19.1

17.30

11.38

89

0.88

4

4

0.62

LLW /L 0.73

max

29.86

18.65

27.55

24.20

15.08

105

1.65

4

13

0.75

mean

26.48

17.24

23.54

20.20

13.03

96

1.21

4

9.5

1.11

3.03

2.26

1.38

5.80

0.25

0

3.02

st. dev. 3.03

0.71

DAS / LA 0.61

0.93

0.85

0.69

0.66

0.89

0.76

0.65

0.05

0.07

0.05

0.03

LA / L

258


“Marginella” giuntellii sp. nov.

L

W

LLW

LA

DAS

AA

DN

NI

NE

W/L

min

3.07

2.13

2.74

1.99

1.24

82

0.15

4

0

0.66

LLW /L 0.84

max

3.48

2.40

3.08

2.42

1.46

100

0.20

4

0

0.71

mean

3.28

2.24

2.90

2.15

1.39

89

0.17

4

0

st. dev. 0.13

0.09

0.12

0.13

0.07

5.67

0.02

0

0


0.61

DAS / LA 0.60

0.93

0.72

0.68

0.69

0.89

0.66

0.65

0.02

0.02

0.03

0.03

LA / L