EOCENE, OLIGOCENE AND MIOCENE NATICID GASTROPODS OF NORTHERN ITALY

Ermanno Quaggiotto

The present paper covers 77 species and six subspecies of naticid gastropods occurring in the Eocene, Oligocene and Miocene units of Northern Italy; all are described and commented on in the systematic account. Forty-two taxa are identified as formerly described ones; of these, 16 are associated with a generic name different from the previous one, and four are ranked at a different taxonomic level. Twenty-four taxa are proposed as new and the rest likely represent undescribed species, but more, better preserved material is required in order to establish their identity and to name them. Of the 83 taxa considered, 17 are assigned to the genus Cochlis Röding, 1798, one belongs in Cryptonatica Dall, 1892, another belongs in Tanea Marwick, 1931, 11 are members of the genus Tectonatica Sacco, 1890, three are included in Ampullonatica Sacco, 1890, one belongs in Eunaticina Fischer, 1885, 20 are assigned to Euspira Agassiz in J. Sowerby, 1837, three belong in Neverita Risso, 1826, five belong in Payraudeautia Bucquoy, Dautzenberg & Dollfus,1883, two agree with the characters of Pliconacca Cossmann & Martin in Martin, 1914, four are members of Polinices Montfort, 1810, two belong in Sigatica Meyer & Aldrich, 1886, four belong in Sigaretotrerma Sacco, 1890, and nine have the characters of the genus Sinum Röding, 1798. All genera are discussed in terms of type species designation, distinctive characters and subfamilial affiliation (when different from the current one). The character analysis has shown that the measurable elements of the protoconch along with its microsculpture (if any) are diagnostic in many species of all subfamilies. The features of the outer surface of the calcareous operculum proved to be diagnostic in the Pliocene and Recent naticine species. In the present case, the operculum of only four Co-chlis species is known, readily differentiating them from one another. The umbilical characters, i.e. umbilicus width and presence/absence and strength of inner spiral structures, proved to be of variable utility in identifying species. The inner umbilical characters are never species-diagnostic in the case of the naticine taxa, particularly those of Co-chlis. The inner structures, primarily presence/absence, number and strength of cordlets or threads on the umbilical bottom, are more effective in distinguishing poliniceine taxa. The umbilical callus can be used diagnostically for a limited number of naticine and poliniceine taxa. The exterior sculpture of the teleoconch (absent in the naticine and in most poliniceine taxa) is well developed in all the sinine species. In the case of Sigaretotrema and Sinum, the sculptural characters proved to be relevant in species recognition. The color pattern is diagnostic in most of the taxa in which this character is preserved. The shell morphology and the apertural characters are relevant only in a few cases. In the absence of diagnostic elements, a combination of all characters can help in recognizing species.

Rivista Italiana di Paleontologia e Stratigraia (Research in Paleontology and Stratigraphy) vol. 122(2): 109-234. July 2016 EOCENE, OLIGOCENE AND MIOCENE NATICID GASTROPODS OF NORTHERN ITALY ELIO ROBBA1, LUCA PEDRIALI2 & ERMANNO QUAGGIOTTO3 1 Università di Milano Bicocca, Dipartimento di Scienze dell’Ambiente e del Territorio e di Scienze della Terra, Sezione Scienze Geologiche e Geotecnologie, 20126 Milano, Italy. E-mail: robba.elio@gmail.com 2 Via S. Pertini 29, 44124 San Martino, Ferrara, Italy. E-mail: lumaca.fe@libero.it 3 Via Secula 13, 36023 Longare, Vicenza, Italy. E-mail: ermanno.quaggiotto@libero.it To cite this article: Robba E., Pedriali L. & Quaggiotto E. (2016) - Eocene, Oligocene and Miocene naticid gastropods of Northern Italy . Riv. It. Paleont. Strat., 122(2): 109-234. Key words: Taxonomy, Gastropoda, Naticinae, Poliniceinae, Sininae, Paleogene, Miocene, Northern Italy. Abstract. The present paper covers 77 species and six subspecies of naticid gastropods occurring in the Eocene, Oligocene and Miocene units of Northern Italy; all are described and commented on in the systematic account. Forty-two taxa are identiied as formerly described ones; of these, 16 are associated with a generic name different from the previous one, and four are ranked at a different taxonomic level. Twenty-four taxa are proposed as new and the rest likely represent undescribed species, but more, better preserved material is required in order to establish their identity and to name them. Of the 83 taxa considered, 17 are assigned to the genus Cochlis Röding, 1798, one belongs in Cryptonatica Dall, 1892, another belongs in Tanea Marwick, 1931, 11 are members of the genus Tectonatica Sacco, 1890, three are included in Ampullonatica Sacco, 1890, one belongs in Eunaticina Fischer, 1885, 20 are assigned to Euspira Agassiz in J. Sowerby, 1837, three belong in Neverita Risso, 1826, ive belong in Payraudeautia Bucquoy, Dautzenberg & Dollfus,1883, two agree with the characters of Pliconacca Cossmann & Martin in Martin, 1914, four are members of Polinices Montfort, 1810, two belong in Sigatica Meyer & Aldrich, 1886, four belong in Sigaretotrerma Sacco, 1890, and nine have the characters of the genus Sinum Röding, 1798. All genera are discussed in terms of type species designation, distinctive characters and subfamilial afiliation (when different from the current one). The character analysis has shown that the measurable elements of the protoconch along with its microsculpture (if any) are diagnostic in many species of all subfamilies. The features of the outer surface of the calcareous operculum proved to be diagnostic in the Pliocene and Recent naticine species. In the present case, the operculum of only four Cochlis species is known, readily differentiating them from one another. The umbilical characters, i.e. umbilicus width and presence/absence and strength of inner spiral structures, proved to be of variable utility in identifying species. The inner umbilical characters are never species-diagnostic in the case of the naticine taxa, particularly those of Cochlis. The inner structures, primarily presence/absence, number and strength of cordlets or threads on the umbilical bottom, are more effective in distinguishing poliniceine taxa. The umbilical callus can be used diagnostically for a limited number of naticine and poliniceine taxa. The exterior sculpture of the teleoconch (absent in the naticine and in most poliniceine taxa) is well developed in all the sinine species. In the case of Sigaretotrema and Sinum, the sculptural characters proved to be relevant in species recognition. The color pattern is diagnostic in most of the taxa in which this character is preserved. The shell morphology and the apertural characters are relevant only in a few cases. In the absence of diagnostic elements, a combination of all characters can help in recognizing species. IntroductIon The present paper, following another three devoted to the Pliocene naticids of Italy (Pedriali & Robba 2005, 2008a, 2009), examines the Eocene through Miocene naticids and aims to complete the treatment of Italian Tertiary naticids. It covers 77 species and six subspecies (11 of them also dealt with in the cited papers on the Pliocene Naticidae). Of these, 21 occur in the Eocene, 13 in the Oligocene, and 29 in the Miocene. The rest range from Received: July 15, 2015; accepted: February 7, 2016 Eocene to Oligocene (one), from Oligocene to Miocene (six), from Miocene to Pliocene (seven), and from Miocene to Pleistocene/Recent (ive); Neverita olla (de Serres, 1829) is the most widely distributed species, collected from Early Oligocene to Pliocene. The Tertiary naticid taxa of Italy identiied so far result to be 103 in total (Pedriali & Robba 2005, 2008a, 2009, and present paper). Other species proposed or referred to in the literature (e.g. Natica achatensis Recluz, 1841, N. canovae Oppenheim, 1901, N. ottiliae Penecke, 1885, N. scapulata Oppenheim, 1901, N. viatrix Vinassa de Regny, 1898) were neither found in the collections examined, nor were 110 Robba E., Pedriali L. & Quaggiotto E. Fig. 1 - Sketch map of Italian naticid localities; locality numbers are those in the Appendix. they recovered during ield work. Further research might increase the total number of naticid taxa indicated above for the Tertiary of Italy. This study is based on about 5.000 specimens collected from 53 localities of Northern Italy, mostly in Piedmont and Veneto, a few in Liguria and Emilia-Romagna (Fig. 1); brief information on the Italian localities is provided in the Appendix. Additional material relevant to the research, from the Eocene of Great Britain and France, along with specimens from the Miocene of France and The Netherlands, was incorporated. Pertinent naticids in the vast Bellardi-Sacco collection (Museo di Geologia e Paleontologia dell’Università di Torino) and in other public and private collections (see the list in the Systematic account) were examined. Despite the fair to poor preservation of the studied shells, several specimens with preserved apical whorls and/or still retaining the color were present in this large collection. Consequently, the shell characters of most species could be observed and analyzed in terms of range of variation and diagnostic value. Forty-two taxa (out of 83) are identiied as formerly described ones, introduced in the course of the 19th and earliest 20th century; of these, 16 are associated with a generic name different from the previous one, and four are ranked at a different taxonomic level. Twenty-four taxa are herein proposed as new and the rest (17) almost certainly represent undescribed species, but more, better preserved material is required in order to establish their identity and to name them. Of the eighty-three taxa considered herein, seventeen are assigned to the genus Cochlis Röding, 1798, one belongs in Cryptonatica Dall, 1892, another belongs in Tanea Marwick, 1931, eleven are members of the genus Tectonatica Sacco, 1890, three are included in Ampullonatica Sacco, 1890, one belongs in Eunaticina Fischer, 1885, twenty are assigned to Euspira Agassiz in J. Sowerby, 1837, three belong in Neverita Risso, 1826, ive belong in Payraudeautia Bucquoy, Dautzenberg & Dollfus,1883, two agree with the characters of Pliconacca Cossmann & Martin in Martin, 1914, four are members of Polinices Montfort, 1810, two belong in Sigatica Meyer & Aldrich, 1886, four belong in Sigaretotrerma Sacco, 1890, and nine have the characters of the genus Sinum Röding, 1798. Morphology and character analysIs For a review of the naticid characters, along with information on the significance accorded to them by different workers, reference can be made to Pedriali & Robba (2005, 2009). The terms indicating the parts of the naticid shells as well as the standard measurements are those adopted by the just cited authors and are shown in Fig. 2. Tables are included in the supplementary file. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 111 diameter of the first half-whorl (mm) 0,40 0,35 0,30 0,25 0,20 0,15 0,10 0,05 0,4 0,6 0,8 1,0 1,2 1,4 protoconch diameter (mm) Protoconch Pedriali & Robba (2005) considered the protoconch small when the average value of the diameter falls within a 0.5-1.0 mm range, and regarded a protoconch of 1.226 mm in diameter as medium-sized. The diameter of the initial halfwhorl was said to be very small (less than 0.1 mm), small (0.1-0.2 mm), medium-sized (0.3-0.4 mm), and large (greater than 0.4 mm). Based on both the Pliocene taxa (Pedriali & Robba 2005, 2008a, 2009) and those covered herein, the above classiication is reined as follows. According to the value of its diameter, the protoconch can be small (0.51.0 mm), medium-sized (1.0-1.5 mm), large (1.52.0 mm), and very large (greater than 2.0 mm). The diameter of the irst half-whorl can be very small (less than 0.1 mm), small (0.1-0.2 mm), medium-sized (0.2-0.4 mm), and large (greater than 0.4 mm). The plots of Figs. 3-8 show the values (mostly average values) of the characteristic elements of the protoconch pertaining to the taxa dealt with in this paper, which have measurable larval shells (70 out of 83). Most of the considered taxa have a small to mediumsized larval shell, a few (Euspira piccolii, Sigatica eleonorae and Sinum cryptostomoides) have a large protoconch (Figs. 4, 5), and only Eunaticina sp. has a very large protoconch (Fig. 4). The diameter of the irst half-whorl ranges from very small to medium-sized, whereas it is large only in Sigatica eleonorae (Figs. 3-5). Eleven taxa 0,30 diameter of the first half-whorl (mm) Fig. 2 - Standard measurements and illustrated glossary of the terms used for parts of naticid shell and operulum (from Pedriali & Robba 2005). Fig. 3 - Relationship between diameter of the irst half-whorl of the protoconch and protoconch diameter in the naticine taxa; solid diamond: Cochlis craccoi; open diamond: C. degrangei; gray-shaded diamond: C. epigloafuniculata; solid circle: C. epiglopardalis; open circle: C. epiglottina epiglottina; gray-shaded circle: C. mortoni; solid triangle: C. neglecta; open triangle: C. pseudovittata; gray-shaded triangle: C. raropunctata raropunctata; solid square: C. sallomacensis; open square: C. separata; grayshaded square: C. tigrina; multiplication sign: Cryptonatica noe; gray-shaded multiplication sign: Tanea dillwyni koeneni; bold multiplication sign: Tectonatica albertii; cross: T. astensis; graydashed cross: T. burtoni altavillensis; bold cross: T. consimilis; asterisk: T. dertomamilla; gray-shaded asterisk: T. miocolligens; bold asterisk: T. pasinii; dash: T. rupeliana; bold dash: T. tectula; large solid triangle: Tectonatica sp. 1; large open triangle: Tectonatica sp. 2. 0,25 0,20 0,15 0,10 0,05 0,4 0,6 0,8 1,0 1,2 1,4 1,6 1,8 2,0 2,2 protoconch diameter (mm) Fig. 4 - Relationship between diameter of the irst half-whorl of the protoconch and protoconch diameter in the poliniceine taxa; solid diamond: Ampullonatica pseudorepressa; open diamond: Ampullonatica sp.; gray-shaded diamond: Eunaticina sp.; solid circle: Euspira gianoi; open circle: E. giuntellii; gray-shaded circle: E. grossularia; solid triangle: E. helicina helicina; open triangle: E. latecallosa; gray-shaded triangle: E. molarensis; solid square: E. notabilis; open square: E. perforata; gray-shaded square: E. piccolii; gray-shaded multiplication sign: E. pulchella; multiplication sign: E. submamillaris; bold multiplication sign: E. subobturata; cross: E. umbilicocarinata; gray-shaded cross: E. umbilicolunata; bold cross: Euspira sp. 2; asterisk: Euspira sp. 4; gray-shaded asterisk: Euspira sp. 5; bold asterisk: Euspira sp. 6; large solid triangle: Neverita maga; large open triangle: N. olla. Robba E., Pedriali L. & Quaggiotto E. diameter of the first half-whorl (mm) 0,60 0,50 0,40 0,30 0,20 0,10 0,00 0,4 0,6 0,8 1,0 1,2 1,4 1,6 1,8 protoconch diameter (mm) Fig. 5 - Relationship between diameter of the irst half-whorl of the protoconch and protoconch diameter in other poliniceine and sinine taxa; solid diamond: Payraudeautia bituberculata; open diamond: P. crassicorda; gray-shaded diamond: P. fasciolata; solid circle: P. zarantonelloi; open circle: Pliconacca plicatulaeformis; gray-shaded circle: P. tortonensis; solid triangle: Polinices proredemptus; open triangle: P. redemptus; gray-shaded triangle: P. submamilla; solid square: Sigatica claudiae; open square: S. eleonorae; gray-shaded square: Sigaretotrema checchii; multiplication sign: S. clathratum deshayesi; gray-shaded multiplication sign: S. michaudi; bold multiplication sign: Sigaretotrema sp.; cross: Sinum borellense; gray-shaded cross: S. cryptostomoides; bold cross: S. patulum; asterisk: S. paviai; gray-shaded asterisk: Sinum sp. 1; large open triangle: Sinum sp. 2; large solid triangle: Sinum sp. 3. (Cochlis epigloafuniculata, C. epiglottina, C. raropunctata raropunctata, C. tigrina, Cryptonatica noe, Tectonatica albertii, T. consimilis, Euspira giuntellii, Neverita maga, Payraudeautia zarantonelloi and Polinices redemptus) have paucispiral protoconchs relecting non-planktotrophic larval development; all the other examined taxa possess a multispiral larval shell of two to over three whorls and their larval development is inferred to be planktotrophic (Figs. 6-8). From the plots published by Pedriali & Robba (2005, 2008a, 2009), it appears 1) that there is a direct correlation between the protoconch diameter and the number of protoconch whorls, and 2) that the size of the initial half-whorl changes inversely with relation to the number of protoconch whorls. The measurements of the specimens considered in the present paper it with those reported by these authors (graphs omitted). These relationships are relected by the direct correlation between the PD/ DHW ratio and the number of protoconch whorls (Figs. 6-8). Pedriali & Robba (2005, 2008a, 2009) pointed out that the protoconch shows little variation within a species, since the range of whorl numbers vari- es by no more than 0.25 whorls and the diameters (protoconch and irst half-whorl) vary by less than 20%. This outcome is conirmed herein by the 95% conidence intervals that could be calculated for 48 of the 83 taxa considered (see data in the systematic account). It is worth noting that in the case of several species (35) the protoconch was missing or only one was measurable. Nevertheless, considering that the cited rule is based on a considerable number of Tertiary species, we consider it likely to be valid for the Naticidae in general. It follows that a difference of at least 20-25% in diameter (protoconch and/or initial half-whorl) as well as a half whorl difference in the number of whorls are suficient to separate species (see also Pedriali & Robba 2005). The matrices in Tabs. 1-12 consider the same taxa of Figs. 3-8 and are mostly based on average values; single measurements were also incorporated. From these matrices, it appears that the difference in the number of protoconch whorls (Tabs. 1, 4, 7, 10) is diagnostic (signiicant values in bold type) for several species pairs as are the percent difference in protoconch diameter (Tabs. 2, 5, 8, 11) and the percent difference in diameter of the irst half-whorl (Tabs. 3, 6, 9, 12). Since genera have quite distinctive characters, the comments below are presented separately for each genus including at least two species/ subspecies. The values of the characteristic elements of the larval shell (Tabs. 1-3) are diagnostic for six taxa of the naticine genus Cochlis (C. craccoi, C. epigloa20 protoconch diameter/diameter of the first half-whorl 112 18 16 14 12 10 8 6 4 2 0 1,0 1,5 2,0 2,5 3,0 3,5 protoconch whorls Fig. 6 - Relationship between protoconch diameter/diameter of the irst half-whorl ratio and protoconch whorls in the naticine taxa; symbols as in Fig. 3. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 18 16 14 12 10 8 6 4 2 1,0 1,5 2,0 2,5 3,0 3,5 protoconch whorls Fig. 7 - Relationship between protoconch diameter/diameter of the irst half-whorl ratio and protoconch whorls in the poliniceine taxa; symbols as in Fig. 4. funiculata, C. epiglottina epiglottina, C. mortoni, C. raropunctata raropunctata and C. tigrina). Exceptions are C. degrangei whose average values for protoconch whorls, protoconch diameter and diameter of the irst half-whorl do not differ signiicantly from those of C. separata; the same happens with the group C. epiglopardalis, C. neglecta and C. pseudovittata, and with the pair C. epiglopardalis-C. sallomacensis. The characteristic elements of the protoconch (Tabs. 4-6) are diagnostic for ive taxa of the naticine genus Tectonatica (T. albertii, T. consimilis, T. dertomamilla, T. pasinii and Tectonatica sp. 1). No signiicant difference resulted for the group T. astensis, T. burtoni altavillensis and T. tectula, for the group T. burtoni altavillensis, T. miocolligens and Tectonatica sp. 2. The protoconch characters fail to distinguish T. miocolligens from T. tectula, T. rupeliana from T. tectula and T. tectula from Tectonatica sp. 2. Tabs. 7-9 (poliniceine taxa) show that Ampullonatica sp. is readily differentiated from A. pseudorepressa by its signiicantly greater diameter of the irst half-whorl. The considered protoconch characters distinguish Euspira piccolii and E. submamillaris from one another and from all the other Euspira taxa. Exceptions are: 1) E. gianoi differs from all except for E. grossularia and E. umbilicolunata; 2) E. giuntellii is differentiated from all, but not from Euspira sp. 4; 3) E. helicina helicina does not differ signiicantly from E. latecallosa, E. perforata, E. umbilicocarinata and Euspira sp. 6; 4) E. molarensis, E. notabilis, E. pulchella and Euspira sp. 5 have similar protoconch cha- racters. Lastly, for the species pair Neverita maga-N. olla, the examined characters of the larval shell distinguish the two species adequately. From Tabs. 10-12, it appears that the examined characters of the larval shell are signiicant elements in distinguishing the species of the poliniceine genera Polinices and Sigatica from one another. The same happens for the species of the genus Payraudeautia (also poliniceine), with the sole exception of the pair P. crassicorda-P. zarantonelloi. The other two poliniceine species Pliconacca plicatulaeformis and P. tortonensis have similar protoconch characters and cannot be differentiated on this basis. For the sinine genus Sigaretotrema, the characters of the larval shell distinguish species eficiently. As regards the genus Sinum, the following remarks can be made: 1) S. borellense differs signiicantly from all the other species except for Sinum sp. 3, 2) S. cryptostomoides also from all the other, but not from S. paviai, 3) S. patulum has protoconch characters that differ from those of S. paviai, Sinum sp. 2 and Sinum sp. 3, and 4) Sinum sp. 1 is differentiated from S. cryptostomoides, S. paviai and Sinum sp. 3. Tab. 13 summarizes the differences between species on the basis of measurable characters of the larval shell. Bandel (1999), dealing with the protoconch of the naticids, afirmed that “the embryonic whorl is normally ornamented with simple tubercles and the larval shell is covered by more or less visible spiral lirae”. He also remarked that “the larval shell 20 protoconch diameter/diameter of the first half-whorl protoconch diameter/diameter of the first half-whorl 20 113 18 16 14 12 10 8 6 4 2 1,0 1,5 2,0 2,5 3,0 3,5 protoconch whorls Fig. 8 - Relationship between protoconch diameter/diameter of the irst half-whorl ratio and protoconch whorls in other poliniceine and sinine taxa; symbols as in Fig. 5. Robba E., Pedriali L. & Quaggiotto E. 114 2,5 14,0 spire height (mm) maximum diameter (mm) 16,0 12,0 10,0 8,0 2,0 1,5 1,0 6,0 0,5 4,0 4,0 6,0 8,0 10,0 12,0 shell height (mm) 14,0 16,0 18,0 2,0 4,0 6,0 8,0 shell height (mm) Fig. 9 - Relationship between maximum diameter and shell height (species of Tectonatica); open circles: T. albertii and T. burtoni altavillensis combined; solid diamonds: T. pasinii. Fig. 10 - Relationship between spire height and shell height (species of Tectonatica); open circles: T. tectula; solid diamond: T. consimilis. consists of quite a lot of organic material and after metamorphosis usually acquires a more or less corroded appearance”; this fact was also reported for many gastropod families by Hickman (2004, p. 206). We recovered several shells of Tertiary naticids with well preserved protoconchs, either smooth or bearing spiral microsculpture, which may consist of threads or rows of granules restricted to the apical whorl, or of threads present on subsequent whorls, throughout or only over the last whorl. Taking into account Bandel’s and Hickman’s remarks cited above, we think that the unsculptured protoconchs are to be considered really smooth when excellently preserved (mostly Pliocene to Recent material), or apparently smooth in all other instances (Eocene to Miocene material). The protoconch microsculpture seems to occur rarely in the species of Cochlis and was never observed in the species of the other naticine genera considered here (Cryptonatica, Tanea, Tectonatica). Instead, it is more common in the poliniceine and sinine species, in particular those of Euspira and Sinum. Pedriali & Robba (2009, pp. 375, 378) stated that the microsculpture of the larval shell (presence/absence, location on the larval shell, morphological details) may constitute an additional distinctive character of the protoconch, useful in distinguishing species. Concerning the present material, the ornamentation of the protoconch is useful in the case of a few Miocene taxa; it was never recorded in the Eocene and Oligocene ones, most probably because of lack of preservation. Tab. 14 lists those taxa whose protoconch retains the microsculpture (or is certainly smooth) and shows the differences between species based on this character. The present study fully conirms the conclu- sion of Pedriali & Robba (2009, p. 378), i.e. that the larval shell is a most relevant element for distinguishing species within each genus, even if not constantly species-diagnostic. The measurable characters of the protoconch are equally effective in all the three subfamilies dealt with in this paper (see Tab. 13). Teleoconch The diagnostic value of the teleoconch characters of naticids was amply discussed by Pedriali & Robba (2005, 2008a, 2009) largely on the basis of statistical analyses. These authors questioned the taxonomic importance accorded to most teleoconch characters in major accounts of naticids published during the last decades (Cernohorsky 1971; Kilburn 1976; Marincovich 1977; Majima 1989). In the following, measurable characters suitable for statistical analysis are considered along with other qualitative (non measurable) features in order to verify once more the conclusions reported on in the papers of Pedriali & Robba cited above. Since genera have quite distinctive characters (see comments in the systematic account), the statistical treatment of species was executed separately for each genus. The coeficients of correlation for each considered pair of characters were signiicant for all the studied taxa (r = 0.74-0.96). For brevity, only the relevant scatters will be presented; pairs or single characters not discussed herein are of scarce relevance in species recognition. Shell shape. As already stated by Pedriali & Robba (2005), the shell shape can be deined statistically on the basis of the relations between 1) Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 115 9,0 3,5 8,0 3,0 spire height (mm) spire height (mm) 7,0 2,5 2,0 1,5 1,0 6,0 5,0 4,0 3,0 2,0 0,5 1,0 0,0 0,0 2,0 4,0 6,0 shell height (mm) 8,0 10,0 0,0 5,0 10,0 15,0 shell height (mm) 20,0 25,0 Fig. 11 - Relationship between spire height and shell height (species of Euspira); solid circles: E. gianoi; open diamonds: E. grossularia. Fig. 13 - Relationship between spire height and shell height (species of Pliconacca); solid squares: P. plicatulaeformis; open circles: P. tortonensis. maximum diameter and shell height, 2) spire height and shell height, and 3) by the values of the spire angle. The relation between maximum diameter and shell height proved to be useless in distinguishing species except for one case. In Tectonatica, the regression line for T. pasinii and that for T. albertii and T. burtoni altavillensis combined have similar slope, but signiicantly different elevation (Fig. 9). This means that, for a given height of the shell, T. pasinii attains a smaller diameter (or has a comparatively taller shell) than both T. albertii and T. burtoni altavillensis. In Tectonatica, the plot of spire height against the height of the shell shows that the regression lines for T. tectula and T. consimilis have a remarkably different slope (Fig. 10). T. tectula can be distinguished from T. consimilis by having a more elevated spire, which grows much faster with increasing shell height. For this same relationship in Euspira, it appears that E. grossularia differs from E. gianoi in that it has a lower spire (Fig. 11). The same occurs if E. grossularia is compared to E. umbilicolunata; moreover, the spire of E. umbilicolunata grows more rapidly with increasing shell height (Fig. 12). In Pliconacca, the plot of spire height against the height of the shell shows that the lines for P. tortonensis and P. plicatulaeformis have a different slope (Fig. 13); the spire of P. plicatulaeformis grows much faster with increasing shell height than that of P. tortonensis, which attains a smaller size. The spire angle varies considerably in the taxa considered herein and can hardly serve to differentiate the species and the subspecies within each genus since respective 95% conidence intervals largely overlap in most cases. From Tab. 15 it will be seen that the spire angle helps in distinguishing Tectonatica miocolligens, T. pasinii and T. rupeliana from the other Tectonatica taxa, but not from one another. In Euspira, E. giuntellii is readily separated on the basis of the spire angle from E. gianoi, E. helicina helicina, E. notabilis, E. pulchella, E. submamillaris and E. umbilicolunata. In Sinum, S. oligopolitum can be distinguished from S. borellense and S. patulum. This character fails to distinguish species of Cochlis, Neverita, Payraudeautia, Pliconacca, Polinices, Sigatica and Sigaretotrema. 6,0 spire height (mm) 5,0 4,0 3,0 2,0 1,0 0,0 2,0 6,0 10,0 14,0 shell height (mm) 18,0 Fig. 12 - Relationship between spire height and shell height (species of Euspira); open circles: E. grossularia; solid triangles: E. umbilicolunata. Aperture. In order to deine quantitatively the aperture, Pedriali & Robba (2005) used the relationships between 1) aperture width and aperture height, 2) aperture height and height of the shell, 3) aperture width and maximum diameter, and 4) the values of the inner lip slope. In Pliconacca, the relationship between aperture width and aperture height distinguishes P. torto- Robba E., Pedriali L. & Quaggiotto E. 12,0 3,5 10,0 3,0 umbilicus width (mm) aperture width (mm) 116 8,0 6,0 4,0 2,5 2,0 1,5 1,0 2,0 0,5 0,0 0,0 2,0 4,0 6,0 8,0 10,0 aperture height (mm) 12,0 14,0 16,0 0,0 5,0 10,0 15,0 maximum diameter (mm) 20,0 Fig. 14 - Relationship between aperture width and aperture height (species of Pliconacca); solid squares: P. plicatulaeformis; open circles: P. tortonensis. Fig. 16 - Relationship between umbilicus width and maximum diameter (species of Pliconacca); open circles: P. tortonensis; solid squares: P. plicatulaeformis. nensis from P. plicatulaeformis since their regression lines have signiicantly different elevations (Fig. 14). For a given height, the width of the aperture is greater in P. tortonensis than in P. plicatulaeformis. The inner lip slope (inclination of the inner lip to the shell axis) appears to be scarcely important in distinguishing species, since the 95% conidence intervals (Tab. 16) in most cases largely overlap and do not allow any reliable separation. However, a few exceptions are to be noted. In Tectonatica, the conidence interval of T. dertomamilla only slightly overlaps those of T. consimilis and of T. pasinii, most specimens of which have values of the inner lip slope greater than those of T. dertomamilla. Something similar occurs in Pliconacca; most specimens of P. plicatulaeformis have values of the inner lip slope greater than those of P. tortonensis. In Sinum, the inclination of the inner lip differentiates both S. borellense and S. oligopolitum from S. patulum and Sinum sp. 3, but not the species of each pair from one another. This character proved to be useful also in the case of two Pliocene species of Sinum (cf. Pedriali & Robba 2009, p. 380). It seems that the inclination of the inner lip could be more effective in separating species in Sinum than in all the other genera considered herein. The parietal callus is of little use in species recognition; it is thick, subrectangular in almost all taxa and the anterior lobe is present in many species, absent in many others. However, the parietal callus is species-diagnostic in the case of Payraudeautia bituberculata, which is readily differentiated from the other species of the genus by its parietal callus with a rounded knob at both ends (see remarks in the systematic account). umbilicus width (mm) 4,5 3,5 2,5 1,5 0,5 2,5 5,0 7,5 10,0 12,5 maximum diameter (mm) 15,0 17,5 Fig. 15 - Relationship between umbilicus width and maximum diameter (species of Payraudeautia); solid circles: P. crassicorda; solid diamonds: P. zarantonelloi. Umbilical characters. The statistical analysis focused on the relationships between 1) umbilicus width and maximum diameter and 2) width of the umbilical callus and maximum diameter. The irst relationship proved to be useful in a few cases. In Payraudeautia, when the umbilicus width is regressed against maximum diameter (Fig. 15), it appears that the regression lines for P. crassicorda and P. zarantonelloi have rather similar slope, but signiicantly different elevation. This means that most specimens of P. crassicorda, for a given size of the shell, have a wider umbilicus than those of P. zarantonelloi. In Pliconacca (Fig. 16), the regression lines for P. tortonensis and P. plicatulaeformis have different slope and markedly different elevation, i.e. P. tortonensis has a signiicantly wider umbilicus than P. plicatulaeformis. The plot of width of the umbilical callus against maximum diameter in species of Cochlis Eocene, Oligocene and Miocene naticid gastropods of Northern Italy width of umbilical callus (mm) 3,5 3 2,5 2 1,5 1 0,5 0 0,0 1,0 2,0 3,0 4,0 5,0 umbilicus width (mm) 6,0 7,0 8,0 Fig. 17 - Relationship between width of umbilical callus and umbilicus width (species of Cochlis); open circles: C. degrangei; open diamonds: C. infelix. shows that C. degrangei can be distinguished from C. infelix by the greater elevation of the regression line (Fig. 17); for a given size of shell, the umbilical callus of C. degrangei is signiicantly broader than that of C. infelix. On the basis of Tab. 17, which summarizes the inner umbilical characters, the following notes can be made. 1. In Cochlis, these characters are never speciesdiagnostic. However, their combination readily differentiates C. mortoni from C. separata. The strength of the funicle can serve for distinguishing C. craccoi, C. epiglottina epiglottina, C. neglecta, C. pseudovittata, C. sallomacensis and Cochlis sp. 1 (funicle cord-like, thick and prominent) from C. degrangei, C. epigloafuniculata, C. infelix, C. rossii and Cochlis sp. 2 (funicle broad and depressed). 2. In Ampullonatica, the inner spiral sculpture of Ampullonatica sp. easily differentiates it from A. pseudorepressa and A. repressa. 3. In Euspira, the inner spiral sculpture separates E. perforata, E. piccolii and Euspira sp. 3 from one another and from all the other Euspira taxa. The characters of the inner spiral furrow are relevant in identifying E. subobturata, E. umbilicocarinata, Euspira sp. 1, Euspira sp. 4 and Euspira sp. 6, but fail to distinguish E. subobturata from Euspira sp. 1 and Euspira sp. 4 from Euspira sp. 6. The funicle is scarcely useful: at most its absence can serve in separating E. molarensis, E. perforata, E. piccolii and Euspira sp. 3 from the other Euspira taxa. 4. In Payraudeautia, the inner umbilical characters combined are distinctive and readily separate the species from one another. 5. In Pliconacca, P. tortonensis is differentiated 117 from P. plicatulaeformis because of its spiral furrow bounded abaxially by a groove. 6. In Sigatica, the number and strength of the inner spiral cordlets serves for differentiating S. claudiae from S. eleonorae. 7. In Sigaretotrema, the pair S. checchii and Sigaretotrema sp. differs from the pair S. clathratum deshayesi and S. michaudi, which are characterized by the presence of inner spiral threads; this same character does not distinguish the taxa of the latter pair from one another. The umbilical callus, absent in all the sinine taxa, is rather thick to thick in the naticine and in the poliniceine species. Focusing on its shape and its demarcation (if any) from the parietal callus (Tab. 18), the following remarks can be made. 1. In Cochlis, C. epigloafuniculata is distinguished from all the other taxa of the genus listed in Tab. 18 because of the shape of its umbilical callus and the absence of demarcation from the parietal callus. The combination of the two cited characters differentiates C. degrangei, C. tigrina and Cochlis sp. 2 from one another and from the other Cochlis taxa. C. craccoi and C. epiglopardalis, which have a semicircular umbilical callus separated from the parietal callus by a shallow notch, are differentiated from C. epiglottina epiglottina, C. mortoni, C. neglecta, C. pseudovittata and Cochlis sp. 1, also with semicircular umbilical callus, but with a deep, narrow notch in between it and the parietal callus. The depressed, arched umbilical callus separated from the parietal callus by a shallow notch distinguishes C. infelix, C. raropunctata raropunctata and C. rossii from the other Cochlis taxa. C. sallomacensis and C. separata, both with a roundly triangular umbilical callus, can be distinguished from one another because of their different notches (broad and shallow in C. sallomacensis, narrow and deep in C. separata); the characters of the umbilical callus differentiate these two taxa also from the others. From the above, it appears that the umbilical callus is species-diagnostic in a few Cochlis taxa. In most cases, it serves for distinguishing a group of taxa from another group. 2. In Tectonatica, the umbilical callus is never species-diagnostic since it varies slightly in shape and extent in most taxa. However, the large, semicircular umbilical callus of T. consimilis and T. tectula readily separates these species from all the others of the genus, but not from one another. Further, T. miocolligens and Tectonatica sp. 2 are the sole taxa 118 Robba E., Pedriali L. & Quaggiotto E. whose umbilical callus is demarcated from the parietal callus by a notch. 3. In Euspira, the umbilical callus, obsolete in a few species, exhibits little difference in shape, being slenderly triangular or subtriangular in most species, roundly triangular in a few; its abapertural outline can be straight, arched, reverse S-shaped, or reverse J-shaped. Accordingly, this character appears to scarcely useful in distinguishing species. The unique exception concerns E. latecallosa, whose unusually wide umbilical callus with convex adaxial outline differentiates it from all the other species of the genus. 4. In Neverita, the umbilical callus appears to be species-diagnostic in the case of the three species considered here. 5. In Payraudeautia, P. bituberculata and P. fasciolata have similar characters of the umbilical callus, which distinguish them from the pair P. crassicorda and P. intricata, also with similar umbilical callus characters. The umbilical callus is diagnostic in the case of P. zarantonelloi. 6. In Pliconacca, the umbilical callus is diagnostic in the case of the two species dealt with here. 7. In Polinices, the subrectangular umbilical callus distinguishes P. redemptus and P. submamilla from P. proredemptus and Polinices sp., which have a subcircular umbilical callus. Outer surface. Taxa of the naticine genera (Cochlis, Cryptonatica, Tanea, Tectonatica) and of most poliniceine genera (Euspira, Neverita, Payraudeautia, Pliconacca, Polinices) have unsculptured shells, whose outer surface bears only growth lines, changing into subsutural wrinkles in a few species. A faint spiral microstriation occurs on the last whorl in some species of Cochlis (C. epigloafuniculata, C. epiglottina epiglottina, C. neglecta, C. raropunctata raropunctata, C. rossii), of Euspira (E. grossularia, E. helicina helicina, E. notabilis, Euspira sp. 6), and of Pliconacca (P. tortonensis). A spiral sculpture of ribbons, cords and threads is always present in the poliniceine genera Eunaticina and Sigatica, and in all the sinine species. In the case of Sigaretotrema and Sinum, the observed differences in ornamentation details proved to be relevant in species recognition (see remarks on species in the Systematic account). Pedriali & Robba (2005, 2008a, 2009), dealing with Pliocene naticids, noted that several species have distinctive background color and/or color patterns. Unfortunately, this character is poorly preserved or not preserved at all in many older, Tertiary taxa; nevertheless, we tried to deine this character of the shell surface whenever possible. In the case of those taxa ranging also into the Pliocene, the respective color patterns cited by Pedriali & Robba have been considered. The results are shown in Tab. 19. As can be seen, the pre-Pliocene taxa Cochlis epiglopardalis, C. epiglottina epiglottina, C. pseudovittata, C. sallomacensis and C. tigrina have distinctive color patterns, which distinguish them from one another. In Tectonatica, T. consimilis can be readily differentiated from the other Tectonatica taxa on the basis of its pale brown, quadrangular spots forming spiral rows. In Euspira, E. grossularia, E. helicina helicina, E. notabilis and E. pulchella, which extend their range subsequent to the Miocene, can be safely identiied by respective color patterns (Pedriali & Robba 2009). In the case of the other Euspira taxa considered here, the color does not help since it is not preserved, or exhibits similar patterns (E. gianoi, E. submamillaris, E. subobturata, Euspira sp. 2). In Neverita, N. antiqua and N. olla, which co-occur in the Early Oligocene, are readily recognized because of their markedly different color patterns. In Polinices, the color pattern unambiguously differentiates P. proredemptus from P. redemptus. The pattern of brown crescent-shaped marks arranged into collabral alignments occurring in P. redemptus seems to be peculiar of this species and likely distinguish it from all the other Polinices taxa described so far. Operculum. Pedriali & Robba (2009) noted that the calcareous operculum is constantly species-diagnostic in the case of the Pliocene taxa of the naticine genera Cochlis and Tectonatica. Concerning the present material, the operculum of most naticine taxa is unknown. Only a few shells of four Cochlis species (C. epiglopardalis, C. neglecta, C. sallomacensis and C. tigrina) were recovered with the operculum still illing the aperture. These four Cochlis species can be readily differentiated from one another and from C. raropunctata raropunctata and C. epigloafuniculata (opercula described by Pedriali & Robba 2005, 2008a) on the basis of respective opercular characters. Our indings seem to agree with the cited statement of Pedriali & Robba (2009), at least as regards the species of Cochlis. From the character analysis reported above, it appears that the concluding remarks on the Eocene, Oligocene and Miocene naticid gastropods of Northern Italy characters of the Pliocene naticids provided by Pedriali & Robba (2009) are largely conirmed and can be extended to the whole Tertiary Naticidae. Basic points are the following. 1. The protoconch shape is useless since it is constant in all naticids. Conversely, its measurable elements (number of whorls, diameter and diameter of the irst half-whorl) along with its eventual microsculpture are valuable characters, species-diagnostic in most cases. The larval shell appears to be equally effective in all naticid subfamilies. The protoconch is diagnostic only at the species level, whereas it does not serve in distinguishing genera because identical protoconchs occur in species of different genera. 2. The use of shell morphology, largely adopted by previous authors in deining species, is a much subjective approach, which has led to the misinterpretation of several taxa. Statistical analysis (Pedriali & Robba 2005, 2008a, 2009; present paper) has shown that the shell shape is of scarce value, since it helps to distinguish species only in a few cases. 3. The apertural proportions, the inner lip slope and the parietal callus are of little use in species recognition; in the case of Sinum, the inclination of the inner lip seems slightly more effective in separating species. 4. The umbilical characters, i.e. umbilicus width and presence/absence and strength of inner spiral structures, proved to be of variable use in identifying species. The width of the umbilicus (related to maximum diameter of the shell) is useful in very few cases, as is the relationship between width of umbilical callus and maximum diameter. The inner umbilical characters are never species-diagnostic in the case of the naticine taxa, particularly those of Cochlis; at most they distinguish one group of species from another. The inner structures, primarily presence/absence, number and strength of cordlets or threads on the umbilical bottom, are more effective in distinguishing poliniceine taxa. The umbilical callus can be used diagnostically for a limited number of naticine and poliniceine taxa. 5. The exterior sculpture of the teleoconch (absent in the naticine and in most poliniceine taxa) is well developed in all the sinine species. In the case of Sigaretotrema and Sinum, the sculptural characters proved to be relevant in species recognition. 6. The color is the other property of the shell 119 surface; in particular, the color pattern is diagnostic in most of the taxa in which this character is preserved. 7. The calcareous operculum is another relevant character in identifying the naticine species. Pedriali & Robba (2009) noted that the operculum is constantly species-diagnostic in the case of Pliocene taxa, in particular those of Cochlis and of Tectonatica. Also the opercula dealt with herein (six species of Cochlis occurring in the Miocene) differ from one another and likely are species-diagnostic. Unfortunately, specimens are seldom recovered with the operculum in situ and this constitutes an obvious constraint in the use of this solid accessory of the shell when Miocene or older material is considered. 8. In summary, the protoconch (all subfamilies) and the operculum (Naticinae) are the most important elements, diagnostic in many species. The other characters can be ranked in descending order of importance as follows: a) sculpture (Sininae), b) color patterns, c) umbilical characters, d) apertural characters (mostly the inner lip slope and the parietal callus), and e) shell morphology. In the absence of the primary characters (lack of preservation), a combination of the others can help in recognizing species. systeMatIc paleontology We follow a traditional (basically typological) classiication. The suprageneric arrangement is that adopted in major accounts of naticids published during the last decades (Kilburn 1976; Marincovich 1977; Majima 1989; Kabat 1991; Torigoe & Inaba 2011), with some modiications. The bulk of the studied material is housed in the Museo di Paleontologia dell’Università, Milano, Italy (MPUM in the following) and in Museo di Archeologia e Scienze Naturali “G. Zannato”, Montecchio Maggiore, Italy (MCZ in the following); the rest is kept as reference material in the authors’ collection (NP), Università di Milano Bicocca, Dipartimento di Scienze dell’Ambiente e del Territorio e di Scienze della Terra, Sezione Scienze Geologiche e Geotecnologie, Milano, Italy. Other abbreviations for institutions cited in the text as repositories of relevant material are as follows: BGR, Bundesanstalt für Geowissenschaften und Rohstoffe, Berlin, Germany; EM, École des Mines collection (including Deshayes’ original fossil specimens) presently curated by Université de Lyon 1, France; IPUM, Museo dell’Istituto di Paleontologia dell’Università di Modena (including Coppi, Doderlein and Foresti collections) presently curated by Dipartimento di Scienze Chimiche e Geologiche, Università di Modena e Reggio Emilia, Italy; MB, Museum für Naturkunde, Berlin, Germany; MCV, Museo Civico D. Dal Lago, Valdagno, Italy; MGC, Museo Geologico G. Cortesi, Castell’Arquato, Italy; MGP-PD, Museo di Geologia e Paleontologia dell’Università di Padova, Italy; MGPT, Museo di Geologia e Paleontologia dell’Università di Torino, Italy (MGPT BS: Bellardi-Sacco collection presently curated by Museo Regionale di Scienze Naturali, Torino); MHNG, Museum d’Histoi- 120 Robba E., Pedriali L. & Quaggiotto E. re Naturelle, Geneva, Switzerland; MPPLF, Museo Paleontologico e Preistorico “P. Leonardi”, Università di Ferrara, Italy; MS, Musei Civici di Imola-Museo Scarabelli, Imola, Italy; MZB, Museo di Zoologia, Università di Bologna, Italy; MNHN, Muséum National d’Histoire Naturelle, Paris, France; NHM (formerly BMNH), Natural History Museum, London, Great Britain; NHMW, Naturhistorisches Museum, Wien, Austria; RGM, Nationaal Naturhistorisch Museum, Naturalis, Leiden, Holland. Valuable additional material was lent by Forli collection (MF), Prato, Italy, Giuntelli collection (PG), Nole (Torino), Italy, Lesport collection (L), Sainte-Hélène, France, Magenes collection (PPMM), Milano, Italy, and Petracci collection (PP), Cesena, Italy. The citations, which are veriiable in that enclose adequate description and/or illustration of species, and other citations referring to material that has been directly examined by the present authors, are included in the synonymies. Other citations, poorly documented or not documented at all, are considered uncertain references and are usually excluded from the synonymies. Several species covered in previous papers devoted to the Pliocene naticids (Pedriali & Robba 2005, 2008a, 2009) were recovered also from Oligocene to Miocene units. For their description and remarks, reference should be made to the cited papers. Unless otherwise stated, the average values of the dimensions have been recalculated on the basis of cumulated measurements of Pliocene and new Oligocene and/or Miocene specimens. Symbols for shell dimensions (see Fig. 2) are: DHW, diameter of the irst half-whorl of the protoconch; PD, diameter of the protoconch; H, height of the shell; D, maximum diameter; SH, height of the spire; AH, height of the aperture; AW, width of the aperture; UW, width of the umbilicus; WUC, width of the umbilical callus; WAD, width of the adapical sulcus; WAB, width of the abapical sulcus; IS, inner lip slope; SA, spire angle. Unless otherwise stated, for each dimension, ranges in the upper row are 95% conidence intervals, igures in the lower row are average values. In the description of species, the aperture length is the distance between adapical and abapical ends measured in the plane of the aperture; obviously, it does not coincide with the aperture height. Family Naticidae Guilding, 1834 The family name Naticidae, currently attributed to Forbes (1838), was validly introduced four years earlier by Guilding (1834), who has the authorship of the name (Kabat 2000, p. 352; see also Huelsken et al. 2012). Subfamily Naticinae Guilding, 1834 The genera included in this subfamily have teleoconchs of variable shape, mostly smooth; the umbilicus is open in most genera, but largely illed by the umbilical callus in some; a variably robust funicle is often present. Most authors concur in considering the presence of a thick, solid calcareous operculum, sometimes with a corneous inner layer, as a distinctive character of the naticine genera (Cernohorsky 1971; Kilburn 1976; Marincovich 1977; Majima 1989; Wilson 1993; Kabat 1998; Pastorino 2005; Huelsken et al. 2008). Genus Cochlis Röding, 1798 Cochlis Röding, 1798, p. 146. Type species by subsequent designation (Hedley 1916, p. 51): Cochlis lammea Röding, 1798 (= Nerita vittata Gmelin, 1791), Pliocene (Italy) to Recent (western Mediterranean, East Atlantic). Remarks. Cernohorsky (1971) and Torigoe & Inaba (2011, p. 69) regarded Cochlis as a synonym of Natica Scopoli, 1777 evidently relying upon the later designation of Cochlis albula Röding, 1798 (= Nerita vitellus Linnaeus, 1758 type species of Natica) as type species of Cochlis made by Iredale (1924, p. 254). Since the earlier type species ixation by Hedley (1916) is valid ((ICZN 1999, Article 69.1 of the Code) and the later ixation by Iredale (1924) is invalid (ICZN 1999, Article 70.2 of the Code), we concur with Oyama (1985) and Kabat (1991) in considering Cochlis as a distinct, valid genus. Kabat (1990, p. 10) thoroughly discussed Natica vitellus and regarded Nerita spadicea Gmelin, 1791 (= Cochlis albula according to Torigoe & Inaba 2011) as another synonym of N. vitellus. The distinctive characters of Cochlis were outlined by Pedriali & Robba (2005) and are reined herein as follows: 1) protoconch low-turbiniform of 1.25-3.5 smooth whorls, protoconch I with spiral rows of ganules in a few species, 2) teleoconch thin to solid, globose to depressed-globose, body whorl moderately expanded, 3) spire rather depressed to moderately elevated, 4) suture adpressed to channeled, 5) parietal callus thin to thick, short in most species, with poorly developed to indistinct anterior lobe, 6) umbilicus rather small to large, 7) funicle present, thread-like to thick, completely illing the umbilicus in a few species, 8) umbilical callus small to broad, separated from the parietal callus by a reverse J-shaped notch in most species, and 9) outer surface of the operculum with two-three marginal ribs. As already noted by Pedriali & Robba (2005), the characters of the parietal callus, of the umbilicus, and of the operculum combined constitute the primary diagnostic elements of Cochlis. The species belonging in the genus Natica are distinguished from those of Cochlis in that they have: 1) the anterior lobe of the parietal callus well developed, hindering the adapical part of the umbilicus at various extent, 2) the funicle absent or vestigial, and 3) the umbilical callus absent to weak. The presence of a well-deined funicle (regardless of its strength) distinguishes Cochlis from Natica. The species of the Eocene, Oligocene and Miocene naticid gastropods of Northern Italy genus Naticarius Duméril, 1806 have the teleoconch similar to that of Cochlis, but their opercula have the outer surface with many ribs instead of two-three. The operculum of most species, herein included in Cochlis on the basis of their umbilical characters (primarily the presence of a well developed funicle), is unknown. Future recovering of specimens with the operculum in situ may prove that some of them deserve a different generic assignment. Cochlis craccoi sp. n. Pl. 1, igs. 1, 2 Derivation of name: The species is named after Gilberto Cracco, who provided material relevant to the present study. Holotype: Borelli: MGPT-PU 135052 (Pl. 1, ig. 1). Paratypes: Borelli: 1 spm., MGPT-PU 135060 (Pl. 1, ig. 2), 1 spm., MGPT-PU 135053, 1 spm., MGPT-PU 135054, 1 spm., MGPT-PU 135055, 1 spm., MGPT-PU 135056, 18 spms., MGPTPU 135057, 1 spm., MPUM 11209, 12 spms., MGPT-PU 135058, 9 spms., MGPT-PU 135059. Preservation: Most specimens are well preserved; a few have minor damage. Type locality: Borelli (see appendix). Diagnosis: Shell globose with moderately elevated spire and globular last whorl. Umbilicus deep wide to medium-sized. Parietal callus wider adapically; anterior lobe indistinct. Funicle cord-like, moderately thick; umbilical callus moderate, semicircular, separated from parietal callus by broad, shallow notch. Vestige of pale reddishbrown background without any color pattern. Description. Protoconch small, depressed turbiniform, of 2.20-2.32 gently convex, apparently smooth whorls, irst half-whorl small. Teleoconch globose, rather thick. Spire broadly conical, moderately elevated, whorls convex, suture ine, adpressed. Last whorl globular, slightly expanded toward aperture; subsutural shelf indistinct, poorly differentiated in a few specimens. Aperture D-shaped, in prosocline plane, length about twice width; outer lip semicircular, basal lip thickened. Parietal callus moderately thick, wider adapically, with concave abapertural outline; anterior lobe indistinct. Umbilicus deep, large to medium-sized, umbilical wall slightly overhanging interior of umbilicus. Funicle cord-like, moderately thick, thick in a few specimens, separated from umbilical wall by narrow spiral furrow. Umbilical callus moderate, rather thick, semicircular in outline, located at abapical one-third of inner lip, separated from parietal callus by broad, shallow notch. Basal fasciole wide, markedly depressed, nearly smooth. Surface with rather dense, 121 thin growth lines. One specimen retains vestige of uniform pale reddish-brown background apparently without any color pattern. No opercula found. Dimensions (mm) DHW PD H D SH AH AW 0.1450.161 0.9151.007 4.3105.802 4.0705.642 0.6561.472 3.4694.541 2.4493.949 0.153 0.961 5.056 4.856 1.064 4.005 3.199 UW WUC WAD WAB IS SA 1.2961.760 0.1710.859 0.2240.888 0.2960.616 21°-29° 108°132° 1.528 0.515 0.556 0.456 25° 120° Remarks. Cochlis craccoi exhibits some resemblance to Cochlis pseudovittata sp. n. (see below) and Cochlis raropunctata raropunctata (Sasso, 1827). C. craccoi differs from C. pseudovittata in that it has: 1) a probably unsculptured protoconch with signiicantly greater diameter of the irst half-whorl (nearly twice the size), 2) a slightly more globular shell, 3) a proportionally wider umbilicus, 4) an umbilical callus demarcated from the parietal callus by a broad, shallow notch (narrow, moderately deep in C. pseudovittata), and 5) a different color without any pattern. C. craccoi is readily differentiated from C. raropunctata raropunctata because of its protoconch of one more whorl, with signiicantly greater diameter and markedly smaller diameter of the initial half-whorl (less than half the size), its smaller size (see the respective 95% conidence intervals calculated for shell height and maximum diameter), and its different color without any pattern. Stratigraphic occurrence. Cochlis craccoi sp. n. was recovered only from Tortonian deposits at the type locality. Cochlis degrangei (Cossmann & Peyrot, 1919) comb. n. Pl. 1, igs. 3-5 1919 Natica degrangei Cossmann & Peyrot, p. 207, pl. 12, igs. 8-10. Type material: Natica degrangei Cossmann & Peyrot, holotype (the shell igured by Cossmann & Peyrot 1919, pl. 12, igs. 8-10), Saucats, Le Péloua (France), not seen. It is included in the Degrange Touzin collection, most probably in the Université de Bordeaux 1 in Talence, or in the Museum of Bordeaux (Didier Merle, personal communication 2014). We obtained excellent photographs of two paratypes housed in MNHN, i.e. F-J05677, Saucats, Le Péloua (France) and F-J05678, Saucats, Pont-Pourquey (France); the latter is igured herein (Pl. 1, ig. 3). Robba E., Pedriali L. & Quaggiotto E. 122 Material examined: Albugnano: 7 spms., PG 84, 2 spms., NP 9926; Borelli: 8 spms., MGPT-PU 135061, 2 spms., MPUM 11210, 3 spms., NP 9929, 50 spms., MGPT-PU 135062, 1 spm., MPUM 11211, 1 spm., NP 9927, 1 spm., MPUM 11212 (Pl. 1, ig. 5); Léognan, Le Coquillat (France): 11 spms., NP 9928, 1 spm., MPUM 11213, 1 spm., PPMM 40818, 2 spms., private collection; Valle Ceppi: 1 spm., MGPT-PU 135268, 4 spms., MGPT-PU 135063, 1 spm., MGPT-PU 135064, 2 spms., PG 85, 1 spm., MGPT-PU 135065, 11 spms., NP 9930, 5 spms., MPUM 11214, 1 spm., MGPT-PU 135066, 1 spm., MGPT-PU 135067, 47 spms., MGPT-PU 135068; Valle Vergnana: 1 spm., MZB 43816 (Pl. 1, ig. 4). Description. Protoconch medium-sized, low-turbiniform, of 3-3.3 gently convex whorls, tip very small; one specimen exhibits remnants of spiral cordlets on last whorl. Teleoconch hardly exceeding 1 cm in height, oval, taller than wide, moderately thick. Spire conical, rather elevated, whorls convex, suture very slightly channeled. Last whorl broadly oval, subsutural shelf indistinct, periphery nearly at midline. Aperture ovately D-shaped, in slightly prosocline plane, length about 1.6 times width; outer lip semicircular, inner lip gently arched. Parietal callus moderately thick, subrectangular, with slightly concave abapertural outline; anterior lobe indistinct. Umbilicus medium-sized, elongated, wider in a few specimens, umbilical wall moderately steep. Funicle broad, rather depressed, extended abapically and largely illing umbilicus, separated from umbilical wall by narrow, comma-shaped spiral furrow. Umbilical callus rather thick, narrow, elongate, with gently arched adaxial outline, separated from parietal callus by narrow, shallow notch. Basal fasciole poorly differentiated, deined by bending of growth markings over slight angulation. Surface with thin growth lines changing into oblique wrinkles subsuturally. No opercula found. low), but respective larval shells are differently sculptured (spiral threads on the last whorl in C. degrangei, spiral rows of granules on protoconch I in C. pseudovittata). Further, C. pseudovittata has a globular teleoconch, a well developed anterior lobe of the parietal callus (absent in C. degrangei), and a smaller umbilicus. C. degrangei is supericially similar to Natica angyglossa Cossmann & Peyrot, 1919, but N. angyglossa has a more globose teleoconch, a wider umbilicus, a weaker funicle, an umbilical callus merging into the parietal callus without any notch in between, and an operculum with one-two weak marginal grooves, similar to that of Tectonatica prietoi (Hidalgo, 1873). On the basis of the opercular characters, N. angyglossa would rather be assigned to the genus Tectonatica Sacco, 1890 than to Cochlis. Cochlis degrangei is readily differentiated from the other Miocene, French and Italian Cochlis species by its taller than wide, oval shell, its very slender, semielliptical umbilical callus separated from the parietal callus by a narrow notch, and its markedly narrow, comma-shaped umbilical channel. Unfortunately, no comparison can be made as regards the opercula, which are unknown. Stratigraphic occurrence. Cochlis degrangei (Cossmann & Peyrot, 1919) is known deinitely from early Burdigalian units of France; its record from an unspeciied location at Saint-Paul-lès-Dax (Cossmann & Peyrot 1919) likely refers to Burdigalian deposits. Italian occurrences are from Burdigalian to Tortonian formations of Piedmont, where the species appears to be less rare than in France. Cochlis epigloafuniculata (Sacco, 1890) Pl. 1, igs. 6, 7 Dimensions (mm) DHW PD H D SH AH AW 0.0690.073 0.9281.100 3.58911.273 3.0199.903 0.8023.938 2.5837.539 1.8516.175 4.013 0.071 1.014 7.431 6.461 2.370 5.061 UW WUC WAD WAB IS SA 0.9124.384 0.5492.837 0.0001.252 0.0000.992 14°26° 87°123° 2.648 1.693 0.544 0.412 20° 105° Remarks. The values of the characteristic elements of the protoconch pertaining to Cochlis degrangei do not differ signiicantly from those measured for Cochlis pseudovittata sp. n. (see be- 1890b Natica (Natica) millepunctata var. epigloafuniculata Sacco, p. 28. 1891 Natica (Natica) millepunctata var. epigloafuniculata - Sacco, p. 48, pl. 2, ig. 9. 2008a Cochlis epigloafuniculata - Pedriali e Robba, p. 99, pl. 1, igs. 1-6; pl. 2, igs. 14, 21, 22; pl. 3, ig. 12 (cum syn.). Type material: Pedriali & Robba (2008a, p. 99) reviewed the type material of Cochlis epigloafuniculata, and described the characters of this species. Material examined: Rio di Bocca d’Asino: 1 spm., MPUM 11215 (Pl. 1, ig. 7), 1 spm., MZB 60069, 1 spm., MZB 60102 (Pl. 1, ig. 6), 4 spms., MZB 60141; Sant’Agata Fossili: 1 spm., MGPT-PU 135069. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Dimensions of Miocene specimens (mm) DHW PD H D SH AH AW 0.1900.238 0.7770.877 9.65517.915 9.40317.987 2.0244.536 7.13113.879 5.11010.810 0.214 0.827 13.785 13.695 3.280 10.505 7.960 UW WUC WAD WAB IS SA 4.0676.123 0.9032.815 0.8552.815 0.6752.175 15°35° 115°131° 5.095 1.835 1.835 1.425 25° 123° Remarks. No signiicant difference was noted between the upper Miocene specimens and the Pliocene ones dealt with by Pedriali & Robba (2008a). For description and comments, see Pedriali & Robba (2008a). Stratigraphic occurrence. Cochlis epigloafuniculata (Sacco, 1890) occurs uncommonly in the Tortonian of northern Italy. Pliocene records were from Zanclean and/or early Piacenzian deposits of Piedmont, Liguria and Tuscany (Pedriali & Robba 2008a). Cochlis epiglopardalis (Sacco, 1890) comb. n. Pl. 1, igs. 8-12 1864 Natica pardalis Doderlein, p. 18 (nomen nudum). 1864 Natica zonata Doderlein, p. 18 (nomen nudum). 1876 Natica millepunctata var. pardalis Coppi, p. 199. 1890b Natica (Natica) epiglopardalis Sacco, p. 29. 1890b Natica (Natica) epiglottina var. zonata Sacco, p. 29 (new synonym). 1891 Natica (Natica) epiglopardalis - Sacco, p. 56, pl. 2, ig. 23. 1891 Natica (Natica) epiglottina var. zonata - Sacco, p. 62, pl. 2, ig. 33. Type material: Natica (Natica) epiglopardalis Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 2, ig. 23) and reigured herein (Pl. 1, ig. 8), IPUM 4477 (Doderlein collection), Montegibbio; 4 paralectotypes, IPUM 4409 (Doderlein collection), Montegibbio. There are no specimens in MGPT BS. Natica epiglottina var. zonata Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 2, ig. 33) and reigured herein (Pl. 1, ig. 9), IPUM 4471 (Doderlein collection), Montegibbio (see remarks below). Material erroneously referred to as Natica redempta Michelotti, 1847 in IPUM: Montegibbio: 2 spms., IPUM 4428. Material erroneously referred to as Natica pardalis Doderlein, 1864 in IPUM: Montegibbio: 27 spms., IPUM 4420, 46 spms., IPUM 4470. Material erroneously referred to as Natica epiglopardalis Sacco, 1890 in IPUM: Tortona: 1 spm., IPUM 8297. Other material examined: Albugnano: 1 spm., PG 88 (Pl. 1, ig. 10), 1 spm., PG 90, 1 spm., PG 91, 1 spm., PG 92, 1 spm., PG 93, 1 spm. (operculated shell), MPUM 11216; Monte dei Cappuccini: 1 spm., MGPTPU 135070, 2 spms., MGPT-PU 25669; Montegibbio: 6 spms., MPUM 11217, 4 spms., MGC 1416, 12 spms., NP 9933, 1 spm., MPUM 11218 (Pl. 1, ig. 11), 13 spms., NP 9934, 1 spm., NP 9935, 1 spm., MPUM 11219 (Pl. 1, ig. 12), 115 spms., private collection, 13 spms., private collection, 38 spms., private collection, 3 spms., private collection; Rio di Bocca d’Asino: 1 spm., PG 94, 1 spm., MPUM 11220, 1 spm., NP 9940; Stazzano: 2 spms., NP 9941. 123 Description. Protoconch medium-sized, turbiniform, of 2.10-2.35 convex, apparently smooth whorls, tip small. Teleoconch globose, rather thick. Spire broadly conical, moderately elevated, earlier whorls gently convex, subsequent ones convex; suture adpressed. Last whorl globose, slightly expanded toward aperture; subsutural shelf lat to gently concave, poorly demarcated. Aperture D-shaped, in slightly prosocline plane, length about 1.9 times width; outer lip semicircular, inner lip thickened and relected toward shell axis abapically. Parietal callus thick, subquadrangular, rather long, moderately expanded adapically; anterior lobe indistinct. Umbilicus medium-sized to small, slitlike in a few specimens; umbilical wall steep. Funicle cord-like and prominent to broad and depressed, separated from umbilical wall by narrow, shallow spiral furrow. Umbilical callus moderate to broad (depending on size of funicle), rather thick, semicircular in outline, ending at abapical one-fourth of inner lip, separated from anterior corner of parietal callus by shallow, reverse J-shaped notch. Basal fasciole broad and markedly depressed. Surface with rather dense growth lines, prosocyrt subsuturally. Many specimens retain remnants of uniform pale brown background with reddish-brown pattern of large, oval to elongately teardrop-shaped spots irregularly arranged into collabral rows. A few specimens (var. zonata Sacco) bear three brownish spiral bands. Operculum moderately thick; central callus short, not exceeding one-third of total height; inner margin slightly arched, with distinct, blunt transverse ridges; inner surface almost lat, nucleus not protruding; outer surface lat to slightly concave, with two marginal grooves and two ridges; outer groove broad, shallow, with nearly lat bottom bearing longitudinal rows of granules; inner groove narrow, attenuated; outer ridge moderately prominent, narrowly round-topped; inner ridge a sharp boundary of outer groove, prostrate to overhang inner groove. Dimensions (mm) DHW PD H D SH AH AW 0.0980.114 0.9170.997 11.84323.827 11.79123.699 2.4347.738 8.91916.579 7.49214.148 0.106 0,957 17.835 17.745 5.086 12.749 10.820 UW WUC WAD WAB IS SA 2.2529.920 0.6595.175 0.5842.756 0.0003.033 16°24° 104°140° 6.086 2.917 1.670 1.499 20° 122° 124 Robba E., Pedriali L. & Quaggiotto E. Remarks. Natica pardalis was introduced by Doderlein (1864) without any description or illustration and is therefore considered a nomen nudum. From Doderlein’s paper, it appears that the name N. pardalis was intended to replace Natica pseudomaculosa, a former name used by Doderlein in a manuscript catalogue, but no explanation for this action was provided. Coppi (1876) irst published a concise diagnosis of N. pardalis (regarded as a variety of Natica millepunctata Lamarck, 1822), thus making the name available, and is to be considered the author of this taxon. However, N. pardalis Coppi is a junior primary homonym of Natica pardalis Philippi, 1852, thence permanently invalid (ICZN 1999, Article 57.2 of the Code). Sacco (1890b), evidently aware of this case of homonymy, introduced the name Natica epiglopardalis, and shortly later (1891) explicitly afirmed that N. epiglopardalis was the replacement name for N. pardalis Doderlein (really of Coppi). Natica zonata is another nomen nudum introduced by Doderlein (1864). The name zonata was made available by Sacco (1890b), who irst diagnosed N. zonata and has the authorship of it. Sacco (1890b, 1891) considered N. zonata as a variety of Natica epiglottina Lamarck, 1804. We examined the shell (IPUM 4471) igured by Sacco (1891) and can state that, except for the color pattern, this specimen conforms in all characters to Cochlis epiglopardalis, with which it co-occurs rarely at Montegibbio. Consequently, we consider Natica (Natica) epiglottina var. zonata Sacco merely as a color-form of C. epiglopardalis and include it in the synonymy of the latter species. Only one specimen labelled Natica epiglottina var. zonata (manuscript writing of Sacco) is present in IPUM. However, since Doderlein (1864) listed two specimens (one probably lost), this shell is not eligible as the holotype of N. epiglottina var. zonata, instead, it can be designated as the lectotype. Cochlis epiglopardalis exhibits a remarkable variability as regards the strength of the funicle (less so of the umbilical callus). We noted shells (rather rare) with strong funicle separated from the umbilical wall by a very narrow, semicircular groove, and shells with slender funicle within a medium-sized, widely open umbilicus. A gradual transition between these two extremes is observable. Cochlis epiglopardalis is closely similar to Cochlis raropunctata raropunctata (Sasso, 1827) and probably has been mistaken for it (there seem to be no citations of C. epiglopardalis subsequent to its introduc- tion). However, Sacco’s species is readily distinguished from C. raropunctata raropunctata in that it has a protoconch of 1 more whorl, with signiicantly greater diameter and smaller diameter of the initial half-whorl (one-third of the size). The operculum of C. epiglopardalis is similar to that of C. raropunctata raropunctata (both have two marginal grooves and two ridges), but the operculum of C. epiglopardalis has a shorter central callus, a lat-bottomed outer groove (concave in C. raropunctata raropunctata), and the inner ridge leaning to overhang the inner groove (always raised in C. raropunctata raropunctata). Moreover, the specimens with a color pattern of teardrop-like spots are quite distinctive. Stratigraphic occurrence. Cochlis epiglopardalis (Sacco, 1890) was hitherto recorded from Langhian to Torto- PLATE 1 Fig. 1 - Cochlis craccoi sp. n. Borelli. Holotype, MGPT-PU 135052; apertural side (height of shell 6.47 mm). Fig. 2 - Cochlis craccoi sp. n. Borelli. Paratype, MGPT-PU 135060; protoconch. Fig. 3 - Cochlis degrangei (Cossmann & Peyrot, 1919). Saucats, PontPourquey (France). Paratype, MNHN-F-J05678; apertural side. Fig. 4 - Cochlis degrangei (Cossmann & Peyrot, 1919). Valle Vergnana. MZB 43816; a, apertural side (height of shell 12.16 mm); b, protoconch. Fig. 5 - Cochlis degrangei (Cossmann & Peyrot, 1919). Borelli. MPUM 11212; a, protoconch; b, detail of protoconch. Fig. 6 - Cochlis epigloafuniculata (Sacco, 1890). Rio di Bocca d’Asino. MZB 60102; apertural side (height of shell 16.87 mm). Fig. 7 - Cochlis epigloafuniculata (Sacco, 1890). Rio di Bocca d’Asino. MPUM 11215; protoconch. Fig. 8 - Cochlis epiglopardalis (Sacco, 1890). Montegibbio. Lectotype (here designated) of Natica (Natica) epiglopardalis Sacco, 1890. IPUM 4477 (Doderlein collection); apertural side (height of shell 26.34 mm). Fig. 9 - Cochlis epiglopardalis (Sacco, 1890). Montegibbio. Lectotype (here designated) of Natica epiglottina var. zonata Sacco, 1890. IPUM 4471 (Doderlein collection); apertural side (height of shell 29.22 mm). Fig. 10 - Cochlis epiglopardalis (Sacco, 1890). Albugnano. PG 88; protoconch. Fig. 11 - Cochlis epiglopardalis (Sacco, 1890). Montegibbio. MPUM 11218; apertural side showing color pattern (height of shell 15.67 mm). Fig. 12 - Cochlis epiglopardalis (Sacco, 1890). Montegibbio. MPUM 11219; a, apertural side of shell with the operculum in situ (height of shell 12.22 mm); b, operculum (height of operculum 7.95 mm). Fig. 13 - Cochlis epiglottina epiglottina (Lamarck, 1804). Grignon (France). Lectotype (here designated) of Natica epiglottina Lamarck, 1804. MNHG GEPI 82538; a, apertural side; b, protoconch. Fig. 14 - Cochlis epiglottina epiglottina (Lamarck, 1804). Cava Boschetto. MPUM 11221; apertural side (height of shell 14.77 mm). Fig. 15 - Cochlis epiglottina epiglottina (Lamarck, 1804). Cava Grola. MPUM 11222; protoconch. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 125 Robba E., Pedriali L. & Quaggiotto E. 126 nian deposits of Piedmont and from the late Miocene of Emilia. Cochlis epiglottina epiglottina (Lamarck, 1804) comb. n. excellently preserved French specimens exhibit microscopic spiral striation on last whorl. One specimen from Cava Grola with well spaced, brown, undulating axial lines. Dimensions (mm) DHW PD H D SH AH AW Pl. 1, igs. 13-15; Pl. 2, igs 1, 2 0.2570.289 0.8891.057 2.68415.996 2.88714.611 0.3674.171 1.85012.090 1.6017.901 1804 Natica epiglottina Lamarck, p. 95. 1806 Natica epiglottina - Lamarck, pl. 62 (14), ig. 6. 1832 Natica epiglottina - Deshayes, p. 165, pl. 20, igs. 5, 6, 11. 1864 Natica epiglottina - Deshayes, p. 56. ? 1864 Natica munda Deshayes, p. 57, pl. 72, igs. 12, 13. 1888 Natica (Natica) epiglottina - Cossmann, p. 159. 1907 Natica epiglottina - Cossmann & Pissarro, pl. 9, ig. 61-1. not 1949 Natica epiglottina - Wrigley, p. 11, igs. 1, 2 (not Lamarck, 1804; see remarks below). 1985 Natica epiglottina - Brigantini, p. 414, pl. 2, ig. 44. 2012 Natica epiglottina - Courville et al., pl. 5, igs. 10, 11, 14. 0.273 0.973 9.340 8.749 2.269 6.970 4.751 Type material: Natica epiglottina Lamarck, lectotype (here designated): the syntype igured by Favre (1918, pl. 5, ig. 65), and reigured herein (Pl. 1, ig. 13), MNHG GEPI 82538, Grignon (France); 6 paralectotypes, MNHG GEPI 46217, Grignon (France). According to Favre (1918), Lamarck’s manuscript catalogue cites 10 specimens, but only 7 of them are present in MNHG. Other material examined: Cava Albanello: 2 spms., MGP-PD 1213R, 1 spm., MGP-PD 1214R, 1 spm., MGP-PD 1215R, 2 spms., MGPPD 1216R, 1 spm., private collection, 3 spms., MCZ 4316; Cava Boschetto: 2 spms., NP 9980, 1 spm., MPUM 11221 (Pl. 1, ig. 14); Cava Grola: 1 spm., private collection, 2 spms., NP 9982, 2 spms., NP 9983, 2 spms., NP 9984, 10 spms., NP 9985, 11 spms., private collection, 3 spms., NP 9986, 1 spm., MPUM 11222 (Pl. 1, ig. 15); Fontenay (France): 3 spms., NP 9987; Monte Merlo: 1 spm., NP 9981, 1 spm., NP 9988; Villiers-Saint-Frédéric (France): 1 spm., MPUM 11223 (Pl. 2, ig. 1), 1 spm., NP 9989, 1 spm., MPUM 11224 (Pl. 2, ig. 2), 6 spms., MPUM 11225, 45 spms., NP 9990. Description. Protoconch small, depressedturbiniform, averaging 1.64 slightly convex whorls, tip medium-sized, rather lattened. Teleoconch globose to globose-oval, thin to rather thick. Spire broadly conical, rather pointed, moderately elevated, less so in a few specimens; whorls very gently convex; suture slightly incised, almost lush. Last whorl broadly oval, somewhat produced toward aperture, subsutural shelf indistinct, periphery above midline. Aperture ovately D-shaped, in slightly prosocline plane, length almost twice width; outer lip nearly semicircular, thin-edged, inner lip very gently arched. Parietal callus subquadrangular, moderately thick, more so adapically; anterior lobe indistinct. Umbilicus deep, medium-sized, small in a few specimens; umbilical wall rather steep. Funicle cord-like, thick and prominent, separated from umbilical wall by shallow spiral furrow of variable breadth. Umbilical callus large, thick, semicircular in outline, located at midabapical part of inner lip, separated from parietal callus by moderately deep, mostly narrow notch. Basal fasciole indistinct. Surface with rather dense, uneven growth lines; UW WUC WAD WAB IS SA 0.5387.430 0.2813.705 0.0941.518 0.0412.411 7°-27° 100°128° 3.984 1.993 0.806 1.185 17° 114° Remarks. Cochlis epiglottina epiglottina is characterized by its thick funicle, which largely ills the umbilicus (almost completely in some specimens); PLATE 2 Fig. 1 - Cochlis epiglottina epiglottina (Lamarck, 1804). Villiers-SaintFrédéric (France). MPUM 11223; apertural side (height of shell 11.32 mm). Fig. 2 - Cochlis epiglottina epiglottina (Lamarck, 1804). Villiers-SaintFrédéric (France). MPUM 11224; protoconch. Fig. 3 - Cochlis epiglottina auriformis (von Koenen, 1891). Case Soghe. MPUM 11262; apertural side (height of shell 12.97 mm). Fig. 4 - Cochlis epiglottina auriformis (von Koenen, 1891). Westeregeln (Germany). Topotype of Natica epiglottina var. auriformis von Koenen, 1891. MB.Ga.12743.1; apertural side. Fig. 5 - Cochlis infelix (Sacco, 1890). Carpenaro. Neotype (here designated) of Natica (Natica) infelix Sacco, 1890. MPUM 11226; apertural side (height of shell 16.18 mm). Fig. 6 - Cochlis infelix (Sacco, 1890). Sassello. MPUM 11227; a, apertural side (height of shell 12.21 mm); b, abapertural side. Fig. 7 - Cochlis mortoni sp. n. Cava Rossi. Holotype, MPUM 11228; a, apertural side (height of shell 21.23 mm); b, apical view. Fig. 8 - Cochlis mortoni sp. n. Cava Albanello. Paratype, MGP-PD 1217R; protoconch. Fig. 9 - Cochlis neglecta (Mayer, 1858). Albugnano. MPUM 11231; apertural side (height of shell 13.19 mm). Fig. 10 - Cochlis neglecta (Mayer, 1858). Le Houga, Carrière (France). Lesport collection 15L0003; operculum (height of operculum 9.30 mm). Fig. 11 - Cochlis neglecta (Mayer, 1858). Lucbardez-et-Bargues, Petit Bargues (France). Lesport collection 15L0002; apertural side (height of shell 17.80 mm). Fig. 12 - Cochlis neglecta (Mayer, 1858). Rio di Bocca d’Asino. MPUM 11233; protoconch. Fig. 13 - Cochlis neglecta (Mayer, 1858). Rio di Bocca d’Asino. PG 87; a, apertural side of shell with the operculum in situ (height of shell 21.12 mm); b, operculum (height of operculum 11.20 mm). Fig. 14 - Cochlis neglecta (Mayer, 1858). Uzeste, Gamachot (France). NP 10010; protoconch. Fig. 15 - Cochlis neglecta (Mayer, 1858). Val Sanfrà. MGPT-PU 135077; apertural side (height of shell 15.33 mm). Fig. 16 - Cochlis pseudovittata sp. n. Borelli. Holotype, MGPT-PU 135079; apertural side (height of shell 5.48 mm). Fig. 17 - Cochlis pseudovittata sp. n. Borelli. Paratype, MGPT-PU 135084; a, protoconch; b, detail of protoconch. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 127 128 Robba E., Pedriali L. & Quaggiotto E. the umbilical channel appears as a rather narrow (very narrow in a few specimens), semicircular groove. Cossmann (1888) regarded Natica munda Deshayes, 1864 as a synonym of the present species. On the basis of the description and the igures of N. munda published by Deshayes, Cossmann’s opinion seems to be correct; however, not having seen the original material of N. munda, we include it in the synonymy of Cochlis epiglottina epiglottina with doubt. The Middle Eocene species Natica epiglottinoides Deshayes, 1864 is rather similar to Cochlis epiglottina epiglottina in teleoconch shape. We examined several shells of Deshayes’ species from the Bartonian of Saint-Witz (Le Guépelle) and can state that N. epiglottinoides (a species of Cochlis) is easily differentiated from C. epiglottina epiglottina in that it has: 1) a protoconch with signiicantly smaller diameter (averaging 0.729 mm) and diameter of the irst halfwhorl (averaging 0.140 mm), 2) a globose last whorl (that of C. epiglottina epiglottina is broadly oval), more depressed in abapertural view, 3) a parietal callus with markedly concave abapertural outline and with a broad, semicircular anterior lobe, and 4) an abapical umbilical sulcus bounded by a distinct angulation (absent in C. epiglottina epiglottina). The characters of the parietal callus were already pointed out by Cossmann (1902, p. 63) as a distinguishing element of N. epiglottinoides. Natica specialis Deshayes, 1864 (also belonging in Cochlis), described from Lutetian deposits of Grignon, is another similar species in shell morphology. Examination of specimens from Fleury-la Rivière and Damery has shown that it differs from C. epiglottina epiglottina because of its protoconch of about 1 more whorl, with signiicantly smaller diameter of the irst half-whorl, its slight subsutural margining developed on later whorls (absent in C. epiglottina epiglottina), and its elongate umbilical callus merging into the anterior corner of the parietal callus and overhanging the most adapical part of the umbilicus. Wrigley (1949) described and illustrated a shell from the Lower Barton beds (bed A3), referred to as Natica epiglottina Lamarck, 1804. We examined some specimens from the same bed A3 in our collection, perfectly conforming to Wrigley’s shell, and could note that they differ from N. epiglottina because of their two-whorled protoconch with signiicantly smaller diameter of the initial half- whorl (half the size), their lower spire with lattish whorls, and their umbilicus bounded by a distinct angulation (the umbilicus of N. epiglottina has a well rounded border). Wrigley afirmed to have recovered some shells with the operculum “in situ in the aperture”; he described the operculum as having the outer surface with “two slightly incised lines equidistant from the outer margin, although in many specimens they are hardly apparent”, and published a poor igure of it (ig. 2). According to this description of the operculum, the British specimens seem to belong in the genus Tectonatica Sacco, 1890 or Cryptonatica Dall, 1892. Stratigraphic occurrence. Cochlis epiglottina epiglottina (Lamarck, 1804) is deinitely known from Lutetian deposits of France and northeastern Italy (Veneto). Cochlis epiglottina auriformis (von Koenen, 1891) comb. n. Pl. 2, igs. 3, 4 1891 Natica epiglottina var. auriformis von Koenen, p. 576, pl. 40, ig. 7. Material examined: Case Soghe: 1 spm., MPUM 11262 (Pl. 2, ig. 3), 2 spms., private collection, 1 spm., NP 9995. Description. Protoconch damaged, apparently small and with medium-sized tip. Teleoconch globose-oval, thick, hardly exceeding 13 mm in height. Spire broadly conical, rather low, whorls gently convex, suture incised, adpressed. Last whorl broadly oval, subsutural shelf indistinct, periphery at midline. Aperture D-shaped, in prosocline plane, length about twice width; outer lip semicircular, inner lip straight, thickened abapically. Parietal callus short, rectangular, thick, more so adapically where a blunt knob may develop; anterior lobe absent. Umbilicus deep, moderately wide, umbilical wall steep. Funicle cord-like, broad and prominent, largely illing umbilicus; umbilical channel rather narrow to very narrow, comma-like. Umbilical callus thick, with arched abapertural outline, located at mid-abapical part of inner lip, separated from anterior end of parietal callus by rather deep, narrow, rounded notch. Basal fasciole indistinct. Surface with remnants of uneven growth lines. German specimens from Westeregeln retain a uniform, pale brown background without any pattern. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy auriformis occurs in lower Oligocene deposits of Northern Germany and Veneto. Dimensions (mm) SH 129 DHW PD H D AH - - 12.970 12.351 3.652 9.319 AW UW WUC WAD WAB IS SA 6.930 6.929 2.840 1.400 2.061 17° 116° Remarks. The examined specimens appear to be strikingly similar to Natica epiglottina var. auriformis, a variety introduced by von Koenen (1891) on the basis of lower Oligocene material from Westeregeln (Northern Germany). The Museum für Naturkunde in Berlin houses ive specimens of N. epiglottina var. auriformis from the same locality cited by von Koenen; they are not identiied as the type material and are not recorded as being part of the von Koenen collection (Martin Aberhan, personal communication 2014). However, these shells from Westeregeln are of much relevance since they are to be regarded as topotypes. We obtained some excellent photographs of specimen MB.Ga.12743.1 (Pl. 2, ig. 4) and could note: 1) that the teleoconch of our specimens is identical to that of N. epiglottina var. auriformis and 2) that their damaged protoconch seems to be closely similar to that of N. epiglottina var. auriformis. Accordingly, we conclude that the examined material can be safely identiied with von Koenen’s taxon, which is assigned to the genus Cochlis Röding, 1798. The var. auriformis and Cochlis epiglottina Lamarck, 1804 are closely similar; the values of the characteristic elements of the protoconch of both taxa are identical, as are their teleoconch shape and umbilical characters. However, the var. auriformis has a slightly shorter spire, a much shorter parietal callus, and a uniform, pale brown background (well spaced, brown, undulating axial lines in C. epiglottina). These differences are not considered suficient for consistent speciic separation. Consequently, we regard the var. auriformis as an allochronous subspecies of C. epiglottina, which was introduced nearly one century earlier. Cochlis epiglottina auriformis is morphologically similar to the coeval Cochlis sp. 2, but differs from it in that it has: 1) a shorter and thicker parietal callus with straight abapertural outline (subangular in Cochlis sp. 2), 2) a larger umbilicus, 3) a narrower, comma-shaped umbilical channel (bean-shaped in Cochlis sp. 2), 4) a more prominent, cord-like funicle, and 5) a differently shaped umbilical callus, separated from parietal callus by a rather deep, narrow and rounded notch (very slight in Cochlis sp. 2). Stratigraphic occurrence. Cochlis epiglottina Cochlis infelix (Sacco, 1890) comb. n. Pl. 2, igs. 5, 6 ig. 2. 1890b Natica (Natica) infelix Sacco, p. 27. 1890b Natica (Natica) infelix var. sasselliana Sacco, p. 27. 1891 Natica (Natica) infelix - Sacco, p. 44, pl. 2, ig. 1. 1891 Natica (Natica) infelix var. sasselliana - Sacco, p. 45, pl. 2, 1984 Natica infelix - Ferrero Mortara et al., p. 27. Type material: Natica (Natica) infelix Sacco, neotype (here designated and illustrated in Pl. 2, ig. 5), MPUM 11226, Carpenaro (see remarks below); 1 syntype, MGPT BS.029.01.001, Sassello, 1 syntype, MGPT BS.029.01.001/01, Cassinelle. Other material examined: Cassinelle: 2 spms., PG 101; Mioglia: 1 spm., MZB 60147, 6 spms., MZB 60148, 3 spms., PG 99; Sassello: 1 spm., MPUM 11227 (Pl. 2, ig. 6), 1 spm., PG 100; Squaneto: 1 spm., PG 98. Description. Protoconch poorly preserved, low-turbiniform, apparently of 2.5 whorls, with small tip. Teleoconch rather small, globose-oval, higher than wide, thick. Spire conical, rather low, somewhat stepped, whorls convex, with maximum curvature at adapical one-third; subsutural shelf narrow, gently sloping, obscurely bounded abaxially; suture ine, adpressed. Last whorl rather tall, broadly oval, slightly produced abapically toward aperture; subsutural shelf indistinct. Aperture D-shaped, in very slightly prosocline plane, length 1.8 times width; outer lip regularly arched, basal lip thickened. Parietal callus thick, subrectangular, slightly wider adapically; anterior lobe indistinct. Umbilicus deep, medium-sized, umbilical wall very steep. Funicle broad, depressed, separated from umbilical wall by shallow, rather narrow spiral depression. Umbilical callus thick, with arched abapertural outline, located at mid-abapical part of inner lip, separated from parietal callus by shallow, reverse J-shaped notch. Basal fasciole indistinct. Surface with rather dense, even growth lines. Dimensions (mm) DHW PD H D SH AH AW - - 9.37717.713 8.11117.507 2.4975.745 4.97613.304 4.4099.797 - - 13.545 12.809 4.121 9.140 7.103 UW WUC WAD WAB IS SA 3.2107.379 1.2283.304 0.4242.168 0.7982.550 10°30° 96°132° 5.296 2.266 1.296 1.674 20° 114° 130 Robba E., Pedriali L. & Quaggiotto E. Remarks. The igured syntypes of Natica (Natica) infelix and of Natica (Natica) infelix var. sasselliana were said to be respectively in the Museo Geologico of Roma and in the Museo Geologico of Genova (see Sacco 1891, explanation of pl. 2). Inquiries there had negative results and the cited syntypes are deemed to be lost. The syntypes presently in MGPT (see above) are quite poorly preserved and can hardly help in clarifying the identity of the species. Another four specimens (syntypes?) in MGPT, labelled N. (Natica) cf. infelix (BS.029.001/02, unspeciied locality), are of no use because of their even worse preservation. In these circumstances, the designation of a neotype is advisable in order to deine Sacco’s taxon objectively (ICZN 1999, Article 75 and Recommendation 75A of the Code). The specimen igured by Sacco (1891) was from Dego; other localities reported by Sacco are Sassello and Cassinelle. We designate as the neotype a specimen from Carpenaro (9.5 km northeast of Dego), which agree with the description provided by Sacco and exhibits all the teleoconch characters unequivocally. The lithostratigraphic unit that yielded this specimen is the same cropping out at the localities cited by Sacco. Obviously, Carpenaro becomes the type locality of N. (Natica) infelix (ICZN 1999, Article 76.3 of the Code). The above description is largely based on the neotype. The few examined specimens show a slight variability concerning the height of the spire (lower than that of the neotype) and the inlation of the last whorl (more globose than that of the neotype). On the basis of its teleoconch characters, Sacco’s species is herein assigned to the genus Cochlis Röding, 1798. The Miocene species Cochlis degrangei (Cossmann & Peyrot, 1919) superficially resembles Cochlis infelix in that it has a taller than wide teleoconch and a broad, depressed funicle. However, C. degrangei differs from C. infelix by its slightly channeled suture, its wider aperture, and its funicle largely filling the umbilicus (Fig. 17). The globose oval, higher than wide teleoconch, the rather low, somewhat stepped spire, and the broad, depressed funicle not filling the umbilicus readily distinguish C. infelix from the other Oligocene Cochlis species. Stratigraphic occurrence. Cochlis infelix (Sacco, 1890) was hitherto recorded from low- er Oligocene deposits of Piedmont and Liguria where it occurs uncommonly. Cochlis mortoni sp. n. Pl. 2, igs. 7, 8 Derivation of name: The species is named after Alan Morton (Penrhyncoch, Aberystwyth), who kindly donated many British Eocene specimens relevant to the present study. Holotype: Cava Rossi, MPUM 11228 (Pl. 2, ig. 7). Paratypes: Cava Albanello: 1 spm., MGP-PD 1217R (Pl. 2, ig. 8); Cava Rossi: 2 spms., MCZ 4317, 1 spm., MPUM 11229, 1 spm., MCZ 4318, 1 spm., MPUM 11230. Other material esamined: Cava Grola: 1 spm., private collection; Cava Rossi: 1 spm., NP 9970, 2 spms., private collection, 1 spm., NP 9971, 1 spm., private collection. Preservation: Moderate to rather fair. Type locality: Cava Rossi (see appendix). Horizon: Volcanoclastic layer of late Ypresian to earliest Lutetian age. Diagnosis: Shell globose; spire rather low, with well convex whorls and narrowly and deeply channeled suture; last whorl globular. Umbilicus deep, medium-sized. Parietal callus broadly rectangular, with rounded anterior edge. Funicle cord-like, thick, largely illing umbilicus in young specimens; umbilical callus large, semicircular, separated from parietal callus by rather narrow and deep notch. Description. Protoconch small, depressedturbiniform, of 2.50-2.85 whorls, tip very small. Teleoconch medium-sized, globose, thick. Spire broadly conical, rather low, whorls convex; suture adpressed on irst whorl, gradually changing into narrowly and deeply channeled on subsequent ones. Last whorl globose, subsutural shelf indistinct, periphery slightly above midline. Aperture D-shaped, in prosocline plane, length averaging 1.8 times width; outer lip semicircular, inner lip straight, thickened abapically. Parietal callus thick, broadly rectangular, with rounded anterior edge covering most adapical part of umbilicus. Umbilicus deep, medium-sized, rather small in a few specimens, umbilical wall steep. Funicle cord-like, thick and prominent, largely illing umbilicus in young specimens, separated from umbilical wall by shallow spiral furrow of variable breadth (rather wide to very narrow), bounded abaxially by obscure step; furrow with 1-2 rough, longitudinal cordlets visible in larger specimens. Umbilical callus large to very large (depending on size of funicle), thick, semicircular in outline, located at mid-abapical part of inner lip and separated from parietal callus by rather narrow, moderately deep notch. Basal fasciole indistinct. Surface with remnants of ine, dense growth lines. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Dimensions (mm) DHW PD H D SH AH AW 0.0780.090 0.6010.689 2.21719.069 2.52218.926 0.2066.182 1.54913.349 1.25611.832 0.084 0.645 10.643 10.724 3.194 7.449 6.544 UW WUC WAD WAB IS SA 0.9416.685 0.6133.477 0.3241.368 0.0002.001 9°-29° 113°129° 3.813 2.045 0.846 0.921 19° 121° Remarks. The umbilical characters of the present new species suggest its assignment to the genus Cochlis Röding, 1798. The narrow, deeply channeled suture and the broad, rectangular parietal callus with rounded anterior edge constitute the main distinguishing elements of Cochlis mortoni and readily separate it from the other Eocene species of the same genus. Stratigraphic occurrence. Cochlis mortoni sp. n. was recovered from upper Ypresian and lower-mid Lutetian units of Veneto. Cochlis neglecta (Mayer, 1858) comb. n. Pl. 2, igs. 9-15 1858 Natica neglecta Mayer, p. 388, pl. 11, ig. 2. 1882 Natica benecki von Koenen, p. 228, pl. 5, igs. 4, 5, 8. 1919 Natica neglecta - Cossmann & Peyrot, p. 199, pl. 11, igs. 11-14. 1952a Natica (Natica) neglecta - Glibert, p. 76, pl. 6, ig. 4. 1952b Natica (Natica) neglecta - Glibert, p. 259, pl. 2, ig. 4. 1969 Natica neglecta - Janssen, p. 173, pl. 5, igs. 18-24; textig. 12 (cum syn.). 1972 Naticarius tigrinus tigrinus - Nordsieck, p. 70, pl. 17, ig. 86 (not the operculum). 1972 Naticarius hoernesi sallomacensis - Nordsieck, p. 71, pl. 17, ig. 88 (not Tournouer, 1873). 1972 Naticarius neglectus - Nordsieck, p. 71, pl. 17, ig. 90. 1984 Natica neglecta - Janssen, p. 202, pl. 57, igs. 5-7, 12. 2001 Natica neglecta - Lozouet et al., p. 43, pl. 18, ig. 2. 2001 Natica neglecta - Wienrich, p. 432, pl. 70, ig. 1; pl. 87, igs. 3, 4; pl. 88, ig. 7. Type material: Natica neglecta Mayer, type material not seen. The Mayer-Eymar collection, housed in Naturhistorisches Museum of Basel (Switzerland), is in a disarray and no types were located so far (Walter Etter, personal communication 2014). Material erroneously referred to as Natica pardalis Doderlein, 1864 in IPUM: Montegibbio: 4 spms., IPUM 4470. Other material examined: Albugnano: 1 spm., MPUM 11231 (Pl. 2, ig. 9), 1 spm., PG 86; Borelli: 7 spms., MGPT-PU 135071; Le Houga, Carrière (France): 1 spm. (operculum), 15L0003 (Pl. 2, ig. 10); Léognan, Le Coquillat (France): 6 spms., NP 9996; Lucbardez-et-Bargues, Petit Bargues (France): 1 spm., 15L0002 (Pl. 2, ig. 11); Mérignac (France): 9 spms., MPUM 11232, 20 spms., NP 9944, 1 spm. (operculum), NP 9997; Monte dei Cappuccini: 1 spm., MGPT-PU 25668, 1 spm., MGPT-PU 25666; Pietracuta: 3 131 spms. NP 9945, 1 spm., PP.PC 03/01; Rio di Bocca d’Asino: 1 spm., MPUM 11233 (Pl. 2, ig. 12), 1 spm. (operculated shell), PG 87 (Pl. 2, ig. 13), 1 spm., PG 6b, 1 spm., PG 12a, 1 spm., PG 13b, 1 spm., MZB 60068, 1 spm., MZB 60101, 1 spm., MZB 60105, 1 spm., MZB 60109; Roquefort (France): 11 spms. MPUM 11234, 2 spms., (operculated shells), MF 115D.1.10, 14 spms., NP 9943; Saint-Martin-d’Oney (France): 2 spms., NP 10006; Sant’Agata Fossili: 1 spm., MGPT-PU 135072, 2 spms., MGPT-PU 135073, 1 spm., PG 18e; Uzeste, Gamachot (France): 6 spms., NP 10008, 1 spm., NP 10010 (Pl. 2, ig. 14); Valle Ceppi: 1 spm., NP 9931, 1 spm., MZB 60138, 1 spm., MGPT 135074, 2 spms., NP 9932, 1 spm., MPUM 11235, 1 spm., MGPT-PU 135075, 1 spm., MGPT-PU 135076; Val Sanfrà: 1 spm., MGPT-PU 135077 (Pl. 2, ig. 15), 1 spm., MGPT-PU 135078. Description. Protoconch small, depressedturbiniform, of 2.5-2.75 gently convex whorls, tip very small. Teleoconch globose, globose-oval in some specimens, thin to rather thick. Spire conical, moderately elevated, less so in a few specimens, whorls convex, suture ine, adpressed. Last whorl globose, not expanded toward aperture; subsutural shelf poorly deined to indistinct. Aperture D-shaped, in very slightly prosocline plane, length averaging 1.7 times width; outer lip semicircular, basal lip thickened. Parietal callus thick, subquadrangular, wider adapically; anterior lobe indistinct. Umbilicus deep, medium-sized, small in a few specimens; umbilical wall rather steep. Funicle cord-like, thin to moderately thick, separated from umbilical wall by shallow spiral furrow of variable breadth. Umbilical callus small to moderate (depending on size of funicle), thick, semicircular in outline, located at abapical one-third of inner lip, separated from parietal callus by moderately deep notch. Basal fasciole poorly differentiated to indistinct. Surface with rather dense growth lines changing into narrow wrinkles subsuturally; microscopic spiral striation present on last whorl. Some specimens retain remnants of uniform pale brown background apparently without any color pattern. Operculum moderately thick; central callus lat, subtriangular, not reaching half-height of operculum; inner margin very slightly convex, smooth; inner surface latly convex, nucleus scarcely protruding; outer surface planar, slightly convex in a few specimens, with three marginal grooves and three ridges; median and outer grooves as well as ridges on gently sloping inward shelf; outer and median grooves narrow and shallow, with concave bottom; inner groove deeper than the other two, slightly wider in a few specimens; ridges subequal, round-topped, median one slightly narrower in a few specimens. Robba E., Pedriali L. & Quaggiotto E. 132 Dimensions (mm) DHW PD H D SH AH AW 0.0920.096 0.9121.012 11.61721.381 10.39919.859 2.6937.373 7.52915.401 5.94612.734 9.340 0.094 0.962 16.499 15.129 5.033 11.465 UW WUC WAD WAB IS SA 2.7377.845 0.9142.410 0.8133.921 0.0002.570 14°34° 91°123° 5.291 1.662 2.367 1.262 24° 107° Remarks. The present species exhibits a moderate variability as regards the height of the spire and the strenght of the funicle. The Italian shells fully conform to the abundant topotypic material in our collection. According to the description and the excellent igures originally published, it appears that the teleoconch and the operculum of Natica benecki von Koenen, 1882 are hardly distinguishable from those of Cochlis neglecta. Consequently, we concur with Glibert (1952a), Janssen (1969) and Wienrich (2001) in considering Koenen’s species a synonym of C. neglecta. Except for a slightly wider aperture, the teleoconch of Cochlis neglecta is strikingly similar to that of Cochlis pseudoepiglottina (Sacco, 1890). However, C. neglecta can be differentiated from Sacco’s species by its protoconch with signiicantly greater diameter; further, the operculum of C. neglecta has three subequal marginal ridges (see also Janssen 1969 and Wienrich 2001) whereas that of C. pseudoepiglottina, also with three marginal ridges, has a sharp, thinner (thread-like in a few specimens) middle ridge (Pedriali & Robba 2005). The threeridged operculum of the Pliocene to Recent similar species Cochlis vittata (Gmelin, 1791) differs from that of C. neglecta in that it has thin, equal outer and inner ridges, and a wider, lat-topped middle ridge (see Pedriali & Robba 2005). Pedriali & Robba (2005, p. 155), on the basis of the teleoconch characters of specimens described and/or illustrated in the literature, considered Cochlis pseudoepiglottina present in the upper Miocene of the Mediterranean area and of Hungary. In the course of the present study, despite the vast upper Miocene material examined, not one specimen proved to belong to C. pseudoepiglottina. Most likely, several upper Miocene shells, with the apical whorls abraded or missing and lacking the operculum, formerly assigned to C. pseudoepiglottina are Cochlis neglecta, which has a closely similar teleo- conch. From the above, it results that the presence of C. pseudoepiglottina in the late Miocene needs to be conirmed by the recovering of specimens retaining the protoconch and/or the operculum. Stratigraphic occurrence. Cochlis neglecta (Mayer, 1858) is known deinitely from the early Miocene of France, The Netherlands, Belgium and Germany, and from the Langhian of Belgium, Germany and Poland. Italian occurrences are from Burdigalian to Tortonian deposits of Piedmont. Cochlis pseudovittata sp. n. Pl. 2, igs. 16, 17; Pl. 3, ig. 1 Derivation of name: From Greek pseudes = false and vittata = the name of the most closely similar species, i.e. Cochlis vittata (Gmelin, 1791). Holotype: Borelli: MGPT-PU 135079 (Pl. 2, ig. 16). Paratypes: Borelli: 66 spms., MGPT-PU 135080, 12 spms., MGPT-PU 135081, 9 spms., MGPT-PU 135082, 9 spms., MGPTPU 135083, 1 spm., MGPT-PU 135084 (Pl. 2, ig. 17), 1 spm., MGPT-PU 135085, 1 spm., MGPT-PU 135086, 1 spm., MGPTPU 135087, 1 spm., MPUM 11239; Rio di Bocca d’Asino: 2 spms., MZB 60139, 1 spm., MZB 60140 (Pl. 3, ig. 1), 1 spm., MPUM 11240; Stazzano: 1 spm., MGPT-PU 135088. Other material examined: Borelli: 3 spms., NP 9949; Rio di Bocca d’Asino: 6 spms., NP 9948. Material referred to as Cochlis vittata (Gmelin, 1791) by Pedriali & Robba (2005): Borelli: 1 spm., authors’ collection (see remarks below). Preservation: Mostly fair. Type locality: Borelli (see appendix). Horizon: Gray, medium sand of Tortonian age. Diagnosis: Protoconch depressed-turbiniform with sculptured initial whorl. Shell globose to globose-oval, with low to moderately elevated spire and globose last whorl. Umbilicus deep rather small to wide. Parietal callus hindering most adapical part of umbilicus, with subangular anterior lobe; funicle thin to moderately thick; umbilical callus small, semicircular to triangular, separated from parietal callus by narrow notch. Color pale brown with darker subsutural band and lower base; reddish median line on last whorl. Description. Protoconch small, depressedturbiniform, of 2.55-2.75 gently convex whorls, tip very small; abapical part of protoconch I with microgranules roughly arranged into spiral rows. Teleoconch globose to globose-oval, rather thick to thick. Spire broadly conical, low to moderately elevated, whorls convex, suture ine, adpressed. Last whorl globose-oval to inlated, moderately produced but not expanded toward aperture; subsutural shelf indistinct. Aperture D-shaped, in prosocline plane, length about 1.6 times width; outer lip semicircular, basal lip thickened. Parietal Eocene, Oligocene and Miocene naticid gastropods of Northern Italy callus rather thick, with concave abapertural outline, hindering most adapical part of umbilicus; anterior lobe subangular, ending below umbilical border. Umbilicus deep, rather small, wide in a few specimens; umbilical wall very steep. Funicle thin to moderately thick, separated from umbilical wall by shallow spiral furrow of variable breadth. Umbilical callus small, moderately thick, semicircular to triangular in outline, located at abapical onethird of inner lip, separated from parietal callus by narrow, moderately deep notch. Basal fasciole poorly differentiated. Surface with rather dense growth lines changing into narrow wrinkles subsuturally. Specimens from Rio di Bocca d’Asino retain pale brown background with darker subsutural band and lower base; vestige of median, reddish spiral line occur on last whorl of a few specimens. Operculum unknown. Dimensions (mm) DHW PD H D SH AH AW 0.0810.097 0.8871.015 4.5338.957 4.4387.770 1.2942.426 3.0716.699 3.0184.778 0.089 0.951 6.745 6.104 1.860 4.885 3.898 UW WUC WAD WAB IS SA 1.0301.786 0.1050.497 0.2450.581 0.4060.982 19°31° 100°132° 1.408 0.301 0.413 0.694 25° 116° Remarks. Cochlis pseudovittata is closely similar to the Pliocene to Recent species Cochlis vittata (Gmelin, 1791) in shell morphology, but can be differentiated from it because of its larval shell having a signiicantly greater diameter and a sculptured initial whorl (protoconch I). Moreover, C. pseudovittata differs from C. vittata in that it has a slightly lower spire with less convex whorls, a different color pattern (C. vittata has a reddishbrown reticulated pattern and two-three spiral rows of brown spots), and attains a smaller size. In a previous paper (Pedriali & Robba 2005), a single shell from the Tortonian of Borelli was assigned to Cochlis vittata. Re-examination of this specimen has proven that it is Cochlis pseudovittata. Consequently, the supposed appearance of C. vittata in the late Miocene (Pedriali & Robba 2005, p. 169) is incorrect. Gmelin’s species appears to have replaced C. pseudovittata by the early Pliocene. Stratigraphic occurrence. Cochlis pseudovittata sp. n. was recovered only from Tortonian deposits of Piedmont. 133 Cochlis raropunctata raropunctata (Sasso, 1827) Pl. 3, igs. 2-4 1827 Natica raro-punctata Sasso, p. 477. 1963 Natica tigrina - Venzo & Pelosio, p. 82, pl. 34, igs. 37, 38 (not Defrance, 1825). 2005 Cochlis raropunctata raropunctata - Pedriali & Robba, p. 155, pl. 3, igs. 10-20; pl. 4, igs. 1-6, 10; pl. 7, igs. 8, 9; pl. 8, igs. 1925; pl. 9, igs. 14-19; pl. 10, igs. 17-21, 23, 24, I-K (cum syn.). Type material: Natica raropunctata Sasso, type material not seen. Pedriali & Robba (2005, p. 156) noted that the syntypes of this species, originally in the Museo Civico di Storia Naturale “G. Doria” in Genova, were not found and are possibly lost. Material examined: Moncucco Torinese: 2 spms., PG 1a; Montegibbio: 1 spm. (operculum), MPUM 11241 (Pl. 3, ig. 2); Passo dei Meloni: 17 spms., (14 shells, 3 opercula), PP.PMO1/03, 20 spms., MPUM 11242, 21 spms., NP 9962; Pietracuta: 1 spm., PP.PC02/01; Rio di Bocca d’Asino: 2 spms., MZB 25987, 8 spms., MZB 45367, 1 spm., MZB 25986, 6 spms., MZB 29712, 14 spms., PG 25c, 5 spms., PG 27, 8 spms., PG 58, 2 spms., PG 6, 2 spms., PG 9, 1 spm., MPUM 11243, 2 spms., PG 11a, 7 spms., PG 13a, 10 spms., NP 9958, 2 spms., MPUM 11244, 1 spm., MPUM 11245 (Pl. 3, ig. 3), 1 spm., MPUM 11246, 1 spm., MPUM 11247 (Pl. 3, ig. 4), 1 spm., NP 9959, 23 spms., NP 9960, 1 spm., MZB 60142, 2 spms., MZB 60064, 2 spms., MZB 25991, 2 spms., MZB 60066, 24 spms., MZB 60067, 4 spms., MZB 60070, 8 spms., MZB 60106, 1 spm., MZB 60107, 7 spms., MZB 60108, 1 spm., MZB 60143, 1 spm., MPUM 11248, 1 spm. (operculated shell), NP 9961; Sant’Agata Fossili: 3 spms., MGPT-PU 23319, 2 spms., PG 14, 2 spms., PG 59a, 1 spm., PG 18a, 1 spm., PG 73c; Stazzano: 1 spm., MZB 29721, 1 spm., MZB 26893, 4 spms., MZB 29737, 6 spms., MGPT-PU 23409, 1 spm., MGPT-PU 23410; Vigoleno: 57 spms., private collection. Dimensions (mm) DHW PD H D SH AH AW 0.3250.389 0.5830.715 9.36531.129 9.64929.833 1.0426.722 7.80224.926 4.81816.282 0.357 0.649 20.247 19.741 3.882 16.364 10.550 UW WUC WAD WAB IS SA 2.49112.667 0.5494.501 0.6676,671 0.2172.549 13°41° 107°146° 7.579 2.525 3.669 1.383 27° 125° Remarks. The upper Miocene specimens listed above are conspeciic with the Pliocene ones dealt with by Pedriali & Robba (2005); for description and comments see the cited authors. Besides the difference in teleoconch characters mentioned by Pedriali & Robba (2005, p. 160), Cochlis raropunctata raropunctata can be distinguished from the lower-middle Miocene species Cochlis tigrina (Defrance, 1825) also by its protoconch of 0,75 fewer whorls, with signiicantly greater diameter of the initial half-whorl (twice the size), and by its operculum with different proportions of both gooves and ridges. As discussed by Pedriali & Robba (2005, p. Robba E., Pedriali L. & Quaggiotto E. 134 160), the name raropunctata Sasso, 1827 is valid and, being based on Pliocene shells from Western Liguria, is the most suitable one for the present species. The use of the name tigrina Defrance, 1825 suggested by Lozouet et al. (2001) for Sasso’s species is unjustiied (see also the remarks on C. tigrina). Stratigraphic occurrence. Cochlis raropunctata raropunctata (Sasso, 1827) appeared in the Tortonian of Italy and spread throughout Southern Europe during the Pliocene. The species occurs rarely in the early Pleistocene and seems to have reached extintion about at the end of this subepoch. As already noted (Pedriali & Robba 2005), the Miocene records outside Italy refer to taxa other than C. raropunctata raropunctata, or need to be conirmed. ally; spiral microstriation observable on last whorl of best preserved specimens. Dimensions (mm) H D SH AH AW UW 15.770 15.379 3.641 12.130 9.631 5.739 9.010 9.358 1.791 7.219 6.510 2.880 WUC WAD WAB IS SA 1.592 2.461 1.690 20° 130° 0.981 0.980 0.921 27° 140° Remarks. Except for two specimens, the rest of the examined material is badly damaged. Its assignment to Oppenheim’s species, based on the original description and rather poor illustration, is made with some reservation. Natica canovae Oppenheim, 1901 (no specimens examined) seems to be Cochlis rossii (Oppenheim, 1901) comb. n. Pl. 3, igs. 5, 6 1901 Natica rossii Oppenheim, p. 200, pl. 3, ig. 10. 1985 Natica rossii - Brigantini, p. 414, pl. 2, ig. 42. Type material: Natica rossii Oppenheim, type material not seen. The Oppenheim collection is housed in the Hebrew University of Jerusalem (Israel). Inquiries there had negative results. Material examined: Valle Organa: 5 spms., MGP-PD 31518, 15 spms., MGP-PD 31520, 1 spm., MGP-PD 31523 (Pl. 3, ig. 5), 1 spm., MGP-PD 31521 (Pl. 3, ig. 6), 1 spm., MGP-PD 31522. Description. Protoconch abraded in all specimens. Teleoconch globose, very thick. Spire obtusely conical, low, less so in a few specimens, earliest whorls nearly lat-sided, subsequent ones gently convex; suture adpressed, almost lush. Last whorl globular, subsutural shelf indistinct, periphery at midline. Aperture Dshaped, in prosocline plane, length about twice width; outer lip regularly arched, abapical part of inner lip and basal lip markedly thickened, everted. Parietal callus thick, rather long, with straight or slightly concave abapertural outline, hindering most adapical part of umbilicus; anterior lobe poorly differentiated to indistinct. Umbilicus deep, rather narrow to narrow, umbilical wall increasingly steep downward. Funicle moderately wide, depressed, ending toward abapical one-fourth of inner lip, separated from umbilical wall by shallow, nearly latbottomed spiral furrow. Umbilical callus, moderately thick, rather narrow, slightly arched in outline, separated from parietal callus by wide, very shallow reverse J-shaped notch. Basal fasciole broad, scarcely prominent, coated with callus layer. Surface with rather dense growth lines changing into narrow wrinkles subsutur- PLATE 3 Fig. 1 - Cochlis pseudovittata sp. n. Rio di Bocca d’Asino. Paratype, MZB 60140; apertural side (height of shell 10.54 mm). Fig. 2 - Cochlis raropunctata raropunctata (Sasso, 1827). Montegibbio. MPUM 11241; operculum (height of operculum 14.38 mm). Fig. 3 - Cochlis raropunctata raropunctata (Sasso, 1827). Rio di Bocca d’Asino. MPUM 11245; protoconch. Fig. 4 - Cochlis raropunctata raropunctata (Sasso, 1827). Rio di Bocca d’Asino. MPUM 11247; apertural side (height of shell 17.18 mm). Fig. 5 - Cochlis rossii (Oppenheim, 1901). Valle Organa. MGP-PD 31523; apertural side (height of shell 15.77 mm). Fig. 6 - Cochlis rossii (Oppenheim, 1901). Valle Organa. MGP-PD 31521; apertural side (height of shell 9.01 mm). Fig. 7 - Cochlis sallomacensis (Tournouër, 1873). Salles (France). Lectotype (here designated) of Natica sallomacensis Tournouër, 1873. MNHN-F-B28542; from left to right: apertural side, lateral side, abapertural side. Fig. 8 - Cochlis sallomacensis (Tournouër, 1873). Monte dei Cappuccini. MGPT-PU 25667; a, apertural side (height of shell 16.14 mm); b, apical view; c, protoconch. Fig. 9 - Cochlis sallomacensis (Tournouër, 1873). Salles, Moulin Débat (France). Lesport collection; a, apertural side of shell with the operculum in situ (15L0005, height of shell 31.50 mm); b, basal view; c, outer surface of operculum (15L0006, height of operculum 22.00 mm); d, inner surface of operculum. Fig. 10 - Cochlis sallomacensis (Tournouër, 1873). Sallespisse, Carré, La Coustillière (France). Lesport collection 15L0004; operculum (height of operculum 9.60 mm). Fig. 11 - Cochlis separata (Deshayes, 1864). Cuise-la-Motte (France). Lectotype (here designated) of Natica separata Deshayes, 1864. EM 32860a; apertural side. Fig. 12 - Cochlis separata (Deshayes, 1864). Cava Boschetto. MPUM 11251; apertural side (height of shell 12.40 mm). Fig. 13 - Cochlis separata (Deshayes, 1864). Cava Boschetto. MPUM 11252; protoconch. Fig. 14 - Cochlis separata (Deshayes, 1864). Cava Grola. MPUM 11253; basal view (height of shell 8.23 mm). Fig. 15 - Cochlis separata (Deshayes, 1864). Cuise-la-Motte (France). MPUM 11254; protoconch. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 135 Robba E., Pedriali L. & Quaggiotto E. 136 the most closely similar species, differing in that has a less globular shell, with higher spire. Stratigraphic occurrence. Cochlis rossii was recovered from Priabonian marly deposits of Veneto. Cochlis sallomacensis (Tournouër, 1873) comb. n. Pl. 3, igs. 7-10 1873a Natica sallomacensis Tournouër, p. 154. 1873b Natica sallomacensis - Tournouër, p. 293, pl. 10, ig. 7. 1919 Natica sallomacensis - Cossmann & Peyrot, p. 194, pl. 11, igs. 6-8. 1952b Natica tigrina f. sallomacensis - Glibert, p. 257, pl. 2, igs. 3c, 3d. 1969 Natica tigrina hoernesi forma sallomacensis - Janssen, p. 173, pl. 7, igs. 19, 20. not 1972 Naticarius hoernesi sallomacensis - Nordsieck, p. 71, pl. 17, ig. 88 (= Natica neglecta Mayer, 1858). 1984 Natica sallomacensis - Janssen, p. 204, pl. 57, ig. 8. 2001 Natica sallomacensis - Wienrich, p. 433, pl. 88, ig. 1. Type material: Natica sallomacensis Tournouër, lectotype (here designated): the shell igured by Tournouër (1873b, pl. 10, ig. 7) and reigured herein (Pl. 3, ig. 7), MNHN-F-B28542, Salles (France); 2 paralectotypes, MNHN-F-A51515, Salles (France). Material erroneously referred to as Natica (Natica) millepunctata var. sismondiana d’Orbigny, 1852 in MGPT: Colli Torinesi: 2 spms., MGPT BS.029.01.003/03. Other material examined: Monte dei Cappuccini: 1 spm., MGPT-PU 25667 (Pl. 3, ig. 8); Montegibbio: 1 spm., NP 9924, 1 spm., MPUM 11249, 1 spm., MPUM 11250, 1 spm., NP 9925; Salles, Argilas (France): 20 spms., NP 10012, 1 spm. (operculated shell), NP 10013, 1 spm. (operculum), NP 10014; Salles, Moulin Débat (France): 1 spm. (operculated shell), 15L0005/15L0006 (Pl. 3, ig. 9); Sallespisse, Carré, La Coustillière (France): 1 spm. (operculum), 15L0004 (Pl. 3, ig. 10). We also examined an excellent photograph of the specimen from Salles illustrated by Cossmann & Peyrot (MNHN-FJ05669); Valle Ceppi: 1 spm., MGPT-PU 25906. Description. Protoconch small, turbiniform, of 2.25-2.40 convex whorls, tip small. Teleoconch medium-sized, globose, rather thick. Spire broadly conical, low to moderately elevated, somewhat stepped; whorls convex, subsutural shelf indistinct; shallowly channeled suture develops by second whorl, sutural channel bounded abaxially by obtuse angulation. Last whorl globular, slightly depressed in a few specimens, scarcely expanded toward aperture. Aperture D-shaped, in prosocline plane, length about 1.82 times width; outer lip semicircular, inner lip straight, basal lip thickened, slightly everted adaxially. Parietal callus moderately thick, trapeziform to rectangular, ending above level of basal fasciole; anterior lobe poorly differentiated to indistinct. Umbilicus deep, moderately wide; umbilical wall very steep; interior of umbilicus coated with callus layer extending over basal fasciole. Funicle cord-like, prominent, separated from umbilical wall by narrow to very narrow spiral furrow. Umbilical callus thick, at abapical one-third of inner lip, roundly triangular, separated from anterior corner of parietal callus by reverse J-shaped, shallow sinuation. Basal fasciole lattened, rather narrow, bounded abaxially by very slight step in larger specimens. Surface with uneven growth markings, coarser subsuturally and slightly bent on border of sutural channel. Some specimens from Montegibbio retain pale brown background, with darker band encircling umbilicus. Operculum (see remarks) moderately thick; central callus tongue-shaped, not prominent at all, reaching halfheight of operculum; inner margin gently arched, smooth or with blunt transverse wrinkles; inner surface nearly lat, with distinct spiral striation, nucleus not protruding; outer surface planar, with three marginal grooves and three latly round-topped ridges; inner and median grooves with concave bottom, equally narrow; outer groove arched in crosssection, constantly twice to three times wider than other two; outer and median ridges robust, about of equal strength; inner ridge also robust, but invariably thinner than other two. Dimensions (mm) DHW PD H D SH AH AW 0.1110.131 0.8270.935 10.28418.672 9.40618.274 1.1205.992 8.84413.000 6.96810.016 0.121 0.881 14.478 13.840 3.556 10.922 8.492 UW WUC WAD WAB IS SA 3.9006.824 0.9183.158 1.7983.138 0.0001.994 11°-31° 113°137° 5.362 2.038 2.468 0.856 21° 125° Remarks. We examined a vast naticid collection from the Serravallian of Sallespisse, largely dominated by Cochlis sallomacensis (300 specimens out of 385). The collection includes several loose opercula, some quite large-sized (up to 25 mm in height), all identical and certainly conspeciic. Among the ive species present in the naticid assemblage, only C. sallomacensis attains a very large size and the aperture of its shell can readily accomodate opercula of the considerable proportions cited. Furthermore, two shells of C. sallomacensis from the Serravallian of Salles were recovered with the operculum within the aperture (Pl. 3, ig. 9); these opercula, identical to those of Sallespisse, perfectly it in the aperture of Eocene, Oligocene and Miocene naticid gastropods of Northern Italy the shell and can be considered in situ. On this basis, we conclude that the opercula from Sallespisse and those from Salles can be safely assigned to C. sallomacensis. The operculum of Cochlis sallomacensis, having the outer surface with three marginal ridges and three grooves, somewhat resembles those of Cochlis neglecta (Mayer, 1858), Cochlis pseudoepiglottina (Sacco, 1890) and Cochlis vittata (Gmelin, 17919), which also have three marginal ridges and three grooves. However, the operculum of C. sallomacensis can be easily differentiated from that of C. neglecta in that it attains a larger size and has: 1) a rather long, tongueshaped central callus (shorter, subtriangular in C. neglecta), 2) the ridges and the grooves in the same plane of the outer surface (outer groove, median groove and ridges on distinctly elevated shelf in C. neglecta), 3) a markedly wider outer groove, and 4) an inner ridge thinner than the other two. The operculum of the Pliocene species C. pseudoepiglottina differs from that of C. sallomacensis by its central callus bending toward the inner margin, its outer and median grooves, and the ridges on an elevated shelf, its sharp median ridge distinctly narrower than the other two, its narrower outer groove, and its inner groove constantly wider and deeper than the other two. The operculum of C. vittata also has the outer and median grooves, and the ridges lying upon an elevated shelf; moreover, it is distinguished from that of C. sallomacensis because of its thin, similar outer and inner ridges, its wider, lat-topped median ridge, and its grooves of same breadth (see Pedriali & Robba 2005). Cochlis sallomacensis is morphologically similar to Natica aquitanica Tournouër, 1873 (a species also belonging in Cochlis Röding, 1798) and the values of the characteristic elements of the protoconch of both taxa do not differ signiicantly. However, C. sallomacensis can be readily distinguished from Tournouër’s species by its channeled suture, its more widely open umbilicus, and its differently shaped inner lip calluses (C. aquitanica has a rectangular, longer parietal callus fused with the small umbilical callus, without any sinuation in between). Natica burdigalensis Mayer, 1864 is another similar species in teleoconch shape, but has a protoconch of 0,5 more whorls, an adpressed suture, and an operculum that is peculiar of the species of the genus Tanea Marwick, 1931. For the relationships with Cochlis tigrina (Defrance, 1825), see the remarks on this species. 137 Stratigraphic occurrence. Cochlis sallomacensis (Tournouër, 1873) is known from Langhian to Serravallian deposits of France, from the Burdigalian of The Netherlands and the Langhian of Germany. Italian occurrences were from the Burdigalian and Langhian of Piedmont (rare) and from the Tortonian of Montegibbio (Emilia). Cochlis separata (Deshayes, 1864) comb. n. Pl. 3, igs. 11-15 1864 Natica separata Deshayes, p. 49, pl. 68, igs. 4-6. 1888 Natica separata - Cossmann, p. 161. 1907 Natica separata - Cossmann & Pissarro, pl. 9, ig. 61-9. Type material: Natica separata Deshayes, lectotype (here designated): the specimen igured by Deshayes (1864, pl. 68, igs. 4-6) and reigured herein (Pl. 3, ig. 11), EM 32860a, Cuise-la-Motte (France); 2 paralectotypes, EM 32860, Cuise-la-Motte (France). Other material examined: Cava Albanello: 6 spms., MGPPD 1218R, 5 spms., MGP-PD 1219R, 1 spm., MGP-PD 1220R, 5 spms., MGP-PD 1221R, 8 spms., NP 9972, 2 spms., NP 9973, 3 spms., MCZ 4319; Cava Boschetto: 1 spm., MPUM 11251 (Pl. 3, ig. 12), 2 spms., NP 9974, 1 spm., MPUM 11252 (Pl. 3, ig. 13); Cava Grola: 1 spm., private collection, 1 spm., private collection, 1 spm., NP 9979, 1 spm., MPUM 11253 (Pl. 3, ig. 14), 1 spm., MCV 15/01, 2 spms., private collection; Cava Rossi: 1 spm., MCZ 4320; Monte Merlo: 1 spm., NP 9976; Cuise-la-Motte (France): 1 spm., MPUM 11254 (Pl. 3, ig. 15), 4 spms., NP 9975; Saint Gobain (France): 1 spm., NP 9978; Villiers-Saint-Frédéric (France): 6 spms., NP 9977. Description. Protoconch small, depressedturbiniform, averaging 2.77 gently convex whorls, tip very small, slightly sunken. Teleoconch globoseoval, globose in some specimens, rather thick. Spire broadly conical, moderately elevated, less so in a few specimens, whorls convex, with poorly deined or indistinct subsutural shelf, suture adpressed. Last whorl globose, not expanded toward aperture; subsutural shelf very slightly concave, obscurely bounded abaxially in larger specimens. Aperture D-shaped, in moderately prosocline plane, length averaging 1.8 times width; outer lip semicircular, inner lip nearly straight. Parietal callus long, rather thick, with subangular abapertural outline; anterior lobe well developed, roundly angular, at level of umbilical border. Umbilicus deep, rather small, umbilical wall steep, umbilical channel a moderately wide, semicircular furrow. Funicle broad, depressed toward interior of umbilicus, separated from umbilical wall by shallow, lat-bottomed spiral furrow forming slight notch on inner lip; furrow bounded abaxially by distinct step. Umbilical callus moderate, Robba E., Pedriali L. & Quaggiotto E. 138 thick, roundly triangular in outline, located at midabapical part of inner lip, separated from anterior lobe of parietal callus by moderately deep, narrow notch. Basal fasciole indistinct. Surface with ine, dense growth lines. Dimensions (mm) DHW PD H D SH AH AW 0.0620.066 0.7560.872 2.89112.919 2.47212.644 0.8213.965 1.6889.336 1.4047.308 0.064 0.814 7.905 7.558 2.393 5.512 4.356 UW WUC WAD WAB IS SA 1.3223.986 0.5922.104 0.3260.802 0.1181.366 7°-27° 102°130° 2.654 1.348 0.564 0.742 17° 116° Remarks. Cochlis separata is remarkably similar to Cochlis epiglottinoides (Deshayes, 1864) in teleoconch characters and could be mistaken for it. However, the protoconch of 0.7 more whorls, with signiicantly smaller diameter of the irst half-whorl (over half the size) unequivocally distinguishes C. separata from C. epiglottinoides. Natica microglossa Deshayes, 1864 (a species of Cochlis) supericially resembles Cochlis separata, but differs from it in that it has a protoconch with signiicantly greater diameter of the initial half-whorl, a spire with less convex whorls, and a spiral furrow separating the funicle from the umbilical wall, which is narrower, V-shaped, and lacks the step-like abaxial border present in C. separata. According to previous authors, the present species exhibits some resemblance also to Natica epiglottina Lamarck, 1804. Actually, both taxa share some teleoconch characters, but C. separata is readily differentiated from N. epiglottina because of its protoconch of one more whorl, with signiicantly smaller diameter of the irst half-whorl (one-fourth of the size). Stratigraphic occurrence. Cochlis separata (Deshayes, 1864) was hitherto known from upper Ypresian and Lutetian units of France. Italian occurrences are from upper Ypresian and lower-mid Lutetian deposits of Veneto. Cochlis tigrina (Defrance, 1825) Pl. 4, igs. 1-5 1825 Natica tigrina Defrance, p. 257. 1828 Natica tigrina - Grateloup, p. 155. 1828 Natica tigrina vars. a, b, c, d Grateloup, p. 156. 1847 Natica tigrina var. A ampullosa Grateloup, pl. 9, igs. 10, 11. 1847 Natica tigrina var. B crassiuscula Grateloup, pl. 9, igs. 12, 13. 1847 Natica tigrina var. C ovata Grateloup, pl. 9, ig. 14. 1847 Natica tigrina var. D millepunctata - Grateloup, pl. 10, igs. 2, 3 (not Lamarck, 1822). 1847 Natica tigrina var. E immaculata Gateloup, pl. 10, ig. 4. 1847 Natica tigrina var. F punctulata Grateloup, pl. 10, ig. 5. ? 1873a Natica aquitanica Tournouër, p. 154. ? 1873b Natica aquitanica - Tournouër, p. 292, pl. 10, ig. 6. 1890b Natica (Natica) millepunctata var. sismondiana - Sacco, p. 28 (not Natica sismondiana d’Orbigny, 1852). 1890b Natica (Natica) millepunctata subvar. miorotunda Sacco, p. 28 (nomen nudum). 1890b Natica (Natica) millepunctata subvar. miolonga Sacco, p. 28 (nomen nudum). 1890b Natica (Natica) millepunctata subvar. miodepressispira Sacco, p. 28 (nomen nudum). ? 1890b Natica (Natica) millepunctata var. miocontorta Sacco, p. 28. 1890b Natica (Natica) millepunctata var. tauropicta Sacco, p. 28. PLATE 4 Fig. 1 - Cochlis tigrina (Defrance, 1825). Léognan, Le Coquillat (Fran ce). MPUM 11255; a, apertural side (height of shell 20.17 mm); b, protoconch. Fig. 2 - Cochlis tigrina (Defrance, 1825). Léognan, Le Coquillat ( Fr a n c e ) . MPUM 11258; operculum. Fig. 3 - Cochlis tigrina (Defrance, 1825). Valle Ceppi. MGPT-PU 108196; protoconch. Fig. 4 - Cochlis tigrina (Defrance, 1825). Val Sanfrà. MGPT-PU 135092; apertural side (height of shell 21.54 mm). Fig. 5 - Cochlis tigrina (Defrance, 1825). Val Sanfrà. MGPT-PU 135093; apertural side (height of shell 24.61 mm). Fig. 6 - Cochlis sp. 1. Cava Grola. MPUM 11260; apertural side (height of shell 12.28 mm). Fig. 7 - Cochlis sp. 1. Ciupio. MGP-PD 11624/a-11664/a; apertural side (height of shell 11.31 mm). Fig. 8 - Cochlis sp. 2. Monte Gloso. MPUM 11261; a, apertural side (height of shell 6.65 mm); b, apical view. Fig. 9 - Cryptonatica noe (d’Orbigny, 1850). Saint-Witz, Le Guépelle (France). Lectotype (here designated) of Natica noe d’Orbigny, 1850. EM 32834a; apertural side. Fig. 10 - Cryptonatica noe (d’Orbigny, 1850). Saint-Witz, Le Guépelle (France). Paralectotype. Lot EM 32834; apertural side of shell with the operculum in situ. Fig. 11 - Cryptonatica noe (d’Orbigny, 1850). Cava Grola. MPUM 11236; apertural side (height of shell 13.37 mm). Fig. 12 - Cryptonatica noe (d’Orbigny, 1850). Cava Grola. MPUM 11237; protoconch. Fig. 13 - Cryptonatica noe (d’Orbigny, 1850). Saint-Witz, Le Guépelle (France). MPUM 11238; protoconch. Fig. 14 - Tanea dillwyni koeneni (Sacco, 1891). Monte dei Cappuccini. Lectotype (here designated) of Natica (Cochlis) dillwyni var. taurominor Sacco, 1904. MGPT BS.029.01.045; apertural side. Fig. 15 - Tanea dillwyni koeneni (Sacco, 1891). Borelli. MGPT-PU 135095; apertural side of shell with the operculum in situ. Fig. 16 - Tanea dillwyni koeneni (Sacco, 1891). Borelli. MGPT-PU 135097; a, apertural side (height of shell 8.84 mm); b, basal view. Fig. 17 - Tanea dillwyni koeneni (Sacco, 1891). Borelli. MGPT-PU 135099; operculum (height of operculum 8.77 mm). Fig. 18 - Tanea dillwyni koeneni (Sacco, 1891). Monte dei Cappuccini. MGPT-PU 135100; a, apertural side (height of shell 11.96 mm); b, apical view. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 139 140 Robba E., Pedriali L. & Quaggiotto E. 1891 Natica (Natica) millepunctata var. sismondiana - Sacco, p. 45, pl. 2, ig. 3 (not Natica sismondiana d’Orbigny, 1852). 1891 Natica (Natica) millepunctata var. miorotunda Sacco, p. 46. 1891 Natica (Natica) millepunctata var. miolonga Sacco, p. 46. 1891 Natica (Natica) millepunctata var. miodepressispira Sacco, p. 46. ? 1891 Natica (Natica) millepunctata var. miocontorta - Sacco, p. 46, pl. 2, ig. 4. 1891 Natica (Natica) millepunctata var. tauropicta - Sacco, p. 46, pl. 2, ig. 5. 1904 Natica (Natica) millepunctata var. miorotunda - Sacco, p. 102, pl. 22, ig. 18. 1904 Natica (Natica) millepunctata var. miolonga - Sacco, p. 102, pl. 22, ig. 19. 1904 Natica (Natica) millepunctata var. miodepressispira - Sacco, p. 102, pl. 22, ig. 20. 1919 Natica pachyope Cossmann & Peyrot, p. 189, pl. 11, igs. 4, 5 (new synonym). 1919 Natica tigrina - Cossmann & Peyrot, p. 190, pl. 11, igs. 1, 9, 10. 1952a Natica tigrina - Glibert, p. 74, pl. 5, ig. 13. 1952b Natica tigrina - Glibert, p. 255 (pars), pl. 2, igs. 3a, b, e, f (not igs. 3c, d = Natica sallomacensis Tournouer, 1873). ? 1960 Natica (N.) tigrina hoernesi - Baldi, p. 64, pl. 2, igs. 2a, 2c. not 1969 Natica tigrina tigrina - Janssen, p. 171, pl. 6, ig. 25; pl. 7, igs. 1-8 (see remarks below). not 1984 Natica tigrina - Janssen, p. 203, pl. 57, igs. 1-4, 10 (see remarks below). 1984 Natica millepunctata var. sismondiana - Ferrero Mortara et al., p. 28 (not Natica sismondiana d’Orbigny, 1852). 1984 Natica millepunctata var. miorotunda - Ferrero Mortara et al., p. 28. 1984 Natica millepunctata var. miolonga - Ferrero Mortara et al., p. 28. 1984 Natica millepunctata var. miodepressispira - Ferrero Mortara et al., p. 28. ? 1984 Natica millepunctata var. miocontorta - Ferrero Mortara et al., p. 28. 1984 Natica millepunctata var. tauropicta - Ferrero Mortara et al., p. 28. not 1987 Natica (Natica) tigrina - Karczewski, p. 131, pl. 34, igs. 4, 7, 9-13 (not Defrance, 1825). 1995 Natica tigrina - Baluk, p. 194, pl. 15, igs. 10-14. 2001 Natica crassiuscula - Lozouet et al., p. 43, pl. 18, ig. 1. 2006 Natica tigrina - Baluk, p. 206, pl. 3, ig. 6. 2007 Natica tigrina - Zunino, p. 124 (pars), pl. 1, igs. 10-12. Type material: Natica tigrina Defrance, type material not seen. Cleevely (1983) stated that the Defrance collection housed in the Musée d’Histoire Naturelle of Caen (France) was destroyed during the 2nd world war. According to Dance (1986), some Defrance’s specimens should be in the Muséum d’Histoire Naturelle of Geneva (Switzerland), but no shells of Natica tigrina are present there (Serret 1986; Lionel Cavin, personal communication 2014). In these circumstances, the shells illustrated by Grateloup (1847) constitute the most relevant reference material; they are curated in the University of Bordeaux 1, but were not located yet (Bruno Cahuzac, personal communication 2014). Natica (Natica) millepunctata var. miorotunda Sacco, lectotype (here designated): the shell igured by Sacco (1904, pl. 22, ig. 18), MGPT BS.029.01.004, Colli Torinesi; 1 paralectotype, MGPT BS.029.01.004/01, Colli Torinesi (other 2 syntypes, also numbered MGPT BS.029.01.004/01, are unidentiiable). Natica (Natica) millepunctata var. miolonga Sacco, lectotype (here designated): the shell igured by Sacco (1904, pl. 22, ig. 19), MGPT BS.029.01.005, Colli Torinesi; 2 paralectotypes, MGPT BS.029.01.005/01, Colli Torinesi. Natica (Natica) millepunctata var. miodepressispira Sacco, lectotype (here designated): the shell igured by Sacco (1904, pl. 22, ig. 20), MGPT BS.029.01.006, Colli Torinesi; 2 paralectotypes, MGPT BS.029.01.006/01, Colli Torinesi (other 2 syntypes, also numbered MGPT BS.029.01.006/01, are unidentiiable. Natica (Natica) millepunctata var. tauropicta Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 2, ig. 5), MGPT BS.029.01.008, Colli Torinesi. Other material examined: Albugnano: 1 spm., PG 96, 1 spm., PG 97; Léognan, Le Coquillat (France): 1 spm., MPUM 11255 (Pl. 4, ig. 1), 1 spm., MPUM 11256, 19 spms., MPUM 11257, 2 spms., MZB 60121, 9 spms., private collection, 6 spms., PPMM 40819, 1 spm. (operculum), PPMM 40820, 13 spms., private collection, 1 spm. (operculum), private collection, 26 spms., private collection, 1 spm. (operculum), private collection, 117 spms., NP 9965, 2 spms. (operculated shells), NP 9998, 8 spms. (opercula), NP 9999, 1 spm. (operculum), MPUM 11258 (Pl. 4, ig. 2), 4 spms. (opercula), MPUM 11259; Martignas-sur-Jalle (France): 1 spm., NP 10005; Mérignac (France): 4 spms., NP 9963; Monte dei Cappuccini: 3 spms., MGPT-PU 25672; Roquefort (France): 1 spm., NP 9964; Saint-Martin-d’Oney (France): 6 spms., private collection; Uzeste, Gamachot (France): 2 spms., NP 10009; Valle Ceppi: 1 spm., MGPT-PU 135089, 1 spm., MGPT-PU 108196 (Pl. 4, ig. 3), 1 spm., MGPT-PU 107637, 1 spm., MGPT-PU 23862, 1 spm., MGPT-PU 107635, 10 spms., MGPT-PU 23668, 4 spms., MZB 60144, 4 spms., MZB 60145, 4 spms., MGPT-PU 23865, 1 spm., MGPT-PU 135090, 1 spm., MGPT-PU 135091, 1 spm., MZB 60146, 1 spm., MGPT-PU 135352; Val Sanfrà: 1 spm., MGPT-PU 107634, 1 spm., MGPT-PU 107630, 15 spms., MGPT-PU 107633, 1 spm., MGPT-PU 135092 (Pl. 4, ig. 4), 2 spms., MGPT-PU 107638, 1 spm., MGPT-PU 135093 (Pl. 4, ig. 5); Valle Vergnana: 1 spm., MZB 44010. Description. Protoconch small, lowturbiniform, of 1.8-2 convex and smooth whorls, tip small. Teleoconch large to very large, globose, thick. Spire broadly conical, low to moderately elevated, whorls convex; lat, gently sloping subsutural shelf, bounded abapically by bending of growth lines, develops by third whorl in most specimens; suture ine, adpressed. Last whorl globose, globoseoval in a few specimens, scarcely expanded toward aperture; subsutural shelf as on spire whorls. Aperture D-shaped, in slightly prosocline plane, length about twice width; outer lip semicircular, inner lip straight, basal lip thickened, everted. Parietal callus thick, more so adapically, trapeziform to rectangular, ending above basal fasciole; anterior lobe absent. Umbilicus deep, moderate to wide; umbilical wall steep; interior of umbilicus coated with callus layer extending over basal fasciole. Funicle cord-like and prominent to broad and markedly depressed, largely illing umbilicus in many specimens, separated from umbilical wall by shallow spiral furrow of variable breadth. Umbilical callus thick, located in mid-abapical part of inner lip, roundly triangular and merging into anterior corner of parietal callus with reverse J-shaped outline (depressed funicle), or semicircular, separated from parietal callus by nar- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy row, shallow notch (prominent funicle). Basal fasciole broad, scarcely prominent, lattened in larger specimens, bounded abaxially by slight step. Surface with rather dense growth lines. Holotype of Natica (Natica) millepunctata var. tauropicta Sacco, 1890 retains pale brown background with lighter pattern of undulating collabral stripes changing into collabral rows of oval spots on inal half-whorl. Operculum slightly thickened; central callus scarcely prominent, tongue-shaped, reaching half-height of operculum; inner margin smooth or with obscure transverse wrinkles; inner surface latly convex, nucleus not protruding; outer surface planar with two marginal grooves and two ridges. Juvenile opercula (about 3 mm in height) with broad, lat-topped marginal ridge and vestige of inner groove; at size of about 5 mm, marginal ridge bifurcates, thence outer groove develops separating narrow and sharp outer ridge from robust, round-topped inner ridge. In fully grown opercula: outer groove rather wide to wide, with concave bottom; inner groove deeper and as wide as outer one or slightly wider, also with concave bottom, bounded adaxially by rounded step in some specimens; outer ridge thin, rather sharp; inner ridge moderately thick, round-topped in most specimens, narrower and overhanging inner groove in a few specimens. Dimensions (mm) DHW PD H D SH AH AW 0.1480.180 0.6360.716 5.89744.461 7.65141.907 1.37611.640 3.94633.570 4.07826.206 0.164 0.676 25.179 24.779 6.508 18.758 15.142 UW WUC WAD WAB IS SA 1.55618.252 0.9796.939 0.0006.721 0.1225.682 11°31° 102°134° 9.904 3.959 3.043 2.902 21° 118° Remarks. Natica tigrina Defrance, 1825 is different from the Recent Indo-West Paciic species Cochlis tigrina Röding, 1798. The species name tigrina Defrance is herein combined with the generic name Cochlis Röding, 1798 and would be a junior secondary homonym of C. tigrina Röding, thence an invalid name (ICZN 1999, Articles 53.3 and 57.3 of the Code). However, since Röding’s species was designated as the type of the genus Paratectonatica Azuma, 1961 (see Kabat 1991 and Torigoe & Inaba 2011), it appears that the two species in question are not congeneric, and that the name tigrina Defrance is a valid name (ICZN 1999, Article 59 of the Code). 141 The name crassiuscula (a variety of N. tigrina Defrance introduced by Grateloup 1847), proposed by Lozouet et al. (2001) for the lower Miocene, French material, was not intended as a replacement name for tigrina Defrance. These authors did not consider the name tigrina Defrance invalid, instead they regarded it as the proper one for the Pliocene specimens, because of its wide use with reference to the fossils of that age in the Mediterranean area. This latter opinion is not acceptable (Pedriali & Robba 2005; present paper). Cochlis tigrina appears to be a rather variable species as regards the elevation of the spire, the amplitude of the umbilicus, the breadth and prominence of the funicle, the extent of the funicle within the umbilicus, and the shape of the umbilical callus. Besides the Italian material, we examined a vast lot of French specimens (including a shell in Cossmann collection, MNHN-F-J05667) and could note that the cited characters exhibit a gradual transition between respective end-forms. Natica pachyope was introduced by Cossmann & Peyrot (1919) on the basis of material collected at Saint-Jean-de-Marsacq (late Burdigalian) and at Saubrigues (Langhian). We have not seen the original material (included in the Degrange Touzin collection), however, from the excellent illustrations published by Cossmann & Peyrot (1919), it appears that the teleoconch of N. pachyope falls within the range of variation of Cochlis tigrina (see above) and the operculum is identical to that of C. tigrina. On the basis of this evidence, we consider N. pachyope a synonym of C. tigrina. Natica aquitanica Tournouër, 1873 was regarded as a synonym of C. tigrina (cited as Natica crassiuscula Grateloup, 1847) by Lozouet et al. (2001). We obtained an excellent photograph of the holotype of Tournouër’s species (MNHN-FB27839) and examined a lot of N. aquitanica from upper Aquitanian deposits of Saint-Martin-d’Oney. The teleoconch characters of N. aquitanica do not warrant speciic separation from C. tigrina and the values of the characteristic elements of the protoconch of both taxa are identical. However, since the operculum of N. aquitanica is unknown, we include the species in the synonymy of C. tigrina with doubt. The lower Badenian (Langhian) shell and the operculum recovered near Szokolya (Hungary), illustrated by Baldi (1960) and referred to as Natica (N.) tigrina hoernesi Fischer & Tournouër, 1873, seem to conform to those of C. tigrina. However, since 142 Robba E., Pedriali L. & Quaggiotto E. we have not examined this material, we prefer to include it in the synonymy of the present species with reservation. Natica sallomacensis Tournouër, 1873 is closely similar to Cochlis tigrina in shell characters, but can be readily distinguished from it in that it has the protoconch with signiicantly smaller diameter of the irst half-whorl, the spire whorls devoid of subsutural shelf, separated by distinctly channeled sutures (those of C. tigrina are adpressed), and the funicle placed more abapically. Cochlis curta (ErünalErentöz, 1958), described from the Serravallian of Turkey, is easily differentiated from C. tigrina by its protoconch with signiicantly smaller diameter of the irst half-whorl and by its depressed, markedly low-spired shell (see Landau et al. 2013). Mayer (1864, p. 166) described the new species Natica burdigalensis from lower Burdigalian deposits of Léognan and Saucats, and remarked that it had been “confondue jusqu’à ce jour avec le N. millepunctata don’t elle est de fait fort voisine” (N. millepunctata referred to by Mayer is Natica tigrina). Cossmann & Peyrot (1919, p. 193) thoroughly discussed the characters that distinguish N. burdigalensis from N. tigrina. Our collection includes several shells of N. burdigalensis from Léognan, two with the operculum still illing the aperture, and some loose opercula. On the basis of this material, we note that Mayer’s species is similar to C. tigrina in shell morphology and that the differentiating characters pointed out by Cossmann & Peyrot (1919) are correct. N. burdigalensis differs from C. tigrina also by its protoconch of 0.75 more whorls, with signiicantly greater diameter and smaller diameter of the initial half-whorl. The operculum with a broad marginal swelling bearing a narrow longitudinal groove demonstrates that N. burdigalensis is not congeneric with C. tigrina, instead it belongs in the genus Tanea Marwick, 1931. Fischer & Tournouër (1873) created the new species Natica hoernesi based on upper Miocene material from Cabrières. Examination of the photograph of syntype MNHN-F-R06714 has shown that the shell of N. hoernesi is more similar to Cochlis raropunctata raropunctata (Sasso, 1827) than to Cochlis tigrina. However, in the lack of information on the protoconch and the operculum of N. hoernesi, we abstain from any conclusion about its relationships. Sacco (1891) igured a shell referred to as Natica (Natica) millepunctata var. sismondiana d’Orbigny, 1852. The comparison of Sacco’s specimens in MGPT (BS.029.01.003, 003/01, 003/02, 003/03) with a syntype of N. sismondiana (MNHN-FA12745) has shown that they differ markedly from d’Orbigny’s taxon. N. sismondiana has an umbilicus largely illed by a massive funicle, an abapical, narrow umbilical channel, and a broad umbilical callus fused with the parietal callus, characters that are reminiscent of those of the genus Polinices Montfort, 1810. Instead, the specimens in MGPT have a widely open umbilicus and perfectly conform to the characters of Cochlis tigrina. It is to be noted that, of the nine specimens in lot BS.029.01.003/03, only six are C. tigrina, whereas one belongs to Pliconacca plicatulaeformis (Kittl, 1887) and two are Cochlis sallomacensis (Tournouër, 1873). The two specimens (MGPT BS.029.01.007 and 007/1) on which Sacco (1890b) based his variety miocontorta of Natica (Natica) millepunctata Lamarck, 1822 could be C. tigrina. However, their poor preservation prevents from a safe speciic identiication; accordingly, they are included in the synonymy of C. tigrina with doubt. The upper Burdigalian, Dutch shells identiied as Natica tigrina by Janssen (1969, 1984) supericially resemble Defrance’s species in teleoconch shape. However, direct examination of specimens RGM 225954 and RGM 225955 (of Janssen 1984) has shown that their opercula have a broad, swollen marginal area bearing a narrow, deep longitudinal groove and are closely similar to that of Natica zelandica Quoy & Gaimard, 1832, type species of the genus Tanea Marwick, 1831. On this basis, we conclude that these Dutch specimens are unlike N. tigrina. Instead, they represent a species belonging in the genus Tanea. Von Koenen (1882, p. 223) introduced the new species Natica beyrichi and excellently illustrated it (pl. 5, igs. 1-3). From the igures published by von Koenen (1882), it appears that also N. beyrichi belongs in Tanea and that it could be a more suitable assignment for the cited Dutch shells. The lower Miocene, Antarctic specimens referred to as Natica (Natica) tigrina by Karczewski (1987) are preserved as internal casts and can hardly be assigned to species. The assumption that these specimens are conspeciic with the European ones of Cochlis tigrina is incorrect. Stratigraphic occurrence. Cochlis tigrina (Defrance, 1825) is deinitely known from Aquitanian to Langhian deposits of France and Belgium, from the Langhian of Poland, and from Burdigalian Eocene, Oligocene and Miocene naticid gastropods of Northern Italy to Serravallian units of Italy (Piedmont). The species probably occurs also in the Langhian of Hungary and in the Serravallian of Belgium. Cochlis sp. 1 143 More, better preserved material retaining the protoconch is required in order to clarify their relationships. Stratigraphic occurrence. Cochlis sp. 1 was recovered from upper Ypresian and lower to middle Lutetian deposits of Veneto. Pl. 4, igs. 6, 7 Material examined: Cava Albanello: 1 spm., MGP-PD 1222R; Cava Grola: 1 spm., MPUM 11260 (Pl. 4, ig. 6), 1 spm., private collection; Cava Rossi: 1 spm., NP 9991, 3 spms., MCZ 4321; Ciupio: 1 spm., MGP-PD 11624/a-11664/a (Pl. 4, ig. 7). Description. Protoconch abraded. Teleoconch globose-oval, globose in a few specimens, thick. Spire broadly conical, rather low, whorls increasingly convex during growth, suture adpressed. Last whorl globose, not expanded toward aperture; larger specimens with very slightly concave subsutural shelf obscurely bounded abaxially. Aperture D-shaped, in moderately prosocline plane, length about twice width; outer lip semicircular, inner lip nearly straight. Parietal callus long, rather thick, with slightly angular abapertural outline; anterior lobe small, roundly angular. Umbilicus deep, rather small, umbilical wall steep. Funicle cord-like, thick and prominent, separated from umbilical wall by shallow spiral depression that is very narrow to moderately wide depending on size of funicle. Umbilical callus thick, semicircular, located at mid-abapical part of inner lip, separated from anterior lobe of parietal callus by moderately deep, narrow notch. Basal fasciole indistinct. Surface with remnants of uneven growth lines. Cochlis sp. 2 Pl. 4, fig. 8 Material examined: Case Soghe: 1 spm., NP 9994; Monte Gloso: 1 spm., MPUM 11261 (Pl. 4, fig. 8). Description. Protoconch poorly preserved, apparently of 2.75 whorls, tip very small. Teleoconch small, globose-oval, thick. Spire broadly conical, rather low, whorls gently convex, suture adpressed. Last whorl globose, not expanded toward aperture; subsutural shelf indistinct. Aperture D-shaped, in moderately prosocline plane, length about 1.7 times width; outer lip semicircular, inner lip very slightly arched. Parietal callus long, moderately thick, with subangular abapertural outline; anterior lobe indistinct. Umbilicus deep, small, crescent-shaped; umbilical wall steep. Funicle broad, increasingly depressed toward interior of umbilicus, separated from umbilical wall by shallow, rather narrow spiral depression. Umbilical callus thick, with arched abapertural outline, located at mid-abapical part of inner lip, separated from anterior end of parietal callus by very slight notch. Basal fasciole indistinct. Surface with remnants of ine growth lines; one specimen retains vestige of dark brown subsutural band and lowermost base. Dimensions (mm) Dimensions (mm) DHW PD H D SH AH AW - - 12.281 11.490 3.200 9.082 6.151 UW WUC WAD WAB IS SA 3.990 1.511 0.959 1.520 25° 111° Remarks. The present specimens resemble Cochlis saparata (Deshayes, 1864) in teleoconch shape, but they differ from Deshayes’ species in having a smaller anterior lobe of the parietal callus, and an abapical sulcus arched in cross section and smoothly merging into the umbilical wall (flat-bottomed and separated from the umbilical wall by a step in C. separata). DHW PD H D SH AH AW ~ 0.071 - 6.651 6.240 1.822 4.830 3.569 UW WUC WAD WAB IS SA 1.921 0.943 0.260 0.719 14° 109° Remarks. The Rupelian species Cochlis infelix (Sacco, 1890) somewhat resembles Cochlis sp. 2 in shell shape, but differs from it in having a wider umbilicus, a narrower, cord-like funicle, and a smaller, semicircular umbilical callus widely separated from the parietal callus. Stratigraphic occurrence. Cochlis sp. 2 appears to be an uncommon element, occurring in Rupelian deposits of Veneto. 144 Robba E., Pedriali L. & Quaggiotto E. Genus Cryptonatica Dall, 1892 Cryptonatica Dall, 1892, p. 362. Type species by subsequent designation (Dall 1909, p. 85): Natica clausa Broderip & Sowerby, 1829 (= Nerita afinis Gmelin, 1791), Recent, North Atlantic. Boreonatica Golikov & Kussakin, 1974, p. 294. Type species by original designation: Natica clausa Broderip & Sowerby, 1829. Sulconatica Golikov & Kussakin, 1974, p. 294. Type species by original designation: Natica janthostoma Deshayes, 1839 (see also Kabat 1991 and Torigoe & Inaba 2011). Dall (1892, p. 362) introduced Cryptonatica with no type species designation. Petit (1986, p. 38) and Kabat (1991, p. 428), evidently relying upon the subsequent designation made by Cossmann (1896, p. 238), indicated Natica (Cryptonatica) loridana Dall, 1892 as the type species of Cryptonatica. However, Bouchet & Waren (1993, p. 758) discussed this type species designation and correctly considered invalid Cossmann’s action. They reinstated Natica clausa Broderip & Sowerby, 1829 (= Nerita afinis Gmelin, 1791) as the type species of Cryptonatica following the designation subsequently made by Dall himself (1909). This opinion was widely accepted by recent authors (cf. Zhang & Wei 2010; Torigoe & Inaba 2011). According to Kabat (1991), Crytonatica Dall, 1921, Cryptonica Cossmann, 1925 and Cryotonatica Oyama, 1969 are uncorrect spellings; Cryptonatica Cossmann, 1925 is an error for Cryptonerita Kittl, 1894. Cryptonatica was considered either as a synonym of Tectonatica Sacco, 1890 (Cossmann 1925; Wenz 1941 in 1938-1944; Wrigley 1949), or as a distinct genus (Marincovich 1977; Oyama 1985; Golikov & Sirenko 1988; Majima 1989; Kabat 1991; Bouchet & Waren 1993; Torigoe & Inaba 2011). The relationships between Tectonatica and Cryptonatica were amply discussed by Pedriali & Robba (2008a, pp. 98, 99), who concluded: 1) that the umbilical characters “hardly provide the ground for separating Cryptonatica from Tectonatica”, and 2) that the operculum is a suitable character in distinguishing the species of Cryptonatica, which have the operculum with smooth outer surface, from those of Tectonatica (outer surface of the operculum with a distinct groove bounding a wide marginal area). The Eocene species Cryptonatica noe dealt with below, along with an unidentiied species from Fleury-la-Rivière (Cave aux Coquillages), France, seem to be the oldest European members of the genus. According to the illustration published by its author (pl. 40, ig. 9b), also the lower Oligocene species Natica semperi von Koenen, 1891 belongs in Cryptonatica. Cryptonatica noe (d’Orbigny, 1850) comb. n. Pl. 4, igs. 9-13 1832 Natica glaucinoides Deshayes, p. 166, pl. 20, igs. 7, 8 (not J. Sowerby, 1812). 1850 Natica noe d’Orbigny, p. 413, n°1437. 1864 Natica noae - Deshayes, p. 55. 1888 Natica noae - Cossmann, p. 160. 1907 Natica noae - Cossmann & Pissarro, pl. 9, ig. 61-5. 1949 Natica noae - Wrigley, p. 12, ig. 3. 2008 Natica noae - Quaggiotto & Mellini, p. 48. Type material: Natica noe d’Orbigny, lectotype (here designated): the specimen igured by Deshayes (1832, pl. 20, igs. 7, 8) as Natica glaucinoides (see remarks below) and reigured herein (Pl. 4, ig. 9), EM 32834a, Saint-Witz, Le Guépelle (France); 9 paralectotypes, EM 32834, Saint-Witz, Le Guépelle (France). Other material examined: Cava Albanello: 2 spms., NP 9967; Cava Grola: 4 spms., private collection, 1 spm., MPUM 11236 (Pl. 4, ig. 11), 9 spms., private collection, 4 spms., private collection, 8 spms., private collection, 4 spms., NP 9968, 1 spm., MPUM 11237 (Pl. 4, ig. 12), 1 spm., private collection, 1 spm., NP 9992; SaintWitz, Le Guépelle (France): 1 spm., NP 9969, 1 spm., MPUM 11238 (Pl. 4, ig. 13). Description. Protoconch small, turbiniform, very depressed, of 1.75-1.90 slightly convex whorls, tip small. Teleoconch globose to globose-oval, rather thin. Spire broadly conical, rather pointed, short, whorls latly convex; suture slightly incised, almost lush. Last whorl broadly oval, subsutural shelf indistinct, periphery above midline. Aperture D-shaped, in prosocline plane, length twice width; outer lip nearly semicircular, thin-edged, inner lip straight, slightly thickened and everted abapically. Parietal callus subtrapezoidal, rather short, thin to moderately thick, with obscure adapical tubercle; anterior lobe indistinct. Umbilicus wide, deep, with angular border more distinct abapically; umbilical wall rather steep. Funicle cord-like, thick and prominent, ending at middle of adapertural border of umbilicus, separated from umbilical wall by shallow spiral furrow. Umbilical callus large, thick, prominently semicircular in outline, located at middle part of inner lip, separated from anterior corner of parietal callus by shallow, reverse J-shaped notch. Basal fasciole indistinct. Surface with uneven and unevenly spaced growth lines. Some specimens from Veneto retain vestige of uniform olive-green background. Calcareous operculum with smooth outer surface. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Dimensions (mm) DHW PD H D SH AH AW 0.1910.207 0.7490.797 5.50513.765 5.34913.821 0.4533.545 4.65910.611 3.0088.512 0.199 0.773 9.635 9.585 1.999 7.635 5.760 UW WUC WAD WAB IS SA 2.0057.697 0.5442.728 0.0402.360 0.9242.932 9°-29° 99°135° 4.851 1.636 1.200 1.928 19° 117° Remarks. Natica noe was introduced by d’Orbigny (1850) as replacement name for Natica glaucinoides Deshayes, 1834, which is a primary homonym of Natica glaucinoides J. Sowerby, 1812 (see also Pacaud 2007, p. 40). The name glaucinoides Deshayes is permanently invalid (ICZN 1999, Article 57.2 of the Code). The specimens dealt with by Deshayes when describing his N. glaucinoides are the syntypes of Natica noe. The original spelling noe is correct (the Latin name Nōe of the patriarch Noah remains unchanged in all cases) and its correction to noae adopted by subsequent workers is unjustiied. The umbilical characters of Natica noe fully agree with those of the species currently included in the genus Cochlis Röding, 1798. However, on the basis of its operculum with smooth outer surface (see Pl. 4, ig. 10), we think that Natica noe is to be assigned to the genus Cryptonatica Dall, 1892. Cryptonatica noe closely resembles Natica microglossa Deshayes, 1864 (a species belonging in Cochlis) in all teleoconch characters, but can be differentiated from it by its protoconch of 0.5 fewer whorls, with a signiicantly greater diameter of the irst half-whorl. C. noe is also similar to Cochlis epiglottina epiglottina (Lamarck, 1804) in shell shape and has sometimes been mistaken for it. However, C. noe is distinguished from C. epiglottina epiglottina because of its protoconch with an inlated initial half-whorl having a signiicantly smaller diameter. Further, C. noe has an angular umbilical border (rounded in C. epiglottina epiglottina), a narrower funicle, thence a wider umbilical channel, and an umbilical callus demarcated from the parietal callus by a wider, shallow and reverse J-shaped notch (moderately deep and narrow in C. epiglottina epiglottina). Obviously, the operculum stands as the main character in distinguishing C. noe from both N. microglossa and C. epiglottina epiglottina. Stratigraphic occurrence. Cryptonatica noe (D’Orbigny, 1850) was hitherto known from Lutetian and Bartonian deposits of France and from the 145 Bartonian of Great Britain. Italian occurrences are from lower and middle Lutetian units of Veneto. Genus Tanea Marwick, 1931 Tanea Marwick, 1931, p. 98. Type species by original designation: Natica zelandica Quoy & Gaimard, 1832, Recent, New Zealand. Pedriali & Robba (2008) listed the following distinctive characters of Tanea: 1) spire moderately to greatly elevated, 2) anterior lobe of the parietal callus small, indistinct in a few species, 3) funicle thick, moderate in a few species, 4) operculum with 1 narrow groove bounding a broad, convex marginal area bearing a median longitudinal furrow. These authors noted that the operculum stands as the primary distinctive element of Tanea. The genus includes several fossil and Recent Indo-Paciic species. The Recent Mediterranean species Tanea dillwyni (Payraudeau, 1826) and the Burdigalian to Zanclean species Tanea dillwyni koeneni (Sacco, 1891) were so far known outside the Indo-Paciic area. The present study demonstrates that other Miocene, European species (Natica burdigalensis Mayer, 1864 and Natica beyrichi von Koenen, 1882), along with Burdigalian, Dutch specimens belong in Tanea; see also the above remarks on Cochlis tigrina (Defrance, 1825). Tanea dillwyni koeneni (Sacco, 1891) Pl. 4, igs. 14-18; Pl. 5, ig. 1 1882 Natica plicatella - von Koenen, p. 229, pl. 5, igs. 6, 7 (not ig. 9: operculum of Natica plicatula Bronn, 1831). 1891 Natica (Natica) epiglottina var. koeneni Sacco, p. 63. 1904 Natica (Cochlis) dillwyni var. taurominor Sacco, p. 103, pl. 22, igs. 43, 44. 1984 Natica dillwyni var. taurominor - Ferrero Mortara et al., p. 31. 2008a Tanea dillwyni koeneni - Pedriali & Robba, p. 104, pl. 1, igs. 9-11; pl. 2, ig. 16; pl. 3, igs. 8, 14 (cum syn.). 2013 Tanea koeneni - Harzhauser et al., p. 360, pl. 1, ig. 9. Type material: Natica (Natica) epiglottina var. koeneni Sacco, type material not seen. The holotype (in Geologisches Institut, Göttingen, Germany) was illustrated by Janssen (1969, pl. 7, ig. 22). Natica (Cochlis) dillwyni var. taurominor Sacco, lectotype (here designated): the shell igured by Sacco (1904, pl. 22, ig. 43) and reigured herein (Pl. 4, ig. 14), MGPT BS.029.01.045, Monte dei Cappuccini; 1 paralectotype, MGPT BS.029.01.046, 20 paralectotypes MGPT BS.029.01.046/01, Monte dei Cappuccini. Other material examined: Albugnano: 3 spms., PG 95; Borelli: 1 spm. (operculated shell), MGPT-PU 135094, 1 spm., MGPTPU 135095 (Pl. 4, ig. 15), 2 spms., MPUM 11263, 2 spms., NP 9947, 9 spms., MGPT-PU 135096, 1 spm., MGPT-PU 135097 (Pl. 4, ig. Robba E., Pedriali L. & Quaggiotto E. 146 16), 26 spms. (opercula), MGPT-PU 135098, 1 spm. (operculum), MGPT-PU 135099 (Pl. 4, ig. 17), 1 spm., PG 22; Monte dei Cappuccini: 1 spm., MGPT-PU 25670, 1 spm., MGPT-PU 135100 (Pl. 4, ig. 18), 31 spms., MGPT-PU 25671, 1 spm., MGPT-PU 135101, 1 spm., MGPT-PU 135102 , 1 spm., MGPT-PU 135103 (Pl. 5, ig. 1); Rio di Bocca d’Asino: 2 spms., NP 9946, 1 spm., PG 6a, 1 spm., PG 24; Sant’Agata Fossili: 1 spm. (operculated shell), MGPT-PU 135104, 3 spms., PG 23; Soprasalmo: 2 spms., private collection. Dimensions of Miocene specimens (mm) DHW PD H D SH AH AW 0.0680.076 1.2501.430 2.51016.914 2.85216.628 0.2803.660 1.97313.509 2.5859.313 5.949 0.072 1.340 9.712 9.740 1.970 7.741 UW WUC WAD WAB IS SA 0.9726.524 0.6962.956 0.0532.633 0.0071.151 9°-29° 107°131° 3.748 1.826 1.343 0.579 19° 119° Remarks. Pedriali & Robba (2008a) noted that the present taxon is closely similar to the Recent species Tanea dillwyni (Payraudeau, 1826) in shell shape, funicle, and opercular characters. They listed as differentiating elements the two-whorled protoconch (that of T. dillwyni has slightly more than three whorls) and the subsutural wrinkles (absent in T. dillwyni). Pedriali & Robba (2008a) concluded that Sacco’s taxon is to be considered as a subspecies of T. dillwyni. The values of the characteristic elements of the protoconch reported by Pedriali & Robba (2008a) were based on a single, partially abraded larval shell. The recovering of new, better preserved material has shown that the protoconch is of 3.25-3.47 whorls (see Pl. 5, ig. 1), thus identical to that of T. dillwyni. It appears that the presence of the subsutural wrinkles constitute the sole character distinguishing Sacco’s taxon from T. dillwyni. This evidence supports their separation, but only at the subspecies level. For the teleoconch characters and the relationships with Natica plicatula Bronn, 1831, see Pedriali & Robba (2008a). Stratigraphic occurrence. Tanea dillwyni koeneni (Sacco, 1891) is deinitely known from upper Burdigalian to Tortonian deposits of the North Sea Basin. Italian occurrences are from upper Langhian to Zanclean units. Pedriali & Robba (2008a) remarked that the subspecies seems to have not survived subsequent to the Zanclean. Tectonatica was introduced as a subgenus of Natica Adanson, 1757 (= Natica Scopoli, 1777), but subsequent workers regarded it as a distinct genus. According to Kabat (1991), Tectonica Carcelles & Williamson, 1951 and Tectonatic Maeda, 1988 are errors for Tectonatica. For the relationships between Tectonatica and Cryptonatica Dall, 1892, see the above remarks on Cryptonatica and also Pedriali & Robba (2008, pp. 98, 99). Besides the grooved calcareous operculum, the species of Tectonatica have mostly small-sized teleoconchs with the umbilicus completely or partly illed by the umbilical callus (in the last case, a broad funicle can be noted) and with the semicircular to subtriangular umbilical callus distinguished from the parietal callus. Except for Tectonatica tectula and Tectonatica astensis (Sacco, 1890), some of the species described below are assigned to Tectonatica only provisionally since the respective opercula are unknown. Future recovering of specimens with the operculum in situ may prove that they belong in Cryptonatica. Tectonatica albertii sp. n. Pl. 5, igs. 2, 3 Genus Tectonatica Sacco, 1890 Derivation of name: The species is named after Riccardo Alberti, who provided material relevant to this study. Holotype: Monte Gloso: MPUM 11264 (Pl. 5, ig. 2). Paratypes: Altavilla Vicentina: 5 spms., MGP-PD 1223R, 2 spms., MPUM 11265, 1 spm., MPUM 11266 (Pl. 5, ig. 3), 1 spm., MPUM 11267, 1 spm., MPUM 11268, 36 spms., MCZ 4322; Case Soghe: 2 spms., MPPLF 5123 pr (Accorsi Benini collection); Monte Gloso: 1 spm., MPUM 11269; Sangonini: 9 spms., MGP-PD 6418/a. Other material examined: Altavilla Vicentina: 2 spms., NP 9908, 6 spms., NP 9909, 1 spm., NP 9910, 1 spm., NP 9912, 1 spm., NP 9913, 1 spm., NP 9914; Monte Gloso: 1 spm., NP 9916, 2 spms., private collection. Preservation: Mostly fair. Type locality: Monte Gloso (see appendix). Horizon: Yellowish-gray, coarse-grained marly sandstone of Rupelian age. Diagnosis. Protoconch paucispiral, depressed-turbiniform. Shell globose, with low to moderately elevated spire and globose last whorl. Umbilicus deep of moderate breadth, comma-like. Parietal callus subrectangular, with very slight, angular anterior lobe; funicle broad and low; umbilical callus thick, with reverse S-shaped adaxial outline, merging into anterior lobe of parietal callus. Color pattern of blackish-brown subsutural band, lower base and collabral stripes over brown background. Tectonatica Sacco, 1890a, p. 205 (nomen nudum). Tectonatica Sacco, 1890b, p. 33. Type species by monotypy: Natica tectula Bonelli, 1826 (MS) = Natica (Tectonatica) tectula Sacco, 1890, Miocene to Pliocene, Italy (see also Kabat 1991). Description. Protoconch medium-sized, depressed-turbiniform, of 1.87-2 gently convex, smooth whorls, tip medium-sized. Teleoconch glo- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy bose, thick. Spire broadly conical, low to moderately elevated, whorls gently convex, latly convex in some specimens, suture adpressed to almost lush. Last whorl globose, scarcely extended toward aperture; subsutural shelf indistinct; periphery above midline. Aperture D-shaped, in prosocline plane, length about 1.60 times width; inner lip straight, outer lip semicircular, basal lip thickened. Parietal callus rather thick, subrectangular, with very slight, angular anterior lobe below border of umbilicus. Umbilicus deep, of variable but moderate breadth, comma-like, wider adapically and gradually narrowing downward. Funicle broad and low, separated from steep umbilical wall by narrow, shallow groove. Umbilical callus long, thick, located at mid-abapical part of inner lip, with reverse S-shaped adaxial outline, merging into anterior lobe of parietal callus. Basal fasciole indistinct. Outer surface with coarse growth markings changing into moderate subsutural wrinkles. Most specimens retain vestige of brown background, with blackish-brown subsutural band, lower base and collabral stripes. Dimensions (mm) DHW PD H D SH 0.3300.414 1.2191.399 5.54913.125 5.58712.707 1.1513.323 0.372 1.309 9.337 9.147 2.237 147 sutures (weakly channeled in N. burtoni) are other minor differences. Stratigraphic occurrence. Tectonatica albertii sp. n. was recovered from lower Oligocene deposits of Veneto. Tectonatica astensis (Sacco, 1890) Pl. 5, igs. 4-6 1890b Natica (Naticina) pulchella var. astensis Sacco, p. 31. 1891 Natica (Naticina) pulchella var. astensis - Sacco, p. 77, pl. 2, ig. 50. 2008a Tectonatica astensis - Pedriali & Robba, p. 106, pl. 1, ig. 14; pl. 2, igs. 1, 2, 17; pl. 3, igs. 1, 2, 15-17 (cum syn.). Type material: Pedriali & Robba (2008a, p. 107) reviewed the type material of Tectonatica astensis, and described the characters of this species. Material examined: Albugnano: 2 spms., PG 79; Borelli: 14 spms., MGPT-PU 135105, 1 spm., MGPT-PU 135106 (Pl. 5, ig. 4), 1 spm., MGPT-PU 135107, 1 spm., MGPT-PU 135108, 1 spm., MGPT-PU 135109, 1 spm., MGPT-PU 135110, 1 spm., MGPT-PU 135111, 1 spm., MGPT-PU 135112, 1 spm., MGPT-PU 135113, 1 spm., MGPT-PU 135114, 2 spms. (opercula), MGPT-PU 135115, 1 spm. (operculum), MGPT-PU 135116 (Pl. 5, ig. 5), 2 spms., MPUM 11270, 2 spms., NP 9907; Rio di Bocca d’Asino: 1 spm., NP 9889; Sant’Agata Fossili: 1 spm., PG 18b, 1 spm., PG 18c, (Pl. 5, ig. 6), 1 spm., PG 18d, 1 spm., PG 73a (operculated shell), 1 spm., NP 9890; Sogliano al Rubicone: 1 spm., PP.SR 01/02. Dimensions of Miocene specimens (mm) AH AW UW IS SA DHW PD H D SH 4.15010.050 3.1968.204 1.6024.490 8°-24° 114°-138° 0.1010.121 1.1491.401 2.6926.664 2.8466.430 0.7951.747 7.100 5.700 3.046 16° 126° 0.111 1.275 4.678 4.638 1.271 AH AW UW IS SA 1.8594.963 1.7684.104 0.9353.215 14°-22° 99°-131° 3.411 2.936 2.075 18° 115° Remarks. The present new species is assigned to the genus Tectonatica Sacco, 1890 on the basis of its umbilical characters, which are closely similar to those of Tectonatica astensis (Sacco, 1890) and Tectonatica prietoi (Hidalgo, 1873). Tectonatica albertii exhibits a striking resemblance to Tectonatica burtoni altavillensis subsp. n. (see below) in shell morphology and color pattern, but differs from it in that it has a larger protoconch with 0.75 fewer whorls and the diameter of the irst half-whorl three times the size. T. albertii differs from Natica burtoni Wrigley, 1949, a species herein assigned to the genus Tectonatica Sacco, 1890 (see the remarks on T. burtoni altavillensis), because of its protoconch with fewer whorls (1.87-2 instead of 2.55-2.75) and signiicantly greater diameter of the initial half-whorl (four times the size). The less convex spire whorls and the adpressed, almost lush Remarks. The specimens listed above fully conform to the Pliocene material dealt with by Pedriali & Robba (2008a). The 95% confidence intervals calculated for the dimension (protoconch and teleoconch) of the Miocene specimens overlap those obtained for the Pliocene shells; in particular, there are no significant differences as regards the values of the characteristic elements of the protoconch. For description and comments, see Pedriali & Robba (2008a). Stratigraphic occurrence. Tectonatica astensis (Sacco, 1890) was known to range from late Miocene (Tortonian) to late Pliocene. The uncommon occurrence in the Serravallian of Robba E., Pedriali L. & Quaggiotto E. 148 Albugnano reported herein pushes back the first known occurrence of the species. Tectonatica burtoni altavillensis subsp. n. Pl. 5, igs. 7, 8 Derivation of name: The name refers to Altavilla Vicentina, which is the type locality. Holotype: Altavilla Vicentina, MPUM 11271 (Pl. 5, ig. 7). Paratypes: Altavilla Vicentina: 1 spm., MPUM 11272 (Pl. 5, ig. 8), 1 spm., MPUM 11273, 9 spms., MGP-PD 1224R; Le Coe: 1 spm., MPUM 11274; Sangonini: 3 spms., MCZ 4323. Other material examined: Altavilla Vicentina: 1 spm., NP 9900, 1 spm., NP 9901, 1 spm., NP 9902, 2 spms., NP 9942, 5 spms., NP 9903; Le Coe: 1 spm., NP 9905, 3 spms., NP 9906, 2 spms., NP 9907; Monte Gloso: 2 spms., private collection, 1 spm., NP 9904. Preservation: Rather fair on the whole. Type locality: Altavilla Vicentina (see appendix). Horizon: Gray-brown marl of Rupelian age. Diagnosis: Protoconch multispiral, depressed-turbiniform. Shell globose, with low to moderately elevated spire and globose last whorl. Umbilicus deep of moderate breadth, comma-like. Parietal callus subrectangular, with very slight, angular anterior lobe; funicle broad and low; umbilical callus thick, with reverse S-shaped adaxial outline, merging into anterior lobe of parietal callus. Color pattern of blackish-brown subsutural band, lower base and collabral stripes over brown background. Description. Protoconch medium-sized, depressed-turbiniform, of 2.75-2.80 gently convex, smooth whorls, tip small. Teleoconch globose, thick. Spire broadly conical, low to slightly elevated, whorls gently convex, latly convex in some specimens, suture adpressed to almost lush. Last whorl globose, scarcely extended toward aperture; subsutural shelf indistinct; periphery above midline. Aperture D-shaped, in prosocline plane, length about 1.60 times width; inner lip straight, outer lip semicircular, basal lip thickened. Parietal callus rather thin to thick, subrectangular, anterior lobe angular, very slight or indistinct. Umbilicus deep, of variable but moderate breadth, comma-like, wider adapically and gradually narrowing downward. Funicle broad, low to markedly depressed, separated from steep umbilical wall by narrow, shallow groove. Umbilical callus long, thick, located at mid-abapical part of inner lip, with attenuated reverse S-shaped adaxial outline, merging into anterior lobe of parietal callus. Basal fasciole indistinct. Outer surface with coarse growth markings changing into slight wrinkles subsuturally. Most specimens retain vestige of brown background, with blackish-brown subsutural band, lower base and undulating collabral stripes. Dimensions (mm) DHW PD H D SH 0.0940.118 0.9341.126 4.96213.502 5.10913.509 1.1134.041 0.106 1.030 9.232 9.309 2.577 AH AW UW IS SA 3.6649.648 3.2338.453 1.6064.294 14°-26° 112°-136° 6.656 5.843 2.950 20° 124° Remarks. We obtained outstanding photographs of specimens in NHM originally grouped by Wrigley (1949) into his new species Natica burtoni. On the basis of the photographs and of the examination of topotypes from the Middle Barton Beds at Barton and from the Upper Bracklesham Beds in our collection, we can state that these specimens are a mixed assortment of species. Since Wrigley (1949, p. 13, ig. 8) designated the holotype (NHM G. 67389), this specimen is to be considered the valid reference for N. burtoni. The holotype of N. burtoni has umbilical characters that agree with those PLATE 5 Fig. 1 - Tanea dillwyni koeneni (Sacco, 1891). Monte dei Cappuccini. MGPT-PU 135103; protoconch. Fig. 2 - Tectonatica albertii sp. n. Monte Gloso. Holotype, MPUM 11264; a, apertural side (height of shell 9.94 mm); b, abapertural side. Fig. 3 - Tectonatica albertii sp. n. Altavilla Vicentina. Paratype, MPUM 11266; protoconch. Fig. 4 - Tectonatica astensis (Sacco, 1890). Borelli. MGPT-PU 135106; apertural side. Fig. 5 - Tectonatica astensis (Sacco, 1890). Borelli. MGPT-PU 135116; operculum. Fig. 6 - Tectonatica astensis (Sacco, 1890). Sant’Agata Fossili. PG 18c; protoconch. Fig. 7 - Tectonatica burtoni altavillensis subsp. n. Altavilla Vicentina. Holotype, MPUM 11271; apertural side (height of shell 12.06 mm). Fig. 8 - Tectonatica burtoni altavillensis subsp. n. Altavilla Vicentina. Paratype, MPUM 11272; protoconch. Fig. 9 - Tectonatica consimilis sp. n. Borelli. Holotype, MGPT-PU 135117; a, apertural side (height of shell 6.62 mm); b, protoconch. Fig. 10 - Tectonatica consimilis sp. n. Borelli. Paratype, MGPT-PU 135120; protoconch. Fig. 11 - Tectonatica dertomamilla (Sacco, 1890). Stazzano. Holotype of Natica (Polinices) dertomamilla Sacco, 1890. MGPT BS.029.06.009; apertural side. Fig. 12 - Tectonatica dertomamilla (Sacco, 1890). Montegibbio. MPUM 11276; a, apertural side (height of shell 18.69 mm); b, protoconch. Fig. 13 - Tectonatica dertomamilla (Sacco, 1890). Rio di Bocca d’Asino. Private collection; apertural side (height of shell 19.12 mm). Fig. 14 - Tectonatica miocolligens (Sacco, 1890). Colli Torinesi. Holotype of Natica (Polinices) miocolligens Sacco, 1890. MGPT BS.029.06.007; apertural side. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 149 Robba E., Pedriali L. & Quaggiotto E. 150 of several species of Tectonatica Sacco, 1890; consequently, Wrigley’s species is herein assigned to this genus. Compared to the holotype of the Eocene species Tectonatica burtoni, the present specimens from the early Oligocene of Altavilla Vicentina appear to have the values of the characteristic elements of the protoconch identical to those measured for Wrigley’s species. They differ from T. burtoni only in having gently to latly convex spire whorls separated by adpressed to almost lush sutures (T. burtoni has well convex whorls and very slightly channeled sutures). These differences are not considered suficient for consistent separation at the species level. Consequently, we regard the studied specimens as representatives of a new allochronous subspecies of T. burtoni and name it altavillensis. Stratigraphic occurrence. Tectonatica burtoni altavillensis subsp. n. was recovered from upper Priabonian and Rupelian units of Veneto. Tectonatica consimilis sp. n. Pl. 5, igs. 9, 10 Derivation of name: From Latin consimilis = closely similar, with reference to the remarkable similarity to Tectonatica tectula (Sacco, 1890). Holotype: Borelli: MGPT-PU 135117 (Pl. 5, ig. 9). Paratypes: Borelli: 1 spm., MGPT-PU 135120 (Pl. 5, ig. 10), 1 spm., MGPT-PU 135121, 29 spms., MGPT-PU 135122, 1 spm., MPUM 11275. Other material esamined: Borelli: 1 spm., NP 9881 Material erroneously referred to as Natica (Polinices) redempta Michelotti, 1847 in MGPT: Borelli: 1 spm., MGPT BS.029.06.014/02. Preservation: Fair on the whole. Type locality: Borelli (see appendix). Horizon: Gray, medium sand of Tortonian age. Diagnosis: Globose shell with depressed to moderately elevated spire and inlated last whorl. Umbilicus completely plugged or faint oblique chink in a few specimens. Parietal callus thick, subquadrangular, expanded adapically; umbilical callus thick, large, semicircular, merging into parietal callus. Basal fasciole broad, depressed. Color pattern of pale brown spots arranged into spiral rows. Description. Protoconch small, lowturbiniform, of 1.75 convex, smooth whorls, tip small. Teleoconch globose, rather thin. Spire broadly conical, depressed to moderately elevated, whorls gently convex, suture very slightly channeled. Last whorl inlated, moderately extended toward aperture; subsutural shelf obscure to indistinct; periphery above midline. Aperture D-shaped, in prosocline plane, length averaging 1.67 times width; inner lip straight, outer lip semicircular. Parietal callus rather thick, subquadrangular, expanded adapically, with concave abapertural outline. Umbilicus completely plugged by umbilical callus; faint oblique chink present in a few specimens. Umbilical callus large, thick, semicircular in outline, overlapping mid-adapical part of basal fasciole, merging into parietal callus (abapertural outline changing from convex to concave). Basal fasciole broad, markedly depressed. Outer surface with dense growth markings prosocyrt adapically. Some specimens retain vestige of pale brown quadrangular spots, arranged into spiral rows. Dimensions (mm) DHW PD H D SH 0.1800.216 0.6930.809 2.6927.016 2.5657.053 0.3521.728 0.198 0.751 4.854 4.809 1.040 AH AW UW = WUC IS SA 2.2545.374 1.4485.308 1.0082.924 17°-33° 116°-140° 3.814 3.378 1.966 25° 128° Remarks. Tectonatica consimilis is morphologically similar to Tectonatica tectula (Sacco, 1890), but differs from it in that it has: 1) a protoconch of one fewer whorl, with a signiicantly smaller diameter and with the diameter of the initial half-whorl two times the size, 2) a less elevated spire (Fig. 10), 3) a slightly channeled suture (adpressed in T. tectula), 4) a distinct basal fasciole (absent in T. tectula), and 5) a different color pattern. The values of the characteristic elements of the protoconch stand as the most relevant distinguishing characters. The present new species is provisionally assigned to Tectonatica Sacco, 1890 because of its very close similarity to T. tectula, type species of the genus. As already noted (see the remarks on Tectonatica), new indings may prove that it belongs in Cryptonatica Dall, 1892. Stratigraphic occurrence. Tectonatica consimilis sp. n. was recovered from Tortonian deposits at the type locality. Tectonatica dertomamilla (Sacco, 1890) comb. n. Pl. 5, igs 11-13 1890b Natica (Polinices) dertomamilla Sacco, p. 36. 1891 Natica (Polinices) dertomamilla - Sacco, p. 93, pl. 2, ig. 70. 1969 Polinices dertomamilla - Janssen, p. 161, pl. 4, ig. 2. 1984 Polinices dertomamilla - Ferrero Mortara et al., p. 37, pl. 3, ig. 12. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Type material: Natica (Polinices) dertomamilla Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 2, ig. 70) and reigured herein (Pl. 5, ig. 11), MGPT BS.029.06.009, Stazzano. Material erroneously referred to as Natica submamillaris var. mioclausa Sacco, 1890 in IPUM: Montegibbio: 2 spms., IPUM 4395. Material erroneously referred to as Natica pardalis Doderlein, 1864 in IPUM: Montegibbio: 3 spms., IPUM 4396. Other material examined: Montegibbio: 1 spm., MPUM 11276 (Pl. 5, ig. 12), 1 spm., MZB 60112; Rio di Bocca d’Asino: 1 spm., private collection (Pl. 5, ig. 13), 1 spm., private collection; Stazzano: 1 spm., MZB 60110, 1 spm., NP 9880. Description. Protoconch small, lowturbiniform, of two gently convex, apparently smooth whorls, tip small. Teleoconch globose, globose-oval in some specimens, moderately thick. Spire broadly conical, pointed, rather elevated, whorls nearly lat-sided. Suture thin, almost lush, adpressed in one specimen, which has gently convex whorls. Last whorl globose to globose-oval, scarcely extended toward aperture; subsutural shelf indistinct; periphery about at midline. Aperture D-shaped in prosocline plane, length averaging 1.84 times width; inner lip straight, outer lip arched, with maximum convexity at abapical one-third. Parietal callus long, thick, with concave abapertural outline, ending at transition between umbilical border and steep umbilical wall; anterior lobe small to indistinct. Umbilicus largely plugged by parietal and umbilical calluses, reduced to narrow, markedly oblique mid-abapical groove exposing thick, broad funicle. Umbilical callus thick, rather short, triangular, with reverse J-shaped adaxial outline, merging into anterior lobe (or corner) of parietal callus. Basal fasciole broad, markedly depressed. Outer surface with ine growth lines prosocyrt adapically. Some specimens retain vestige of uniform pale brown background, with dark brown subsutural band. Dimensions (mm) DHW PD H D SH 0.1650.189 0.8830.995 15.99226.032 13.56025.312 0.9307.818 0.177 0.939 21.012 19.436 4.374 AH AW UW IS SA 12.55820.718 7.23017.358 2.308-7.804 7°-19° 103°-119° 16.638 12.294 5.056 13° 111° Remarks. The present species was assigned by the authors to Polinices Montfort, 1810. We think that the narrow, adaxial, markedly oblique umbilical 151 groove and the umbilical callus well distinct from the parietal callus, as commonly noted in species of Tectonatica Sacco, 1890, e.g. Tectonatica astensis (Sacco, 1890), account for an assignment to Tectonatica rather than to Polinices (in the latter genus the umbilical callus is fused with the parietal callus and is hardly distinguishable from it). As regards the relationships with Tectonatica miocolligens (Sacco, 1890), the most closely related species, see below (remarks on T. miocolligens). Stratigraphic occurrence. Tectonatica dertomamilla (Sacco, 1890) occurs uncommonly only in Tortonian deposits of Piedmont and Emilia. Tectonatica miocolligens (Sacco, 1890) comb. n. Pl. 5, ig. 14; Pl. 6, igs. 1-6 1890b Natica (Polinices) submamillaris var. praenuntia Sacco, p. 35 (new synonym). 1890b Natica (Polinices) miocolligens Sacco, p. 36. 1890b Natica (Polinices) miocolligens var. pseudomamilla Sacco, p. 36. 1891 Natica (Polinices) submamillaris var. praenuntia - Sacco, p. 90, pl. 2, ig. 63. 1891 Natica (Polinices) miocolligens - Sacco, p. 93, pl. 2, ig. 68. 1891 Natica (Polinices) miocolligens var. pseudomamilla - Sacco, p. 93, pl. 2, ig. 69. 1919 Natica (Polinices) dertomamilla - Cossmann & Peyrot, p. 219, pl. 12, igs. 16-18 (not Natica dertomamilla Sacco, 1890). 1952a Polynices (Polynices) submamillaris f. dertomamilla - Glibert, p. 73, pl. 5, ig. 12 (not Natica dertomamilla Sacco, 1890). 1969 Polinices (Polinices) miocolligens - Janssen, p. 161 (pars), pl. 4, igs. 1, 3, 9 (not igs. 4-8). 1984 Polinices submamillaris var. praenuntia - Ferrero Mortara et al., p. 36. 1984 Polinices miocolligens - Ferrero Mortara et al., p. 36, pl. 4, ig. 4. 1984 Polinices miocolligens var. pseudomamilla - Ferrero Mortara et al., p. 36. not 1984 Polinices (Polinices) miocolligens - Janssen, p. 199, pl. 56, igs. 1, 2 (not Sacco, 1890). ? 2001 Polinices (Polinices) miocolligens - Wienrich, p. 430, pl. 69, ig. 4; pl. 86, ig. 9. 2007 Polinices miocolligens - Zunino, p. 120, pl. 1, ig. 6. Type material: Natica (Polinices) miocolligens Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 2, ig. 68) and reigured herein (Pl. 5, ig. 14), MGPT BS.029.06.007, Colli Torinesi. Natica (Polinices) miocolligens var. pseudomamilla Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 2, ig. 69) and reigured herein (Pl. 6, ig. 1), MGPT BS.029.06.008, Colli Torinesi. Natica (Polinices) submamillaris var. praenuntia Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 2, ig. 63) and reigured herein (Pl. 6, ig. 2), MGPT BS.029.06.003, Cassinelle; 1 paralectotype, MGPT BS.029.06.003/01, Cassinelle. Other material examined: Albugnano: 1 spm., PG 48; Case Soghe: 1 spm., MPUM 11277, 1 spm., NP 9878, 2 spms., private collection; Mioglia: 2 spms., MZB 60099; Monte dei Cappuccini: 3 Robba E., Pedriali L. & Quaggiotto E. 152 spms., MGPT-PU 25662; Sangonini: 1 spm., MGP-PD 15948/a, 1 spm., MGP-PD 15924 (Pl. 6, ig. 3), 1 spm., MGP-PD 31536; Valle Ceppi: 1 spm., NP 9879, 1 spm., MGPT-PU 25269, 1 spm., MGPTPU 23858, 2 spms., MGPT-PU 135123, 1 spm., MGPT-PU 25902, 1 spm., MGPT-PU 135124 (Pl. 6, ig. 4), 1 spm., PG 47, 1 spm., MPUM 11278, 2 spms., MZB 60029, 1 spm., MZB 60150, 1 spm., MGPT-PU 135125, 1 spm., MPUM 11279, 1 spm., NP 9828; Val Sanfrà: 1 spm., MGPT-PU 135126, 8 spms., MGPT-PU 107626, 1 spm., MGPT-PU 135127 (Pl. 6, ig. 5), 2 spms., MGPT-PU 23663, 1 spm., MGPT-PU 107629, 4 spms., MGPT-PU 135128; Valle Vergnana: 1 spm., MZB 60028 (Pl. 6, ig. 6), 2 spms., PG 46. Description. Protoconch small, lowturbiniform, of 2.65-2.70 gently convex, apparently smooth whorls, tip small. Teleoconch globose, globose-pyriform in some specimens, rather thick. Spire broadly conical, pointed, moderately elevated, slightly more so in a few specimens, whorls lat-sided. Suture thin, almost lush, adpressed in a few specimens, which have gently convex whorls. Last whorl globose to globose-oval, only slightly extended toward aperture; subsutural shelf indistinct, very slightly concave but poorly deined in a few larger specimens; periphery above midline. Aperture D-shaped in prosocline plane, length averaging 1.60 times width; inner lip straight, outer lip semicircular. Parietal callus thick, subrectangular, ending in slight anterior lobe below border of umbilical area. Umbilicus largely plugged by umbilical callus; narrow, markedly oblique mid-abapical groove separates umbilical callus from umbilical wall and exposes thick funicle. Umbilical callus at abapical half of inner lip, thick, subtriangular, with arched adaxial outline, demarcated from anterior lobe of parietal callus by very slight sinus. Basal fasciole indistinct. Outer surface with dense growth lines prosocyrt adapically. Dimensions (mm) DHW PD H D SH 0.1070.115 0.9250.981 8.77023.626 7.67821.210 0.3465.322 0.111 0.953 16.198 14.444 2.834 AH AW UW IS SA 6.47119.591 4.59112.535 2.8865.882 9°-33° 85°-113° 13.031 8.563 4.384 21° 99° Remarks. The present species was regarded as belonging in Polinices Montfort, 1810 by previous authors. Actually, it has the characters of Tectonatica Sacco, 1890 and is herein assigned to this genus (see also the remarks on Tectonatica dertomamilla Sacco, 1890). Tectonatica miocolligens is morphologically sim- ilar to Tectonatica dertomamilla, but differs from it in that it has: 1) a protoconch with 0.5 more whorls and with signiicantly smaller diameter of the irst half-whorl, 2) a wider aperture with different curvature of the outer lip, 3) a shorter parietal callus, 4) a longer umbilical callus with arched adaxial outline, demarcated from parietal callus by a slight sinus (the umbilical callus is triangular, with reverse J-shaped outline in T. dertomamilla), and 5) an indistinct basal fasciole (present, broad and low in T. dertomamilla). Janssen (1969) included in the synonymy of Polinices miocolligens the specimens dealt with by Glibert (1952a) and referred to as Polynices (Polynices) submamillaris f. dertomamilla Sacco, 1891. We have not seen Glibert’s material, however, considering: 1) that the shell igured by Glibert appears to agree with the characters of Tectonatica miocolligens, and 2) that the age cited by Glibert for his specimens is late Burdigalian to Langhian, we retain the Belgian material in the synonymy of the present species. Concerning Sacco’s variety pseudomamilla of the present species (see the above synonymy), we examined the holotype in MGPT and can state that it cannot be distinguished from T. miocolligens consistently. The variety praenuntia of Natica (Polinices) submamillaris Sacco,1890 was proposed by Sacco (1890b) on the basis of scarce material from the early Oligocene of Piedmont. Wheras the typical form belongs in Euspira Agasiz in J. Sowerby, 1837 (see the species of Euspira described below), the variety praenuntia exhibits the characters of T. miocolligens and is herein included in the synonymy of it. The lower Miocene specimens recorded by Wienrich (2001) and referred to as Polinices (Polinices) miocolligens Sacco, 1891 may be this species in that they have a 2.5-whorled protoconch, a close teleoconch shape and a rather similar umbilical callus (the umbilicus slightly more widely open than in the typical specimens of Piedmont being the sole difference). Since the values of the other characteristic elements of the protoconch (diameter and diameter of the irst half-whorl) are not known, we include these shells in the synonymy of the present species only doubtfully. The specimens igured by Janssen (1984) and referred to as Polinices (Polinices) miocolligens Sacco, 1891 have umbilical characters that are unlike those of Sacco’s species and are herein excluded from the synonymy Eocene, Oligocene and Miocene naticid gastropods of Northern Italy of Tectonatica miocolligens; likely, these Dutch shells belong to a species of the genus Cochlis Röding, 1798. Stratigraphic occurrence. Tectonatica miocolligens (Sacco, 1890) is known from the early Oligocene of Piedmont, Liguria and Veneto (Natica submamillaris var. praenuntia Sacco) and from the Burdigalian and the Langhian of Piedmont; it occurs also in the late Aquitanian to Langhian of Belgium. The species was recovered at Winterswijk-Miste in The Netherlands (Janssen 1969) from deposits of late Burdigalian age (Kowalewski et al. 2002). Cossmann & Peyrot (1919) described it as Natica dertomamilla (see the above synonymy) from deposits in the Saubrigues area said to be of Tortonian age, but actually of Langhian age (Cahuzac & Poignant 1993; Cahuzac et al. 1995). We recovered only one specimen from Serravallian deposits of Albugnano. In summary, T. miocolligens ranges from early Oligocene to Serravallian, being extremely rare in the Serravallian. Tectonatica pasinii (Bayan, 1870) Pl. 6, igs. 7-9 ig. 11. 1870a Natica pasinii Bayan, p. 481. 1870b Natica pasinii - Bayan, p. 23, pl. 3, ig. 6. 1896 Natica pasinii var. zagaropsis De Gregorio, p. 56, pl. 5, 1901 Natica pasinii - Oppenheim, p. 259. 1953 Polynices (Naticina) pasinii - Szőts, p. 178, pl. 5, igs. 12, 13. 1985 Polinices pasinii - Brigantini, p. 413, pl. 2, igs. 28, 29. 2008 Polinices pasinii - Quaggiotto & Mellini, p. 48, pl. 3, ig. 12. Type material: Natica pasinii Bayan, type material not seen. Bayan’s material is curated at the Université de Lyon 1, France, but the syntypes of Natica pasinii were not identiied yet and are presently unavailable (Robert Emmanuel, personal communication 2013). Material examined: Roncà: 2 spms., NP 9917, 5 spms., NP 9911, 8 spms., NP 9915, 2 spms., NP 9918, 5 spms., MPUM 11280, 1 spm., MPUM 11281 (Pl. 6, ig. 7), 1 spm., MPUM 11282 (Pl. 6, ig. 8), 1 spm., MPUM 11283 (Pl. 6, ig. 9), 15 spms., MCZ 4324. Description. Protoconch small, depressedturbiniform, of 2.25 gently convex, apparently smooth whorls, tip small. Teleoconch globose-oval to elongate-oval, moderately thick. Spire cyrtoconoid, rather elevated, less so in a few specimens, whorls latly convex, suture almost lush. Last whorl globose to oval, scarcely extended toward aperture; subsutural shelf indistinct; periphery at midline. Aperture D-shaped, in prosocline plane, length averaging twice width; inner lip straight, outer lip 153 arched, basal lip slightly thickened. Parietal callus thick, subrectangular, with obtuse adapical ridge, roundly expanded abapically, rather long to very long, hindering umbilicus at variable extent. Umbilicus rather narrow and deep to very narrow. Funicle broad and thick, separated from umbilical wall by narrow groove. Umbilical callus subtriangular, short to very short, thick, restricted to abapical part of inner lip in most specimens, with straight or attenuated reverse S-shaped adaxial outline, merging into anterior expansion of parietal callus, demarcated from parietal callus by shallow transverse depression in a few specimens. Basal fasciole indistinct. Outer surface with rather coarse growth markings. A few specimens retain vestige of uniform brown background. Dimensions (mm) DHW PD H D SH 0.141* 0.672* 9.38016.992 6.39916.935 1.6555.503 * 1 protoconch measurable 13.186 11.667 3.579 AH AW UW IS SA 7.41911.795 5.146-7.486 2.3243.792 18°-30° 88°-116° 9.607 6.316 3.058 24° 102° Remarks. The present species is rather variable as regards the teleoconch shape, the length of the parietal callus and the size of the umbilical opening. The elongate-oval shells (var. zagaropsis De Gregorio, 1896) do not warrant separation from the typical form with which they co-occur in the type locality (Roncà). Natica pasinii was currently included in the genus Polinices Montfort, 1810 by later authors. However, the umbilical characters of Bayan’s species do not agree with those of Polinices species, whereas they appear to be more similar to the characters occurring in the species belonging in Tectonatica Sacco, 1890. Thus, we concur with Cossmann (1925, p. 121), who assigned Natica pasinii to Tectonatica. Tectonatica pasinii is readily distinguished from the other Italian Paleogene taxa Tectonatica albertii sp. n. and Tectonatica burtoni altavillensis subsp. n. in that it has a more elongated teleoconch (Fig. 9) with higher spire, a longer parietal callus, and a smaller umbilicus. Moreover, it is differentiated from T. albertii by its protoconch with signiicantly smaller diameter and diameter of the initial half-whorl (less than half the size), and from T. burtoni altavillensis because of its protoconch with 0.5 fewer whorls, with signiicantly Robba E., Pedriali L. & Quaggiotto E. 154 smaller diameter. Stratigraphic occurrence. Tectonatica pasinii (Bayan, 1870) is known deinitely from middle Eocene (Bartonian) deposits of Veneto. It occurs also in the early and middle Eocene of Hungary. Tectonatica rupeliana sp. n. Pl. 6, igs. 10-12 Derivation of name: The name refers to the Rupelian age of the unit that yielded the bulk of the material. Holotype: Case Soghe: MPUM 11284 (Pl. 6, ig. 10). Paratypes: Case Soghe: 2 spms., MPUM 11285, 1 spm., MPUM 11286 (Pl. 6, ig. 11), 1 spm., MPUM 11287, 3 spms., MPUM 11288, 1 spm., MPPLF 5157 pr, 1 spm., MCZ 4325; Sangonini: 1 spm., MGP-PD 31537. Other material examined: Cà Sella: 1 spm., NP 9891; Case Soghe: 3 spms., NP 9892, 3 spms., NP 9893, 2 spms., NP 9895, 2 spms., private collection, 1 spm., NP 9897, 1 spm., private collection (Pl. 6, ig. 12), 1 spm., private collection, 7 spms., private collection, 2 spms., NP 9898; Chiuppano: 1 spm., NP 9899; Lerma: 1 spm., PG 80; Monte Gloso: 2 spms., private collection, 1 spm., NP 9896. Preservation: Most specimens are rather well preserved. Type locality: Case Soghe (see appendix). Horizon: Gray to yellowish-brown clayey silt with small pebbles of Rupelian age. Diagnosis: Globose-oval shell with depressed, broadly conical spire and oval, very tall last whorl. Umbilicus a narrow, oblique crescentic chink, more widely open in a few specimens. Parietal callus thick, subquadrangular, wider abapically; umbilicus largely illed by thick funicle; umbilical callus large, rather thick, with convex adaxial outline, merging into anterior corner of parietal callus. Basal fasciole indistinct. Description. Protoconch small, lowturbiniform, of 3.15 convex, smooth whorls, tip very small. Teleoconch globose-oval, slightly higher than wide, rather thick. Spire low, broadly conical, whorls latly convex, suture almost lush. Last whorl oval, very tall (about nine-tenths of total height), inlated; subsutural shelf indistinct; periphery slightly above midline. Aperture D-shaped, in prosocline plane, length averaging two times width; outer lip asymmetrically arched; inner lip very slightly bent and thick abapically. Parietal callus thick to very thick, subquadrangular, expanded abapically, with concave abapertural outline; anterior lobe poorly deined. Umbilicus largely illed by thick funicle separated from umbilical wall by narrow, oblique crescentic groove, more widely open in a few specimens. Umbilical callus large, rather thick, with convex adaxial outline, merging into anterior corner of parietal callus, narrover, subtriangular in specimens with more widely open umbilicus. Basal fasciole in- distinct. Outer surface with dense growth markings prosocyrt adapically. Dimensions (mm) DHW PD H D SH 0.079* 0.943* 6.57118.815 2.99218.588 1.0695.629 *1 protoconch measurable 12.693 10.790 3.349 AH AW UW IS SA 5.39013.298 2.93410.838 1.8995.803 14°-26° 87°-111° 9.344 6.886 3.851 20° 99° Remarks. Tectonatica rupeliana is closely similar to Tectonatica pasinii (Bayan, 1870) in shell shape. However, it can be readily distinguished from T. pasinii because of its protoconch with about one more whorl and signiicantly greater diameter and smaller diameter of the initial half-whorl, its shorter parietal PLATE 6 Fig. 1 - Tectonatica miocolligens (Sacco, 1890). Colli Torinesi. Holotype of Natica (Polinices) miocolligens var. pseudomamilla Sacco, 1890. MGPT BS.029.06.008; apertural side. Fig. 2 - Tectonatica miocolligens (Sacco, 1890). Cassinelle. Lectotype (here designated) of Natica (Polinices) submamillaris var. praenuntia Sacco, 1890. MGPT BS.029.06.003; apertural side. Fig. 3 - Tectonatica miocolligens (Sacco, 1890). Sangonini. MGP-PD 15924; apertural side (height of shell 20.16 mm). Fig. 4 - Tectonatica miocolligens (Sacco, 1890). Valle Ceppi. MGPT-PU 135124; protoconch. Fig. 5 - Tectonatica miocolligens (Sacco, 1890). Val Sanfrà. MGPT-PU 135127; apertural side (height of shell 15.04 mm). Fig. 6 - Tectonatica miocolligens (Sacco, 1890). Valle Vergnana. MZB 60028; protoconch. Fig. 7 - Tectonatica pasinii (Bayan, 1870). Roncà. MPUM 11281; protoconch. Fig. 8 - Tectonatica pasinii (Bayan, 1870). Roncà. MPUM 11282; apertural side (height of shell 10.79 mm). Fig. 9 - Tectonatica pasinii (Bayan, 1870). Roncà. MPUM 11283; apertural side (height of shell 11.35 mm). Fig. 10 - Tectonatica rupeliana sp. n. Case Soghe. Holotype, MPUM 11284; apertural side (height of shell 11.77 mm). Fig. 11 - Tectonatica rupeliana sp. n. Case Soghe. Paratype, MPUM 11286; protoconch. Fig. 12 - Tectonatica rupeliana sp. n. Case Soghe. Private collection; apertural side (height of shell 17.41 mm). Fig. 13 - Tectonatica tectula (Sacco, 1890). Borelli. MGPT-PU 135129; protoconch. Fig. 14 - Tectonatica tectula (Sacco, 1890). Borelli. MGPT-PU 135134; apertural side (height of shell 4.71 mm). Fig. 15 - Tectonatica tectula (Sacco, 1890). Borelli. MGPT-PU 135137; operculum. Fig. 16 - Tectonatica sp. 1. Cava Albanello. MGP-PD 1225R; a, apertural side (height of shell 8.49 mm); b, basal view; c, protoconch. Fig. 17 - Tectonatica sp. 2. Cava Grola. MGP-PD 1227R; apertural side (height of shell 14.57 mm). Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 155 Robba E., Pedriali L. & Quaggiotto E. 156 callus devoid of adapical ridge (present in T. pasinii), and its larger, differently shaped umbilical callus (that of T. pasinii is subtriangular instead of semicircular). T. rupeliana differs from the coeval Tectonatica albertii sp. n. (see above) by its protoconch with one more whorl and signiicantly smaller diameter and diameter of the irst half-whorl, its slenderer, ovate teleoconch, and its narrower umbilicus. Finally, the present species is differentiated from the also coeval Tectonatica burtoni altavillensis subsp. n. in that it has a protoconch with signiicantly smaller diameter of the irst half-whorl, a less globose teleoconch, a narrower umbilicus, and a different outline of the umbilical callus. Stratigraphic occurrence. Tectonatica rupeliana sp. n. was recovered only from Rupelian deposits of Veneto and Piedmont. intervals calculated for the dimensions (protoconch and teleoconch) of the Miocene specimens overlap those pertaining to the Pliocene shells; in particular, there is no signiicant difference as regards the values of the characteristic elements of the protoconch. For description and comments, see Pedriali & Robba (2008a). Stratigraphic occurrence. Combined reliable records from European countries and Italy (Pedriali & Robba 2008a) show that Tectonatica tectula (Sacco, 1890) ranges from the Burdigalian to the late Pliocene; the species is also present in the Serravallian of Turkey. According to Pedriali & Robba (2008a), the rare citations from the Pleistocene probably refer to Cryptonatica ilosa (Philippi,1845). Tectonatica sp. 1 Tectonatica tectula (Sacco, 1890) Pl. 6, ig. 16 Pl. 6, igs. 13-15 1890b Natica (Tectonatica) tectula Sacco, p.33. 1891 Natica (Tectonatica) tectula - Sacco, p. 81, pl. 2, ig. 53. 2008a Tectonatica tectula - Pedriali & Robba, p. 110, pl. 2, igs. 7-9, 19, 20; pl. 3, igs. 9, 22, 23 (cum syn.). 2013 Tectonatica tectula - Landau et al., p. 102, pl. 11, ig. 6. Type material: Pedriali & Robba (2008a, p. 111) listed the type material of Tectonatica tectula. Material examined: Borelli: 1 spm., MGPT-PU 135129 (Pl. 6, ig. 13), 1 spm., MGPT-PU 135130, 1 spm., MGPT-PU 135131, 1 spm., MGPT-PU 135132, 9 spms., MGPT-PU 135133, 1 spm., MGPT-PU 135134 (Pl. 6, ig. 14), 9 spms., MGPT-PU 135135, 6 spms., MPUM 11289, 3 spms. (opercula), MGPT-PU 135136, 1 spm. (operculum), MGPT-PU 135137 (Pl. 6, ig. 15), 5 spms., PG 22d, 1 spm., NP 9843, 3 spms., NP 9906; Orthez, Le Paren (France): 1 spm., NP 10011; Passo dei Meloni: 1 spm., PP.PM 01/01; Rio di Bocca d’Asino: 2 spms., NP 9887; Sogliano al Rubicone: 4 spms., PP.SR 01/01; Valle Ceppi: 5 spms., NP 9888, 3 spms., MPUM 11290, 4 spms., MGPT-PU 135138, 1 spm., MGPT-PU 135139. Dimensions of Miocene specimens (mm) DHW PD H D SH 0.0800.100 1.0191.195 3.3045.956 3.2895.453 0.4352.059 Material examined: Cava Albanello: 1 spm., MGP-PD 1225R (Pl. 6, ig. 16), 1 spm., MGP-PD 1226R, 1 spm., MCZ 4326. Description. Protoconch medium-sized, lowturbiniform, of two whorls, tip medium-sized, slightly sunken. Teleoconch globose, rather thin. Spire moderately elevated, broadly conical, whorls convex, suture very slightly channeled. Last whorl globular; subsutural shelf indistinct; periphery above midline. Aperture D-shaped, in slightly prosocline plane, length about 1.6 times width; outer lip semicircular; inner lip straight. Parietal callus thin, rectangular, wider adapically, with straight abapertural outline; anterior lobe indistinct. Umbilicus rather wide, deep. Funicle broad, markedly depressed, obsolescent toward interior of umbilicus. Umbilical callus thick, narrowly elongate, with gently convex adaxial outline, merging into anterior end of parietal callus. Basal fasciole indistinct. Outer surface with remnants of ine growth lines. Dimensions (mm) 0.090 1.107 4.630 4.371 1.247 AH AW UW IS SA DHW PD H D SH AH AW 2.7134.053 2.2203.336 1.3482.304 10°-22° 106°-126° 0.230 1.143 8,490 8.211 2.200 6.290 4.849 UW IS SA 3.383 2.778 1.826 16° 2.702 18° 116° 116° Remarks. The Miocene shells listed above are identical to the Pliocene ones described by Pedriali & Robba (2008a). The 95% conidence Remarks. See below for the relationships with Tectonatica sp. 2. Tectonatica sp. 1 supericially resembles the Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Eocene to lower Oligocene species Tectonatica pasinii (Bayan, 1870), but differs from it in that it has: 1) a protoconch with signiicantly greater diameter and diameter of the initial half-whorl, 2) a more globose teleoconch, 3) a spire with more convex whorls (latly convex in T. pasinii), 4) a shorter aperture, 5) a parietal callus lacking the obtuse adapical ridge (present in T. pasinii), 6) a narrower umbilicus, and 7) a narrowly elongate umbilical callus (subtriangular, short in T. pasinii). The present specimens probably represent a new species, but a decision in this respect must await that more, better preserved material be recovered. Stratigraphic occurrence. Tectonatica sp. 1 was found to occur in lower Lutetian deposits of Veneto. Tectonatica sp. 2 Pl. 6, ig. 17; Pl. 7, ig. 1 Material examined: Cava Albanello: 1 spm., MCZ 4327; Cava Grola: 1 spm., MGP-PD 1227R (Pl. 6, ig. 17), 1 spm., MCV 15/04 (Pl. 7, ig. 1). 157 Remarks. Tectonatica sp. 2 is closely similar to Tectonatica sp. 1 (see above) in teleoconch morphology, but can be differentiated from it because of its protoconch of 0.8 more whorls, with significantly smaller diameter of the irst half-whorl, its adpressed suture (slightly channeled in Tectonatica sp. 1), its parietal callus markedly thickened at both ends, with concave abapertural outline (thin, rectangular in Tectonatica sp. 1), its narrower, crescent-shaped umbilicus, its more prominent funicle, and its large, semicircular umbilical callus (narrowly elongate in Tectonatica sp. 1). The protoconch of 0.5 more whorls, with signiicantly greater diameter, the shorter spire with more convex whorls, and the different characters of the inner lip calluses readily distinguish Tectonatica sp. 2 from Tectonatica pasinii (Bayan, 1870). The three examined shells likely represent a new species, but naming it requires more material. Stratigraphic occurrence. Tectonatica sp. 2 was recovered from middle Lutetian deposits of Veneto. Subfamily Poliniceinae Finlay & Marwick, 1937 Description. Protoconch small, lowturbiniform, of 2.80 whorls, tip small. Teleoconch globose, rather thick. Spire moderately elevated, broadly conical, whorls convex, suture adpressed. Last whorl globose, very slightly higher than wide; subsutural shelf very slightly concave, poorly deined; periphery above midline. Aperture D-shaped, in moderately prosocline plane, length 1.8 times width; outer lip regularly arched; inner lip straight, thickened abapically. Parietal callus thick, more so at both ends, with concave abapertural outline; anterior lobe rounded, scarcely produced. Umbilicus deep, crescent-shaped; umbilical wall moderately steep. Funicle broad, rather thick to thick, separated from umbilical wall by rather narrow, shallow spiral depression. Umbilical callus large and thick, at mid-abapical part of inner lip, with convex adaxial outline, separated from anterior lobe of parietal callus by shallow notch. Basal fasciole indistinct. Outer surface with dense growth markings prosocyrt adapically. The species of the Poliniceinae have teleoconch types similar to those occurring in the Naticinae. The umbilicus is either widely open or completely covered by the umbilical callus; the funicle is absent to weak, or thick. The outer surface is mostly smooth, but some genera are spirally sculptured (e.g. Eunaticina Fischer, 1885 and Sigatica Meyer & Aldrich, 1886). The Poliniceinae were said to be characterized by their corneous operculum (Cernohorsky 1971; Kilburn 1976; Marincovich 1977; Majima 1989; Wilson 1993; Kabat 1998; Pastorino 2005; Huelsken et al. 2008). However, Marincovich (1977, p. 212) remarked that Eunaticina insculpta (Carpenter, 1865) has a partially calciied operculum. A similar mixed composition of the operculum was cited by Pastorino (2005, p. 226) concerning the poliniceine species Euspira falklandica (Preston, 1913) and Euspira patagonica (Philippi, 1845). Obviously, the capacity for preservation in the fossil record of these only partly calciied opercula is strongly reduced. Genus Ampullonatica Sacco, 1890 Dimensions (mm) DHW PD H D SH AH AW 0.110 0.912 12.238 12.080 3.621 8.619 7.230 UW IS SA 4,370 23° 125° Ampullonatica Sacco, 1890a, p. 208 (nomen nudum). Ampullonatica Sacco, 1890b, p. 40. Type species by subsequent designation (Cossmann 1893, p. 740): Ampullaria ambulacrum Sowerby, 1822, Eocene, Great Britain and France (see also Kabat 1991 and Caze et al. 2011). Robba E., Pedriali L. & Quaggiotto E. 158 Ampullonatica was introduced as a subgenus of Sigaretus Lamarck, 1799 (= Sinum Röding, 1798). Kabat (1991) treated Ampullonatica as a junior subjective synonym of Euspira Agassiz in J. Sowerby, 1837, implicitly allocating it in the Naticidae. Cossmann (1925) regarded Ampullonatica as a subgenus of Ampullina Lamarck in Defrance, 1821 (= Ampullina Bowdich, 1822; see Kabat 1991, pp. 426, 427). Several subsequent authors considered Ampullonatica as a member of the family Ampullinidae Cossmann, 1918 (Wrigley 1946; Pacaud & Le Renard 1995 among others). Caze et al. (2011) included Ampullonatica in a group of “ampullinid-like gastropods of uncertain afinity”. Recently, the Ampullinidae were regarded as a family unrelated to the Naticidae, removed from the Naticoidea and included in the superfamily Ampullinoidea Cossmann, 1918 (Bandel 2006; Harzhauser et al. 2009) or in the superfamily Campaniloidea Douvillé, 1904 (Bouchet et al. 2005; Beu & Marshall 2011; Caze et al. 2011). According to Kase & Ishikawa (2003) and Beu & Marshall (2011), the distinctive morphological characters of the Ampullinidae are: 1) a rather elevated, pointed, conical protoconch with inlated and smooth whorls, different from that of the Naticidae, which is low-turbiniform, with obtuse, often sculptured tip, 2) a layer of callus (termed sheath by Wrigley 1946) around the umbilical area, overlain by the inner lip calluses, and 3) an umbilicus (when present) devoid of funicle. Both Ampullonatica ambulacrum, type species of Ampullonatica, and Ampullonatica repressa Sacco, 1890 (described below) have the protoconch shaped like that of the naticids and lack the sheath; further, a vestigial funicle was noted in some species. On this basis, we are inclined to consider Ampullonatica better allocated in the Naticidae than in the Ampullinidae, at least provisionally. The deep sutural channel with subangular to sharply angular abaxial border is markedly different from the less deep sutural channel with rounded abaxial border, occurring in a few Euspira species (e.g. Euspira magenesi Pedriali & Robba, 2001; Euspira giuntellii sp. n., herein), and easily separates Ampullonatica from Euspira. Ampullonatica pseudorepressa sp. n. Pl. 7, igs. 2, 3 Derivation of name: From Greek pseudes = false and repressa = the name of the similar species Ampullonatica repressa (Sacco, 1890). Holotype: Monte Gloso: MPUM 11291 (Pl. 7, ig. 2). Paratypes: Cà Sella: 1 spm., MPUM 11292; Cassinelle: 1 spm., MPUM 11293 (Pl. 7, ig. 3); Mioglia: 1 spm., MZB 60100; Sangonini; 1 spm., MGP-PD 15948/b. Other material examined: Cà Sella: 1 spm., NP 9922; Cassinelle: 3 spms., PG 83; Mioglia: 1 spm., NP 9923. Preservation: Most specimens are rather well preserved. Type locality: Monte Gloso (see appendix). Horizon: Yellowish-gray, coarse-grained marly sandstone of Rupelian age. Diagnosis: Medium-sized, globose shell with moderately elevated to low spire; earlier whorls largely covered by succeeding whorl; suture widely channeled, with subangular abaxial border. Last whorl globose. Umbilicus deep, rather wide; funicle short, markedly depressed, rapidly vanishing toward interior of umbilicus. Parietal callus thick; umbilical callus slender to very slender, merging into anterior end of parietal callus. Description. Protoconch medium-sized, turbiniform, of 2.8 convex whorls, tip small. Teleoconch medium-sized, globose, rather thick. Spire moderately elevated to low; whorls gently convex, quickly increasing in diameter, earlier ones largely covered by succeeding whorl. Suture widely channeled, with subangular abaxial border. Last whorl globose to depressed-globose, scarcely expanded toward aperture; subsutural shelf indistinct; latly rounded periphery slightly above or at midline. Aperture D-shaped in moderately prosocline plane, length about 1.7 times width; outer lip semicircular, PLATE 7 Fig. 1 - Tectonatica sp. 2. Cava Grola. MCV 15/04; a, apertural side (height of shell 12.24 mm); b, protoconch. Fig. 2 - Ampullonatica pseudorepressa sp. n. Monte Gloso. Holotype, MPUM 11291; a, apertural side (height of shell 18.11 mm); b, apical view; c, basal view. Fig. 3 - Ampullonatica pseudorepressa sp. n. Cassinelle. Paratype, MPUM 11293; protoconch. Fig. 4 - Ampullonatica repressa (Sacco, 1890). Albugnano. Lectotype (here designated) of Sigaretus (Ampullonatica) repressa Sacco, 1890. MGPT BS.029.10.001; a, apertural side (height of shell 32.01 mm); b, apical view. Fig. 5 - Ampullonatica repressa (Sacco, 1890). Cascina Pianiorito. PG 81; a, apertural side (height of shell 18.96 mm); b, apical view; c, basal view. Fig. 6 - Ampullonatica sp. Monte Gloso. Private collection; apertural side showing color pattern (height of shell 10.65 mm). Fig. 7 - Ampullonatica sp. Monte Gloso. MPUM 11294; a, apertural side (height of shell 13.38 mm); b, apical view; c, basal view; d, protoconch. Fig. 8 - Eunaticina sp. Cava Albanello. MPUM 11295; a, apertural side (height of shell 13.17 mm); b, apical view; c, protoconch. Fig. 9 - Euspira gianoi sp. n. Rio di Bocca d’Asino. Holotype, MPUM 11296; a, apertural side (height of shell 7.49 mm); b, protoconch. Fig. 10 - Euspira gianoi sp. n. Rio di Bocca d’Asino. Paratype, MPUM 11298; a, protoconch; b, detail of protoconch. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 159 Robba E., Pedriali L. & Quaggiotto E. 160 inner lip straight, slightly thickened abapically. Parietal callus rather thick; anterior lobe small to indistinct, ending below umbilical border. Umbilicus deep, rather wide, asymmetrically teardrop-shaped; umbilical wall steeply sloping, with coarse growth ridges. Funicle a short, depressed adaxial thickening of inner lip, soon vanishing toward interior of umbilicus. Umbilical callus slender to very slender, triangular, merging into anterior end of parietal callus. Basal fasciole broad, scarcely prominent, with coarse growth ridges. Outer surface with uneven and unevenly spaced, straight growth lines. Dimensions (mm) DHW PD H D SH 0.118* 1.116* 14.05719.053 14.34421.592 2.0337.017 * 1 protoconch measurable 16.555 17.968 4.525 AH AW UW IS SA 10.49813.562 7.183-13.315 5.1617.489 11°-31° 115°-147° 12.030 10.249 6.325 21° 131° Remarks. The present species differs from Ampullonatica repressa (Sacco, 1890) in that it has: 1) a globose instead of globose-oval shell, 2) a differently coiled spire (earlier whorls largely covered by subsequent whorl), with greater spiral angle, 3) a wider, less deep sutural channel, with roundly angular abaxial border (sharply angular in A. repressa), and 4) a more distinct funicle (vestigial to indistinct in A. repressa). No comparison of respective larval shells can be made since all the examined specimens of A. repressa have the apical whorls abraded at various extent. The Eocene species Natica brongniarti Deshayes, 1864 also belongs in Ampullonatica (Pacaud & Le Renard 1995; Caze et al. 2011). Deshayes’ species is strikingly similar to Ampullonatica pseudorepressa in overall shell shape, but can be differentiated from it by its very narrow and very deep sutural channel, and by its umbilicus without any trace of funicle. Stratigraphic occurrence. Ampullonatica pseudorepressa sp. n. was recovered from lower Oligocene deposits of Piedmont, Liguria and Veneto. Ampullonatica repressa (Sacco, 1890) Pl. 7, igs. 4, 5 ig. 2. 1890b Sigaretus (Ampullonatica) repressa Sacco, p. 40. 1891 Ampullonatica repressa - Sacco, p. 105, pl. 1, ig. 75. 1984 Ampullonatica repressa - Ferrero Mortara et al., p. 39, pl. 4, Type material: Sigaretus (Ampullonatica) repressa Sacco, lectotype (here desinated): the unique syntype presently existing in MGPT, igured by Ferrero Mortara et al. (1984, pl. 4, ig. 2) and reigured herein (Pl. 7, ig. 4), MGPT BS.029.10.001, Albugnano (see remarks below). Other material examined: Cascina Pianiorito: 1 spm., PG 81 (Pl. 7, ig. 5), 2 spms., PG 82. Description. Protoconch abraded. Teleoconch medium-sized, globose-oval, rather thick. Spire moderately elevated, somewhat stepped; whorls convex, roundly subangular at adapical onethird, with obscure subsutural shelf. Suture deeply and rather widely channeled, with sharply angular abaxial border. Last whorl widely oval to globose, scarcely expanded toward aperture; subsutural shelf indistinct; periphery slightly below or at midline. Aperture D-shaped in prosocline plane, length about 1.9 times width; outer lip semicircular, inner lip straight, slightly thickened abapically. Parietal callus rather thick, with concave abapertural outline; anterior lobe short, subangular, ending at level of umbilical border. Umbilicus deep, moderately wide, subrectangular, angular adapically, rounded abapically; umbilical wall very steep, with coarse growth ridges. Funicle vestigial to indistinct. Umbilical callus slenderly triangular, slightly overhanging adapical part of umbilicus and merging into anterior lobe of parietal callus. Basal fasciole indistinct. Outer surface with unevenly spaced, straight growth lines. One specimen retains vestige of uniform, pale brown background. Dimensions (mm) DHW PD H D SH AH AW - - 18.961 17.649 6.570 12.392 10.270 UW IS SA 5.598 20° 121° Remarks. The present species was created by Rovasenda and named Natica repressa on a manuscript label. Subsequently, the name repressa was merely listed only by Sacco (1890a). Sacco (1890b) irst published a concise diagnosis of N. repressa (see the above synonymy), thus making the name available, and is to be considered the author of this species. The Bellardi-Sacco collection in MGPT encloses a single syntype of Ampullonatica repressa (cf. Ferrero Mortara et al. 1984). Since Sacco (1891) afirmed that some specimens (more than one) have been collected, it is supposed that other syntypes existed in the collection, but were probably lost. In Eocene, Oligocene and Miocene naticid gastropods of Northern Italy these circumstances, the unique syntype remaining (MGPT BS.029.10.001) is not eligible as the holotype of the species, instead, it can be designated as the lectotype. Ampullonatica ambulacrum (Sowerby, 1822), type species of Ampullonatica, is rather similar to Ampullonatica repressa in shell shape. We examined some topotypes of A. ambulacrum in our collection and can state that Sowerby’s species is readily differentiated from A. repressa in that it has: 1) a higher, pointed, more stepped spire, 2) a wider, lat-bottomed sutural channel with abaxial border subrounded and slightly overhanging the channel, 3) a last whorl more expanded toward the aperture, with the periphery above midline (at midline in A. repressa), 4) a remarkably thin parietal callus, and 5) a smaller umbilicus. Stratigraphic occurrence. Ampullonatica repressa (Sacco, 1890) appears to be a rare species, hitherto recovered only from the Langhian Baldissero Formation near Albugnano (Piedmont). Ampullonatica sp. Pl. 7, igs. 6, 7 Material examined: Monte Gloso: 1 spm., private collection, 1 spm., private collection (Pl. 7, ig. 6), 1 spm., MPUM 11294 (Pl. 7, ig. 7), 1 spm., NP 9950. Description. Protoconch medium-sized, lowturbiniform, of 2.5-2.75 gently convex whorls, tip small. Teleoconch small, globose and thick. Spire broadly conical, moderately elevated; whorls gently convex, roundly subangular at adapical one-third, with obscure subsutural shelf. Suture rather shallowly channeled; sutural channel V-shaped in cross section, with subangular abaxial border. Last whorl globose, slightly expanded toward aperture; subsutural shelf poorly deined; periphery well above midline. Aperture D-shaped in prosocline plane, length about 1.7 times width; outer lip regularly arched, inner lip straight, thickened abapically. Parietal callus thickened at both ends, with concave abapertural outline; anterior lobe indistinct. Umbilicus deep, moderately wide, angular adapically, rounded abapically; umbilical wall very steep, interior of umbilicus with rather even spiral cordlets. Funicle a short, depressed adaxial thickening of inner lip, soon vanishing toward interior of umbilicus. Umbilical callus triangular, merging into anterior corner of parietal callus at 161 level of abaxial border of basal fasciole. Basal fasciole rather broad, markedly depressed, enclosed within obscure spiral angulations. Outer surface with unevenly spaced, straight growth lines, slightly prosocyrt subsuturally. One specimen with dark brown, well spaced axial lines. Dimensions (mm) DHW PD H D SH AH AW 0.172 1.053 13.380 12.571 3.779 9.601 7.432 UW IS SA 3.860 25° 112° Remarks. The protoconch of these specimens is poorly preserved, thence the above values of its characteristic elements are reported with reservation. We have been uncertain about the generic assignment of the present taxon. The inner spiral cordlets of the umbilicus point toward the genus Euspira Agassiz in J. Sowerby, 1837, whereas the other shell characters (especially the V-shaped sutural channel) are much alike those of Ampullonatica. The inal decision has been in favor of the allocation in Ampullonatica, at least provisionally. Ampullonatica sp. has teleoconch characters intermediate between Ampullonatica repressa (Sacco, 1890) and Ampullonatica pseudorepressa sp. n. The inner spiral cordlets of the umbilicus distinguish it from both A. repressa and A. pseudorepressa. Probably, Ampullonatica sp. can be differentiated from A. pseudorepressa also in that it has the protoconch with a signiicantly greater diameter of the initial half-whorl. Ampullonatica sp. attains a size smaller than that of both A. pseudorepressa and A. repressa. Euspira bartonensis Wrigley, 1949 (topotypes examined) is closely similar in shell shape and likewise has the umbilicus with inner spiral cordlets, but its adpressed, slightly incised suture is a primary differentiating element and denotes that Wrigey’s species deinitely belongs in Euspira. Most likely, the present specimens represent a new species, which, however, requires more material to be named. Stratigraphic occurrence. Ampullonatica sp. was recovered only from lower Oligocene deposits of Monte Gloso (Veneto). Genus Eunaticina Fischer, 1885 Naticina Gray, 1840, p. 147 (nomen nudum). Naticina Gray, 1847, p. 150. Type species by original designa- 162 Robba E., Pedriali L. & Quaggiotto E. tion: Nerita papilla Gmelin, 1791, Pliocene to Recent, Indo-West Paciic (junior homonym of Naticina Guilding, 1834). Eunaticina Fischer, 1885, p. 768. Replacement name for Naticina Gray, 1847, preoccupied. Heliconatica Dall, 1924, p. 90. Type species by original designation: Eunaticina (Heliconatica) margaritaeformis Dall, 1924 (see the remarks on Sigatica Meyer & Aldrich 1886). Pervisinum Iredale, 1931, p. 216. Type species by original designation: Pervisinum dingeldii Iredale, 1931 (=Nerita papilla Gmelin, 1791). records were so far from the early and late Miocene of Europe, from middle Miocene to Pleistocene deposits of West Paciic countries, and from the Pleistocene of Baja California, Mexico. The Eocene species described below constitutes the irst record of the genus in Italy and predates the appearance of Eunaticina. The status of Nerita papilla was thoroughly discussed by Beu (2004, p. 206), who provided a detailed synonymy. Kabat (1991) considered Pervisinum Iredale, 1931, Propesinum Iredale, 1924 and Sigaretotrema Sacco, 1890 as junior subjective synonyms of Eunaticina. We concur with Kabat as concerns Pervisinum (see also Kilburn 1976 and Beu 2004), but regard Sigaretotrema as a distinct genus and place Propesinum in the synonymy of it (see comments on Sigaretotrema). Eunaticina was originally introduced as a subgenus of Sigaretus Lamarck, 1799 (= Sinum Röding, 1798). Fischer (1885, p. 768) published the following concise diagnosis for Eunaticina: “Coquille ombiliquée, ovale oblongue, naticiforme, mince, venture; spire acuminée; tours striés ou sillonnés; ouverture oblongue; opercula de Sigaretus”. Examination of the type species (material in our collection; igures of Beu 2004) and of Narica insculpta Carpenter, 1865 has shown that Eunaticina is distinguished by the following combination of characters: 1) protoconch of 2.5-3 smooth whorls; 2) teleoconch elongate-oval, with moderately elevated spire; 3) spire whorls rather convex, meeting at channeled sutures; 4) last whorl extended abapically; 5) aperture prosocline, with length almost twice the width; 6) umbilicus open and deep, devoid of funicle; 7) umbilical callus indistinct to very narrow; 8) sculpture of spiral cords. Eunaticina has been currently included in the subfamily Sininae (Kilburn, 1976; Marincovich, 1977; Majima, 1989). Recent research supports its allocation in Poliniceinae (Aronowsky 2000). Eunaticina is a poorly speciose genus at present occurring widely throughout the Indo-West Paciic (Eunaticina papilla) and in the Eastern Paciic, from Southern Gulf of California to Ecuador (Eunaticina insculpta). E. papilla has recently entered the Mediterranean (one specimen found in the Iskenderum Bay, Turkey), probably by means of larval stages in ballast water (Öztürk & Bitlis 2013). Fossil Eunaticina sp. Pl. 7, ig. 8 Material examined: Cava Albanello: 1 spm., MPUM 11295. Description. Protoconch very large, lowturbiniform, of 2.25 convex whorls, tip mediumsized. Teleoconch oval, rather thick. Spire cyrtoconoid, moderately elevated, whorls convex. Suture channeled. Last whorl inlate-oval, higher than wide; subsutural shelf indistinct; periphery above midline. Aperture D-shaped in prosocline plane, length 1.8 times width. Outer lip arched; inner lip almost straight, slightly relected over adapical part of umbilicus. Parietal callus rectangular, rather thin, angular at level of umbilical border. Umbilicus deep, rather widely open; umbilical border rounded; umbilical wall steep, with prominent growth ridges. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture of rather even, lat-topped spiral cords, approximately 50 on last whorl, wider than interspaces, iner and more widely spaced over base; spirals crossed by dense growth markings. Dimensions (mm) DHW PD H D SH AH AW 0.258 2.114 13.171 11.890 2.513 10.662 6.670 UW IS SA 3.13 14° 102° Remarks. This small Eocene shell is unlike any known Eunaticina species and likely belongs to a previously undescribed species. However, we refrain from proposing a new species on the basis of a single, probably not fully grown specimen, whose preservation is not perfect. The upper Miocene to Recent Indo-West Paciic species Eunaticina papilla (Gmelin, 1791) differs from Eunaticina sp. in that has: 1) a protoconch with signiicantly smaller diameter and diameter of the irst half-whorl, 2) a higher last whorl and a higher aperture, and 3) wider, ribbon-like spirals separated Eocene, Oligocene and Miocene naticid gastropods of Northern Italy by narrow grooves, less attenuated over the base and continuing on the umbilical wall (Eunaticina sp. has the umbilical wall only with growth markings). Eunaticina (Eunaticina) visenda, introduced by Beets (1984, p. 13) on the basis of Preangerian (middle Miocene) material from Kalimantan (Indonesia), differs by its more globose shell with more deeply channeled sutures and wider umbilicus, and by its sculpture of uneven spiral bands. The upper Miocene to Recent West American species Eunaticina insculpta (Carpenter, 1865) is readily distinguished in having a globose shell, also with wider spirals and “sharply incised”, narrower interspaces (cf. Marincovich 1977). Lozouet et al. (2001) regarded Natica sulcata and Natica striatella, both introduced by Grateloup (1828), as synonyms; they used the name striatella as the valid one and assigned the species to Eunaticina. We examined the specimens illustrated by Cossmann & Peyrot (1919), referred to as Sigaretus (Sigaretotrema) striatellus (MNHN-F-J03843) and Sigaretus (Sigaretotrema) sulcatus (MNHN-F-J05702), another shell of Eunaticina striatella from Mérignac in our collection and concur with the opinion of the cited authors. Eunaticina striatella appears to differ from Eunaticina sp. in having a protoconch with signiicantly smaller diameter and diameter of the irst half-whorl, a more globose teleoconch with wider umbilicus, and a sculpture of spiral ribbons. Sigaretus gouldianus Recluz, 1851, described from Eocene deposits of Paris area, was later assigned to Eunaticina Fischer, 1885 (cf. Cossmann 1888; Cossmann & Pissarro 1907 and Pacaud & Le Renard 1995). From the original description (Recluz 1851, p. 188), it is evident that this species is devoid of umbilicus, which is said to be “réduit à un tout petit trou, équivalant à une piqùre d’épingle”. On the basis of this character, it results that: 1) Recluz’s species differs markedly from Eunaticina sp., and 2) it is unlikely to belong in Eunaticina. The Tortonian Bulgarian shell referred to as Sinum (Sigaretotrema) michaudi (Michelotti, 1847) by Kojumdgieva & Strachimirov (1960) seems better allocated in Eunaticina (see comments to Sigaretotrema michaudi below). According to the igure published by Kojumdgieva & Strachimirov, the Bulgarian specimen appears to have the teleoconch similar in shape to that of Eunaticina sp., but differs from it in having less convex whorls, thicker inner lip and parietal callus, and the umbilical wall with spiral cords. 163 Stratigraphic occurrence. Eunaticina sp. was recovered from lower Lutetian deposits of Cava Albanello (Veneto). Genus Euspira Agassiz in J. Sowerby, 1837 Euspira Agassiz in J. Sowerby, 1837, p. 14. Type species by subsequent designation (Bucquoy, Dautzenberg & Dollfus 1883, p. 143): Natica glaucinoides J. Sowerby, 1812, Eocene, Great Britain. Lunatia Gray, 1847, p. 149. Type species by original designation: Natica ampullaria Lamarck, 1822 (= Natica heros Say, 1822). Labellinacca Cossmann in Cossmann & Peyrot, 1919, p. 188. Type species by subsequent designation (Cossmann 1925, p. 137): Natica labellata Lamarck, 1804. The genus was briely discussed in terms of type species designation and authorship by Kabat (1991). Cox (1930) considered Natica glaucinoides as a junior synonym of Natica labellata Lamarck, 1804 and this conclusion was implicitly or explicitly agreed upon by several workers (e.g. Glibert 1933; MacNeil 1960; Murphy & Rodda 1960; Kilburn 1976; Torigoe & Inaba 2011), who indicated N. labellata as the type species of Euspira. Wrigley (1949) redescribed N. glaucinoides and N. labellata (both included in Euspira) and afirmed them to be distinct species. Wrigley’s opinion was accepted by other authors (e.g. Glibert 1963; Kanno 1971; Majima 1989; Pastorino 2005; Pedriali & Robba 2009; Huelsken et al. 2012; Garvie 2013; Landau et al. 2013). We examined some topotypes of N. glaucinoides and of N. labellata and could note that these species share many teleoconch characters. However, N. labellata differs from N. glaucinoides in that it has: 1) a greater protoconch of about one more whorl, with significantly smaller diameter of the initial half-whorl, 2) a narrow, gently sloping subsutural shelf (indistinct in N. glaucinoides), 3) a vestigial to absent funicle (broad, markedly depressed in N. glaucinoides), and 4) an abapical sulcus with subangular abaxial border (rounded in N. glaucinoides). On this basis, it is concluded that N. glaucinoides is the type species of Euspira. Natica labellata is the type species of Labellinacca Cossmann, 1919. Since it is evidently congeneric with Natica glaucinoides, we agree with the opinion of Kabat (1991) and Torigoe & Inaba (2011), who considered Labellinacca a junior synonym of Euspira. Cossmann (in Cossmann & Peyrot 1919, p. 188) introduced Labellinacca as a section of Lunatia and wrote “Section s’appliquant aux forms qui, telles que N. labellata par exemple”. This statement nei- 164 Robba E., Pedriali L. & Quaggiotto E. ther is designation (ICZN 1999, Article 67.5 of the Code), nor is ixation by monotypy (ICZN 1999, Article 68.3 of the Code). The irst valid designation was subsequently made by Cossmann (1925, p. 137), who explicitly indicated N. labellata as the type species of Labellinacca. Torigoe & Inaba (2011, p. 13) reinstated Lunatia (type species Natica ampullaria Lamarck, 1822 = Natica heros Say, 1822) as a valid genus, distinct from Euspira. However, the characters of Natica heros and of Natica glaucinoides demonstrate that these species are congeneric. Accordingly, we concur with Kabat (1991) in considering Lunatia as a junior synonym of Euspira. Besides Labellinacca and Lunatia, Kabat (1991) regarded Laguncula Benson, 1842, Bensonia Gray, 1847, Ampullonatica Sacco, 1890, Dallitesta Mansield, 1930, Scarlatia Schileyko, 1977, and Pseudopolinices Golikov & Sirenko, 1983 as synonyms of Euspira. Concerning Ampullonatica, we note that the remarkably deep sutural channel characterizing the species of this genus constitutes a reliable element distinguishing Ampullonatica from Euspira (see also the above comments on Ampullonatica). In a previous paper (Pedriali & Robba 2009, p. 387), Pseudopolinices was included in the synonymy of Polinices Montfort, 1810 (instead of Euspira) on the basis of the umbilical and parietal callus characters of Natica nana Möller, 1842 (type species of Pseudopolinices). This conclusion is retained herein. Laguncula, Bensonia, Dallitesta and Scarlatia appear to be unrelated to Euspira. Euspira was regarded as a subgenus of Polinices Montfort, 1810 by some authors (e.g. Marwick 1924; Kilburn 1976; Marincovich 1977; Garvie 2013), but recently Huelsken et al. (2012), on the basis of molecular and shell morphology analyses, concluded that “Euspira is not a subgenus of Polinices but represents a valid genus”. Pedriali & Robba (2009) listed the diagnostic characters of Euspira as follows: 1) protoconch lowturbiniform, of 1.25-3 convex whorls, initial halfwhorl with spiral microsculpture in some species; 2) teleoconch thin to moderately thick, globose or globose-elongate; 3) spire depressed to moderately elevated, somewhat stepped in some species; 4) suture almost lush to adpressed, channeled in a few species; 5) parietal callus moderately thick, slender, with more-or-less distinct anterior lobe overhanging the adapical part of the umbilicus, slightly in some species; 6) umbilicus open and deep, narrow to wide, with an inner spiral furrow in most species, spirally sculptured in some species; 7) funicle markedly depressed to absent; 8) umbilical callus slender to indistinct, better developed in a few species, merging into the anterior lobe of the parietal callus or demarcated from it by a weak groove. Pedriali & Robba (2009) remarked that Euspira is a distinctive cosmoplolitan genus useful to accommodate poli-niceine species with an open (not slitlike) umbilicus virtually devoid of a funicle and with a reduced to absent umbilical callus. Euspira gianoi sp. n. Pl. 7, igs. 9, 10 Derivation of name: The species is named after Giano Della Bella, who provided lot of material relevant to the present study. Holotype: Rio di Bocca d’Asino: MPUM 11296 (Pl. 7, ig. 9). Paratypes: Borelli: 1 spm., MGPT-PU 135140, 1 spm., MGPT-PU 135141, 1 spm., MGPT-PU 135142, 1 spm., MGPT-PU 135143, 1 spm., MGPT-PU 135144, 11 spms., MGPT-PU 135145, 3 spms., MPUM 11297, 2 spms., NP 9872; Monte dei Cappuccini: 3 spms., MGPT-PU 135146; Rio di Bocca d’Asino: 1 spm., MPUM 11298 (Pl. 7, ig. 10), 4 spms., MPUM 11299; Valle Ceppi: 2 spms., MGPT-PU 135147, 1 spm., MGPT-PU 135148. Other material examined: Albugnano: 5 spms., PG 64, 1 spm., PG 74; Pietracuta: 4 spms., PP.PC02/04; Rio di Bocca d’Asino: 5 spms., NP 9860, 1 spm., PG 62, 1 spm., PG 63; Sant’Agata Fossili: 1 spm., PG 73b. Preservation: Rather fair on the whole. Type locality. Rio di Bocca d’Asino (see appendix). Horizon: Resedimented sandstones and conglomerates forming lenticular bodies into the lower member of the S. Agata Fossili Formation; the age is Tortonian. Diagnosis: Small, globose-oval shell with rather elevated spire and broadly oval last whorl. Umbilicus deep, narrowly beanshaped, narrower and comma-shaped in a few specimens; funicle markedly depressed. Parietal callus thick, with anterior lobe covering adapical part of umbilicus; umbilical callus subtriangular, slightly overhanging umbilical opening and merging into anterior lobe of parietal callus. Vestige of pale brown background with darker subsutural band and lower base. Description. Protoconch medium-sized, low-turbiniform, of 3-3.12 convex whorls, tip very small; protoconch I with evenly spaced spiral rows of interconnected, axially elongated and variously shaped microprotuberances forming a markedly irregular, somewhat reticulated pattern. Teleoconch small, globose-oval, moderately thick. Spire conical, rather elevated, whorls gently convex; suture adpressed. Last whorl broadly oval, scarcely produced abapically toward aperture; subsutural shelf moderately wide, somewhat concave, bounded by obtuse Eocene, Oligocene and Miocene naticid gastropods of Northern Italy but distinct angulation, with slight subsutural margining in a few specimens; periphery above midline. Aperture D-shaped in slightly prosocline plane, length about 1.7 times width; outer lip semicircular, inner lip nearly straight. Parietal callus thick, less so medially, with concave abapertural outline and slight adapical tubercle; anterior lobe subangular covering adapical part of umbilicus. Umbilicus deep, narrowly bean-shaped, narrower and comma-shaped in a few specimens; umbilical wall steep, bounded downward by very slight groove. Funicle markedly depressed. Umbilical callus subtriangular, with straight or attenuated reverse S-shaped abapertural outline, slightly overhanging umbilical opening and merging into anterior lobe of parietal callus. Basal fasciole weak to indistinct. Outer surface with ine growth lines obliquely prosocyrt over subsutural shelf. Specimens from Rio di Bocca d’Asino retain vestige of pale brown background with darker subsutural band and lower base. Dimensions (mm) DHW PD H D SH 0.0740.086 1.2171.473 1.6538.201 1.6007.352 0.3512.799 0.080 1.345 4.927 4.476 1.575 AH AW UW IS SA 1.0795.623 0.8164.576 0.3131.849 17°-29° 81°-109° 3.351 2.696 1.081 23° 95° 165 from it because of its protoconch of about 0.5 more whorls, with signiicantly greater diameter and smaller diameter of the initial half-whorl; further, the rather elaborated sculpture of protoconch I is different from the simpler one observed in E. helicina helicina (spiral cordlets). E. notabilis is another similar species, differing from E. gianoi in having: 1) a protoconch of 0.8 fewer whorls, with signiicantly smaller diameter and with the irst whorl bearing nodulose spiral cordlets, 2) a more globose teleoconch with lower spire, and 3) a crescent-shaped umbilicus. E. pulchella is similar to E. gianoi in teleoconch shape and size, but has a more widely open umbilicus, an obsolete funicle, and a larval shell signiicantly smaller in diameter. E. umbilicocarinata is easily distinguished from E. gianoi because of its protoconch with signiicantly greater diameter of the irst half-whorl, its globose shell with lower spire, and its wider umbilicus, whose interior is bounded by a keel-like angulation. E. grossularia, E. helicina helicina and E. notabilis attain a size larger than that of E. gianoi. Stratigraphic occurrence. Euspira gianoi sp. n. was recovered from Burdigalian to Tortonian deposits of Piemont. Euspira giuntellii sp. n. Pl. 8, igs. 1-4 Remarks. The present new species can be compared only with Euspira grossularia (MarcheMarchad, 1957), Euspira helicina helicina (Brocchi, 1814), Euspira notabilis (Jeffreys, 1885), Euspira pulchella (Risso, 1826) and Euspira umbilicocarinata sp. n. (see below), taxa that also occur in the Miocene and are closely or supericially similar to Euspira gianoi. The values of the characteristic elements of the protoconch measured for E. gianoi do not differ signiicantly from those pertaining to E. grossularia and the sculpture of protoconch I is similar in both species. However, E. gianoi is readily distinguished from E. grossularia in that it has: 1) a higher, more narrowly coiled spire (Fig. 11), 2) an oval, less inlated last whorl with distinct subsutural shelf (indistinct in E. grossularia), and 3) a slight groove bounding downward the umbilical wall (absent in E. grossularia). E. gianoi resembles E. helicina helicina in teleoconch shape, umbilical characters and color pattern, but can be differentiated Derivation of name: The species is named after Piero Giuntelli, who provided a wealth of material relevant to the present study. Holotype: Borelli: MGPT-PU 135149 (Pl. 8, ig. 1). Paratypes: Borelli: 1 spm., MGPT-PU 135150 (Pl. 8, ig. 2), 1 spm., MGPT-PU 135151 (Pl. 8, ig. 3), 1 spm., MGPT-PU 135152 (Pl. 8, ig. 4), 1 spm., MGPT-PU 135153, 1 spm., MGPT-PU 135154, 29 spms., MGPT-PU 135155. Preservation: Most specimens have minor damages. Type locality: Borelli (see appendix). Horizon: Gray, medium sand of Tortonian age. Diagnosis: Small, depressed-globose shell with scarcely elevated spire and somewhat depressed last whorl. Suture slightly channeled. Umbilicus deep, narrowly oval to comma-shaped, funicle markedly depressed to indistinct. Parietal callus thickened at both ends, thin medially; umbilical callus slender, with attenuated reverse S-shaped or straight outline, merging into anterior lobe of parietal callus. Description. Protoconch small, lowturbiniform, of 1.75-1.85 convex whorls, last halfwhorl with even spiral cordlets, tip small. Teleoconch small, depressed-globose, slightly thick. Spire broadly conical, scarcely elevated, moderately taller in a few specimens, whorls convex. Suture slightly channeled. Last whorl large, globose, somewhat de- Robba E., Pedriali L. & Quaggiotto E. 166 pressed, expanding and produced abapically toward aperture; subsutural shelf poorly deined; periphery well above midline. Aperture ovately D-shaped in slightly prosocline plane, length about 1.6 times width; outer lip semicircular, inner lip very slightly arched adapically. Parietal callus thin medially, thickened at both ends, with concave abapertural outline; anterior lobe tongue-shaped covering adapical part of umbilicus, poorly differentiated in a few specimens. Umbilicus deep, narrowly oval, narrower and comma-shaped in a few specimens; umbilical wall steep, bounded downward by slight groove. Funicle markedly depressed to indistinct. Umbilical callus slender, with straight or attenuated reverse S-shaped outline, merging into anterior lobe (or corner) of parietal callus. Basal fasciole weak to indistinct. Outer surface with ine growth lines slightly bent subsuturally. Vestige of yellowish-brown color present in a few specimens. Type material: Pedriali & Robba (2009, p. 391) reviewed the type material of Euspira grossularia and described the characters of this species. Material examined: Borelli: 1 spm., MGPT-PU 135156 (Pl. 8, ig. 5), 1 spm., MGPT-PU 135157, 1 spm., MGPT-PU 135158, 5 spms., MGPT-PU 135159, 1 spm., MGPT-PU 135160 (Pl. 8, ig. 6); 2 spms., MPUM 11300, 4 spms., NP 9869; Monte dei Cappuccini: 2 spms., MGPT-PU 135161, 8 spms., MGPT-PU 135162, 26 spms., MGPT-PU 135163, 1 spm., MGPT-PU 135164 (Pl. 8, ig. 7); Rio di Bocca d’Asino: 1 spm., NP 9855, 1 spm., MPUM 11301; Sant’Agata Fossili: 1 spm., NP 9856; Valle Ceppi: 2 spms., NP 9857, 1 spm., MGPT-PU 135165. Dimensions (mm) DHW PD H D SH 0.0710.091 1.2901.606 2.4246.564 2.7226.138 0.4031.563 0.081 1.448 4.494 4.430 0.983 AH AW UW IS SA 1.9455.077 1.5193.667 0.2751.563 18°-34° 109°-133° 3.511 2.593 0.919 26° 121° Dimensions (mm) DHW PD H D SH 0.1780.210 0.6820.802 2.8825.786 2.9636.091 0.5151.259 0.194 0.742 4.334 4.527 0.887 AH AW UW IS SA 2.1554.739 1.9733.873 0.6281.776 14°-30° 124°-144° 3.447 2.923 1.202 22° 134° Remarks. Euspira giuntellii is closely similar to the Pliocene species Euspira magenesi Pedriali & Robba, 2001 in teleoconch shape and there is no signiicant difference as regards the values of the characteristic elements of the protoconch. However, E. giuntellii is readily distinguished from E. magenesi in that it has: 1) a protoconch with spirally sculptured last half-whorl (the protoconch of E. magenesi has smooth whorls), 2) a less deeply channelled suture, 3) a narrowly oval, distinctly smaller umbilicus (that of E. magenesi is round and wide), and 4) an interior of the umbilicus lacking the spiral cords, which are present in E. magenesi. Stratigraphic occurrence. Euspira giuntellii sp. n. was recovered only from Tortonian deposits at the type locality. Euspira grossularia (Marche-Marchad, 1957) Pl. 8, igs. 5-7 1957 Polynices grossularia Marche-Marchad, p. 201, pl. 1, ig. 3. 2009 Euspira grossularia - Pedriali & Robba, p. 391, pl. 1, igs. 8, 9; pl. 3, ig. 6; pl. 4, igs. 4-6 (cum syn.). PLATE 8 Fig. 1 - Euspira giuntellii sp. n. Borelli. Holotype, MGPT-PU 135149; a, apertural side; b, apical view. Fig. 2 - Euspira giuntellii sp. n. Borelli. Paratype, MGPT-PU 135150; protoconch. Fig. 3 - Euspira giuntellii sp. n. Borelli. Paratype, MGPT-PU 135151; protoconch. Fig. 4 - Euspira giuntellii sp. n. Borelli. Paratype, MGPT-PU 135152; a, apertural side (height of shell 4.35 mm); b, apical view. Fig. 5 - Euspira grossularia (Marche-Marchad, 1957). Borelli. MGPTPU 135156; a, protoconch; b, detail of protoconch. Fig. 6 - Euspira grossularia (Marche-Marchad, 1957). Borelli. MGPTPU 135160; apertural side (height of shell 3.98 mm). Fig. 7 - Euspira grossularia (Marche-Marchad, 1957). Monte dei Cappuccini. MGPT-PU 135164; apertural side (height of shell 5.03 mm). Fig. 8 - Euspira helicina helicina (Brocchi, 1814). Montegibbio. MPUM 11302; apertural side (height of shell 14.76 mm). Fig. 9 - Euspira helicina helicina (Brocchi, 1814). Valle Ceppi. MPUM 11305; protoconch. Fig. 10 - Euspira helicina helicina (Brocchi, 1814). Valle Ceppi. MGPTPU 135171; apertural side (height of shell 14.44 mm). Fig. 11 - Euspira latecallosa sp. n. Borelli. Holotype, MGPT-PU 135180; apertural side (height of shell 4.93 mm). Fig. 12 - Euspira latecallosa sp. n. Borelli. Paratype, MGPT-PU 135182; protoconch. Fig. 13 - Euspira latecallosa sp. n. Borelli. Paratype, MGPT-PU 135183; protoconch. Fig. 14 - Euspira molarensis sp. n. Molare. Holotype, MPUM 11307; apertural side (height of shell 12.77 mm). Fig. 15 - Euspira molarensis sp. n. Mioglia. Paratype, MPUM 11308; protoconch. Fig. 16 - Euspira notabilis (Jeffreys, 1885). Borelli. MGPT-PU 135185; detail of protoconch. Fig. 17 - Euspira notabilis (Jeffreys, 1885). Borelli. MGPT-PU 135186; protoconch. Fig. 18 - Euspira notabilis (Jeffreys, 1885). Borelli. MGPT-PU 135188; apertural side (height of shell 6.48 mm). Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 167 Robba E., Pedriali L. & Quaggiotto E. 168 Remarks. Except for a smaller size, the examined specimens are identical to the Pliocene ones dealt with by Pedriali & Robba (2009). Their larval shells have a slightly greater diameter, but the difference (+ 7%) is far from being signiicant. The 95% conidence intervals calculated for each dimension of the teleoconch overlap the lower part of those obtained for the Pliocene material. For description and comments, see Pedriali & Robba (2009). Stratigraphic occurrence. Euspira grossularia (Marche-Marchad, 1957) was hitherto known to range from late Miocene to Recent (see Pedriali & Robba 2009). The new records from Piedmont (Monte dei Cappuccini, Valle Ceppi) show that the species was already present during the Burdigalian (rare) and the Langhian (rather abundant). 1 spm., MZB 60045; Valle Ceppi: 1spm., MGPT-PU 135167, 3 spms., MGPT 26046, 12 spms., MGPT-PU 135168, 3 spms., MGPT-PU 135169, 1 spm., MGPT-PU 25904, 1 spm., MGPT-PU 26144, 37 spms., MZB 60118, 1 spm., MZB 60042, 5 spms., MZB 60050, 47 spms., MZB 60051, 1 spm., MZB 60052, 25 spms., MZB 60055, 2 spms., MGPT-PU 107766, 11 spms., MGPT-PU 107767, 33 spms., MGPT-PU 108196, 1 spm., MGPT-PU 135170, 3 spms., MGPT-PU 107940, 1 spm., MPUM 11304, 1 spm., MPUM 11305 (Pl. 8, ig. 9), 1 spm., MPUM 11306, 24 spms., PG 71, 45 spms., PG 21b, 25 spms., MGPT-PU 23667, 1 spm., MGPT-PU 135171 (Pl. 8, ig. 10), 34 spms., MGPT-PU 25265, 3 spms., PG 55, 16 spms., MGPT-PU 23861, 1 spm., MGPT-PU 25907, 1 spm., MGPT-PU 26146, 1 spm., MGPT-PU 135172, 3 spms., MGPTPU 135173, 1 spm., MGPT-PU 135174, 1 spm., MGPT-PU 135175, 1 spm., MGPT-PU 135176, 3 spms. NP 9852; Val Sanfrà: 27 spms., MGPT-PU 107942, 1 spm., MGPT-PU 135177, 1 spm., MGPT-PU 135178; Valle Vergnana: 4 spms., MZB 19144, 3 spms., MZB 43806, 6 spms., MZB 60040, 1 spm., MZB 60041, 16 spms., MZB 60047, 6 spms., MZB 60048, 5 spms., MZB 60049, 1 spm., NP 9851; Villa Allason: 1 spm., MGPT-PU 107072, 12 spms., MGPT-PU 108783; Villa Bertini: 1 spm., MGPT-PU 135179, 37 spms., MGPT-PU 108782. Dimensions (mm) Euspira helicina helicina (Brocchi, 1814) Pl. 8, igs. 8-10 ig. 1. 1814 Nerita helicina Brocchi, p. 297, pl. 1, ig. 10. 2008b Nerita helicina - Pedriali & Robba, pp. 173-175, text- 2009 Euspira helicina helicina - Pedriali & Robba, p. 393, pl. 1, igs. 12, 13, 16-18; pl. 3, ig. 8; pl. 4, ig. 10 (cum syn.). 2013 Euspira helicina helicina - Landau et al., p. 103, pl. 11, ig. 9; pl. 12, igs. 1-3; pl. 62, igs. 6, 7. Type material: Pedriali & Robba (2009, p. 396, 397) reviewed the type material of Euspira helicina helicina, and described the characters of this subspecies. Material erroneously referred to as Natica submamillaris var. mioclausa Sacco, 1890 in IPUM: Montegibbio: 1 spm., IPUM 4395. Material erroneously referred to as Natica pardalis Doderlein, 1864 in IPUM: Montegibbio: 4 spms., IPUM 4420, 4 spms., IPUM 4470. Other material examined: Albugnano: 20 spms., PG 52, 4 spms., PG 70, 2 spms., PG 74a; Grazzano Badoglio: 2 spms., PG 54, 2 spms., PG 76a; Moncucco Torinese: 1 spm., PG 1; Monte dei Cappuccini: 1 spm., MGPT-PU 135196, 10 spms., MGPT-PU 25661, 1 spm., MGPT-PU 135166, 27 spms., MGPT-PU 25663, 3 spms., MGPT-PU 25664; Montegibbio: 1 spm., MPUM 11302 (Pl. 8, ig. 8), 2 spms., MGC 1417, 34 spms., private collection, 12 spms., private collection, 42 spms., private collection, 144 spms., private collection, 1 spm., private collection; Pietracuta: 23 spms., PP.PC 02/03, 4 spms., PP.PC 03/03; Rio di Bocca d’Asino: 19 spms., MZB 60034, 14 spms., MZB 60035, 27 spms., MZB 60036, 1 spm., MZB 60037, 17 spms., MZB 60038, 3 spms., MZB 60039, 4 spms., MZB 60053, 10 spms., NP 9847, 5 spms., PG 2, 2 spms., PG 5, 2 spms., PG 53, 3 spms., PG 7a, 2 spms., PG 8, 2 spms., PG 11, 12 spms., PG 12, 4 spms., PG 13, 22 spms., PG 25a, 2 spms., PG 26, 3 spms., PG 27a, 13 spms., PG 58a, 19 spms., NP 9846, 3 spms., NP 9848, 38 spms., private collection; Sant’Agata Fossili: 4 spms., MGPT-PU 23321, 1 spm., NP 9849, 10 spms., PG 18, 3 spms., PG 23a, 7 spms., PG 59b, 2 spms., PG 73; Stazzano: 6 spms., MGPT-PU 23408, 5 spms., NP 9850, 1 spm., MPUM 11303, 1 spm., MZB 60033, 26 spms., private collection; Termofourà: 2 spms., MGPT-PU 23518; Tetti Civera: DHW PD H D SH 0.1080.136 0.8731.001 9.31125.467 8.38922.077 1.6688.272 0.122 0.937 17.389 15.233 4.970 AH AW UW IS SA 6.93217.996 4.23011.450 1.619-5.159 15°-35° 94°-122° 12.464 7.840 3.389 25° 108° Remarks. The specimens listed above agree in all characters with those described by Pedriali & Robba (2009); for extensive description and remarks, reference to the cited authors. The specimen 4680 in Museo Civico di Storia Naturale di Milano is the neotype of Nerita helicina Brocchi, 1814 (International Commission on Zoological Nomenclature 2010, Opinion 2247). Stratigraphic occurrence. Euspira helicina helicina (Brocchi, 1814) ranges continuously from the Burdigalian to the Pliocene of Italy. Other reliable occurrences are from the Burdigalian of France and North Sea Basin, from the Serravallian of Turkey, and from the Pliocene of France. Pedriali & Robba (2009) pointed out that it was replaced by Euspira helicina fusca (Blainville, 1825) subsequent to the Pliocene. Euspira latecallosa sp. n. Pl. 8, igs. 11-13 Derivation of name. From Latin late = widely and callosus, a, um = provided with a callus, with reference to the umbilical callus unusually wide for the genus. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Holotype: Borelli: MGPT-PU 135180 (Pl. 8, fig. 11). Paratypes: Borelli: 1 spm., MGPT-PU 135181, 1 spm., MGPT-PU 135182 (Pl. 8, fig. 12), 1 spm., MGPT-PU 135183 (Pl. 8, fig. 13), 8 spms., MGPT-PU 135184. Preservation: All specimens are rather well preserved. Type locality: Borelli (see appendix). Horizon: Gray, medium sand of Tortonian age. Diagnosis: Small, globose shell with scarcely elevated spire and globose last whorl. Umbilicus deep, narrowly crescentshaped, funicle a broad, depressed adaxial thickening of inner lip. Parietal callus thin medially; umbilical callus wide and thick, with convex abapertural outline, overhanging mid-adapical part of umbilicus and merging into anterior lobe of parietal callus. Description. Protoconch medium-sized, low-turbiniform, of 2.87-3.06 convex, smooth whorls, tip small. Teleoconch globose, thin, hardly exceeding six mm in height. Spire broadly conical, scarcely elevated, whorls gently convex; suture adpressed, nearly flush. Last whorl globose to globose-oval, approximately nine-tenth of total height, slightly produced abapically toward aperture; subsutural shelf indistinct; periphery well above midline. Aperture ovately D-shaped in prosocline plane, length about 1.6 times width; outer lip nearly semicircular, inner lip subangular in the middle. Parietal callus thin medially, slightly thickened at both ends, with concave abapertural outline, anterior lobe small, at level of umbilical border, poorly differentiated in a few specimens. Umbilicus deep, crescentshaped, very narrow, slightly less so in a few specimens; umbilical wall steep. Funicle a broad, depressed adaxial thickening of inner lip, soon obsolete toward interior of umbilicus. Umbilical callus wide and thick, with convex abapertural outline, expanded to overhang mid-adapical part of umbilicus, merging into anterior lobe of parietal callus; weak oblique depression at adapical end of umbilical callus present in most specimens. Basal fasciole indistinct. Outer surface with fine, incised growth lines. Dimensions (mm) DHW PD H D SH 0.0960.112 1.1301.190 2.5046.408 2.3906.054 0.4491.357 0.104 1.160 4.456 4.222 0.903 AH AW UW IS SA 1.9795.127 1.4663.886 0.8091.981 15°-27° 104°-136° 3.553 2.676 1.395 21° 120° Remarks. Euspira latecallosa exhibits teleo- 169 conch characters strikingly similar to those of the Burdigalian to Serravallian species Euspira submamillaris (Sacco, 1890). The two taxa can be differentiated from one another on the basis of their larval shells: that of E. submamillaris has a significantly smaller diameter of the initial halfwhorl. Further, Sacco’s species attains a larger size and has a moderately wide umbilical opening. Euspira latecallosa somewhat resembles Euspira grossularia (Marche-Marchad, 1957) in shell shape and the respective values of the characteristic elements of the protoconchs do not differ significantly. However, E. grossularia is readily distinguished from E. latecallosa in that it has: 1) the protoconch I spirally sculptured, 2) a more inflated last whorl slightly expanded abapically toward the aperture, 3) a small, oval umbilicus, 4) a triangular umbilical callus; it is also worth noting that E. grossularia attains a larger size. The Tortonian to Recent species Euspira notabilis (Jeffreys, 1885) and Euspira pulchella (Risso, 1826) are also similar in teleoconch shape. E. notabilis differs from Euspira latecallosa because of its protoconch with fewer whorls (2.12-2.25 instead of 2.87-3.06), spirally sculptured tip and significantly smaller diameter and diameter of the first half-whorl, its more widely open umbilicus, and its narrower umbilical callus. E. pulchella (Risso, 1826) is distinguished from E. latecallosa in having a protoconch with significantly smaller diameter and sculptured tip, a more elevated spire, a wider umbilicus, an obsolete funicle, and a narrowly subtriangular umbilical callus. The values of the characteristic elements of the protoconch of Euspira latecallosa do not differ significantly from those measured for the Burdigalian to Pliocene subspecies Euspira helicina helicina (Brocchi, 1814), but E. latecallosa can be distinguished easily from E. helicina helicina because of its smooth protoconch (that of E. helicina helicina has a spirally sculptured tip), its lower spire, its smaller to poorly differentiated anterior lobe of the parietal callus, its wider, abaperturally convex umbilical callus, and its smaller size. Stratigraphic occurrence. Euspira latecallosa sp. n. was recovered only from Tortonian deposits at the type locality. Robba E., Pedriali L. & Quaggiotto E. 170 Euspira molarensis sp. n. Pl. 8, igs. 14, 15 Derivation of name: The name refers to the village of Molare, which is the type locality. Holotype: Molare: MPUM 11307 (Pl. 8, ig. 14). Paratypes. Mioglia: 1 spm., MPUM 11308 (Pl. 8, ig. 15); Molare: 1 spm., MPUM 11309; Monte Gloso: 1 spm., MPUM 11310. Other material examined: Madonna delle Rocche: 1 spm., PG 49; Molare: 3 spms., PG 50; Monte Gloso: 1 spm., NP 9839, 1 spm., private collection. Preservation: Most specimens have minor damages, some are rather well preserved. Type locality: Molare (see appendix). Horizon: Gray, coarse sandstone belonging to the upper part of the Molare Formation; the age is late Rupelian. Diagnosis: Globose-oval shell with moderately elevated spire and oval last whorl. Umbilicus deep, rather wide to wide, subcircular to subquadrangular, funicle absent. Parietal callus thick, truncated abapically, with blunt, transverse adapical bulge separated from outer lip by narrow groove; umbilical callus obsolete, short and narrowly triangular in some specimens. Description. Protoconch small, lowturbiniform, of 2.5 convex, apparently smooth whorls, tip very small. Teleoconch globose-oval, slightly higher than wide, rather thick. Spire broadly conical, moderately elevated, whorls gently convex; suture adpressed, almost lush on earlier whorls. Last whorl oval, approximately nine-tenth of total height; subsutural shelf indistinct; periphery nearly at midline. Aperture D-shaped in slightly prosocline plane, length about two times width; outer lip regularly arched, adapical half of inner lip very slightly concave. Parietal callus thick, subrectangular, obliquely truncated abapically at level of umbilical border, expanded adapically toward interior of aperture and forming a blunt transverse bulge separated from outer lip by long, narrow furrow; anterior lobe indistinct. Umbilicus deep, subcircular to subquadrangular, moderately wide to wide; umbilical wall steep, with coarse growth markings. Funicle absent. Umbilical callus obsolete, short, very narrowly triangular in a few specimens. Basal fasciole indistinct. Outer surface with ine, dense growth lines. Dimensions (mm) DHW PD H D SH 0.078* 0.674* 7.01624.892 7.85321.285 0.6978.629 * 1 protoconch measurable 15.954 14.569 4.663 AH AW UW IS SA 5.58516.997 4.69112.187 1.8929.196 11°-27° 81°-129° 11.291 8.439 5.544 19° 105° Remarks. The small protoconch, the characters of the parietal callus, the widely open umbilicus devoid of funicle, and the obsolete umbilical callus combined distinguish Euspira molarensis from the other Euspira species covered in the present study. Stratigraphic occurrence. Euspira molarensis sp. n. was recovered from lower Oligocene deposits of Piedmont, Liguria and Veneto. Euspira notabilis (Jeffreys, 1885) Pl. 8, figs. 16-18 1885 Natica notabilis Jeffreys, p. 31, pl. 4, figs. 1, 1a. 2009 Euspira notabilis - Pedriali & Robba, p. 400, pl. 2, figs. 1, 2; pl. 3, figs. 11, 12; pl. 4, figs. 12-14 (cum syn). Type material: See Pedriali & Robba (2009, p. 400). Material examined: Borelli: 1 spm., MGPT-PU 135185 (Pl. 8, fig. 16), 1 spm., MGPT-PU 135186 (Pl. 8, fig. 17) 1 spm., MPUM 11311, 3 spms., MPUM 11312, 3 spms., NP 9870, 11 spms., MGPT-PU 135187, 1 spm., MGPT-PU 135188 (Pl. 8, fig. 18); Rio di Bocca d’Asino: 2 spms., NP 9854, 1 spm., MZB 60103, 1 spm., MZB 60126. Dimensions (mm) DHW PD H D SH 0.0700.086 0.7250.817 1.72518.413 1.59817.130 0.2624.310 0.078 0.771 10.069 9.364 2.286 AH AW UW IS SA 1.15214.412 1.2668.954 0.5074.095 11°-27° 95°-119° 7.782 5.110 2.301 19° 107° Remarks. The small upper Miocene specimens listed above perfectly conform to the Pliocene ones dealt with by Pedriali & Robba (2009); the values of the characteristic elements of the protoconch are identical and the 95% confidence intervals obtained for Pliocene and upper Miocene protoconchs fully overlap one another. Pedriali & Robba (2009) described the protoconch as smooth throughout. The new indings from Borelli show that the protoconch I is sculptured with unevenly noded, thin and well spaced spiral threads (Pl. 8, fig. 16). Also Euspira grossularia (Marche-Marchad, 1957) has the protoconch I similarly sculptured, but the spirals are coarser and more closely set. For full description and other comments, see Pedriali & Robba (2009). Stratigraphic occurrence. Euspira notabilis (Jeffreys, 1885) was recorded from Torto- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy nian to Pleistocene deposits of Italy. Presently, it occurs in the western Mediterranean and in the eastern Atlantic, from Portugal to Mauritania. Euspira perforata (Deshayes, 1864) Pl. 9, figs. 1-3 1864 Natica perforata Deshayes, p. 46, pl. 72, figs. 9-11. 1888 Natica perforata - Cossmann, p. 161. 1902 Natica perforata - Cossmann, p. 63, pl. 7, figs. 5, 6. 1907 Natica perforata - Cossmann & Pissarro, pl. 9, fig. 61-11. 2011 Payraudeautia perforata - Caze et al., fig. 11B. 2012 Payraudeautia perforata - Caze et al., p. 36, pl. 13, figs. G, H. 2012 Natica perforata - Courville et al., pl. 5, fig. 17. Type material: Natica perforata Deshayes, lectotype (here designated): the shell figured by Deshayes (1864, pl. 72, figs. 9-11) and refigured herein (Pl. 9, fig. 1), EM 32699, Grignon (France). From Deshayes’ text, it appears that Natica perforata was introduced on the basis of specimens from many localities (possible paralectotypes), but these specimens were not identified so far (Robert Emmanuel, personal communication 2014). Other material examined Cava Grola: 1 spm., private collection, 1 spm., NP 9886, 1 spm., MPUM 11313 (Pl. 9, fig. 2); Cava Rossi: 1 spm., MCZ 4328; Damery: 13 spms., NP 9882, 1 spm., NP 9883, 1 spm., MPUM 11314, 3 spms., NP 9884, 1 spm., MPUM 11315; Villiers-Saint-Frédéric (France): 3 spms., NP 9885, 1 spm., MPUM 11316 (Pl. 9, fig. 3), 3 spms., MPUM 11317. Description. Protoconch small, depressed turbiniform, of 2.12-2.25 gently convex, smooth whorls, tip small. Teleoconch globose, moderately thick. Spire rather elevated, low in smaller specimens, almost lat-topped, whorls well convex; suture adpressed, slightly incised. Last whorl globular, slightly produced abapically; subsutural shelf indistinct; periphery above midline. Aperture Dshaped in prosocline plane, length about 1.5 times width; outer lip semicircular, inner lip almost straight. Parietal callus rather thin, with straight or gently concave abapertural outline; anterior lobe indistinct. Umbilicus medium-sized, deep; umbilical wall vertical. Interior of umbilicus with 4-9 uneven, rather coarse spiral cordlets crossed by growth markings, abapical one markedly broader, depressed to moderately prominent, obsolescent in a few specimens. Funicle absent. Umbilical callus moderately thick, triangular, relected over adapical part of umbilicus and merging into anterior corner of parietal callus. Basal fasciole indistinct. Outer surface with uneven growth markings. 171 Dimensions (mm) DHW PD H D SH 0.1080.128 0.8911.011 7.94410.056 8.0309.886 1.7613.313 0.118 0.951 9.000 8.958 2.537 AH AW UW IS SA 5.8877.039 4.5286.136 2.292-2.932 16°-32° 110°-130° 6.463 5.332 2.612 24° 120° Remarks. The four shells from Veneto have the protoconch abraded, but the teleoconch, identical to that of the French specimens listed above, makes their identiication certain. Natica perforata was recently included in Payaraudeautia Bucquoy, Dautzenberg & Dollfus, 1883 by some authors (Pacaud & Le Renard 1995; Caze et al. 2011; Caze et al. 2012). This generic allocation of N. perforata was probably based on its lowermost, broad spiral cordlet in the interior of the umbilicus, reminiscent of the inner spiral ridge or cord occurring abapical to the funicle in the species of Payraudeautia. However, the lack of a true (adapical or mid-adapical) funicle and the presence of additional spiral cordlets within the umbilicus (never present in Payraudeautia) make N. perforata unlike Payraudeautia species and, instead, support its assignment to the genus Euspira Agassiz in J. Sowerby, 1837. Stratigraphic occurrence. Euspira perforata (Deshayes, 1864) is deinitely known from the Early and Middle Eocene of France; it occurs rarely also in the middle Lutetian of Veneto (present paper). Euspira piccolii sp. n. Pl. 9, igs. 4, 5 Derivation of name: The species is named after the late Prof. Giuliano Piccoli (University of Padova), who extensively dealt with the Paleogene fossils of Veneto. Holotype: Cava Rossi, MPUM 11318 (Pl. 9, ig. 4). Paratypes: Cava Boschetto: 1 spm., MPUM 11319 (Pl. 9, ig. 5); Cava Rossi: 2 spms., MPUM 11320, 3 spms., MCZ 4329, 2 spms., MGP-PD 31524. Other material examined: Cava Grola: 1 spm., NP 9836; Cava Rossi: 3 spms., NP 9838, 6 spms., private collection; Le Coe: 1 spm., NP 9837. Preservation: Most specimens are rather well preserved. Type locality: Cava Rossi (see appendix). Horizon: Volcanoclastic layer of late Ypresian to earliest Lutetian age. Diagnosis: Depressed-globose shell with moderately elevated to low spire and globose last whorl. Umbilicus deep, medium- Robba E., Pedriali L. & Quaggiotto E. 172 sized, teardrop-shaped, with over 20 inner spiral cordlets; funicle absent. Parietal callus rather thin; anterior lobe slight to indistinct. Umbilical callus moderately thick, triangular, slender in most specimens, relected over adapical part of umbilicus, merging into anterior corner of parietal callus. Description. Protoconch large, lowturbiniform, of three convex, apparently smooth whorls, tip small, slightly sunken. Teleoconch depressed-globose, moderately thick. Spire broadly conical, moderately elevated, low in a few specimens, whorls well convex; suture narrowly and shallowly channeled. Last whorl globose, slightly wider than tall, markedly depressed in abapertural view, somewhat expanded toward aperture; subsutural shelf narrow, poorly deined; periphery above midline. Aperture D-shaped in slightly prosocline plane, length about 1.66 times width; outer lip regularly arched, inner lip almost straight. Parietal callus rather thin, with straight or gently concave abapertural outline; anterior lobe slight to indistinct. Umbilicus medium-sized, deep, asymmetrically teardropshaped; umbilical wall vertical. Interior of umbilicus with numerous (more than 20), rather even spiral cordlets crossed by growth markings. Funicle absent. Umbilical callus moderately thick, triangular, slender in most specimens, relected over adapical part of umbilicus and merging into anterior corner of parietal callus. Basal fasciole indistinct. Outer surface with remnants of ine growth lines, more evident over lowermost base. Dimensions (mm) DHW PD H D SH 0.1230.131 1.5742.022 11.06422.512 12.47022.262 2.0719.379 0.127 1.798 16.788 17.366 5.725 AH AW UW IS SA 8.21313.913 5.41112.855 3.580-7.472 8°-28° 99°-135° 11.063 9.133 5.526 18° 117° Remarks. Euspira piccolii is morphologically similar to the British, Bartonian species Euspira bartonensis Wrigley, 1949. We examined some topotypes of E. bartonensis from Middle Barton Beds at Barton and can state that Wrigley’s species differs from E. piccolii in that it has: 1) the protoconch half the size of that of E. piccolii, 2) the interior of the umbilicus bounded by a sharp ridge (absent in E. piccolii), and 3) the umbilical callus nearly semicircular, demarcated from the parietal callus by a distinct transver- se depression. Euspira perforata (Deshayes, 1864) is another similar species in shell shape, but differs from E. piccolii by its protoconch of 0.75 fewer whorls, with significantly smaller diameter, and its umbilicus with less numerous, coarser inner spiral cordlets. Stratigraphic occurrence. Euspira piccolii sp. n. was recovered from upper Ypresian to Priabonian units of Veneto, being more common in the earliest Lutetian. Euspira pulchella (Risso, 1826) Pl. 9, figs. 13, 14 1826 Natica pulchella Risso, p. 148, pl. 4, fig. 42. 2009 Euspira pulchella - Pedriali & Robba, p. 401, pl. 2, figs. 3, 4; pl. 3, fig. 13; pl. 4, figs. 15, 16 (cum syn.). Type material: Pedriali & Robba (2009, p. 401) reviewed the type material of Euspira pulchella, and described the characters of this species. Material examined: Borelli: 1 spm., MGPT-PU PLATE 9 Fig. 1 - Euspira perforata (Deshayes, 1864). Grignon (France). Lectotype (here designated) of Natica perforata Deshayes, 1864. EM 32699; apertural side. Fig. 2 - Euspira perforata (Deshayes, 1864). Cava Grola. MPUM 11313; apertural side (height of shell 9.30 mm). Fig. 3 - Euspira perforata (Deshayes, 1864). Villiers Saint Frédéric (France). MPUM 11316; protoconch. Fig. 4 - Euspira piccolii sp. n. Cava Rossi. Holotype, MPUM 11318; a, apertural side (height of shell 13.34 mm); b, apical view; c, basal view. Fig. 5 - Euspira piccolii sp. n. Cava Boschetto. Paratype, MPUM 11319; protoconch. Fig. 6 - Euspira submamillaris (Sacco, 1890). Colli Torinesi. Holotype of Natica (Naticina) catena var. proboniensis Sacco, 1890. MGPT BS.029.02.008; apertural side (scale bar 4 mm). Fig. 7 - Euspira submamillaris (Sacco, 1890). Colli Torinesi. Lectotype (here designated) of Natica (Polinices) submamillaris Sacco, 1890. MGPT BS.029.06.001; apertural side (scale bar 4 mm). Fig. 8 - Euspira submamillaris (Sacco, 1890). Albugnano. MPUM 11321; a, protoconch; b, detail of protoconch. Fig. 9 - Euspira submamillaris (Sacco, 1890). Albugnano. PG 44; protoconch. Fig. 10 - Euspira submamillaris (Sacco, 1890). Valle Ceppi. MPUM 11323; apertural side (height of shell 16.40 mm). Fig. 11 - Euspira submamillaris (Sacco, 1890). Valle Ceppi. MPUM 11324; apertural side (height of shell 14.95 mm). Fig. 12 - Euspira submamillaris (Sacco, 1890). Valle Vergnana. MZB 60022; protoconch. Fig. 13 - Euspira pulchella (Risso, 1826). Borelli. MGPT-PU 135197; apertural side (height of shell 7.20 mm). Fig. 14 - Euspira pulchella (Risso, 1826). Rio di Bocca d’Asino. MPUM 11326; a, protoconch; b, detail of protoconch. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 173 Robba E., Pedriali L. & Quaggiotto E. 174 135197 (Pl. 9, ig. 13), 19 spms., MGPT-PU 135198, 1 spm., MGPTPU 135199, 1 spm., MGPT-PU 135200, 1 spm., MGPT-PU 135201, 1 spm., MGPT-PU 135202, 1 spm., MGPT-PU 135203, 2 spms., MPUM 11325, 4 spms., NP 9871, 6 spms., PG 22a; Rio di Bocca d’Asino: 1 spm., MPUM 11326 (Pl. 9, ig. 14), 6 spms., PG 22, 2 spms., NP 9840; Sogliano al Rubicone: 1 spm., PP.PM 01/02. Dimensions (mm) DHW PD H D SH 0.0720.092 0.7140.804 1.37212.204 0.95011.042 0.6573.191 0.082 0.738 6.788 5.996 1.899 AH AW UW IS SA 0.4569.344 0.6165.940 0.296-3.048 12°-24° 77°-121° 4.900 3.278 1.672 18° 99° Remarks. The examined upper Miocene specimens conform to the Pliocene and Recent ones dealt with by Pedriali & Robba (2009) in terms of values of the characteristic elements of the protoconch, sculpture of protoconch I, teleoconch shape, umbilical characters and color pattern. For full description and comments see Pedriali & Robba (2009). Stratigraphic occurrence. Euspira pulchella (Risso, 1826) was known to range from Pliocene to Recent. The new recoverings listed above show that the species was already present in the Tortonian of Piedmont and Emilia. Euspira pulchella presently is distributed from Iceland and northern Norway and southward into the Mediterranean. Euspira submamillaris (Sacco, 1890) comb. n. Pl. 9, igs. 6-12 1847 Natica mamillaris - Sismonda, p. 51 (not Natica mamillaris Lamarck, 1822 = Albula hepatica Röding, 1798). 1852 Natica submamillaris d’Orbigny, p. 38 (nomen nudum). 1890b Natica (Naticina) catena var. proboniensis Sacco, p. 30 (new synonym). 1890b Natica (Polinices) submamillaris Sacco, p. 35. 1890b Natica (Polinices) submamillaris var. mioinlata Sacco, p. 35. 1890b Natica (Polinices) submamillaris var. mioaperta Sacco, p. 36. 1890b Natica (Polinices) submamillaris var. mioclausa Sacco, p. 36. 1891 Natica (Naticina) catena var. proboniensis - Sacco, p. 70, pl. 2, ig. 42. 1891 Natica (Polinices) submamillaris - Sacco, p. 90, pl. 2, ig. 62. 1891 Natica (Polinices) submamillaris var. mioinlata - Sacco, p. 91, pl. 2, ig. 64. 1891 Natica (Polinices) submamillaris var. mioaperta - Sacco, p. 91, pl. 2, ig. 65. 1891 Natica (Polinices) submamillaris var. mioclausa - Sacco, p. 91, pl. 2, ig. 66. not 1956 Polynices (Polynices) submamillaris - Banke Rasmussen, p. 60, pl. 5, ig. 1 (not Sacco, 1890). not 1958 Natica (Naticina) submamillaris - Sorgenfrei, p. 192, pl. 36, ig. 124 (not Sacco, 1890). 1984 Naticina catena var. probononiensis (sic) - Ferrero Mortara et al., p. 32. 1984 Polinices submamillaris - Ferrero Mortara et al., p. 36 (not BS.029.06.002 = Natica (Polinices) submamilla Sacco, 1891). 1984 Polinices submamillaris var. mioinlata - Ferrero Mortara et al., p. 36. 1984 Polinices submamillaris var. mioaperta - Ferrero Mortara et al., p. 36. 1984 Polinices submamillaris var. mioclausa - Ferrero Mortara et al., p. 36. 2007 Polinices submamillaris - Zunino, p. 121, pl. 1, ig. 7. Type material: Natica (Polinices) submamillaris Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 2, ig. 62) and reigured herein (Pl. 9, ig. 7), MGPT BS.029.06.001, Colli Torinesi. Natica (Polinices) submamillaris var. mioinlata Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 2, ig. 64), MGPT BS.029.06.004, Colli Torinesi; 11 paralectotypes, MGPT BS.029.06.004/01, Colli Torinesi (other 2 syntypes are unidentiiable, 1 could be a species of Cochlis, 5 are Natica (Polinices) proredempta Sacco, 1890, all also numbered MGPT BS.029.06.004/01). Natica (Polinices) submamillaris var. mioaperta Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 2, ig. 65), MGPT BS.029.06.005, Colli Torinesi; 17 paralectotypes, MGPT BS.029.06.005/01, Colli Torinesi. Natica (Polinices) submamillaris var. mioclausa Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 2, ig. 66), MGPT BS.029.06.006, Colli Torinesi; 11 paralectotypes, MGPT BS.029.06.006/01, Colli Torinesi. Natica (Naticina) catena var. proboniensis Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 2, ig. 42) and reigured herein (Pl. 9, ig. 6), MGPT BS.029.02.008, Colli Torinesi. Other material examined: Albugnano: 1 spm., MPUM 11321 (Pl. 9, ig. 8), 1 spm., PG 44 (Pl. 9, ig. 9), 3 spms., MPUM 11322, 11 spms., PG 42; Mioglia: 1 spm., PG 43; Miste (The Netherlands): 42 spms., NP 9826; Monte dei Cappuccini: 1 spm., MGPTPU; Termofourà: 1 spm., MGPT-PU 135190; Tetti Civera: 1 spm., MZB 15799; Valle Ceppi: 26 spms., MZB 60119, 5 spms., MGPTPU 25264, 6 spms., MGPT-PU 23860, 1 spm., MGPT-PU 26143, 15 spms., PG 45, 15 spms., MGPT-PU 107939, 15 spms., MGPTPU 23664, 1 spm., MZB 28566, 1 spm., MZB 60026, 7 spms., MZB 60027, 3 spms., MZB 60021, 1 spm., MPUM 11323 (Pl. 9, ig. 10), 1 spm., MPUM 11324 (Pl. 9, ig. 11), 2 spms., NP 9827, 8 spms., MZB 60056, 1 spm., MZB 60057, 1 spm., MZB 60058, 10 spms., MZB 60059, 1 spm., MZB 60061, 1 spm., MGPT-PU 135191, 1 spm., MGPT-PU 135192, 1 spm., MGPT-PU 135193, 1 spm., MGPT-PU 135194, 1 spm., PG 75; Val Sanfrà: 2 spms., MGPT-PU 107621, 3 spms., MGPT-PU 135195; Valle Vergnana: 2 spms., MZB 23757, 6 spms., MZB 43239, 2 spms., MZB 60054, 1 spm., MZB 60023, 1 spm., MZB 60022 (Pl. 9, ig. 12), 6 spms., MZB 60024, 1 spm., MZB 60025, 2 spms., PG 77; Villa Allason: 1 spm., MGPT-PU 107620, 2 spms., MGPT-PU 107623, 5 spms., MGPT-PU 107954; Villa Bertini: 2 spms., MGPT-PU 107622. Description. Protoconch almost mediumsized, low-turbiniform, of 3-3.25 gently convex whorls, tip very small; one specimen from Albugnano with remnants of straight collabral riblets (Pl. 9, ig. 8b). Teleoconch globose-oval to oval, thick. Spire conical to broadly conical, pointed, moder- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy ately elevated, less so in a few specimens; whorls gently convex to lat-sided; suture thin, almost lush. Last whorl globose-oval, globose in a few specimens, moderately extended toward aperture; subsutural shelf indistinct, very slightly concave but poorly deined in specimens with more elevated spire; periphery above midline. Aperture somewhat bean-shaped in prosocline plane, length about twice width; outer lip arched, inner lip markedly thick, slightly bent abapically. Parietal callus thick, subrectangular, rather short, ending in slight anterior lobe below border of umbilical area. Umbilicus deep, narrow to semicircular and largely open, umbilical wall subvertical, with coarse growth ridges. Funicle a broad, depressed adaxial thickening of inner lip, soon obsolete toward interior of umbilicus. Umbilical callus at submedian part of inner lip, thick, subtriangular, with straight or reverse S-shaped adaxial outline, overhanging adapical part of umbilicus and merging into anterior lobe of parietal callus. Basal fasciole indistinct. Outer surface with dense growth lines prosocyrt adapically. Some specimens retain vestige of brown subsutural band. Dimensions (mm) DHW PD H D SH 0.0680.076 0.9291.057 10.08319.251 9.41217.224 0.6273.311 0.072 0.993 14.667 13.318 1.969 AH AW UW IS SA 8.78416.604 4.5899.157 2.1084.776 9°-29° 97°-121° 12.694 6.873 3.442 19° 109° Remarks. Sismonda (1847, p. 51) used the binomen Natica mamillaris Lamarck, 1822 for Miocene naticids from the Torino Hills; he did not provide any description or illustration of his material. Since N. mamillaris is a Recent Caribbean species, d’Orbigny (1852, p. 38) introduced the replacement name submamillaris for the fossil shells dealt with by Sismonda. D’Orbigny just listed (n° 566) a Natica submamillaris d’Orbigny, 1847, but we failed to locate this reference. The name N. submamillaris was reported as of d’Orbigny, 1852 by all subsequent authors. However, from the 1852 compilation, it appears that the name submamillaris is a nomen nudum and, thence, unavailable. Sacco (1890b, p. 35), also dealing with Miocene naticids from the Torino Hills, irst published a concise diagnosis of N. submamillaris, thus making the name available, and is to be considered 175 the author of the species. N. mamillaris is currently regarded as a synonym of Albula hepatica Röding, 1798 (cf. Warmke & Tucker Abbott 1975; Kabat et al. 1997). N. submamillaris was currently included in Polinices Montfort, 1810 by the authors. Instead, we think that it fully agrees with the characters of the genus Euspira Agassiz in J. Sowerby, 1837. Natica submamillaris appears to be a variable species in teleoconch shape, spire height and amplitude of the umbilical opening. This fact induced Sacco (1890b, 1891) to propose the varieties praenuntia, submioclausa, mioinlata, mioaperta and mioclausa. The vast material examined has shown that the varieties mioinlata, mioaperta and mioclausa (of Sacco, 1890b) cannot be separated consistently from one another and from the typical form because of gradual transitions among them. Thus, they are included in the synonymy of N. submamillaris. The variety praenuntia Sacco, 1890 has a narrow, oblique umbilical opening and an arched adaxial outline of the umbilical callus, characters that are more similar to those of Tectonatica miocolligens (Sacco, 1890). On this basis, we consider the variety praenuntia as a synonym of T. miocolligens. Nothing can be reliably said about the variety submioclausa Sacco, 1891, reported to occur rarely in lower Oligocene units of Piedmont, since no specimens originally assigned to it were found in MGPT. Sacco (1890b) briely described the new variety proboniensis of Natica catena (da Costa, 1778) from lower Miocene deposits of Colli Torinesi (Torino Hills). Examination of the unique specimen (holotype) of the variety proboniensis in MGPT demonstrated that it is unrelated to N. catena because of its smaller umbilicus devoid of inner spiral cordlets (present and numerous in N. catena). Consequently, the variety proboniensis is herein ranked at species level and considered distinct from N. catena. The comparison of the original material of Natica submamillaris with the holotype of Natica proboniensis has shown that these two taxa are congeneric and that no reliable difference exists between them. We conclude that N. submamillaris and N. proboniensis are synonyms. Both N. submamillaris and N. proboniensis were published simultaneously in 1890 (see the above synonymy). The valid name is submamillaris, which was proposed at higher rank (ICZN 1999, Article 24 of the Code). It is also worth noting that the name submamillaris is that in current usage. Euspira submamillaris supericially resembles Euspira helicina helicina (Brocchi, 1814) in shell shape, Robba E., Pedriali L. & Quaggiotto E. 176 but can be differentiated from it because of its protoconch with signiicantly smaller diameter of the initial half-whorl, and its almost lat-sided spire whorls (convex in Brocchi’s taxon). The Miocene specimens from Denmark referred to as Polynices (Polynices) submamillaris or as Natica (Naticina) submamillaris (see the above synonymy) are similar to E. submamillaris, but are distinguished from it in that they have a protoconch of 0.5 fewer whorls (cf. Sorgenfrei 1958), a more inlated teleoconch with depressed, obtuse spire and a different course of the inner lip. Stratigraphic occurrence. Euspira submamillaris Sacco, 1890 occurs abundantly in the Burdigalian of Piedmont, less so in the Serravallian of Piedmont; we have only one specimen from the early Oligocene of Liguria. The species is also present in the Burdigalian of Miste, The Netherlands (specimens in our collection). Euspira subobturata (Sacco, 1891) stat. n., comb. n. Pl. 10, igs. 1-4 1890b Natica (Naticina) catena var. helicina subvar. subopturata Sacco, p. 30 (nomen nudum). 1891 Natica (Naticina) catena var. subobturata Sacco, p. 72. 1904 Naticina catena var. subobturata - Sacco, p. 103, pl. 22, igs. 31, 32. 1984 Naticina catena var. subopturata (sic) - Ferrero Mortara et al., p. 32. Type material: Natica (Naticina) catena var. subobturata Sacco, lectotype (designated by Pedriali & Robba 2009, p. 396): the shell igured by Sacco (1904, pl. 22, ig. 32) and reigured herein (Pl. 10, ig. 1), MGPT BS.029.02.011, Savona; 1 paralectotype, MGPT BS.029.02.010, Colli Torinesi; 28 paralectotypes, MGPT BS.029.02.011/04, Savona. Other material examined: Albugnano: 1 spm., MPUM 11327 (Pl. 10, ig. 2), 1 spm., MPUM 11328, 3 spms., PG 56; Valle Ceppi: 1 spm., NP 9853, 1 spm., MZB 60123, 1 spm., MZB 60151 (Pl. 10, ig. 3), 1 spm., MZB 60124, 1 spm., MZB 60125 (Pl. 10, ig. 4), 1 spm., MGPT-PU 135205; Val Sanfrà: 5 spms., MGPT-PU 135206. Description. Protoconch medium-sized, lowturbiniform, of 2.08-2.25 convex, apparently smooth whorls, tip small. Teleoconch globose to globose-oval, rather thick. Spire broadly conical, pointed, moderately elevated; whorls gently convex to nearly lat-sided; suture thin, adpressed, almost lush in some specimens. Last whorl globose, subsutural shelf indistinct, periphery well above midline. Aperture D-shaped in prosocline plane, length about 1.8 times width; outer lip regularly arched, inner lip straight, thickened abapically. Parietal callus thick, subrectangular, extended abapically and largely covering the umbilicus; anterior lobe expanded adaxially, broad, obtusely rounded, ending at level of upper umbilical wall. Umbilicus deep, small to very small, narrowly oval; umbilical wall subvertical, bounded downward by shallow groove. Funicle only partially exposed, broad, depressed, obsolescent toward interior of umbilicus. Umbilical callus at mid-abapical part of inner lip, very short, thick, subtriangular, with straight or reverse J-shaped adaxial outline, merging into anterior lobe of parietal callus. Basal fasciole broad and markedly depressed to poorly differentiated, producing abapical tickening of inner lip. Outer surface with dense growth lines prosocyrt adapically. Some specimens retain vestige of brown subsutural band. Dimensions (mm) DHW PD H D SH 0.1740.214 1.1171.193 7.92815.724 6.70815.660 1.3533.973 0.194 1.155 11.826 11.184 2.663 AH AW UW IS SA 6.36111.981 4.3068.854 1.2713.675 15°-31° 98°-134° 9.171 6.580 2.473 23° 116° PLATE 10 Fig. 1 - Euspira subobturata (Sacco, 1891). Savona. Lectotype (here designated) of Natica (Naticina) catena var. subobturata Sacco, 1891. MGPT BS.029.02.011; apertural side (scale bar 4 mm). Fig. 2 - Euspira subobturata (Sacco, 1891). Albugnano. MPUM 11327; protoconch. Fig. 3 - Euspira subobturata (Sacco, 1891). Valle Ceppi. MZB 60151; apertural side (height of shell 13.07 mm). Fig. 4 - Euspira subobturata (Sacco, 1891). Valle Ceppi. MZB 60125; apertural side (height of shell 14.04 mm). Fig. 5 - Euspira umbilicocarinata sp. n. Borelli. Holotype, MGPT-PU 135207; a, apertural side (height of shell 5.61 mm); b, protoconch. Fig. 6 - Euspira umbilicocarinata sp. n. Rio di Bocca d’Asino. Paratype, MPUM 11329; protoconch. Fig. 7 - Euspira umbilicolunata sp. n. Valle Ceppi. Holotype, MPUM 11331; apertural side (height of shell 15.98 mm). Fig. 8 - Euspira umbilicolunata sp. n. Valle Ceppi. Paratype, MPUM 11333; protoconch. Fig. 9 - Euspira sp. 1. Cava Albanello. NP 9851; apertural side (height of shell 22.30 mm). Fig. 10 - Euspira sp. 2. Valle Vergnana. MZB 43250; apertural side (height of shell 11.18 mm). Fig. 11 - Euspira sp. 2. Valle Vergnana. MZB 60132; protoconch. Fig. 12 - Euspira sp. 3. Case Soghe. MPUM 11335; a, apertural side (height of shell 12.34 mm); b, apical view. Fig. 13 - Euspira sp. 4. Cassinelle. MPUM 11336; apertural side (height of shell 6.59 mm). Fig. 14 - Euspira sp. 4. Monte Gloso. MPUM 11337; protoconch. Fig. 15 - Euspira sp. 5. Albugnano. MPUM 11338; a, apertural side; b, protoconch. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 177 Robba E., Pedriali L. & Quaggiotto E. 178 Remarks. Sacco (1890b) listed the subvar. subopturata of Natica (Naticina) catena var. helicina (Brocchi, 1814) without description or illustration; the name subopturata results to be nomen nudum. One year later (1891), Sacco published a brief description of this form using the name subobturata, hence making this name available, with the authorship of Sacco, 1891. Sacco (1891) regarded subobturata as a variety of Natica (Naticina) catena (da Costa, 1778), distinct from the variety helicina. Da Costa’s and Brocchi’s taxa are currently assigned to the genus Euspira Agassiz in J. Sowerby, 1837. In a recent paper (Pedriali & Robba 2009), Natica (Naticina) catena var. subobturata was included in the synonymy of Euspira helicina helicina (Brocchi, 1814). Re-examination of the original material in MGPT and of additional Miocene specimens retaining the protoconch induced us to consider Sacco’s taxon as a distinct species. Euspira subobturata differs from E. helicina helicina in that it has: 1) a protoconch with signiicantly greater diameter, 2) a lower spire with less convex whorls, 3) a longer parietal callus covering the mid-adapical part of the umbilicus, and 4) a signiicantly shorter umbilical callus located more abapically than that of E. helicina helicina. The lower Miocene, French species Euspira benoisti (Cossmann & Peyrot, 1919) is similar to Euspira subobturata in shell shape, inner lip calluses and umbilical characters. We examined some specimens from Lariey and noted that E. benoisti is readily distinguished from E. subobturata by its protoconch with signiicantly smaller diameter of the sculptured irst half-whorl. Stratigraphic occurrence. Euspira subobturata (Sacco, 1891) is known from Burdigalian, Langhian and Piacenzian units of northwestern Italy. Euspira umbilicocarinata sp. n. Pl. 10, igs. 5, 6 Derivation of name: From Latin umbilicus = umbilicus and carinatus = shaped like a keel, with reference to the keel-like angulation bounding the interior of the umbilicus. Holotype: Borelli: MGPT-PU 135207 (Pl. 10, ig. 5). Paratypes: Borelli: 1 spm., MGPT-PU 135208, 1 spm., MGPT-PU 135209, 1 spm., MGPT-PU 135210, 1 spm., MGPTPU 135211, 48 spms., MGPT-PU 135212, 1 spm., NP 9873; Monte dei Cappuccini: 2 spms., MGPT-PU 135213, 16 spms., MGPT-PU 135214; Rio di Bocca d’Asino: 1 spm., MPUM 11329 (Pl. 10, ig. 6), 1 spm., MPUM 11330; Sant’Agata Fossili: 5 spms., MGPT-PU 135215. Other material examined: Albugnano: 6 spms., PG 61; Rio di Bocca d’Asino: 1 spm., PG 60, 1 spm., NP 9858, 2 spms., NP 9859. Preservation: Most specimens are rather well preserved; some have minor damages to the outer lip. Type locality: Borelli (see appendix). Horizon: Gray, medium sand of Tortonian age. Diagnosis: Protoconch I with nodulose spiral cordlets. Small, globose shell with scarcely elevated spire and globose last whorl. Umbilicus deep, rather wide in most specimens, subrectangular; umbilical wall bounded downward by keel-like angulation; funicle markedly depressed. Parietal callus thick, with small, subangular anterior lobe. Umbilical callus short, thick, subtriangular; faint transverse depression between umbilical and parietal calluses present in some specimens. Description. Protoconch small, lowturbiniform, of 2.75-3 convex whorls, tip small; protoconch I with remnants of irregularly nodulose spiral cordlets. Teleoconch small-sized, globose, moderately thick. Spire broadly conical, scarcely elevated, slightly higher in a few specimens; whorls gently convex; suture thin, adpressed. Last whorl globose, subsutural shelf indistinct, periphery well above midline. Aperture D-shaped in slightly prosocline plane, length exceeding twice width; outer lip semicircular, inner lip straight, thickened abapically. Parietal callus rather thick, less so medially, with concave, subangular abapertural outline; anterior lobe small, subangular, ending at level of upper umbilical wall. Umbilicus deep, rather wide, less so in a few specimens, obliquely subrectangular; umbilical wall subvertical, bounded downward by distinct, keel-like angulation. Funicle markedly depressed, obsolescent toward interior of umbilicus, separated from keel-like angulation by broad, shallow depression forming lat sinus on abapical part of inner lip. Umbilical callus short, thick, subtriangular, with straight or reverse S-shaped adaxial outline, slightly overhanging umbilicus and merging into anterior lobe of parietal callus; faint transverse depression may occur between umbilical and parietal calluses. Basal fasciole indistinct. Outer surface with ine growth lines prosocyrt adapically. Specimens from Rio di Bocca d’Asino retain vestige of pale brown background. Dimensions (mm) DHW PD H D SH 0.1020.126 1.1191.203 0.92012.492 0.94411.800 0.0403.156 0.114 1.161 6.706 6.372 1.598 AH AW UW IS SA 0.8619.233 0.3337.885 0.030-3.710 12°-32° 98°-130° 5.047 4.109 1.870 22° 114° Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Remarks. The present new species is characterized by the keel-like angulation forming a distinct step that demarcates the umbilical wall from the innermost part of the umbilicus. Euspira latecallosa sp. n. (see above) is the most closely similar species in shell shape and the values of the characteristic elements of the protoconch are identical to those measured for Euspira umbilicocarinata. However, E. umbilicocarinata is readily differentiated from E. latecallosa because of its rather wide, subrectangular umbilicus (that of E. latecallosa is narrower, crescent-shaped), its narrower umbilical callus with straight or reverse S-shaped adaxial outline (that of E. latecallosa has a well convex adaxial outline), and, principally, its umbilical wall bounded downward by a distinct keel-like angulation. The relationships with Euspira gianoi sp. n. have been already dealt with (see above). Other similar species are Euspira grossularia (Marche-Marchad, 1957), Euspira helicina helicina (Brocchi, 1814), Euspira notabilis (Jeffreys, 1885) and Euspira pulchella (Risso, 1826). Euspira umbilicocarinata differs from E. grossularia in having a protoconch with signiicantly greater diameter of the irst half-whorl, a spire with less convex whorls, a less globular last whorl, and the umbilicus with an inner step (absent in E. grossularia). Concerning E. helicina helicina, the values of the characteristic elements of the protoconch do not differ signiicantly from those measured for E. umbilicocarinata, but the teleoconch characters of E. umbilicocarinata distinguish it easily from E. helicina helicina. E. umbilicocarinata differs from E. helicina helicina by its smaller size, its lower-spired shell, its smaller anterior lobe of the parietal callus, its rather wide, subrectangular umbilicus with the innermost part bounded by a distinct step, and its noticeably wider depression separating the funicle from the base of the umbilical wall. E. umbilicocarinata is distinguished from E. notabilis because of its protoconch with over 0.5 more whorls and signiicantly greater diameter and diameter of the initial hal-whorl, its lower spire, and its different umbilical characters (E. notabilis has a crescent-shaped instead of subrectangular umbilicus, devoid of any inner step). The protoconch with signiicantly greater diameter and diameter of the first half-whorl is the primary element differentiating E. umbilicocarinata from E. pulchella. 179 Stratigraphic occurrence. Euspira umbilicocarinata sp. n. was recovered from Langhian to Tortonian deposits of Piedmont. Euspira umbilicolunata sp. n. Pl. 10, igs. 7, 8 Derivation of name: From Latin umbilicus = umbilicus and lunatus = crescentic, with reference to the narrow, crescent-shaped umbilicus. Holotype: Valle Ceppi: MPUM 11331 (Pl. 10, ig. 7). Paratypes: Grazzano Badoglio: 2 spms., MPUM 11332, 4 spms., MZB 60031, 13 spms., MZB 29800; Valle Ceppi: 1 spm., MGPT-PU 135216, 1 spm., MPUM 11333 (Pl. 10, ig. 8), 1 spm., MZB 60030, 8 spms., MGPT-PU 135217, 2 spms., MGPT-PU 135218, 1 spm., MGPT-PU 135219, 1 spm., MGPT-PU 135220, 6 spms., MGPT-PU 135221, 1 spm., MGPT-PU 135222, 4 spms., MZB 60120; Tetti Civera: 2 spms., MZB 60032; Val Sanfrà: 1 spm., MGPTPU 135223; Villa Allason: 23 spms., MGPT-PU 135224; Villa Bertini: 21 spms., MGPT-PU 108782. Other material examined: Grazzano Badoglio: 6 spms., PG 51, 1 spm., NP 9842, 4 spms., PG 76b; Valle Ceppi: 2 spms., NP 9841. Preservation: Most specimens are rather well preserved; some have minor damages. Type locality: Valle Ceppi (see appendix). Horizon: Coarse, gray sand (Termofourà Formation) of Burdigalian age. Diagnosis: Globose to globose-oval shell with rather elevated spire and globose last whorl. Umbilicus narrow to very narrow, crescent-shaped; umbilical wall with coarse growth markings; funicle broad, markedly depressed. Parietal callus rather thick; anterior lobe slight to indistinct. Umbilical callus slender, merging into anterior corner of parietal callus; slight transverse depression between umbilical and parietal calluses present in some specimens. Uniform pale brown color throughout. Description. Protoconch medium-sized, low-turbiniform, of 2.85-3 convex, apparently smooth whorls, tip very small, slightly sunken. Teleoconch globose to globose-oval, moderately thick. Spire conical, rather elevated, less so in a few specimens, whorls convex. Suture adpressed; slight subsutural margining present in some specimens. Last whorl globose, depressed in abapertural view, scarcely produced toward aperture; subsutural shelf poorly deined to absent; periphery about at midline. Aperture D-shaped in slightly prosocline plane, length about 1.76 times width; outer lip regularly arched, basal lip slightly thickened, inner lip almost straight. Parietal callus rather thick, with concave abapertural outline; anterior lobe slight to indistinct. Umbilicus narrow to very narrow, crescent-shaped, somewhat wider in a few larger specimens; umbilical wall with coarse growth markings. Funicle broad, Robba E., Pedriali L. & Quaggiotto E. 180 markedly depressed. Umbilical callus rather slender, with gently convex to convex adaxial outline, merging into anterior corner of parietal callus; slight transverse depression in between umbilical callus and parietal callus present in some specimens. Basal fasciole indistinct. Outer surface with ine growth lines slightly bent subsuturally. Best preserved specimens retain uniform pale brown color throughout. Dimensions (mm) DHW PD H D SH 0.0730.089 1.2291.265 8.25916.103 7.29714.485 2.2436.023 0.081 1.247 12.181 10.891 4.133 AH AW UW IS SA 5.61610.396 3.8008.788 1.9293.977 13°-29° 88°-112° 8.006 6.294 2.953 21° 100° Remarks. The teleoconch of Euspira umbilicolunata bears a close resemblance to that of Euspira pulchella (Risso, 1826). However, E. umbilicolunata can be distinguished easily from E. pulchella because of its signiicantly greater protoconch diameter, its narrower umbilicus, and its better developed funicle (obsolete in E. pulchella) not separated from the umbilical wall by a furrow. The values of the characteristic elements of the protoconch of E. umbilicolunata do not differ signiicantly from those measured for Euspira grossularia (Marche-Marchad, 1957), but the teleoconch characters of E. umbilicolunata readily distinguish it from E. grossularia. E. umbilicolunata differs from E. grossularia by its more elevated spire (Fig. 12), its less inlated last whorl, its more widely open umbilicus, its obsolescent funicle, and its more slender, subtriangular umbilical callus. Euspira notabilis (Jeffreys, 1885) is another similar species. E. umbilicolunata differs from it in having the protoconch of 2.85-3 whorls (2.12-2.25 in E. notabilis), with signiicantly greater diameter and no groove separating the funicle from the umbilical wall. E. umbilicolunata might be confused with specimens of E. helicina helicina (Brocchi, 1814), in particular with those having a very narrow, crescentic umbilicus. The most relevant characters distinguishing E. umbilicolunata from Brocchi’s taxon are: 1) the larger protoconch with signiicantly smaller diameter of the initial half-whorl, and 2) the absence of any spiral furrow separating the funicle from the umbilical wall (present and deep in E. helicina helicina). Stratigraphic occurrence. Euspira umbili- colunata appears to be restricted to the BurdigalianLanghian time span. Euspira sp. 1 Pl. 10, ig. 9 Material examined: Cava Albanello: 1 spm., NP 9851 (Pl. 10, ig. 9); Cava Grola: 1 spm., private collection, 1 spm., NP 9852, 1 spm., MPUM 11334, 1 spm., private collection, 2 spms., private collection. Description. Protoconch poorly preserved, medium-sized, of 2.25 convex whorls, tip missing. Teleoconch medium to large, depressed-globose, rather thin. Spire obtusely conical, moderately elevated, whorls convex; suture shallowly channeled. Last whorl large, globose, somewhat depressed, very slightly expanded toward aperture; subsutural shelf rather narrow, poorly deined; periphery above midline. Aperture ovately D-shaped in prosocline plane, length about 1.6 times width; outer lip semicircular, inner lip straight. Parietal callus moderately thick, rectangular; anterior corner (right angle) at level of umbilical border. Umbilicus deep, moderately wide, crescent-shaped; umbilical wall steep, bounded downward by slight groove ending at most abapical part of inner lip; interior of umbilicus with spiral cordlets crossed by growth markings. Funicle obsolete. Umbilical callus very slender, with reverse J-shaped outline, ending in middle of abapical margin of parietal callus. Basal fasciole weak to indistinct. Outer surface with vestige of growth lines more evident on lower base. Dimensions (mm) DHW PD H D SH - - 7.00026.288 8.06627.958 1.8046.732 - - 16.644 18.012 4.268 AH AW UW IS SA 4.82019.932 5.15816.138 2.6708.966 10°-22° 122°-138° 12.376 10.648 5.818 16° 130° Remarks. The Eocene Euspira piccolii sp. n. (see above) appears to be the most closely similar species in shell morphology, but differs from Euspira sp. 1 in that it has a protoconch of 0.75 more whorls, a comparatively wider, teardrop-shaped umbilicus, and a better developed, triangular umbilical callus ending into the anterior corner of the parietal Eocene, Oligocene and Miocene naticid gastropods of Northern Italy callus, not in the middle of the anterior margin of the parietal callus as in Euspira sp. 1. The British, Bartonian species Euspira bartonensis Wrigley, 1949 is also closely similar (topotypes examined). However, E. bartonensis differs from Euspira sp. 1 because of its interior of the umbilicus bounded by a sharp ridge instead of a groove as in Euspira sp. 1 and its umbilical callus nearly semicircular, demarcated from the parietal callus by a distinct transverse depression. The Tortonian species Euspira giuntellii sp. n. (see above) is also similar, but is readily distinguished from Euspira sp. 1 in having the umbilicus devoid of spirals; moreover, it attains a larger size. The present specimens almost certainly represent an undescribed species, but more material retaining the protoconch is required in order to name it. Stratigraphic occurrence. Euspira sp. 1 was recovered only from Lutetian deposits of Veneto. Euspira sp. 2 Pl. 10, igs. 10, 11 Material examined: Valle Ceppi: 1 spm., MZB 60130;Valle Vergnana: 1 spm., MZB 43250 (Pl. 10, ig. 10), 1 spm., MZB 60132 (Pl. 10, ig. 11), 1 spm., MZB 60131. Description. Protoconch medium-sized, of 2.16-2.25 convex, apparently smooth whorls, tip small. Teleoconch medium-sized, depressed-globose, rather thick. Spire broadly conical, scarcely to moderately elevated, whorls gently convex to convex; suture, ine, adpressed. Last whorl inlated, somewhat depressed, very slightly expanded toward aperture; subsutural shelf indistinct; periphery about at midline. Aperture D-shaped in prosocline plane, length about 1.9 times width; outer lip semicircular, inner lip straight, thickened abapically. Parietal callus thick, with markedly concave abapertural outline; anterior lobe poorly differentiated, subangular to rounded, at level of adapical umbilical border. Umbilicus deep, crescent-shaped, rather narrow to moderately wide; umbilical wall slightly overhanging interior of umbilicus, bounded downward by shallow, rather narrow spiral depression forming slight sinus on abapical part of inner lip. Funicle broad, depressed, obsolescent toward interior of umbilicus. Umbilical callus thick, with well curved, reverse S-shaped adaxial outline and merging into anterior lobe of parietal callus. Basal fasciole indis- 181 tinct. Outer surface with ine growth lines prosocyrt subsuturally. All specimens retain vestige of pale brown background, with darker subsutural band and lowermost base. Dimensions (mm) DHW PD H D SH 0.1780.194 1.1001.132 9.97416.002 9.38915.701 1.5654.621 0.186 1.116 12.988 12.545 3.093 AH AW UW IS SA 8.11311.677 6.41510.191 1.9485.972 12°-20° 112°-128° 9.895 8.303 3.960 16° 120° Remarks. Euspira sp. 2 appears to be closely similar to the Tortonian to Recent species Euspira notabilis (Jeffreys, 1885) if the teleoconch characters and the size are considered. However, it can be differentiated from E. notabilis on the basis of its protoconch, which has signiicantly greater diameter and diameter of the initial half-whorl. Euspira sp. 2 is also similar to the coeval species Euspira submamillaris (Sacco, 1890) and Euspira umbilicolunata sp. n (see above). It differs from E. submamillaris in having the protoconch of about one fewer whorl, with a signiicantly greater diameter of the irst half-whorl, and the teleoconch depressed-globose instead of globose-oval to oval as in E. submamillaris. Euspira sp. 2 is readily distinguished from E. umbilicolunata in that it has: 1) a protoconch of about one fewer whorl, with signiicantly greater diameter of the irst half-whorl, 2) a depressed-globose teleoconch (that of E. umbilicolunata is more elongate, globoseoval), 3) a lower spire with less convex whorls, and 4) a distinctly wider umbilicus. Most probably the examined specimens represent a new species, but additional material is required to assess them. Stratigraphic occurrence. Euspira sp. 2 seems to be an uncommon element, occurring in Burdigalian deposits of the Torino Hills. Euspira sp. 3 Pl. 10, ig. 12 Material examined: Case Soghe: 1 spm., MPUM 11335. Description. Protoconch abraded. Teleoconch medium-sized, globose, rather thick. Spire broadly conical, scarcely elevated, whorls latly Robba E., Pedriali L. & Quaggiotto E. 182 convex; suture, ine, adpressed. Last whorl inlated, slightly produced abapically toward aperture; subsutural shelf indistinct; periphery above midline. Aperture D-shaped in prosocline plane, length about twice width; outer lip semicircular, inner lip almost straight. Parietal callus expanded adapically, thick at both ends, rather thin medially, with concave abapertural outline; anterior lobe poorly differentiated, subangular, hindering most adapical part of umbilicus. Umbilicus deep, broadly semicircular; umbilical wall and interior of umbilicus with uneven spiral cordlets crossed by growth markings. Funicle absent. Umbilical callus obsolete. Basal fasciole indistinct. Outer surface with remnants of growth lines. Dimensions (mm) DHW PD H D SH AH AW - - 12.340 11.681 1.762 10.580 8.339 UW IS SA 5.320 14° 126° Remarks. The present form seems unlike any other Euspira species dealt with in the literature and likely represents a new species, which, however, requires more material retaining the protoconch to be named. Euspira sp. 3 can be compared with the Eocene species Euspira piccolii sp. n. and Euspira sp. 1 (see above) that also have the interior of the umbilicus spirally sculptured. E. piccolii differs from Euspira sp. 3 in that it has: 1) a more depressed teleoconch, 2) a more elevated spire with well convex whorls and narrowly channeled sutures, 3) a rather wide, teardrop-shaped umbilicus (semicircular in Euspira sp. 3), and 4) a distinct, triangular umbilical callus. Euspira sp. 1 is readily distinguished from Euspira sp. 3 on the basis of its depressed-globose teleoconch, its convex spire whorls and channeled suture, its narrower, crescent-shaped umbilicus, and its narrow but distinct umbilical callus (obsolete in Euspira sp. 3). Stratigraphic occurrence. Euspira sp. 3 seems to be a very rare taxon, recovered from Rupelian deposits at Case Soghe (Veneto). Euspira sp. 4 Pl. 10, igs. 13, 14 Material examined: Cassinelle: 1 spm., MPUM 11336 (Pl. 10, ig. 13); Monte Gloso: 1 spm., MPUM 11337 (Pl. 10, ig. 14); Sangonini: 1 spm., MCZ 4330. Description. Protoconch small, depressedturbiniform, of 2.25 whorls, tip small. Teleoconch small, globose-oval, thick. Spire moderately elevated, whorls convex; suture, ine, adpressed. Last whorl broadly oval; subsutural shelf indistinct; periphery nearly at midline. Aperture Dshaped in prosocline plane, length about twice width; outer lip semicircular, inner lip almost straight, thickened abapically. Parietal callus rather thin, less so at both ends, with concave abapertural outline; anterior lobe short, subrounded, hindering most adapical part of umbilicus. Umbilicus deep, broadly semicircular, umbilical wall steep. Funicle broad, markedly depressed, separated from umbilical wall by shallow spiral depression bounded abaxially by distinct step. Umbilical callus rather thick, slenderly subtriangular, merging into anterior lobe of parietal callus. Basal fasciole broad, poorly deined. Outer surface with ine, dense growth lines. Dimensions (mm) DHW PD H D SH AH AW 0.173 0.802 6.590 6.038 2.221 4.370 3.061 UW IS SA 2.640 13° 117° Remarks. Euspira sp. 4 differs from the coeval species Euspira sp. 3 by its more elevated spire with well convex whorls (latly convex in Euspira sp. 3), its better developed anterior lobe of the parietal callus, its umbilicus devoid of inner spiral cordlets (present in Euspira sp. 3), and 4) its broad funicle (absent in Euspira sp. 3). The latter two characters constitute the most relevant distinguishing elements. Euspira sp. 4 can be readily differentiated from the other coeval species Euspira molarensis sp. n. in that it has a protoconch with signiicantly greater diameter of the irst half-whorl, a markedly different parietal callus, and a distinct funicle (absent in E. molarensis). The Langhian to Tortonian new species Euspira umbilicocarinata (see above) shares many characters with Euspira sp. 4, but is distinguished from it by having a protoconch with signiicantly greater diameter and smaller diameter of the initial half-whorl, a more globose shell, and an indistinct basal fasciole. Stratigraphic occurrence. Euspira sp. 4 was found in Rupelian deposits of Piedmont and Veneto. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Euspira sp. 5 Pl. 10, ig. 15 Material examined: Albugnano: 1 spm., MPUM 11338 (Pl. 10, ig. 15), 1 spm., PG 105. Description. Protoconch small, depressedturbiniform, of 2.56 whorls, tip very small. Teleoconch small, globose, rather thin. Spire broadly conical, somewhat elevated, whorls gently convex; suture, ine, adpressed. Last whorl globose-oval, tall; subsutural shelf indistinct; periphery slightly above midline. Aperture ovately D-shaped in prosocline plane, length about 1.5 times width; outer lip nearly semicircular, inner lip very slightly arched. Parietal callus rather thick medially, slightly more so at both ends, with concave abapertural outline; anterior lobe small, subrounded, at level of umbilical border. Umbilicus deep, rather wide, nearly semicircular, umbilical wall moderately sloping, more steep downward. Funicle a broad, depressed adaxial thickening of inner lip, soon obsolete toward interior of umbilicus. Umbilical callus rather wide, thick, with convex abapertural outline, expanded to overhang mid-adapical part of umbilicus, merging into anterior lobe of parietal callus. Basal fasciole indistinct. Outer surface with ine, incised, uneven and unevenly spaced growth lines. Dimensions (mm) DHW PD H D SH AH AW 0.068 0.755 5.290 5.110 1.601 3.689 2.840 UW IS SA 1.920 17° 117° Remarks. Euspira sp. 5 is closely similar to the Tortonian new species Euspira latecallosa (see above) in teleoconch morphology, but can be distinguished from it because of its protoconch with signiicantly smaller diameter and diameter of the initial half-whorl. Moreover, Euspira sp. 5 has a wider, semicircular umbilicus, whereas that of E. latecallosa is crescent-shaped and very narrow. Stratigraphic occurrence. Euspira sp. 5 was recovered from deposits near Albugnano (Piedmont) said to be of Serravallian age. Euspira sp. 6 Pl. 11, igs. 1, 2 Material examined: Valle Ceppi: 1 spm., MPUM 11339 (Pl. 11, ig. 1), 1 spm., MPUM 11340 (Pl. 11, ig. 2), 2 spms., NP 9993, 1 spm., MZB 60149. 183 Description. Protoconch small, depressedturbiniform, of 2.05-2.40 whorls, tip small. Teleoconch small, globose-oval, moderately thick. Spire broadly conical, rather elevated, nearly straight-sided; whorls latly convex, suture, ine, almost lush. Last whorl broadly oval; subsutural shelf indistinct; periphery slightly above midline. Aperture ovately D-shaped in moderately prosocline plane, length about 1.5 times width; outer lip semicircular, inner lip very slightly arched. Parietal callus moderately thick, long, subrectangular; anterior lobe very small to indistinct. Umbilicus deep, small; umbilical wall regularly sloping. Funicle a broad, depressed adaxial thickening of inner lip, obsolescent toward interior of umbilicus, separated from umbilical wall by narrow, shallow depression bounded abaxially by slight step more distinct in small specimens. Umbilical callus thick, triangular, hindering midadapical part of umbilicus, merging into anterior corner of parietal callus; umbilical channel obliquely elliptical, narrow to very narrow. Basal fasciole poorly differentiated. Outer surface with uneven and unevenly spaced growth lines; faint spiral striation occurs on last whorl. Two specimens retain vestige of uniform, light brown background. Dimensions (mm) DHW PD H D SH 0.1040.120 0.7680.828 4.39111.495 3.76910.317 0.3913.855 0.112 0.798 7.943 7.043 2.123 AH AW UW IS SA 3.6408.000 1.7226.698 0.9332.835 9°-21° 97°-137° 5.820 4.210 1.893 15° 117° Remarks. Euspira sp. 6 appears to be closely similar to the Burdigalian to Piacenzian species Euspira subobturata (Sacco, 1891) in teleoconch characters and probably has been mistaken for it. However, Euspira sp. 6 has a protoconch with significantly smaller diameter and diameter of the first half-whorl. The protoconch stands as the main differentiating element. The examined specimens likely represent a new species, but more material is required in order to name it. Stratigraphic occurrence. Euspira sp. 6 was collected from coarse sandy deposits belonging to the mid-Burdigalian Termofourà Formation. 184 Robba E., Pedriali L. & Quaggiotto E. Genus Neverita Risso, 1826 Neverita Risso, 1826, p. 149. Type species by monotypy: Neverita josephinia Risso, 1826, Recent, Mediterranean. Mamillaria Swainson, 1840, p. 345. Type species by subsequent designation (Hedley 1924, p. 154): Mamillaria lactea Swainson, 1840 (= Nerita peselephanti Link, 1807). Kabat (1991, p. 434) regarded Poliniciella Petuch, 1988 as synonym of Neverita. We note that Poliniciella marylandica Petuch, 1988 (type species of Poliniciella by monotypy) does not agree with the characters of Neverita because of its markedly different umbilical features. Accordingly, we consider Kabat’s opinion untenable. Neverita was either considered as a subgenus of Polinices Montfort, 1810 (Cernohorsky 1972; Kilburn 1976) or as a distinct genus (Marincovich 1977; Majima 1989; Kabat 1990, 1991). Huelsken et al. (2012, p. 372), on the basis of molecular data, rejected the concept of Neverita as a subgenus of Polinices, and concluded that “the taxa Neverita and Polinices each form statistically well-supported monophyletic clades”. The diagnostic characters of Neverita were listed by Pedriali & Robba (2009). These characters are herein redefined as follows: 1) shell rather thick to thick, depressed-globose, globose-pyriform in a few species, wider than high in most species; 2) spire very low to moderately elevated, whorls slightly convex to convex, with obscure subsutural shelf in a few species, suture almost flush to adpressed; 3) last whorl greatly enlarged, depressed in most species; 4) aperture strongly prosocline, D-shaped to ovately Dshaped; 5) parietal callus rather thick to thick, rather short, with slender anterior lobe encircling adapical part of umbilical callus in most species; 6) umbilicus broad, largely or completely filled by umbilical callus, with excavated, rather wide, abapical or mid-abapical umbilical channel present in a few species; 7) funicle thick and prominent, covered by umbilical callus in most species; 8) umbilical callus massive, semicircular, subtriangular in a few species, separated from parietal callus by a slight notch, overlapped from and fused to parietal callus in a few species. The depressed shell somewhat Sinum-like and the large, button-like umbilical callus are the most obvious distinctive characters of Neverita. Neverita antiqua (Sacco, 1890) stat. n. Pl. 11, igs. 3, 4 1890b Natica (Neverita) josephinia var. antiqua Sacco, p. 34. 1891 Natica (Neverita) josephinia var. antiqua - Sacco, p. 85, pl. 2, ig. 55. 1984 Natica (Neverita) josephinia var. antiqua - Ferrero Mortara et al., p. 35. Type material: Natica (Neverita) josephinia var. antiqua Sacco, lectotype (designated by Pedriali & Robba 2009, p. 407): the shell igured by Sacco (1891, pl. 2, ig. 55) and reigured herein (Pl. 11, ig. 3), MGPT BS.029.05.003, Cassinelle; 2 paralectotypes, MGPT BS.029.05.003/01 and MGPT BS.029.05.003/02, Dego. Other material examined: Cassinelle: 2 spms., PG 17; Colle del Giovo Ligure: 1 spm., NP 9861; Madonna delle Rocche: 1 spm., PG 65; Mioglia: 10 spms., MZB 60127, 1 spm., MZB 60128; Molare, Torrente Orba: 2 spms., PG 66; Pareto: 1 spm., PG 67; Monte Gloso: 1 spm., MPUM 11341 (Pl. 11, ig. 4), 1 spm., NP 9862. Description. Protoconch abraded in all specimens. Teleoconch globose-conical, depressed, wider than high, moderately thick. Spire broadly cyrtoconoid, moderately elevated, low in some PLATE 11 Fig. 1 - Euspira sp. 6. Valle Ceppi. MPUM 11339; apertural side (height of shell 9.46 mm). Fig. 2 - Euspira sp. 6. Valle Ceppi. MPUM 11340; protoconch. Fig. 3 - Neverita antiqua (Sacco, 1890). Cassinelle. Lectotype of Natica (Neverita) josephinia var. antiqua Sacco, 1890. MGPT BS.029.05.003; a, Fig. 4 - Neverita antiqua (Sacco, 1890). Monte Gloso. MPUM 11341; a, apertural side showing color pattern (height of shell 17.27 mm); b, abapertural side; c, basal view. Fig. 5 - Neverita maga (De Gregorio, 1880). Cava Boschetto. MPUM 11342; protoconch. Fig. 6 - Neverita maga (De Gregorio, 1880). Cava Grola. MCZ 4331; a, apertural side (height of shell 22.84 mm); b, abapertural side; c, basal view. Fig. 7 - Neverita olla (de Serres, 1829). Borelli. MGPT-PU 135225; protoconch. Fig. 8 - Neverita olla (de Serres, 1829). Valle Ceppi. MGPT-PU 135238; a, apertural side (height of shell 20.24 mm); b, abapertural side. Fig. 9 - Payraudeautia bituberculata sp. n. Borelli. Holotype, MGPT-PU 135240; a, apertural side; b, protoconch. Fig. 10 - Payraudeautia bituberculata sp. n. Borelli. Paratype, MGPT-PU 135241; apertural side. Fig. 11 - Payraudeautia crassicorda sp. n. Cava Albanello. Holotype, MPUM 11350; a, apertural side (height of shell 8.52 mm); b, basal view. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 185 Robba E., Pedriali L. & Quaggiotto E. 186 specimens; whorls very slightly convex; suture almost flush. Last whorl expanded, rather depressed; subsutural shelf indistinct; periphery nearly at midline; base almost flat. Aperture Dshaped in markedly prosocline plane, length about 1.70 width; inner lip very slightly concave. Parietal callus rather thick to thick; anterior lobe slender, sealing adapical part of umbilical groove. Umbilicus broad, largely illed by thick, prominent funicle; umbilical wall very steep, with coarse growth ridges; umbilical channel well developed, deeply excavated, of variable breadth, restricted to abapical part of umbilicus in most specimens, extended to middle part in some. Umbilical callus massive, roundly triangular, adapically overlapped by and fused to parietal callus. Outer surface with dense growth lines, prosocyrt adapically. One specimen retains vestige of brown color over umbilical wall and of reddishbrown collabral lines throughout last whorl. Dimensions (mm) DHW PD H D SH - - 8.19317.289 9.51521.327 0.9725.648 - - 12.741 15.421 3.310 AH AW WUC IS SA 6.23312.629 3.36611.030 4.38110.297 12°-32° 115°-139° 9.431 7.198 7.339 22° 127° Remarks. In a previous paper (Pedriali & Robba 2009), Natica (Neverita) josephinia var. antiqua Sacco, 1890 was included with doubt in the synonymy of Neverita olla (de Serres, 1829). Examination of additional material has convinced us to treat Sacco’s variety as a species, distinct from N. olla. The deeply excavated, rather wide umbilical channel and the subtriangular umbilical callus distinguish Neverita antiqua from N. olla, which, instead, has a semicircular umbilical callus and a narrower to indistinct umbilical channel. These morphological characters, supported by molecular data, were used by Huelsken et al. (2006) in separating Neverita delessertiana (Récluz in Chenu, 1843) from Neverita duplicata (Say, 1822). Further, N. olla exhibits a uniform pale brown background with suprasutural darker stripe (Pedriali & Robba 2009), a color pattern different from that occurring in N. antiqua (see above). Stratigraphic occurrence. Neverita antiqua (Sacco, 1890) appears to be restricted to the early Oligocene of Piedmont, Liguria and Veneto. Neverita maga (De Gregorio, 1880) Pl. 11, igs. 5, 6 1880 Natica maga De Gregorio, p. xiii, pl. 3, igs. 24, 25; pl. 6, ig. 69. 2008 Neverita maga - Quaggiotto & Mellini, p. 48, pl. 3, igs. 24, 25; pl. 6, ig. 69 (plates reproduced from De Gregorio 1880). Type material: Natica maga De Gregorio, type material not seen. De Gregorio collection, stored in the Istituto di Geologia dell’Università di Palermo (presently Dipartimento di Geologia e Geodesia), was heaped up by American troops landed in Sicily during the second World War and the material is still awaiting to be identiied (Dolin & Pacaud 2009). Material examined: Cava Albanello: 1 spm., NP 9919, 1 spm., NP 9920; Cava Boschetto: 1 spm., MPUM 11342 (Pl. 11, ig. 5), 1 spm., NP 9921; Cava Grola: 1 spm., MCZ 4331 (Pl. 11, ig. 6); Cava Main: 1 spm., private collection; Cava Rossi: 1 spm., MPUM 11343, 1 spm., private collection, 1 spm., private collection. Description. Protoconch small, depressed turbiniform, of 1.9 gently convex whorls, tip small. Teleoconch large, globose-conical, depressed, wider than high, moderately thick. Spire broadly conical, rather low, somewhat stepped; whorls slightly convex, roundly subangular at adapical one-third, with poorly deined subsutural shelf; suture adpressed. Last whorl expanded, rather depressed; subsutural shelf indistinct; narrowly rounded periphery nearly at midline; base slightly concave. Aperture Dshaped in markedly prosocline plane, length about twice width. Parietal callus moderately thick, short; anterior lobe slender, encircling adapical part of umbilical callus. Umbilicus broad, completely illed by umbilical callus; umbilical wall slightly concave, gently sloping, with growth markings; umbilical channel wanting. Umbilical callus large, semicircular, thick and prominent medially, somewhat excavated abapically (more so in smaller specimens), adapically overlapped by and fused to parietal callus. Outer surface with dense, prosocyrt growth lines. Dimensions (mm) DHW PD H D SH 0.189* 0.975* 12.33826.578 15.68733.039 2.3158.151 * 1 protoconch measurable 19.458 24.363 5.233 AH AW WUC IS SA 8.54519.905 8.81920.851 8.76117.985 9°-21° 107°-147° 14.225 14.835 13.373 15° 127° Remarks. The Eocene, Nigerian species Neverita amekiensis, introduced by Eames (1957, p. 39), is closely similar to Neverita maga in teleoconch shape, Eocene, Oligocene and Miocene naticid gastropods of Northern Italy but differs from it because of its less concave base and its umbilical callus not completely illing the umbilicus. The species Natica calvimontana Deshayes, 1864, Natica lineolata Deshayes, 1832 and Natica semiclausa Deshayes, 1864, described from Eocene deposits of the Paris Basin, were subsequently assigned to Neverita Risso, 1826 (Cossmann 1888; Cossmann & Pissarro 1907; Le Renard & Pacaud 1995; Pacaud & Le Renard 1995). These three French species exhibit a supericial resemblance to N. maga in that they have a globose shell, a higher and pointed spire, and an umbilical callus distinctly smaller than that of N. maga. We have not seen the original material of the cited Deshayes’ species, but, according to the excellent illustration of them provided by Deshayes (1864) and Cossmann & Pissarro (1907), we are doubtful concerning their assignment to Neverita. The name calvimontensis, used by Cossmann (1888) and Cossmann & Pissarro (1907) instead of the original name calvimontana, is an unjustiied emendation. Stratigraphic occurrence. Neverita maga (De Gregorio, 1880) is known from upper Ypresian to middle Lutetian deposits of Veneto. Neverita olla (de Serres, 1829) Pl. 11, igs. 7, 8 1829 Natica olla de Serres, p. 102, pl. 1, igs. 1, 2. 1866 Natica (Neverita) josephinia - Speyer, p. 27, pl. 3, ig. 2 (not Risso, 1826). 1896 Neverita josephinia var. bellunensis Vinassa de Regny, p. 206, pl. 5, ig. 4 (new synonym). 1917 Natica josephinia - Stefanini, p. 98, pl. 2, ig. 20 (not Risso, 1826). 1937 Natica (Neverita) bellunensis - Venzo, p. 46, pl. 2, ig. 37. 1985 Neverita bellunensis - Brigantini, p. 413, pl. 2, ig. 30. 2007 Neverita olla - Pister & Wegmüller, p. 102, pl. 18, igs. 17-24; pl. 19, igs. 1-7. 2007 Neverita josephinia - Zunino, p. 123, pl. 1, ig. 9 (not Risso, 1826). 2009 Neverita olla - Pedriali & Robba, p. 404, pl. 2, igs. 5-9; pl. 3, igs. 14, 15; pl. 4, ig. 17 (cum. syn.). 2013 Neverita olla - Landau et al., p. 107, pl. 11, ig. 8; pl. 62, ig. 8 (cum syn.). Type material: Pedriali & Robba (2009, p. 407) reviewed the type material of Neverita olla, and described the characters of this species. Material referred to as Natica subglaucinoides d’Orbigny, 1852 in MGPT: Termofourà: 10 spms., MGPT-PU 23520. Material examined.: Albugnano: 1 spm., PG 68; Borelli: 1 spm., MGPT-PU 135225 (Pl. 11, ig. 7), 1 spm., MGPT-PU 135226, 1 spm., MGPT-PU 135227, 1 spm., MGPT-PU 135228, 1 spm., MGPTPU 135229, 1 spm., MGPT-PU 135230, 1 spm., MGPT-PU 135231, 27 spms., MGPT-PU 135232, 2 spms., MPUM 11344, 4 spms., NP 187 9868, 7 spms., PG 22b; Léognan, Le Coquillat (France): 10 spms., private collection, 6 spms., PPMM 40821, 140 spms., private collection, 40 spms., private collection, 258 spms., MPUM 11345, 802 spms., NP 9966; Martignas-sur-Jalle (France): 1 spm., NP 10004; Mérignac (France): 10 spms., NP 9957; Monte dei Cappuccini: 2 spms., MGPTPU 25665; Montegibbio: 2 spms., MGC 1418, 1 spm., private collection, 11 spms., private collection, 25 spms., private collection, 1 spm., private collection; Orthez, Le Paren (France): 10 spms., NP 10007; Passo dei Meloni: 1 spm., NP 9953; Pauvrelay: 1 spm., NP 10001; Rio di Bocca d’Asino: 25 spms., MZB 60063, 1 spm., MZB 60111, 1 spm., MZB 60104, 11 spms., PG 25, 1 spm., NP 9864; Salles, Argilas (France): 26 spms., NP 10015; Saucats, Le Péloua (France): 6 spms., MPUM 11346, 1 spm., MPUM 11347, 1 spm., NP 9863, 1 spm., NP 9865, 3 spms., NP 10002; Stazzano: 2 spms., MGPT-PU 23407; Termofourà: 10 spms., MGPT-PU 23520; Valle Ceppi: 3 spms., MGPT-PU 135204, 3 spms., MGPT-PU 107768, 5 spms., MGPT-PU 135233, 3 spms., MGPT-PU 135234, 4 spms., MGPT-PU 135235, 1spm., MGPT-PU 135236, 6 spms., MGPT-PU 135237, 1 spm., MPUM 11348, 11 spms., PG 21, 35 spms., PG 69, 7 spms., MGPT-PU 107862, 1 spm., MGPTPU 107023, 1 spm., MGPT-PU 107024, 3 spms., MGPT-PU 23864, 7 spms., MGPT-PU 25905, 1 spm., MGPT-PU 135238 (Pl. 11, ig. 8), 39 spms., MGPT-PU 25267, 1 spm., MGPT-PU 26048, 2 spms., MGPTPU 26049, 1 spm., MGPT-PU 26145, 44 spms., MGPT-PU 108041, 5 spms., MGPT-PU 135239, 33 spms., MZB 60062, 1 spm., MZB 60129, 1 spm., NP 9866, 4 spms., NP 9867; Val Sanfrà: 6 spms., MGPT-PU 23666, 3 spms., MGPT-PU 107619, 1 spm., MGPT-PU 107627; Valle Vergnana: 1 spm., MZB 60133, 1 spm., MZB 44000; Vigoleno: 9 spms., private collection; Villa Allason: 9 spms., MGPT-PU 107617; Villa Bertini: 5 spms., MGPT-PU 107618; locality unknown: 2 spms., MGP-PD 2764z (specimens dealt with by Briganini 1985; see remarks below). Dimensions of Miocene specimens (mm) DHW PD H D SH 0.0920.112 0.7540.938 3.19529.727 5.64835.408 0.7627.186 0.102 0.846 16.461 20.528 3.974 AH AW WUC IS SA 1.49123.503 2.47817.262 4.10910.097 10°-26° 118°-142° 12.497 9.870 7.103 18° 130° Remarks. The examined Miocene specimens have the protoconch slightly larger than that of the Pliocene ones dealt with by Pedriali & Robba (2009), but the difference (+ 8% in protoconch diameter) is not significant. The protoconch of Neverita olla was described as smooth by Pedriali & Robba (2009); we noted that the protoconch of one specimen from the Tortonian of Borelli shows vestige of spiral rows of granules on the first half-whorl. For complete description, relationships with Neverita josephinia Risso, 1826, and other comments, see Pedriali & Robba (2009). Vinassa de Regny (1896) introduced the variety bellunensis of Neverita josephinia on the basis of a single Oligocene shell (holotype by monotypy) characterized by a broad and prom- 188 Robba E., Pedriali L. & Quaggiotto E. inent umbilical callus. The variety bellunensis was regarded later on as a distinct species (Venzo 1937; Brigantini 1985). The holotype of Neverita bellunensis was not located. However, we examined the two specimens in MGPPD dealt with by Brigantini (1985), referred to as N. bellunensis, and conforming to the short diagnosis published by Vinassa de Regny (1896). We could note that they are hardly distinguishable from Neverita olla, a species with a much variable extent and thickness of the umbilical callus. Moreover, the specimen illustrated by Brigantini (1985, pl. 2, fig. 30) has the protoconch of 2.60 whorls, with values of the diameter and diameter of the initial half-whorl that fall within the ranges calculated for N. olla (see above). Accordingly, we consider N. bellunensis as a synonym of N. olla. It is worth noting that the two specimens of N. bellunensis in MGPPD were said to had been collected from upper Eocene deposits of Valle Organa. This age assignment is inconsistent with the calcareous nannoplankton present in the matrix filling the shells, which demonstrates an age not older than the Miocene (Maria Triantaphyllou, personal communication 2014). It is likely that the specimens were accidentally coupled with the wrong label during movings of the material. Stratigraphic occurrence. Neverita olla (de Serres, 1829) ranges from early Oligocene to Pliocene. According to Pedriali & Robba (2009), it was replaced by Neverita josephinia Risso, 1826 by the Pleistocene. Genus Payraudeautia Bucquoy, Dautzenberg & Dollfus, 1883 Payraudeautia Bucquoy, Dautzenberg & Dollfus, 1883, p. 149. Type species by original designation: Natica intricata Donovan, 1803, Recent, Mediterranean. Payraudeautia was introduced as a subgenus of Natica Adanson, 1757 (= Natica Scopoli, 1777). Kabat (1991) considered Payraudeautia to be a junior subjective synonym of Natica Scopoli, 1777. Kabat’s conclusion is untenable since the type species of Payraudeautia has a corneous operculum denoting a poliniceine instead of a naticine allocation of the genus. Huelsken et al. (2008, 2012), on the basis of molecular data, have shown that Payraudeautia intricata exhibits a high genetic similarity with species of Euspira Agassiz in J. Sowerby, 1837. Thus, Huelsken et al. (2012) concluded that “synonymisation of Payraudeautia with Euspira is therefore appropriate”. We provisionally retain Payraudeautia as valid on account of its distinctive open umbilicus with an inner spiral ridge or cord abapical to the funicle. Payraudeautia appears to be a poorly speciose genus in that it includes, besides the type species, only Payraudeautia fasciolata (Sacco, 1890) and Payraudeautia nubila (Dall, 1889). The Eocene species Natica perforata Deshayes, 1864 and the Recent, eastern Atlantic species Natica gruveli Dautzenberg, 1910 (Payraudeautia esterias Bernard, 1983 is a synonym) were assigned to Payraudeautia, but have a single funicle (N. perforata has a funicle-like abapical cord) and deserve a different generic allocation. Presently, Payraudeautia ranges in the Mediterranean (P. intricata) and in the Caribbean Sea (P. nubila). Fossil occurrences of Payraudeautia were so far from late Miocene to Pleistocene of Italy. The two Eocene species described below are the earliest records of the genus and push back the irst occurrence of Payraudeautia. Payraudeautia bituberculata sp. n. Pl. 11, igs. 9, 10 Derivation of name: From Latin bis = two times and tuberculata = provided with a tubercle, with reference to the knob present at both ends of the parietal callus. Holotype: Borelli: MGPT-PU 135240 (Pl. 11, ig. 9). Paratypes: Borelli: 1 spm., MGPT-PU 135241 (Pl. 11, ig. 10), 4 spms., MGPT-PU 135242, 2 spms., MPUM 11349. Other material examined: Borelli: 2 spms., NP 9831. Preservation. Rather fair. Type locality: Borelli (see appendix). Horizon. Gray, medium sand of Tortonian age. Diagnosis: Globose, moderately depressed shell with scarcely elevated spire, somewhat depressed last whorl. Umbilicus very large with broad, depressed funicle and abapical inner spiral ridge. Parietal callus with rounded knob at both ends; umbilical callus thick, subtriangular, demarcated from anterior knob of parietal callus by slight transverse groove. Description. Protoconch medium-sized, turbinate with nearly lat-topped spire, of 2.90-3.05 convex and smooth whorls, tip very small. Teleoconch globose, moderately depressed, rather solid. Spire broadly conical, scarcely elevated, whorls gently convex; suture thin, adpressed. Last whorl inlated, somewhat depressed, slightly expanded toward aperture; subsutural shelf indistinct; pe- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy riphery above midline. Aperture D-shaped in slightly prosocline plane, length approximately 1.5 times width. Parietal callus rather narrow, thin medially, with markedly concave abapertural outline and with rounded knob at both ends, abapical one slightly more prominent nearly at level of basal fasciole. Umbilicus deep, very large; umbilical border narrowly rounded; umbilical wall concave, bearing prominent spiral ridge that overhangs interior of umbilicus and terminates in subtriangular, asymmetric plug on lowermost part of inner lip. Funicle broad and rather depressed, separated from inner spiral ridge by wide groove whose abaxial side ascends to form sharp angle with top of ridge. Umbilical callus rather thick, subtriangular, demarcated from anterior knob of parietal callus by slight transverse groove. Basal fasciole poorly differentiated, rather broad and blunt. Surface with very ine growth lines subangular adapically. Dimensions (mm) DHW PD H D SH 0.0540.062 1.0801.084 1.5035.855 1.7065.686 0.4091.253 0.058 1.082 3.679 3.696 0.831 AH AW UW IS SA 1.0834.611 0.7673.831 0.4672.227 12°-24° 115°-127° 2.847 2.299 1.347 18° 121° Remarks. Payraudeautia bituberculata is very close to Payraudeautia fasciolata (Sacco, 1890) in teleoconch shape and umbilical characters, but differs from Sacco’s species in that has: 1) a protoconch with significantly smaller diameter of the first half-whorl, and 2) a knob at both ends of the parietal callus. The features of the parietal callus constitute the most obvious distinguishing character of Payraudeautia bituberculata. Payraudeautia intricata (Donovan, 1803) is another morphologically similar species, differing from P. bituberculata because of its protoconch with fewer whorls (1.50-1.70 instead of 2.90-3.05) and significantly greater diameter of the first half-whorl, its parietal callus devoid of knobs and with distinct anterior lobe, and its narrower funicle. It is also worth noting that P. bituberculata hardly exceeds five mm in height, whereas P. fasciolata and P. intricata attain a larger size. Stratigraphic occurrence. Payraudeautia 189 bituberculata sp. n. was recovered only from Tortonian deposits at the type locality. Payraudeautia crassicorda sp. n. Pl. 11, fig. 11; Pl. 12, fig. 1 Derivation of name: From Latin crassus = thick and corda = cord, with reference to the thick, cord-like inner ridge of the umbilicus. Holotype: Cava Albanello: MPUM 11350 (Pl. 11, fig. 11). Paratypes: Cava Albanello: 1 spm., MPUM 11351 (Pl. 12, fig. 1), 2 spms., MPUM 11352, 48 spms., MGP-PD 1228R, 10 spms., MCZ 4332; Ciupio: 1 spm., MGP-PD 11624/b-11664/b. Other material examined: Cava Albanello: 20 spms., NP 9832; Cava Boschetto: 1 spm., NP 9833, 1 spm., NP 9834, 1 spm., private collection; Cava Grola: 2 spms., private collection. Preservation: All specimens have the surface somewhat abraded; a few are broken. Type locality: Cava Albanello (see appendix). Horizon: Rather coarse, gray tuff forming the lower part of a thick, fining upward volcanoclastic bed of early Lutetian age. Diagnosis: Globose, rather thin shell with scarcely elevated spire and inflated, somewhat depressed last whorl. Umbilicus deep, large, with thick, cord-like spiral ridge ending in semicircular plug on lowermost inner lip; funicle broad, markedly depressed, adjacent to adapical part of umbilical wall. Parietal callus thick; umbilical callus thick, narrowly subtriangular, merging into anterior angle of parietal callus. Description. Protoconch medium-sized, low-turbiniform, of 1.95-2.05 convex and apparently smooth whorls, tip medium-sized. Teleoconch globose, slightly wider than high, rather thin. Spire low-conical, scarcely elevated, whorls very gently convex; suture almost lush. Last whorl inlated, somewhat depressed, slightly produced toward aperture; subsutural shelf indistinct; periphery above midline. Aperture D-shaped in slightly prosocline plane, length approximately 1.7 times width. Parietal callus thick, subrectangular, with gently concave abapertural outline, ending at level of umbilical border. Umbilicus deep, large; umbilical border broadly rounded; umbilical wall steeply sloping; interior of umbilicus bearing thick, cord-like spiral ridge that terminates in semicircular plug on lowermost part of inner lip. Funicle markedly depressed, broad, adjacent to adapical part of umbilical wall, separated from inner spiral ridge by narrow, deeply incised groove. Umbilical callus thick, narrowly subtriangular, with oblique, straight or reverse S-shaped outline, merging into anterior angle of parietal callus. Basal fasciole in- Robba E., Pedriali L. & Quaggiotto E. 190 distinct. Surface with vestige of ine growth lines faintly arched subsuturally. Material examined: Borelli: 1 spm., MGPT-PU 135243 (Pl. 12, fig. 2); Montegibbio: 1 spm., MZB 31669 (Pl. 12, fig. 3). Dimensions (mm) DHW PD H D SH 0.2760.292 1.1351.207 3.4989.602 3.6159.959 0.6442.496 0.284 1.171 6.550 6.787 1.570 AH AW UW IS SA 2.6467.314 2.2206.056 0.9873.319 14°-26° 106°-126° 4.980 4.138 2.153 20° 116° Remarks. Payraudeautia crassicorda sp. n. is characterized by its large umbilicus with unusually thick, cord-like inner spiral ridge and markedly depressed funicle; these two structures occupy almost entirely the straight adapertural margin of the umbilicus. P. crassicorda is morphologically similar to the lower and middle Lutetian species Payraudeautia zarantonelloi sp. n. (see below) and the values of the characteristic elements of the protoconch of both taxa do not display any significant difference. However, P. crassicorda can be distinguished from P. zarantonelloi by its wider umbilicus (Fig. 15), with a thick inner spiral ridge and a markedly depressed funicle. The reverse occurs in P. zarantonelloi that has a smaller umbilicus with a narrower, subangular spiral ridge and a rather prominent funicle. This difference is deemed to warrant specific separation. Stratigraphic occurrence. Payraudeautia crassicorda sp. n. was recovered from lower Lutetian deposits of Veneto. Payraudeautia fasciolata (Sacco, 1890) Pl. 12, igs. 2, 3 1890b Natica (Payraudeautia) intricata var. fasciolata Sacco, p. 33. 1891 Natica (Payraudeautia) intricata var. fasciolata - Sacco, p. 80, pl. 2, ig. 52. 1984 Payraudeautia intricata var. fasciolata - Ferrero Mortara et al., p. 35. 1996 Payraudeautia intricata - Pedriali, p. 14 (pars), pl. 4, igs. 5, 6 (not Donovan, 1803). 2009 Payraudeautia fasciolata - Pedriali & Robba, p. 410, pl. 2, igs. 11, 12; pl. 3, ig. 17; pl. 4, igs. 19, 20. Type material: Pedriali & Robba (2009, p. 411) listed the type material of Payraudeautia fasciolata, and described the characters of this species. Dimensions (mm) DHW PD H D SH 0.0930.101 0.9571.117 1.55511.827 1.35611.784 0.4711.891 0.097 1.037 6.691 6.570 1.181 AH AW UW IS SA 1.1279.855 1.1665.462 0.4304.782 11°-23° 111°-135° 5.491 3.314 2.606 17° 123° Remarks. For description, dimensions and comparison with Payraudeautia intricata (Donovan, 1803), see Pedriali & Robba (2009). An unidentiied Payraudeautia species from the Pliocene Lillo Formation of Belgium (specimens in our collection) appears to be similar to Payraudeautia fasciolata in teleoconch morphology, but differs from it in that it has the protoconch of less than two whorls (2.60-2.65 in P. fasciolata), with signiicantly greater diameter of the initial half-whorl and the umbilicus with a broad, markedly depressed abapical spiral ridge (prominent in P. fasciolata). PLATE 12 Fig. 1 - Payraudeautia crassicorda sp. n. Cava Albanello. Paratype, MPUM 11351; protoconch. Fig. 2 - Payraudeautia fasciolata (Sacco, 1890). Borelli. MGPT-PU 135243; a, apertural side (height of shell 3.67 mm); b, basal view. Fig. 3 - Payraudeautia fasciolata (Sacco, 1890). Montegibbio. MZB 31669; protoconch. Fig. 4 - Payraudeautia intricata (Donovan, 1803). Rio di Bocca d’Asino. NP 9835; a, apertural side; b, basal view. Fig. 5 - Payraudeautia zarantonelloi sp. n. Cava Grola. Holotype, MPUM 11353; a, apertural side (height of shell 15.26 mm); b, basal view; c, protoconch. Fig. 6 - Payraudeautia zarantonelloi sp. n. Cava Grola. Paratype, MPUM 11355; a, apertural side (height of shell 12.95 mm); b, basal view. Fig. 7 - Pliconacca plicatulaeformis (Kittl, 1887). Orlau (Czech Republic). Lectotype (here designated) of Natica plicatulaeformis Kittl, 1887. NHMW I.26; a, apertural side; b, apical view. Fig. 8 - Pliconacca plicatulaeformis (Kittl, 1887). Orlau (Czech Republic). Paralectotype. NHMW I.25; apertural side. Fig. 9 - Pliconacca plicatulaeformis (Kittl, 1887). Colli Torinesi. Lectotype (here designated) of Natica (Naticina) catena var. prohelicina Sacco, 1890. MGPT BS.029.02.001; apertural side (scale bar 4 mm). Fig. 10 - Pliconacca plicatulaeformis (Kittl, 1887). Valle Ceppi. MZB 60013; protoconch. Fig. 11 - Pliconacca plicatulaeformis (Kittl, 1887). Valle Ceppi. MGPTPU 135259; a, apertural side (height of shell 17.58 mm); b, apical view; c, basal view. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 191 Robba E., Pedriali L. & Quaggiotto E. 192 Stratigraphic occurrence. Payraudeautia fasciolata (Sacco, 1890) appears to be very rare in the Tortonian of Piedmont and Emilia. It is an uncommon element in the Pliocene of Piedmont, Liguria, Emilia and Tuscany. There are no records subsequent to the early Piacenzian. Payraudeautia intricata (Donovan, 1803) Pl. 12, ig. 4 1803 Nerita intricata Donovan, pl. 167 (short description in the explanation to the plate). 2009 Payraudeautia intricata - Pedriali & Robba, p. 411, pl. 2, igs. 13, 14; pl. 3, ig. 18; pl. 4, ig. 21 (cum syn.). Type material: Pedriali & Robba (2009, p. 412) reviewed the type material of Payraudeautia intricata, and described the characters of this species. Material examined: Rio di Bocca d’Asino: 1 spm., NP 9835. Remarks. For description, dimensions and comments, see Pedriali & Robba (2009). Stratigraphic occurrence. Payraudeautia intricata (Donovan, 1803) is extremely rare in the Tortonian of Piedmont. The species was recorded from Pliocene deposits in Piedmont, Emilia, Tuscany and Latium. Pleistocene occurrences are from Tuscany and Calabria. Payraudeautia intricata ranges nowadays throughout the Mediterranean, being more common in the western part of it. Payraudeautia zarantonelloi sp. n. Pl. 12, igs. 5, 6 Derivation of name: The species is named after Giannino Zarantonello, who donated material relevant to the present study. Holotype: Cava Grola: MPUM 11353 (Pl. 12, ig. 5). Paratypes: Cava Albanello: 1 spm., MGP-PD 1229R, 2 spms., MGP-PD 1230R, 1 spm., MCZ 4333; Cava Boschetto: 1 spm., MPUM 11354; Cava Grola: 1 spm., MPUM 11355 (Pl. 12, ig. 6), 1 spm., MCV 15/02; Cava Rossi: 1 spm., MCZ 4334. Other material examined: Cava Boschetto: 1 spm., NP 9829; Cava Grola: 2 spms., NP 9830, 3 spms., private collection, 3 spms., private collection. Preservation: All specimens have the surface somewhat abraded. Type locality: Cava Grola (see appendix). Horizon: Greenish-gray volcanoclastic sandstone of middle Lutetian age. Diagnosis: Globose, rather thin shell with moderately elevated spire and globose last whorl very slightly expanded toward aperture. Umbilicus deep, moderate, with low, subangular spiral ridge that terminates in subtriangular, asymmetric plug on lowermost part of inner lip. Funicle moderately prominent. Parietal callus narrow, with straight abapertural outline; umbilical callus thick, triangular, demarcated from anterior angle of parietal callus by sinuation of various width. Description. Protoconch medium-sized, low-turbiniform, of 1.80 convex and apparently smooth whorls, tip medium-sized. Teleoconch globose, nearly as high as wide, rather thin. Spire obtusely conical, moderately elevated, whorls convex; suture adpressed. Last whorl globose, somewhat depressed in abapertural view, very slightly expanded toward aperture; subsutural shelf indistinct; periphery about at midline. Aperture D-shaped in slightly prosocline plane, length approximately 1.6 times width. Parietal callus narrow, rather thick, with straight abapertural outline, ending at level of umbilical border. Umbilicus deep, moderate; umbilical border broadly rounded; umbilical wall steeply sloping; interior of umbilicus bearing low, subangular spiral ridge that terminates in subtriangular, asymmetric plug on lowermost part of inner lip. Funicle moderately prominent, separated from inner spiral ridge by rather narrow groove. Umbilical callus thick, triangular, demarcated from anterior angle of parietal callus by sinuation of various breadth. Basal fasciole indistinct. Surface with vestige of ine growth lines. Dimensions (mm) DHW PD H D SH 0.242* 1.129* 4.14716.323 4.73816.174 0.8865.078 * 1 protoconch measurable 10.235 10.456 2.982 AH AW UW IS SA 3.03111.475 3.04010.052 1.1654.717 14°-34° 97°-133° 7.253 6.546 2.941 24° 115° Remarks. For discussion and comparison with Payraudeautia crassicorda sp. n., see the remarks on this species. Stratigraphic occurrence. Payraudeautia zarantonelloi sp. n. was recovered from upper Ypresian, lower and middle Lutetian deposits of Veneto. Genus Pliconacca Cossmann & Martin in Martin, 1914 Pliconacca Cossmann & Martin in Martin, 1914, p. 171. Type species by monotypy: Natica (Pliconacca) trisulcata Martin, 1914, middle Eocene, Java. The type species was excellently illustrated by Leloux & Wesselingh (2009, pls. 204, 205). Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Pliconacca was originally proposed as a subgenus of Natica Lamarck, 1799 (according to Kabat 1991, Natica Lamarck is a junior homonym of Natica Scopoli, 1777, equivalent to Naticarius Dumeril, 1806). Subsequent to its introduction, Pliconacca was regarded either as a subgenus/section of Polinices Montfort, 1810 (cf. Van Winkle Palmer 1937), or as a distinct genus (cf. Ladd 1977; Majima 1989). We concur with the latter workers in considering Pliconacca a full genus. Pliconacca is distinguished by the following characters: 1) teleoconch globose to oval, 2) inner lip callus with one-three variously prominent obtuse transverse folds separated by shallow depressions, 3) umbilicus largely covered by the umbilical callus in most species, less so in some, and 4) outer surface with subsutural collabral wrinkles in most species. The genus has been sometimes compared with Glossaulax Pilsbry, 1929, which differs markedly in having a rather depressed teleoconch with the umbilical callus (broad in most species) divided medially by a deep and narrow transverse groove (there are no folds). Some New Zealand species formerly assigned to the genus Uberella Finlay, 1928 (Uberella cicatricella Marwick, 1965; Uberella cicatrix Marwick, 1931; Natica denticulifera Marwick, 1924) have been correctly transferred to Pliconacca in most recent years (cf. Beu & Raine 2009) because of their umbilical callus provided with transverse folds. Uberella somewhat resembles Pliconacca in terms of shell shape and umbilical characters, but the transverse callus folds are absent from Uberella species. The New Zealand species Pliconacca denticulifera (Marwick, 1924) seems to be the sole Recent species of the genus. Fossil occurrences of Pliconacca were from the middle Eocene of southern United States and Java (Indonesia), from the late Eocene of Japan, from lower to upper Miocene units of Indonesia, New Zealand and Fiji Islands, from the Pliocene of Japan and New Zealand, and from the Pleistocene of Japan and New Zealand. The Burdigalian Polinices cerovaensis Harzhauser, 2011, the Burdigalian and Langhian Natica (Naticina) catena var. prohelicina Sacco, 1890, and the middle Miocene Natica plicatulaeformis Kittl, 1887 agree with the characters of Pliconacca and are the representatives of the genus so far described from Europe. 193 Pliconacca plicatulaeformis (Kittl, 1887) comb. n. Pl. 12, igs. 7-11 1887 Natica plicatulaeformis Kittl, p. 257, pl. 8, igs. 23, 24. 1890b Natica (Naticina) catena var. prohelicina Sacco, p. 30 (new synonym). 1890b Natica (Naticina) catena var. prohelicina subvar. tauroumbilicata Sacco, p. 30 (nomen nudum). 1890b Natica (Naticina) catena var. ampullinoides Sacco, p. 30 (new synonym). 1890b Natica (Naticina) catena var. cyclostomoides Sacco, p. 30 (new synonym). 1890b Natica (Naticina) catena var. cyclostomoides subvar. scalarioides Sacco, p. 30 (nomen nudum). 1890b Natica (Naticina) catena var. cyclostomoides subvar. subhemiclausa Sacco, p. 30 (nomen nudum). 1891 Natica (Naticina) catena var. prohelicina - Sacco, p. 67, pl. 2, ig. 38. 1891 Natica (Naticina) catena var. tauroumbilicata Sacco, p. 67 (new synonym). 1891 Natica (Naticina) catena var. ampullinoides - Sacco, p. 67, pl. 2, ig. 39. 1891 Natica (Naticina) catena var. cyclostomoides - Sacco, p. 68, pl. 2, ig. 40. 1891 Natica (Naticina) catena var. scalarioides Sacco, p. 69 (new synonym). 1891 Natica (Naticina) catena var. subhemiclausa Sacco, p. 69 (new synonym). 1891 Natica (Naticina) catena var. varians - Sacco, p. 69, pl. 2, ig. 41 (not Dujardin, 1837). 1904 Natica (Naticina) catena var. tauroumbilicata - Sacco, p. 102, pl. 22, ig. 27. 1904 Natica (Naticina) catena var. scalarioides - Sacco, p. 102, pl. 22, ig. 28. 1904 Natica (Naticina) catena var. subhemiclausa - Sacco, p. 103, pl. 22, ig. 30. not 1952b Polynices (Euspira) cyclostomoides - Glibert, p. 247, pl. 2, ig. 1 (not Sacco, 1890). 1984 Naticina catena var. prohelicina - Ferrero Mortara et al., p. 32. 1984 Naticina catena var. tauroumbilicata - Ferrero Mortara et al., p. 32. 1984 Naticina catena var. cyclostomoides - Ferrero Mortara et al., p. 32. 1984 Naticina catena var. scalarioides - Ferrero Mortara et al., p. 32. 1984 Naticina catena var. subhemiclausa - Ferrero Mortara et al., p. 32. 1984 Naticina catena var. varians - Ferrero Mortara et al., p. 32 (not Dujardin, 1837). 2007 Euspira catena - Zunino, p. 119, pl. 1, ig. 5 (not da Costa, 1778). 2009 Euspira catena - Zunino & Pavia, pl. 1, ig. 6 (not da Costa, 1778). Type material: Natica plicatulaeformis Kittl, lectotype (here designated): the shell igured by Kittl (1887, pl. 8, ig. 24) and reigured herein (Pl. 12, ig. 7), NHMW I.26, Orlau (Czech Republic); 1 paralectotype: the shell igured by Kittl (1887, pl. 8, ig. 23) and reigured herein (Pl. 12, ig. 8), NHMW I.25, Orlau (Czech Republic); other 118 syntypes cited by Kittl (1887) were not examined. Natica (Naticina) catena var. prohelicina Sacco, lectotype (here designated): the 194 Robba E., Pedriali L. & Quaggiotto E. shell igured by Sacco (1891, pl. 2, ig. 38) and reigured herein (Pl. 12, ig. 9), MGPT BS.029.02.001, Colli Torinesi; 26 paralectotypes, MGPT BS.029.02.001/01, Villa Forzano (other 3 syntypes are Nerita helicina Brocchi, 1814, 1 is Natica submamillaris d’Orbigny, 1852 and 6 were not identiied, all also numbered BS.029.02.001/01); 3 paralectotypes, MGPT BS.029.02.001/03, Colli Torinesi; another 64 syntypes, MGPT BS.029.02.001/02, Colli Torinesi were not examined. Natica (Naticina) catena var. tauroumbilicata Sacco, holotype (by monotypy): the shell igured by Sacco (1904, pl. 22, ig. 27), MGPT BS.029.02.002, Colli Torinesi. Natica (Naticina) catena var. ampullinoides Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 2, ig. 39), MGPT BS.029.02.003, Colli Torinesi; 8 paralectotypes, MGPT BS.029.02.003/01, Colli Torinesi (another syntype, also numbered BS.029.02.003/01, was not identiied). Natica (Naticina) catena var. cyclostomoides Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 2, ig. 40), MGPT BS.029.02.004, Colli Torinesi; 23 paralectotypes, MGPT BS.029.02.004/01, Colli Torinesi (other 10 syntypes are Nerita helicina Brocchi, 1814, 8 are Euspira umbilicolunata sp. n., all also numbered BS.029.02.004/01). Natica (Naticina) catena var. scalarioides Sacco, lectotype (here designated): the shell igured by Sacco (1904, pl. 22, ig. 28), MGPT BS.029.02.005, Colli Torinesi; in MGPT there is only another syntype numbered BS.029.02.005/01, which is Nerita helicina Brocchi, 1814. Natica (Naticina) catena var. subhemiclausa Sacco, lectotype (here designated): the shell igured by Sacco (1904, pl. 22, ig. 30), MGPT BS.02.029.006, Colli Torinesi; 5 paralectotypes, MGPT BS.029.02.006/01, Colli Torinesi (other 8 syntypes, also numbered BS.029.02.006/01, are Nerita helicina Brocchi, 1814). Material erroneously referred to as Natica (Natica) millepunctata var. sismondiana d’Orbigny, 1852 in MGPT: Colli Torinesi: 1 spm., MGPT BS.029.01.003/03. Other material examined: Albugnano: 3 spms., PG 29; Termofourà: 3 spms., MGPT-PU 135244; Tetti Civera: 4 spms., MZB 15773, 24 spms., MZB 60044, 2 spms., MGPT-PU 135245; Valle Ceppi: 1 spm., MGPT-PU 135246, 1 spm., MGPT-PU 135247, 4 spms., MGPT-PU 135248, 1 spm., MGPT-PU 135249, 4 spms., MGPT-PU 135250, 1 spm., MGPT-PU 135251, 3 spms., MGPT-PU 135252, 2 spms., MGPT-PU 135253, 4 spms., MGPT-PU 135254, 1 spm., MGPT-PU 135255, 1 spm., MGPT-PU 135256,14 spms., MPUM 11356, 1 spm., MZB 60122, 6 spms., MZB 60043, 216 spms., MZB 60117, 61 spms., MZB 60005, 159 spms., MZB 60006, 3 spms., MZB 60007, 12 spms., MZB 60008, 1 spm., MZB 60009, 5 spms., MZB 60010, 57 spms., MZB 60011, 1 spm., MZB 60012, 1 spm., MZB 60013 (Pl. 12, ig. 10), 1 spm., MZB 60014, 14 spms., MZB 60020, 5 spms., MGPT-PU 135257, 16 spms., MGPT-PU 135258, 1 spm., MGPT-PU 135259 (Pl. 12, ig. 11), 29 spms., MGPTPU 135260, 1 spm., MGPT-PU 25903, 5 spms., MGPT-PU 135261, 3 spms., MGPT-PU 135262, 3 spms., MGPT-PU 135263, 2 spms., MGPT-PU 135264, 1 spm., MGPT-PU 135265, 1 spm., MGPT-PU 135266, 2 spms., MGPT-PU 135267, 3 spms., MGPT-PU 135269, 1 spm., MGPT-PU 135270, 3 spms., MGPT-PU 135271, 2 spms., MGPT-PU 135272, 1 spm., MGPT-PU 135273, 1 spm., MGPT-PU 135274, 2 spms., MGPT-PU 135275, 2 spms., MGPT-PU 135276, 38 spms., PG 28, 1 spm., NP 9809, 6 spms., NP 9825; Val Sanfrà: 1 spm., MGPT-PU 107636, 19 spms., MGPT-PU 135277; Valle Vergnana: 4 spms., MZB 23754, 1 spm., MZB 23773, 4 spms., MZB 60046, 4 spms., MZB 43255, 2 spms., MZB 43795, 2 spms., MZB 43940, 1 spm., MZB 43972, 1 spm., MZB 44020, 1 spm., MZB 60019; Villa Allason: 3 spms., MGPT-PU 135278; Villa Bertini: 13 spms., MGPTPU 135279. Description. Protoconch small, lowturbiniform, of 2.70-2.90 convex, apparently smooth whorls, tip also small. Teleoconch globose to globose-pyriform, moderately thick. Spire conical, rather elevated in some specimens, whorls convex. Suture thin, adpressed. Last whorl globose to globose-oval, only slightly extended toward aperture; subsutural shelf poorly deined; periphery at midline. Aperture D-shaped in gently prosocline plane, length about 1.65 times width. Parietal callus thick, subrectangular, with concave abapertural outline; anterior lobe well developed, rather broad, with rounded to squarish edge; two transverse abapical folds occur at level of anterior lobe, lower rather prominent, other slightly above, narrower and shorter in most specimens, of same strength or obsolescent in some. Umbilicus deep, narrow to very narrow, more widely open in some specimens, margin broadly rounded, hence demarcation from base poorly deined; umbilical wall subvertical; inner surface with coarse growth markings. Funicle obsolete. Umbilical callus rather thick, narrowly subtriangular, with oblique reverse J-shaped outline, merging into anterior lobe of parietal callus and largely covering adapical part of umbilicus in most specimens, less so in some. Basal fasciole indistinct. Surface with subsutural prosocline collabral wrinkles. Dimensions (mm) DHW PD H D SH 0.1030.111 0.9371.013 9.89520.231 8.95317.537 1.7167.008 0.107 0.975 15.063 13.245 4.362 AH AW UW IS SA 7.42113.993 4.9989.870 1.0712.823 20°-40° 70°-118° 10.707 7.434 1.947 30° 94° Remarks. The present species is herein assigned to the genus Pliconacca Cossmann & Martin in Martin, 1914 on account of its subsutural collabral wrinkles and its transverse folds on the abapical part of the parietal callus. It appears to be rather variable as regards the teleoconch shape, the elevation of the spire, the width of the umbilical opening, and the development of the folds on the parietal callus. Sacco (1890b) introduced the variety prohelicina of Natica (Naticina) catena (da Costa, 1778), a species currently included in the genus Euspira Agassiz in J. Sowerby, 1837. The variety prohelicina is morphologically similar to Euspira catena, but dif- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy fers from da Costa’s taxon because of its transverse folds on the abapical part of the parietal callus (absent in Euspira catena). We conclude that this difference is adequate to warrant speciic separation from E. catena and that the cited character irmly points toward an assignment of Sacco’s taxon to the genus Pliconacca. Examination of the igured syntypes of Pliconacca plicatulaeformis in NHMW revealed that Pliconacca prohelicina does not differ signiicantly from P. plicatulaeformis and, therefore, we consider it as a synonym of Kittl’s species, which has priority. The same conclusion proved true as regards other Sacco’s varieties of Natica (Naticina) catena (see the above synonymy). Polinices cerovaensis Harzhauser, 2011, described from Burdigalian deposits of Slovak Republic, also belongs in Pliconacca. According to Harzhauser (personal communication 2013), it differs from P. plicatulaeformis in that it lacks the subsutural collabral wrinkles. Stratigraphic occurrence. Pliconacca plicatulaeformis (Kittl, 1887) occurs in Burdigalian and Langhian units of Piedmont, being particularly abundant in the early Burdigalian of Valle Ceppi, and in the Badenian (middle Miocene) of Central Paratethys. Pliconacca tortonensis sp. n. Pl. 13, igs. 1-3 Derivation of name: The name refers to the Tortonian age of the type horizon. Holotype: Borelli: MGPT-PU 135280 (Pl. 13, ig. 1). Paratypes: Borelli: 1 spm., MGPT-PU 135281 (Pl. 13, ig. 2), 1 spm., MGPT-PU 135282 (Pl. 13, ig. 3), 1 spm., MGPT-PU 135283, 1 spm., MGPT-PU 135284, 1 spm., MGPT-PU 135285, 22 spms., MGPT-PU 135286, 1 spm., MPUM 11357. Other material examined: Borelli: 2 spms., NP 9823; Stazzano: 1 spm., NP 9824, 1 spm., MZB 60015, 1 spm., MZB 60016. Preservation: All specimens are rather well preserved. Type locality: Borelli (see appendix). Horizon: Gray, medium sand of Tortonian age. Diagnosis: Globose shell with slightly elevated spire, rather depressed last whorl, small to very small umbilicus; inner lip callus with three transverse folds, one on uppermost umbilical callus, two at level of anterior lobe of parietal callus. Surface with subsutural prosocline collabral wrinkles. Description. Protoconch medium-sized, low-turbiniform, of 2.70-2.84 convex, apparently smooth whorls, tip small. Teleoconch globose, of moderate thickness. Spire broadly conical, slightly elevated, whorls convex. Suture thin, adpressed. Last whorl globose, rather depressed, scarcely 195 extended toward aperture; subsutural shelf poorly deined to indistinct; periphery at midline. Aperture D-shaped in gently prosocline plane, length about 1.8 times width. Parietal callus thick, subrectangular, with concave abapertural outline; anterior lobe well developed, rather broad, with rounded to squarish edge; two transverse abapical folds occur at level of anterior lobe, lower rather prominent, other some distance above, shorter in most specimens, narrower and extending into aperture in some. Umbilicus deep, small to very small, margin broadly rounded, hence demarcation from base poorly deined; umbilical wall subvertical, bounded downward by slight groove. Funicle obsolete. Umbilical callus rather thick, narrowly subtriangular, with oblique reverse J-shaped outline and with 1 transverse fold where it merges into anterior lobe of parietal callus. Basal fasciole indistinct. Surface with subsutural prosocline collabral wrinkles; last whorl with faint spiral microstriation throughout. Some specimens retain traces of pale reddish-brown color. Dimensions (mm) DHW PD H D SH 0.1150.131 0.9701.142 2.80412.728 2.29611.368 0.8863.018 0.123 1.056 7.766 6.832 1.952 AH AW UW IS SA 1.8189.850 1.2707.934 0.3972.765 12°-28° 105°-125° 5.834 4.602 1.581 20° 115° Remarks. The present new species has a quite constant teleoconch shape, whereas it exhibits a moderate variability concerning the development of the transverse folds on the parietal callus. The values of the characteristic elements of the protoconch of Pliconacca tortonensis do not differ signiicantly from those measured for Pliconacca plicatulaeformis (Kittl, 1887), but the two species can be readily distinguished on the basis of respective teleoconch characters. P. tortonensis differs from P. plicatulaeformis in that it has: 1) a lower spire (Fig. 13) with greater mean spiral angle, 2) a more depressed last whorl, 3) a wider aperture (Fig. 14), 4) a wider umbilicus (Fig. 16), 5) an umbilical wall bounded downward by a slight but distinct groove (absent in P. plicatulaeformis), and 6) a transverse fold on the adapical end of the umbilical callus that is never present in P. plicatulaeformis. The three-folded inner lip callus stands as the main differentiating character. 196 Robba E., Pedriali L. & Quaggiotto E. Stratigraphic occurrence. Pliconacca tortonensis sp. n. was recovered from Tortonian deposits of Piedmont. Genus Polinices Montfort, 1810 Polinices Montfort, 1810, p. 222. Type species by original designation: Polinices albus Montfort, 1810 (= Nerita mammilla Linnaeus, 1758), Recent, Indo-Pacific. Kabat (1990) selected the lectotype of Nerita mammilla, designated it as the neotype of Polinices albus, and redescribed Linnaeus’ species (see also Bouchet & Waren 1993). Albula Röding, 1798, p. 20. Type species by subsequent des-ignation (Winckworth 1945, p. 137): Nerita mammilla Linnaeus, 1758. Eucaryorum Ehrenberg, 1831, p. 46. Type species by mono-typy: Nerita mammilla Linnaeus, 1758 (fide Kabat 1991). Naticina Guilding, 1834, p. 30. Type species by original des-ignation: Naticina lactea Guilding, 1834. Naticella Swainson, 1840, p. 345. Type species by monotypy: “N. aurantia Martini” Swainson, 1840 (= Albula aurantium Röding, 1798). Uber Gray, 1847, p. 149 (Uber Humphrey, 1797 unavailable). Type species by subsequent designation (Philippi 1853, p. 497): Nerita mammilla Linnaeus, 1758. Mamma H. & A. Adams, 1853, p. 210 (Mamma Klein MS.; Mamma Klein in Mörch, 1852 merely listed in the synonymy of Polinices). Type species by original designation: Nerita mammilla Linnaeus, 1758. Pseudopolinices Golikov & Sirenko, 1983, p. 1339. Type species by original designation: Natica nana Möller, 1842. Kabat (1991) regarded Pseudopolinices as a synonym of Euspira Agassiz in J. Sowerby, 1837. However, Natica nana (type species of Pseudopolinices) has the umbilicus completely filled by a rather broad umbilical callus fused with the parietal callus (cf. Golikov & Sirenko 1983, fig. 1: 20), a character that is distinctive of Nerita mammilla. Consequently, we regard Pseudopolinices as another synonym of Polinices. Polinices was discussed by Pedriali & Robba (2009, p. 388), who listed the following distinctive characters of the genus: 1) shell usually thick, pyriform to slightly globular, 2) spire moderately el-evated to low, sutures almost flush, 3) parietal callus thick and broad, 4) umbilicus completely plugged or a narrow and deep abapical groove, and 5) umbilical callus thick, broad in most species, fused with the parietal callus. The cited authors considered the umbilical callus completely filling the umbilicus or nearly so, and its smooth merging with the parietal callus as the main distinguishing characters of Polinices. Polinices proredemptus (Sacco, 1890) Pl. 13, igs. 4-6 ? 1856 Natica redempta - Hoernes, p. 522, pl. 47, ig. 3. 1890b Natica (Polinices) proredempta Sacco, p. 36. 1890b Natica (Polinices) proredempta subvar. scalariformis Sacco, p. 36 (nomen nudum). 1890b Natica (Polinices) proredempta var. subnaticoides Sacco, p. 36. 1890b Natica (Polinices) proredempta var. tauromamilla Sacco, p. 36. 1891 Natica (Polinices) proredempta - Sacco, p. 93, pl. 2, ig. 71. 1891 Natica (Polinices) proredempta var. scalariformis Sacco, p. 94. 1891 Natica (Polinices) proredempta var. subnaticoides - Sacco, p. 94, pl. 2, ig. 72. 1891 Natica (Polinices) proredempta var. tauromamilla - Sacco, p. 94, pl. 2, ig. 73. 1904 Natica (Polinices) proredempta var. scalariformis - Sacco, p. 104, pl. 23, ig. 6. ? 1919 Natica (Polinices) proredempta - Cossmann & Peyrot, p. 221, pl. 12, igs. 23-26. ? 1923 Natica redempta var. Friedberg, p. 433, pl. 26, igs. 6, 7. PLATE 13 Fig. 1 - Pliconacca tortonensis sp. n. Borelli. Holotype, MGPT-PU 135280; a, apertural side (height of shell 12.65 mm); b, abapertural side; c, apical view; d, basal view. Fig. 2 - Pliconacca tortonensis sp. n. Borelli. Paratype, MGPT-PU 135281; protoconch. Fig. 3 - Pliconacca tortonensis sp. n. Borelli. Paratype, MGPT-PU 135282; a, apertural side (height of shell 6.07 mm); b, basal view. Fig. 4 - Polinices proredemptus (Sacco, 1890). Colli Torinesi. Lectotype (here designated) of Natica (Polinices) proredempta Sacco, 1890. MGPT BS.029.06.010; a, apertural side (scale bar 4 mm); b, basal view. Fig. 5 - Polinices proredemptus (Sacco, 1890). Valle Ceppi. MZB 60134; protoconch. Fig. 6 - Polinices proredemptus (Sacco, 1890). Val Sanfrà. MGPT-PU 135287; apertural side (height of shell 24.08 mm). Fig. 7 - Polinices redemptus (Michelotti, 1847). Stazzano. Neotype (here designated) of Natica redempta Michelotti, 1847. MGPT BS.029.06.014; a, apertural side (scale bar 4 mm); b, abapertural side. Fig. 8 - Polinices redemptus (Michelotti, 1847). Montegibbio. NP 10000; protoconch. Fig. 9 - Polinices redemptus (Michelotti, 1847). Rio di Bocca d’Asino. PG 102; a, apertural side (height of shell 38.19 mm); b, abapertural side; c, apical view. Fig. 10 - Polinices redemptus (Michelotti, 1847). Rio di Bocca d’Asino. PG 103; abapertural side showing color pattern (height of shell 31.81 mm). Fig. 11 - Polinices submamilla (Sacco, 1891). Colli Torinesi. Lectotype (here designated) of Natica (Polinices) submamilla Sacco, 1891. MGPT BS.029.06.002; apertural side (scale bar 4 mm). Fig. 12 - Polinices submamilla (Sacco, 1891). Colli Torinesi. Paralectotype. MGPT BS.029.06.002/02a; protoconch. Fig. 13 - Polinices submamilla (Sacco, 1891). Valle Ceppi. MGPT-PU 135288; apertural side (height of shell 16.62 mm). Fig. 14 - Polinices submamilla (Sacco, 1891). Valle Ceppi. PG 104; apertural side (height of shell 12.00 mm). Fig. 15 - Polinices sp. Valle Organa. MGP-PD 31519; apertural side (height of shell 16.53 mm). Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 197 198 Robba E., Pedriali L. & Quaggiotto E. ? 1923 Natica staszici Friedberg, p. 435, pl. 26, igs. 8, 9. ? 1925 Natica (Polynices) proredempta - Cossmann, p. 126, pl. 2, igs. 17, 18. ? 1966 Natica (Polynices) redempta - Strausz, p. 231 (pars), pl. 47, igs. 29, 30 (not igs. 19-28, 31, 32 = Natica pseudoredempta Friedberg, 1923). 1966 Polynices (Lunatia) ex gr. proredempta - Kόkai, p. 55, pl. 7, ig. 7. 1984 Polinices proredempta - Ferrero Mortara et al., p. 37, pl. 4, ig. 1. 1984 Polinices proredempta var. scalariformis - Ferrero Mortara et al., p. 37. 1984 Polinices proredempta var. subnaticoides - Ferrero Mortara et al., p. 37. 1984 Polinices proredempta var. tauromamilla - Ferrero Mortara et al., p. 37. ? 1995 Polinices redemptus - Baluk, p.198, pl. 15, igs. 7, 8. 2007 Polinices proredemptus - Zunino, p. 122, pl. 1, ig. 8. 2009 Polinices proredemptus - Zunino & Pavia, pl. 1, ig. 7. Type material: Natica (Polinices) proredempta Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 2, ig. 71) and reigured herein (Pl. 13, ig. 4), MGPT BS.029.06.010, Colli Torinesi; 7 paralectotypes, MGPT BS.029.06.010/01, Colli Torinesi (other 2 syntypes, also numbered MGPT BS.029.06.010/01, are unidentiiable). Natica (Polinices) proredempta var. scalariformis Sacco, lectotype (here designated): the shell igured by Sacco (1904, pl. 23, ig. 6), MGPT BS.029.06.011, Colli Torinesi; 1 paralectotype, MGPT BS.029.06.011/01, Colli Torinesi (other 2 syntypes, also numbered MGPT BS.029.06.011/01, are Natica tigrina Defrance, 1825). Natica (Polinices) proredempta var. subnaticoides Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 2, ig. 72), MGPT BS.029.06.012, Colli Torinesi; 17 paralectotypes, MGPT BS.029.06.012/01, Colli Torinesi (other 4 syntypes, also numbered MGPT BS.029.06.012/01, are unidentiiable or belong to another species). Natica (Polinices) proredempta var. tauromamilla Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 2, ig. 73), MGPT BS.029.06.013, Colli Torinesi. Material erroneously referred to as Natica (Polinices) submamillaris var. mioinlata Sacco, 1890 in MGPT: Colli Torinesi: 5 syntypes, MGPT BS.029.06.004/01. Other material examined: Salles, Argilas (France): 22 spms., NP 10016; Salles, Largileyre (France): 3 spms., NP 10003; Valle Ceppi: 1 spm., MGPT-PU 23859, 1 spm., MZB 60135, 1 spm., MZB 60134 (Pl. 13, ig. 5); Val Sanfrà: 1 spm., MGPT-PU 135287 (Pl. 13, ig. 6), 15 spms., MGPT-PU 23665, 1 spm., MGPT-PU 107628; Valle Vergnana: 1 spm., MZB 60136, 1 spm., MZB 60137, 1 spm., MZB 60152. Description. Protoconch medium-sized, low-turbiniform, of 2.45 convex, apparently smooth whorls, tip small. Teleoconch globose, wider than high in most specimens, sometimes globosepyriform, moderately thick. Spire broadly conical, pointed and low, more elevated in a few specimens (globose-pyriform), whorls latly convex. Suture thin, almost lush, adpressed in a few specimens that have more convex whorls. Last whorl inlated, somewhat depressed, only slightly extended toward aperture; subsutural shelf poorly deined to indi- stinct; periphery slightly above midline. Aperture D-shaped in moderately prosocline plane, length averaging 1.85 times width. Parietal callus thick. Umbilicus completely plugged or a narrow abapical groove in a few specimens. Umbilical callus rather thick, subcircular, expanded to overlap basal fasciole, slightly excavated in most specimens, fused with parietal callus. Abapertural outline of inner lip callus (parietal + umbilical) concave, straight in a few specimens. Basal fasciole broad, scarcely prominent. Many specimens retain vestige of uniform pale brown background with darker subsutural and lowermost basal bands. Dimensions (mm) DHW PD H D SH 0.137* 1.029* 8.55031.990 9.02030.680 1.6207.020 * 1 protoconch measurable 20.276 19.858 4.326 AH AW WUC IS SA 6.52025.380 5.40017.800 2.66011.280 18°-34° 99°-131° 15.955 11.600 6.975 26° 115° Remarks. Sacco (1890b) introduced Natica (Polinices) proredempta on the basis of material from the Torino Hills (Colli Torinesi) said to be of “Helvetian” age, and pointed out the characters that distinguish this “Helvetian” species from the Tortonian Natica (Polinices) redempta Michelotti, 1847. Glibert (1952b) regarded the distinguishing characters cited by Sacco (1890b) as “peu apparents” and, in order to clarify the relationships between the “Helvetian” species and the Tortonian one, compared some middle Miocene shells from France and Austria to specimens from the Tortonian of Montegibbio “représentant P. redempta typique”. Glibert concluded that the middle Miocene French and Austrian specimens were not distinguishable from those of Montegibbio. Consequently, he rejected the separation of Polinices proredemptus from Polinices redemptus made by Sacco. Glibert’s opinion was agreed upon subsequently by some workers (Strausz 1954; Ruggieri & Davoli 1984; Baluk 1995; Landau et al. 2013). However, considering 1) that Glibert (1952b) did not examine specimens from the “Helvetian” of Piedmont, and 2) that recent research has shown that the “Helvetian” fossils dealt with by Sacco were collected from coarse sandstone levels of the Termofourà Formation, demonstrated to be of early Miocene (Burdigalian) age (see Zunino & Pavia 2009), Eocene, Oligocene and Miocene naticid gastropods of Northern Italy we think that Glibert’s conclusion is not acceptable since it is based on specimens of different age, but all belonging to P. redemptus, which appears to occur also in the middle Miocene (Serravallian). In our opinion, Polinices proredemptus is a distinct species differing from Polinices redemptus in that it has: 1) a protoconch with 0.5 more whorls and with the diameter of the irst half-whorl half of the size, 2) a teleoconch attaining a smaller size, 3) a nearly lat-sided spire (that of P. redemptus is somewhat stepped, with more convex whorls), and 4) a quite different color pattern. Furthermore, P. proredemptus lacks the subsutural groove that is always present in P. redemptus. Specimens either assigned to Polinices proredemptus or to Polinices redemptus were illustrated from middle Miocene deposits of France (Cossmann & Peyrot 1919; Cossmann 1925) and eastern Europe (Hoernes 1856; Friedberg 1923; Strausz 1954, 1966; Kojumdgieva & Strachimirov 1960; Kόkai 1966; Baluk 1995). We have not seen the material dealt with by the cited authors, and it is dificult to decide any species assignment on the basis of the igures published by them. However, it seems that the French specimens (Cossmann & Peyrot 1919; Cossmann 1925) could be P. proredemptus. Of the eastern European shells, those illustrated by Hoernes (1856), Friedberg (1923), Strausz (1966, in part) and Baluk (1995) probably are P. proredemptus (see the above synonymy); that igured by Kόkai (1966) is P. proredemptus. Finally, we note that several Badenian (Langhian to early Serravallian) shells from Austria, Poland, Hungary and Ukraine in our collection fully agree with the characters of P. proredemptus, whereas others conform to P. redemptus. In summary, P. proredemptus is well documented in the early Miocene (Burdigalian) of Italy as is P. redemptus in the late Miocene (Tortonian) also of Italy; so far, there are no records of these species from the Italian middle Miocene. Both taxa result to have co-occurred in the middle Miocene of other European countries. Natica staszici Friedberg, 1923 appears to be closely similar to Polinices proredempta and could be a synonym of it. The var. revoluta Friedberg, 1923 of Natica staszici seems to be a different species belonging in the genus Euspira Agassiz in J. Sowerby, 1837. Stratigraphic occurrence. Polinices proredemptus (Sacco, 1890) is known deinitely from the Burdigalian Termofourà Formation in the Torino 199 Hills (Piedmont). Sacco (1891) quoted his variety subnaticoides of Natica (Polinices) proredempta as very rare in the Tortonian deposits of Sant’Agata Fossili (Piedmont). However, since no specimens of this variety in MGPT are from the cited locality, the presence of Polinices proredemptus in the Tortonian of Piedmont is to be conirmed. The species occurs also in the middle Miocene of eastern Europe and, probably, of France. Polinices redemptus (Michelotti, 1847) Pl. 13, igs. 7-10 1847 Natica redempta Michelotti, p. 157, pl. 6, igs. 6, 61. 1890b Natica (Polinices) redempta - Sacco, p. 36. 1890b Natica (Polinices) redempta subvar. subalbula Sacco, p. 36 (nomen nudum). 1890b Natica (Polinices) redempta subvar. elliptica Sacco, p. 36 (nomen nudum). 1890b Natica (Polinices) redempta var. dertoconvexa Sacco, p. 36. 1891 Natica (Polinices) redempta - Sacco, p. 95, pl. 2, ig. 74. 1891 Natica (Polinices) redempta var. subalbula Sacco, p. 95. 1891 Natica (Polinices) redempta var. elliptica Sacco, p. 96. 1891 Natica (Polinices) redempta var. dertoconvexa - Sacco, p. 96, pl. 2, ig. 75. 1903 Natica (Polinices) redempta - Dollfus et al., p. 18, pl. 35, ig. 1. 1904 Natica (Polinices) redempta var. elliptica - Sacco, p. 104, pl. 23, ig. 7. ? 1917 Natica redempta - Stefanini, p. 99. 1952b Polynices (Polynices) redempta - Glibert, p. 252, pl. 2, igs. 5a-i. 1963 Polinices (Polinices) redemptus - Venzo & Pelosio, p. 83, pl. 34, igs. 39-42. 1968 Polinices (Polinices) redemptus - Robba, p. 527. 1973 Polinices (Polinices) redemptus - Marasti, p. 87, pl. 20, igs. 6, 7. 1984 Polinices redempta - Ferrero Mortara et al., p. 37. 1984 Polinices redempta var. subalbula - Ferrero Mortara et al., p. 37. 1984 Polinices redempta var. elliptica - Ferrero Mortara et al., p. 37. 1984 Polinices redempta var. dertoconvexa - Ferrero Mortara et al., p. 37. 1984 Polinices (Polinices) redemptus - Ruggieri & Davoli, p. 55, pl. 10 [1], ig. 19. 2011 Euspira redempta - Caprotti, p. 53, igs. 2M-P. 2013 Polinices redemptus - Landau et al., p. 105, pl. 11, ig. 7; pl. 79, ig. 7. Type material: Natica redempta Michelotti, neotype (here designated): the specimen igured by Sacco (1891, pl. 2, ig. 74) and reigured herein (Pl. 13, ig. 7), MGPT BS.029.06.014, Stazzano (see remarks below). Natica (Polinices) redempta var. subalbula Sacco, lectotype (here designated): the specimen MGPT BS.029.06.015, Stazzano; 4 paralectotypes, MGPT BS.029.06.015/01, Sant’Agata Fossili. Natica (Polinices) redempta var. elliptica Sacco, lectotype (here designated): the shell igured by Sacco (1904, pl. 23, ig. 7), MGPT BS.029.06.016, Stazzano; 7 paralectotypes, MGPT BS.029.06.016/01, Stazzano. Natica (Polinices) redempta var. dertoconvexa Sacco, lectotype (here des- 200 Robba E., Pedriali L. & Quaggiotto E. ignated): the shell igured by Sacco (1891, pl. 2, ig. 75), MGPT BS.029.06.017, Stazzano; 5 paralectotypes, MGPT BS.029.06.017/01, Sant’Agata Fossili, 1 paralectotype, MGPT BS.029.06.017/02, Stazzano, 5 paralectotypes, MGPT BS.029.06.017/03, Stazzano. Material erroneously referred to as Natica submamillaris var. mioclausa Sacco, 1890 in IPUM: Montegibbio: 1 spm., IPUM 4395. Other material examined: Colli Torinesi: 1 spm., MGPT BS.029.06.014/01; Colli Tortonesi: 5 spms., MGPT BS.029.06.014/07; Moncucco Torinese: 1 spm., PG 1c; Montegibbio: 1 spm., MZB 005036, 2 spms., MZB 005724, 1 spm., MZB 60113, 1 spm., MZB 60014, 1 spm., MZB 60115, 3 spms., MGC 1419, 6 spms., private collection, 2 spms., private collection, 12 spms., NP 9811, 1 spm., NP 10000 (Pl. 13, ig. 8), 7 spms., private collection, 6 spms., private collection, 14 spms., private collection, 3 spms., private collection; Passo dei Meloni: 1 spm., PP.PM 01/04, 3 spms., NP 9954; Pietracuta: 1 spm., MS 8940,0; Rio di Bocca d’Asino: 1 spm., MGPT BS.029.06.014/06, 1 spm., MZB 60001, 3 spms., MZB 60002, 2 spms., MZB 29718, 11 spms., MZB 29726, 9 spms., MZB 29759, 12 spms., MZB 29801, 3 spms., MZB 25955, 6 spms., MZB 60003, 8 spms., MZB 60004, 19 spms., MZB 45451, 10 spms., PG 25b, 1 spm., PG 102 (Pl. 13, ig. 9), 1 spm., PG 103 (Pl. 13, ig. 10), 11 spms., NP 9812, 2 spms., MPUM 11358, 6 spms., private collection; Sant’Agata Fossili: 1 spm., MGPT BS.029.06.014/03, 1 spm., MGPT-PU 23320, 2 spms., NP 9822, 2 spms., MPUM 11359; Scipione Ponte: 1 spm., private collection; Sogliano al Rubicone: 3 spms., MS 8917,0; Stazzano: 6 spms., MGPT BS.029.06.014/04, 4 spms., MGPT BS.029.06.014/05, 3 spms., MGPT-PU 23405, 1 spm., MGPT-PU 23406, 2 spms., MPUM 11360, 5 spms., MPUM 11361, 2 spms., MS 12824, 1 spm., NP 9810, 8 spms., NP 9813, 2 spms., NP 9821, 11 spms., private collection; Vigoleno: 12 spms., private collection. Description. Protoconch medium-sized, low turbiniform, of 1.75-1.90 moderately convex whorls, tip medium-sized, rather inlated. Teleoconch globose-ovate, slightly higher than wide, robust. Spire moderately elevated, somewhat stepped, whorls convex, with distinct, gently sloping subsutural shelf in most specimens. Suture thin, adpressed. Last whorl inlated, somewhat depressed, slightly expanded toward aperture; subsutural shelf less deined than on spire whorls, but distinct in all specimens; periphery about at midline. Aperture D-shaped in moderately prosocline plane, length averaging 1.83 times width. Parietal callus thick, subrectangular, slightly expanded adapically to meet suture. Umbilicus completely plugged or a narrow abapical groove in a few specimens. Umbilical callus also thick, subrectangular with arched abapical outline, touching basal fasciole, fused with parietal callus. Abapertural outline of inner lip callus (parietal + umbilical) straight, very slightly concave in a few specimens. Basal fasciole broad, scarcely prominent, bounded by very slightly incised groove in most specimens. Outer surface with subsutural spiral groove separating from suture by second or third (more frequently) spire whorl, often appearing as breakage line of shell. Many specimens retain vestige of uniform pale brown background with darker crescent-shaped marks roughly arranged into collabral alignments, observable on the last 1.5 whorls. Dimensions (mm) DHW PD H D SH 0.2710.295 0.9381.110 11.17541.939 9.98140.681 1.9729.896 0.283 1.024 26.557 25.331 5.934 AH AW WUC IS SA 8.63332.623 5.84322.063 2.05016.606 13°-33° 95°-115° 20.623 13.953 9.328 23° 105° Remarks. Inquiries about the location of Michelotti collection had negative results and that collection is to be considered lost (Giulio Pavia, personal communication 2013). Since Sacco (1891, p. 95), in the remarks to Natica (Polinices) redempta, explicitly afirmed to have examined Michelotti’s type specimens, his specimens in MGPT, collected from the same area (“environs de Tortone”), stand as the most relevant reference material for Polinices redemptus. Thus, it is advisable to designate the shell igured by Sacco (1891, pl. 2, ig. 74) as the neotype of P. redemptus, in order to clarify and ix the characters of Michelotti’s species (ICZN 1999, Article 75 of the Code). Polinices redemptus exhibits a considerable variability in shell shape, relected by the varieties proposed by Sacco (1890b, 1891). However, the diagnostic characters (protoconch, inner lip callus, subsutural groove and color pattern) are rather stable. The lower Messinian specimens from Vigoleno differ from the rest of the examined material only in that they have a wider abapical groove (umbilical channel); this difference is not considered suficient to warrant separation from the typical form. The relationships with Polinices proredemptus (Sacco, 1890) have been already dealt with in the remarks on that species (see above). Sacco (1890b) introduced Natica (Polinices) redemptoaurantia on the basis of extremely rare material from the Tortonian of Mioglia di Montaldo in the Torino Hills, said to be in the Museo Geologico of Roma. Inquiries there had no answer and no specimens of this species are present in MGPT. According to the igures and the brief descriptive remarks published by Sacco (1891, Eocene, Oligocene and Miocene naticid gastropods of Northern Italy p. 96, pl. 2, ig. 76), N. (Polinices) redemptoaurantia seems to be strikingly close to Polinices redemptus, in particular to its variety elliptica herein included in the synonymy of the typical form. Likely Sacco’s species is a synonym of the variable P. redemptus, but a decision in this respect would require that the original material (probably a single specimen) be examined. Pister & Wegmüller (2007) described and figured specimens referred to as Polinices cf. redemptus. Because of the preservation (mostly internal casts), an attempt to assign them to species can be hardly made. However, considering the Burdigalian age of these specimens, they could be referred to as Polinices cf. proredemptus. Stratigraphic occurrence. Polinices redemptus (Michelotti, 1847) was recorded from middle Miocene deposits of France, Paratethys and Turkey. It occurs in the Tortonian and in the early Messinian of Italy and Portugal (according to Studencka & Zieliński 2013, the age of the Portuguese Cacela Formation is late Tortonian to early Messinian). There are no reliable records of this species prior to middle Miocene and subsequent to the early Messinian. Polinices submamilla (Sacco, 1891) Pl. 13, igs. 11-14 1847 Natica mamilla - Sismonda, p. 51 (not Natica mamilla Lamarck, 1822 = Nerita mammilla Linnaeus, 1758). 1852 Natica submamilla d’Orbigny, p. 38 (nomen nudum). 1861 Natica submamilla - Michelotti, p. 87. 1890b Natica (Polinices) submamilla - Sacco, p. 36 (nomen nudum). 1891 Natica (Polinices) submamilla Sacco, p. 92, pl. 2, ig. 67. not 1903 Natica (Polinices) submamilla - Dollfus et al., p. 19, pl. 35, ig. 8 (not Sacco, 1891). 1984 Polinices submamillaris - Ferrero Mortara et al., p.36 (only BS.029.06.002, see remarks below). Type material: Natica (Polinices) submamilla Sacco, lectotype (here designated); the shell igured by Sacco (1891, pl. 2, ig. 67) and reigured herein (Pl. 13, ig. 11), MGPT BS.029.06.002, Colli Torinesi; 1 paralectotype (Pl. 13, ig. 12), MGPT BS.029.06.002/02a, Colli Torinesi; 15 paralectotypes, MGPT BS.029.06.002/02b, Colli Torinesi; 6 paralectotypes, MGPT BS.029.06.002/01, Villa Forzano (another syntype, also numbered MGPT BS.029.06.002/01, is Natica olla de Serres, 1829). Other material examined: Valle Ceppi: 1 spm., MGPT-PU 135288 (Pl. 13, ig. 13); 1 spm., PG 104 (Pl. 13, ig. 14). Description. Protoconch small, low turbiniform, of 2.75 moderately convex, apparently smooth whorls, tip very small. Teleoconch oval, higher than wide, rather thin. Spire elevated, broadly cyr- 201 toconoid, whorls very slightly convex, lat-sided in a few specimens. Suture thin, adpressed to almost lush. Last whorl oval, subsutural shelf indistinct, periphery about at midline. Aperture D-shaped in moderately prosocline plane, length twice width. Parietal callus thick, subrectangular. Umbilicus completely plugged. Umbilical callus very thick, subrectangular with arched abapical outline, overlapping basal fasciole, fused with parietal callus. Abapertural outline of inner lip callus (parietal + umbilical) straight, very slightly concave in a few specimens. Basal fasciole broad, scarcely prominent, bounded by slight step in most specimens. Outer surface with rather distant growth markings. A few better preserved specimens have yellowish-white, shining surface. Dimensions (mm) DHW PD H D SH AH AW 0.069 0.742 16.620 13.971 3.022 13.601 10.062 UW IS SA 8.220 24° 106° Remarks. Sismonda (1847) applied the name Natica mamilla Lamarck, 1822 (= Nerita mammilla Linnaeus, 1758) to Miocene fossils from the Torino Hills, which were neither described, nor illustrated. Since Nerita mammilla is a Recent Indo-Paciic species, d’Orbigny (1852, p. 38) introduced the replacement name submamilla for the fossil shells dealt with by Sismonda. D’Orbigny just listed (n° 563) a Natica submamilla d’Orbigny, 1847, but this reference was not located. N. submamilla was later on attributed to d’Orbigny, 1852 by the authors. However, from the 1852 work, it appears that the name submamilla is a nomen nudum and, thence, unavailable. Subsequent to its introduction, N. submamilla d’Orbigny, 1852 was only cited by Michelotti (1861, p. 87) and Sacco (1890b, p. 36). Sacco (1891) is to be considered the irst reference author. He published a concise description, some remarks, and igured the species. Thus, the name submamilla was made available by Sacco (1891), who is to be considered the author of this species. It is to be noted that in the Catalogue of the Bellardi e Sacco collection (Ferrero Mortara et al. 1984, p. 36), probably because of an oversight, the whole material of Polinices submamilla (registration numbers BS.029.06.002, BS.029.06.002/01 and BS.029.06.002/02) was listed as Polinices submamillaris (d’Orbigny, 1852). Robba E., Pedriali L. & Quaggiotto E. 202 Polinices albus Montfort, 1810 (= Nerita mammilla Linnaeus, 1758) differs from Polinices submamilla because of its protoconch with 0.5 fewer whorls and signiicantly smaller diameter, and its more inlated, ovate-pyriform to pyriform teleoconch (see Huelsken et al. 2012 for the protoconch of Nerita mammilla). The shell igured by Dollfus et al. (1903) and referred to as Natica (Polinices) submamilla does not agree with the characters of Polinices submamilla in that it has a globose teleoconch and a somewhat different abapical outline of the umbilical callus. Stratigraphic occurrence. Polinices submamilla (Sacco, 1891) was hitherto recorded from Burdigalian deposits exposed in the Torino Hills. Polinices sp. Pl. 13, ig. 15 Material examined: Valle Organa: 1 spm., MGP-PD 31519. Descripion. Protoconch poorly preserved, apparently of two whorls. Teleoconch globose, nearly as wide as high, thick, damaged abapically. Spire broadly conical, pointed, scarcely elevated, whorls very slightly convex; suture adpressed, almost lush. Last whorl globular, subsutural shelf indistinct, periphery at midline. Aperture D-shaped in slightly prosocline plane. Parietal callus thick, more so adapically. Umbilicus largely plugged, appearing as narrow, crescent-shaped abapical groove. Umbilical callus rather thick, subcircular, expanded against umbilical border, fused with parietal callus. Abapertural outline of inner lip callus (parietal + umbilical) concave. Basal fasciole not observable, likely present. Outer surface with uneven growth markings, more prominent and prosocyrt subsuturally. Dimensions (mm) DHW PD H D SH AH AW 16.130 4.742 11.790 9.759 - - 16.531 UW IS SA - 21° 134° Remarks. The preservation of the present specimen, erroneously labeled Natica canovae Oppenheim, 1900 in MGP-PD, prevents any reliable comparison with named species. Stratigraphic occurrence. Polinices sp. was recovered from Priabonian marly deposits of Veneto. Genus Sigatica Meyer & Aldrich, 1886 Sigatica Meyer & Aldrich, 1886, p. 42. Type species by monoypy: Sigaretus (Sigatica) boettgeri Meyer & Aldrich, 1886, Eocene, United States. Gennaeosinum Iredale, 1929, p. 279. Type species by original designation: Gennaeosinum peleum Iredale, 1929. Glyptanatica Gardner, 1947, p. 554. Type species by original designation: Sigatica (Glyptanatica) euglypta Gardner, 1947. Nerinatica Olsson, 1930, p. 68. Type species by original designation: Natica (Nerinatica) paytensis Olsson, 1930. Kabat (1991, p. 430) regarded Heliconatica Dall, 1924 as a subjective synonym of Sigatica. However, Eunaticina (Heliconatica) margaritaeformis Dall, 1924 (type species of Heliconatica) is devoid of funicle (present in Sigatica) and appears to agree closely with the characters of Eunaticina Fischer, 1885. On this basis, we concur with Torigoe & Inaba (2011) in considering Heliconatica a synonym of Eunaticina. According to Kabat (1991), Sigaticus Aldrich, 1887 is an unjustiied emendation. Sigatica was originally proposed as a subgenus of Sigaretus Lamarck, 1799 (= Sinum Röding, 1798). Meyer & Aldrich (1886, p. 42) considered the globose shell, the wide umbilicus, the inner lip without callus, the spiral striation of basal and upper part of the whorls, and of the interior of the umbilicus as distinguishing characters of Sigatica. Examination of the type species and of those conidently assigned to Sigatica allowed us to deine the diagnostic characters of this genus as follows: 1) shell globose in most species; 2) umbilicus wide and spirally sculptured in most species; 3) funicle present, variously developed; 4) umbilical callus triangular, demarcated from the parietal callus by a groove, or prominent, rounded, separated from the parietal callus by a distinct sinus in some species (see also Kabat 2000, igs. 41, 42); 5) spiral sculpture restricted to subsutural and/or basal part of the whorls, extending across the entire last whorl in some species. The spiral sculpture within the umbilicus, present in most species of Sigatica, occurs also in species of Euspira Agassiz in J. Sowerby, 1837, e.g. Euspira magenesi Pedriali & Robba, 2001 and Euspira nysti (d’Orbigny, 1852). However, this character, combined with the spiral sculpture of the teleoconch, is another useful distinguishing element of Sigatica. Steven Tracey (NHM) collected from the London Clay (Div. C1, Kingsclere) a shell of Sigatica hantoniensis (Pilkington, 1804) with the operculum still illing the aperture. We obtained from Alan Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Fig. 18 - Outer surface of the operculum of fossil Sigatica species: a, Sigatica hantoniensis (Pilkington, 1804), Early Eocene, London Clay, Division C1, Kingsclere, United Kingdom; b, Sigatica clarkeana (Aldrich, 1887), Early Eocene, Hatchetigbee Formation (Upper member), Hatchetigbee Bluff, Tombigbee River, Washington County, Alabama, USA. Morton an excellent photograph of this operculum (Fig. 18a) and could note that it has the whole outer surface sculptured with coalescent beads roughly arranged into radial rows, and that it seems to be solid and calciied throughout. Other opercula of S. hantoniensis, with identical external sculpture, were igured by Deshayes (1864, pl. 68, igs. 29, 30), von Koenen (1891, pl. 40, igs. 4a, b) and Gürs (1983, pl. 4, igs. 10a, b). Also the operculum of the lower Eocene, US species Sigatica clarkeana (Aldrich, 1887) exhibits a closely similar outer sculpture and seems to be calciied throughout (Fig. 18b). The presence of a solid, calciied operculum would imply a subfamilial afiliation of Sigatica with the Naticinae. However, Marincovich (1977, p. 212) afirmed that Eunaticina insculpta (Carpenter, 1865), a poliniceine species (see remarks herein on the genus Eunaticina Fischer, 1885), has a partially calciied operculum, and wrote “whereas the inner surface of the operculum is entirely corneous, most of the outer surface has a thin calcareous coating formed of radial rows of minute, coalescent beads”. It appears that the external sculpture of the operculum of both S. hantoniensis and E. insculpta is remarkably similar. On this basis, Sigatica is herein provisionally included in the subfamily Poliniceinae. Only future recovering of in situ, fully calciied opercula of other Sigatica species (especially Recent species) will demonstrate that Sigatica is to be moved to the Naticinae. Eunaticina Fischer, 1885 is another spirally sculptured poliniceine genus, which differs from Sigatica in that its species have an oval, more elongate teleoconch with the open umbilicus devoid of funicle. Sigaretopsis Cossmann, 1888 (type species Natica 203 infundibulum Watelet, 1853 by original designation), originally introduced as a “section” of Natica Adanson, 1757, was later on regarded either as a synonym of Natica Scopoli, 1777 (Newton 1891), or as a synonym of Polinices Montfort, 1810 (Dall 1909), or as a subgenus of Gyrodes Conrad, 1860 (Cossmann 1925; Murphy & Rodda 1960), or as a subgenus of Sigatica (Forney & Nitecki 1976; Pacaud & Le Renard 1995). Actually, Sigaretopsis appears to be related to Sigatica because of its shell shape, its open umbilicus with inner spirals, and its subsutural spiral cords. We examined the specimen referred to as Natica (Sigaretopsis) infundibulum by Cossmann & Pissarro (1907) in MNHN (F-J02184) and can note that the absence of funicle stands as the main difference between Sigatica and Sigaretopsis. On the basis of the cited difference, we are inclined to consider Sigaretopsis as a poliniceinae genus distinct from Sigatica. Kabat (1991, p. 436) regarded Sigaretopsis as a synonym of Mammilla Schumacher, 1817. Kabat’s conclusion is untenable because of the many important differences between Natica infundibulum and Albula mammata Röding, 1798, type species of Mammilla. Sigatica was hitherto known from Eocene to Holocene deposits of United States, from the Eocene of Northern Europe, and from the Miocene and the Holocene of Japan; the Eocene Sigatica species described below stand for the irst sure record of the genus from Italy. Recent species of Sigatica occur in the tropical Atlantic and Indo-Paciic oceans. Sigatica claudiae sp. n. Pl. 14, igs. 1, 2 Derivation of name: The species is named for Claudia Savegnago, wife of the late collector Isidoro Rossi, who donated her husband’s collection to MGP-PD. Holotype: Cava Boschetto: MGP-PD 31525 (Pl. 14, ig. 1). Paratypes: Cava Albanello: 3 spms., MGP-PD 1231R; Cava Boschetto: 1 spm., MPUM 11362 (Pl. 14, ig. 2), 1 spm., NP 9815; Cava Rossi: 1 spm., MCZ 4335. Preservation: All specimens have the surface abraded at various extent. Type locality: Cava Boschetto (see appendix). Horizon: Greenish-gray tuff of early Lutetian age. Diagnosis: Globose, low-spired shell with deep, very large umbilicus, robust, moderately prominent funicle and semicircular umbilical callus, separated from parietal callus by slight sinus. Umbilical wall with two-four spiral cords. Surface of last whorl with traces of spiral cords over base. Description. Protoconch small, low-turbiniform, of two slightly convex whorls, tip medium- Robba E., Pedriali L. & Quaggiotto E. 204 sized. Teleoconch small, globose, rather thick. Spire low-conical, moderately depressed, whorls very slightly convex. Suture adpressed (almost lush). Last whorl inlated, slightly depressed; subsutural shelf indistinct; periphery at midline. Aperture D-shaped, prosocline, length about twice width. Parietal callus thick, rectangular, ending at level of umbilical border. Umbilicus deep, very large; umbilical border rounded, overhanging umbilical wall; umbilical wall slightly concave, bounded upward by groove and bearing two to four spiral cords crossed by growth lines. Funicle robust, moderately prom-inent, in midadapical part of umbilicus. Umbilical callus rather thick, semicircular, separated from parietal callus by slight sinus. Basal fasciole indistinct. Surface of last whorl with traces of spiral cords over basal part. Dimensions (mm) DHW PD H D SH 0.227* 0.773* 5.5977.473 4.8916.519 0.9432.063 * 1 protoconch measured 6.535 5.705 1.503 AH AW UW IS SA 4.2895.777 2.594-3.122 1.4423.274 17°-25° 103°-119° 5.033 2.858 2.358 21° 111° Remarks. The present form is herein assigned to the genus Sigatica Meyer & Aldrich, 1886 on account of its teleoconch shape, its widely open umbilicus with inner spiral cords and thick funicle, and its basal spiral cords. Sigatica claudiae sp. n. is closely similar to Sigatica eleonorae sp. n. (see below) in shell shape and adapical location of the funicle, but differs from it in that it has: 1) a protoconch with signiicantly smaller diameter and diameter of the irst half-whorl, 2) an umbilical wall bounded by a distinct groove and overhung by the umbilical border, and 3) a more prominent funicle; it is also worth noting that S. claudiae attains a smaller size. Stratigraphic occurrence. Sigatica claudiae sp. n. was recovered from upper Ypresian-lower Lutetian deposits of Veneto. Sigatica eleonorae sp. n. Pl. 14, igs. 3, 4 ? 1870a Natica hantoniensis - Bayan, p. 461. ? 1870 Natica hantoniensis - Fuchs, p. 195. ? 1894 Natica hantoniensis - Oppenheim, pp. 442, 445. 1985 Sigatica hantoniensis - Brigantini, p. 414, pl. 2, ig. 34 (not Pilkington, 1804). 2008 Sigatica hantoniensis - Quaggiotto & Mellini, p. 48, pl. 6, ig. 56 (not Pilkington, 1804). Derivation of name: The species is named for Eleonora, daughter of one of the present authors (L.P.). Holotype: Cava Grola: MCZ 4336 (Pl. 14, ig. 3). Paratypes: Cava Albanello: 1 spm., MGP-PD 31526, 1 spm., MCZ 4337; Cava Grola: 1 spm., MCZ 4338 (Pl. 14, ig. 4); Ciupio: 1 spm., MGP-PD 11624/c-11664/c; Monte Merlo: 1 spm., MGP-PD 31527. Other material examined: Cava Albanello: 3 spms., private collection, 1 spm., private collection; Cava Grola: 2 spms., private collection, 1 spm., private collection; Cava Rossi: 1 spm., private collection; Monte Merlo: 1 spm., NP 9814. Preservation: All specimens have the surface abraded at various extent. Type locality: Cava Grola (see appendix). Horizon: Greenish-gray volcanoclastic sandstone of middle Lutetian age. Diagnosis: Globose, low-spired shell with deep, large to very large umbilicus, robust, moderately depressed funicle and semicircular umbilical callus, separated from parietal callus by deep sinus. Umbilical wall with coarse spiral cords. Surface of last whorl with traces of uneven, low spiral cords throughout. Description. Protoconch large, low-turbiniform, of 2-2.10 slightly convex whorls, tip large. Teleoconch globose, rather thin. Spire low-conical, moderately depressed, whorls very slightly convex; suture adpressed (almost lush). Last whorl inlated, moderately depressed, somewhat expanded toward aperture in larger specimens; subsutural shelf in- PLATE 14 Fig. 1 - Sigatica claudiae sp. n. Cava Boschetto. Holotype, MGP-PD 31525; a, apertural side; b, abapertural side. Fig. 2 - Sigatica claudiae sp. n. Cava Boschetto. Paratype, MPUM 11362; protoconch. Fig. 3 - Sigatica eleonorae sp. n. Cava Grola. Holotype, MCZ 4336; a, apertural side (height of shell 23.67 mm); b, basal view; c, protoconch. Fig. 4 - Sigatica eleonorae sp. n. Cava Grola. Paratype, MCZ 4338; a, apertural side (height of shell 20.29 mm); b, apical view; c, basal view. Fig. 5 - Sigaretotrema checchii sp. n. Cava Grola. Holotype, MPUM 11363; a, apertural side (height of shell 18.13 mm); b, abapertural side. Fig. 6 - Sigaretotrema checchii sp. n. Cava Albanello. Paratype, MGP-PD 31528; protoconch. Fig. 7 - Sigaretotrema clathratum deshayesi (Michelotti, 1847). Colli Torinesi. MGPT BS.029.08.003; apertural side (scale bar 4 mm). Fig. 8 - Sigaretotrema clathratum deshayesi (Michelotti, 1847). Colli Torinesi. MGPT BS.029.08.003/01; a, apertural side (scale bar 4 mm); b, protoconch. Fig. 9 - Sigaretotrema michaudi (Michelotti, 1847). Colli Torinesi. Neotype (here designated) of Sigaretus michaudi Michelotti, 1847. MGPT BS.029.07.001; a, apertural side; b, basal view (scale bars 4 mm). Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 205 Robba E., Pedriali L. & Quaggiotto E. 206 distinct; periphery above midline. Aperture D-shaped in slightly prosocline plane, length about twice width. Parietal callus thick, rectangular, ending at level of umbilical border. Umbilicus deep, large to very large; umbilical border rounded; umbilical wall concave, bearing coarse, rough spiral cords crossed by growth lines. Funicle robust, moderately depressed, in mid-adapical part of umbilicus. Umbilical callus rather thick, semicircular, separated from parietal callus by deep sinus, shallower to obsolescent in larger specimens. Basal fasciole indistinct. Surface of last whorl with traces of uneven, low spiral cords throughout, better discernible over basal part. Dimensions (mm) DHW PD H D SH 0.440-0.552 1.5661.714 12.35126.771 10.46527.193 0.9734.969 0.496 1.640 19.561 18.829 2.971 AH AW UW IS SA 10.30222.878 6.62316.775 4.84513.673 16-24° 112°-140° 16.590 11.699 9.259 20° 126° Remarks. Four European species have been hitherto included in Sigatica, i.e. Natica abducta Deshayes, 1864, Nerita hantoniensis Pilkington, 1804, Natica obovata Sowerby, 1850 (Wrigley 1949; Pacaud & Le Renard 1995), and Natica repanda Deshayes, 1864 (Glibert 1963; Pacaud & Le Renard 1995). Natica abducta and Natica repanda were thoroughly described by Deshayes (1864), who unambiguously stated that both species have the umbilicus devoid of funicle and a smooth outer surface (except for growth markings). We examined the specimen of N. abducta igured by Cossmann & Pissarro (1907; MNHN-F-J02173) and noted that it lacks the funicle and has spiral sculpture neither on the outer surface nor within the umbilicus. On this basis, it is apparent that neither N. abducta nor N. repanda can be assigned to Sigatica and a comparison with Sigatica eleonorae sp. n. is therefore unnecessary. Nerita hantoniensis fully matches the characters of Sigatica. The original material of Pilkington’s species is not in NHM (Consuelo Sendino, personal communication 2012) and we were unable to locate it elsewhere. Consequently, for comparison with Sigatica eleonorae, we make reference to two relevant specimens, i.e. that illustrated by Wrigley (1949, ig. 38; NHM PI G 67417) and that illustrated by Cossmann & Pissarro (1907, pl. 9, ig. 61-21; MNHN-F-J02171). Examination of these specimens and of another from the Barton Beds shows that S. eleonorae is morphologically similar to N. hantoniensis, but differs from it in that it has: 1) a protoconch with one fewer whorl and with the diameter of the irst half-whorl over three times greater, 2) a parietal callus devoid of anterior lobe, 3) a better developed funicle, and 4) a semicircular umbilical callus demarcated from the parietal callus by a sinus, whereas it is subtriangular, separated from the parietal callus by a transverse groove in N. hantoniensis. It is dificult to compare the respective sculptural characters since the considered specimens have an abraded surface; however, there seems to be no signiicant difference, the spirals being manifest subsuturally and basally, less so in the middle of last whorl in both species. No reliable evaluation of the number of spirals can be done because of abrasion. Sigatica hantoniensis seems to be restricted to Paris and North Sea Basins. The records of Natica hantoniensis from Eocene units of Vicenza area made by Bayan (1870a), Fuchs (1870) and Oppenheim (1894) likely refer to Sigatica eleonorae, but without examination of the material dealt with by these authorities nothing can be stated deinitely in this respect. Natica obovata also belongs in Sigatica since it has the umbilicus with a low but distinct funicle and ine spiral grooves on the subsutural area and around the umbilicus. Its assignment to the genus Pliconacca Martin, 1914, assumed by Majima (1989, p. 67), is untenable because Natica (Pliconacca) trisulcata Martin, 1914, type species of Pliconacca, has an unsculptured shell and the inner lip callus with prominent transverse folds. N. obovata differs markedly from Sigatica eleonorae in that it has: 1) a oval shell with more elevated spire, 2) a narrower umbilicus devoid of inner spirals, and 3) a subtriangular umbilical callus separated from the parietal callus by a transverse groove. The specimen igured by Wrigley (1949; NHM PI G 67418) has ine spiral grooves observable only subsuturally and basally. Its protoconch is abraded, but the diameter of the irst half-whorl seems deinitely smaller than that of the protoconch of S. eleonorae. Stratigraphic occurrence. Sigatica eleonorae sp. n. was recovered from upper Ypresian, lower and middle Lutetian deposits of Veneto. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Subfamily Sininae Woodring, 1928 The subfamily Sininae includes genera characterized by spirally sculptured teleoconchs (mostly the last two whorls). The teleoconch is: 1) auriculate, variably lattened, with very depressed spire and with the umbilicus closed or reduced to a faint chink (Sinum Röding, 1798), or 2) globose, lowspired, with open umbilicus (Sigaretotrema Sacco, 1890). The operculum of the sinine species was said to be corneous, reduced to vestigial (Kilburn 1976; Kabat 1998). Huelsken et al. (2012, p. 373), on the basis of molecular data, stated that the Sininae should be “a true subfamilial taxon independent of Polinicinae”. Genus Sigaretotrema Sacco, 1890 Sigaretotrema Sacco, 1890b, p. 38 . Type species by monotypy: Sigaretus michaudi Michelotti, 1847, Oligocene to early Miocene, Piedmont. Propesinum Iredale, 1924, p. 256. Type species by original designation: Natica umbilicata Quoy & Gaimard, 1832. Sigaretotrema was originally proposed as a subgenus of Sigaretus Lamarck, 1799 (= Sinum Röding, 1798). Kabat (1991, p. 436) regarded Sigaretotrema as a junior subjective synonym of Eunaticina Fischer, 1885. Sigaretotrema is herein considered as distinct from Eunaticina since the species of Eunaticina have more elongate, oval shells with taller spire, more convex whorls separated by shallowly channeled sutures (those of Sigaretotrema species are adpressed), and the apertures that are more prosocline, more produced abapically, with length almost twice the width. According to Kabat (1991, p. 435), Propesinum is another junior subjective synonym of Eunaticina. However, the type species of Propesinum, is more similar to Sigaretus michaudi than to the type species of Eunaticina (Nerita papilla Gmelin, 1791). Consequently, we concur with Finlay & Marwick (1937) in considering Propesinum as a synonym of Sigaretotrema. We are uncertain about the suprageneric allocation of Sigaretotrema and provisionally follow Pacaud & Le Renard (1995), which included it in the subfamily Sininae. Sacco (1890b, p. 38) cited the conical-subglobose shell, the slightly convex whorls and the visible, variously deep umbilicus as the main distinguishing characters of Sigaretotrema. Examination of the type species (described herein) and of several European 207 species certainly belonging in Sigaretotrema enabled us to deine the diagnostic characters of the genus as follows: 1) protoconch depressed-turbiniform, smooth or spirally sculptured; 2) teleoconch globose in most species; 3) spire low, conical to domeshaped; 4) whorls very slightly convex, sutures adpressed; 5) umbilicus open, deep, narrow in most species; 6) aperture broad, nearly as wide as high in most species; 7) sculpture of spiral cords, with intervening cordlets and/or threads in most species. Sigaretotrema should be present in Eocene to Oligocene deposits of United States; the species Sigaretus bilix Conrad, 1833, Sigaretus declivis Conrad, 1833, Sigaretus arctatus Conrad, 1833 and Naticina obliqua Gabb, 1864 seem to belong in Sigaretotrema (see the igures published by Van Winkle Palmer 1937, pl. 15 and by Marincovich 1977, pl. 33). The genus occurs in the Eocene of France, Italy and New Zealand, as well as in the early and middle Miocene of some European localities and, possibly, of Australia (Natica subinfundibulum Tate, 1893). Modern occurrences are from South Australia and China (Sigaretus inlatus Tesch, 1920). Sigaretotrema checchii sp. n. Pl. 14, igs. 5, 6 Derivation of name: The species is named after Andrea Checchi, who provided lot of material relevant to the present study. Holotype: Cava Grola: MPUM 11363 (Pl. 14, ig. 5). Paratypes: Cava Albanello: 1 spm., MGP-PD 31528 (Pl. 14, ig. 6), 1 spm., MGP-PD 1232R, 1 spm., MGP-PD 1233R, 2 spms., MCZ 4339; Cava Grola: 1 spm., MGP-PD 31529, 1 spm., MCV 15/03; Cava Main: 1 spm., MCZ 4340. Other material examined: Cava Albanello: 1 spm., private collection; Cava Boschetto: 1 spm., NP 9816; Cava Grola: 1 spm., NP 9817, 4 spms., private collection, 1 spm., private collection, 2 spms., private collection. Preservation: Most specimens have the surface somewhat abraded. Type locality: Cava Grola (see appendix). Horizon: Greenish-gray volcanoclastic sandstone of middle Lutetian age. Diagnosis: Globose to oval, low-spired shell with deep, rather narrow umbilicus. Funicle and umbilical callus absent. Aperture oval, slightly prosocline. Sculpture starting with 16-17 rounded, closely set spiral cords becoming uneven and unevenly spaced, slightly undulating, iner over base. Description. Protoconch small, low-turbiniform, of two moderately convex, apparently smooth whorls, tip small. Teleoconch globose, oval in larger specimens, thin to moderately thick. Spire broadly conical, rather depressed, whorls Robba E., Pedriali L. & Quaggiotto E. 208 very slightly convex. Suture adpressed (almost flush). Last whorl globose-ovate, moderately depressed, slightly extended toward aperture, much so in fully grown specimens; subsutural shelf indistinct; periphery above midline. Aperture oval in slightly prosocline plane, length about 1.7 times width. Mid-abapical part of inner lip rather thick, gently arched. Parietal callus rather thick, with concave abapertural outline, ending with very slight lobe at level of umbilical border or slightly below it. Umbilicus deep, rather narrow; umbilical border broadly rounded; umbilical wall steep. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture of 16-17 rounded, closely set spiral cords on first quarter of first teleoconch whorl, then slightly undulating, nearly flat-topped, uneven and unevenly spaced, increasingly finer over base; spirals crossed by dense growth markings. Last whorl of one specimen retains traces of rectangular spots arranged into two spiral rows, respectively halfway between suture and periphery and peripheral. Dimensions (mm) DHW PD H D SH 0.161* 0,707* 10.04019.076 10.59418.474 1.1154.011 * 1 protoconch measurable 14.558 14.534 2.563 AH AW UW IS SA 8.37515.615 6.22413.552 1.6333.429 25°-53° 108°-128° 11.995 9.888 2.531 39° 118° Remarks. The present new species is characterized by its globose-ovate, moderately depressed last whorl noticeably extended toward the aperture. Sigaretotrema checchii somewhat resembles the Eocene species Sigaretotrema clathratum (Gmelin, 1791), but is readily distinguished from it because of its probably smooth protoconch of 0.7 fewer whorls and with signiicantly greater diameter of the irst half-whorl, its less globose teleoconch, its umbilical wall devoid of spirals, and its sculpture of spiral cords with intervening threads or cordlets. The lower Miocene species Sigaretotrema michaudi (Michelotti, 1847) is supericially similar, but differs in that it has: 1) a protoconch of 0.5 more whorls and with the diameter of the irst half-whorl nearly two thirds of the size, 2) a more globose teleoconch, 3) a wider aperture less produced abapically, 4) an umbilical wall with ine spi- ral threads, and 5) a sculpture of narrower, more distant spiral cords with one-two threads or one cordlet in their interspaces. Stratigraphic occurrence. Sigaretotrema checchii sp. n. was recovered from lower and middle Lutetian deposits of Vicenza area. Sigaretotrema clathratum deshayesi (Michelotti, 1847) comb. n. Pl. 14, igs. 7, 8 1847 Sigaretus deshayesi Michelotti, p. 158. 1891 Sigaretus (Sigaretus) aquensis var. deshayesi - Sacco, p. 99, pl. 1, ig. 61. ? 1952a Sigaretus (Sigaretus) aquensis f. deshayesi - Glibert, p. 78, pl. 6, ig. 7. 1971 Sigaretus (Sigaretus) aquensis subsp. deshayesi - Accorsi Benini, p. 245, pl. 4, igs. 4, 4a. 1984 Sigaretus aquensis var. deshayesi - Ferrero Mortara et al., p. 38. Type material: Sigaretus deshayesi Michelotti, type material not seen. Michelotti’s collection was not located and is possibly lost. Material examined: Colli Torinesi: 1 spm., MGPT BS.029.08.003 (Pl. 14, ig. 7) [other 3 spms. are Sigaretus cryptostomoides Sacco, 1890 and 5 spms. are Sigaretus haliotideus var. patula Grateloup, 1847, all also numbered BS.029.08.003], 1 spm., MGPT BS.029.08.003/01 (Pl. 14, ig. 8); Case Soghe: 2 spms., MPPLF 5110 pr. (Accorsi Benini collection). Description. Protoconch medium-sized, lowturbiniform, of 2.75 convex whorls, last 1.3 whorls with even, latly rounded spiral cordlets, tip small. Teleoconch globose, rather thin. Spire broadly conical, depressed, whorls slightly convex. Suture adpressed. Last whorl globose, slightly depressed, somewhat expanded toward aperture; subsutural shelf indistinct; periphery above midline. Aperture oval in moderately prosocline plane, length about 1.3 times width. Abapical part of inner lip gently arched, thickening upward, slightly bent to merge into parietal callus. Parietal callus rather thin, with concave abapertural outline, ending with very slight lobe below level of umbilical border. Umbilicus deep, narrow; umbilical border broadly rounded; umbilical wall rather steep, with very ine spiral threads. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture of 16-18 rounded, closely set spiral cords on irst quarter of irst teleoconch whorl, then slightly undulating, lat-topped, wider or as wide as interspaces, without threads in interspaces or with one ine thread in some, iner over base; spirals crossed and made somewhat rough by dense growth markings. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy Dimensions (mm) DHW PD H D SH 0.0970.121 1.2541.618 8.14012.440 7.79512.915 0.6843.456 0.109 1.436 10.290 10.355 2.070 AH AW UW IS SA 7.4768.984 4.8678.743 1.1521.888 45°-61° 108°-128° 8.220 6.805 1.520 53° 118° Remarks. Michelotti (1847), introducing Sigaretus deshayesi, published a quite concise diagnosis that reads “S. testa ovato-depressa, transversim undulato-striata; aperture valde concava; umbilico subtecto”. Michelotti also afirmed that his material was from “la colline de Turin”. From this diagnosis, it appears that the umbilicus of Sigaretus deshayesi is only partially covered, thence narrow but visible. Since Michelotti’s collection is lost, we are forced to make reference to the material in MGPT studied by Sacco (1891) and collected from the same area (Colli Torinesi = la colline de Turin). We have selected from Sacco’s specimens labeled Sigaretus (Sigaretus) aquensis var. deshayesi (Michelotti) those having a narrow, open umbilicus and consider them to be representative of Michelotti’s taxon. It is to be noted that Sigaretus deshayesi agrees with the characters of Sigaretotrema Sacco, 1890 and is unrelated to Sigaretus aquensis Récluz, 1851 (= Sigaretus patulus Grateloup, 1847, see below), which has a more depressed, imperforated teleoconch and belongs in Sinum Röding, 1798. Sigaretus deshayesi is strikingly similar to Nerita clathrata Gmelin, 1791 (also belonging in Sigaretotrema). In order to clarify the relationships between these two taxa, the shells of Gmelin’s species igured by Cossmann & Pissarro (1907, pl. 10, ig. 62-1; MNHN-F-J02185), by Wrigley (1949, ig. 45; NHM PI G 67423), and others from Middle Barton Beds (Great Britain) and from Oise (France) in our collection were examined. We note that the characteristic elements of the protoconch of both taxa are identical, as are the shell shape, the aperture and the bulk of sculptural characters. However, S. deshayesi has the protoconch with nearly lat, wider and less numerous spirals, attains a smaller size and has few or no threads at all in between the spiral cords. These differences are not considered to be suficient for consistent separation at the species level. Consequently, we regard the Miocene S. deshayesi as an allochronous subspecies of 209 the Eocene Sigaretotrema clathratum. Sigaretotrema clathratum deshayesi is closely similar to Sigaretotrema michaudi (Michelotti, 1847), but differs from it in having the sculptured protoconch with signiicantly greater diameter and diameter of the irst half-whorl, and the teleoconch with iner spiral cords with scarce or no threads in the interspaces. Glibert (1952a) igured a shell referred to as Sigaretus (Sigaretus) aquensis f. deshayesi Michelotti, 1847. From Glibert’s igure, that shell seems to have a narrow umbilicus and could be Michelotti’s taxon. However, having not seen the specimen, we include it in the synonymy of Sigaretus deshayesi with reservation. Stratigraphic occurrence. Sigaretotrema clathratum deshayesi (Michelotti, 1847) is surely known from the early Oligocene of Veneto and from the early Miocene (Burdigalian) of Piedmont. Sigaretotrema michaudi (Michelotti, 1847) Pl. 14, ig. 9; Pl. 15, ig. 1 1847 Sigaretus michaudi Michelotti, p. 158, pl. 6, igs. 16, 18. ? 1862 Sigaretus michaudi - Doderlein, p. 18. 1890b Sigaretus (Sigaretotrema) michaudi var. clausula Sacco, p. 38. 1890b Sigaretus (Sigaretotrema) michaudi var. eunaticinoides Sacco, p. 38. 1890b Sigaretus (Sigaretotrema) michaudi var. pseudoaquensis Sacco, p. 38. 1891 Sigaretus (Sigaretotrema) michaudi - Sacco, p. 97, pl. 1, ig. 55. 1891 Sigaretus (Sigaretotrema) michaudi var. clausula - Sacco, p. 97, pl. 1, ig. 56. 1891 Sigaretus (Sigaretotrema) michaudi var. eunaticinoides Sacco, p. 98, pl. 1, ig. 57. 1891 Sigaretus (Sigaretotrema) michaudi var. pseudoaquensis Sacco, p. 98, pl. 1, ig. 58. not 1960 Sinum (Sigaretotrema) michaudi - Kojumdgieva & Strachimirov, p. 122, pl. 34, ig. 1 (not Michelotti, 1847). 1984 Sigaretus michaudi - Ferrero Mortara et al., p. 37. 1984 Sigaretus michaudi var. clausula - Ferrero Mortara et al., p. 37. 1984 Sigaretus michaudi var. eunaticinoides - Ferrero Mortara et al., p. 38. 1984 Sigaretus michaudi var. pseudoaquensis - Ferrero Mortara et al., p. 38. 2007 Eunaticina michaudi - Zunino, p.126, pl. 1, ig. 13. Type material: Sigaretus michaudi Michelotti, neotype (here designated): the specimn igured by Sacco (1891, pl. 1, ig. 55) and reigured herein (Pl. 14, ig. 9), MGPT BS.029.07.001, Colli Torinesi (see remarks below). Sigaretus (Sigaretotrema) michaudi var. clausula Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 1, ig. 56), MGPT BS.029.07.002, Colli Torinesi. Sigaretus (Sigaretotrema) michaudi var. eunaticinoides Sacco, holotype (by monotypy): the shell Robba E., Pedriali L. & Quaggiotto E. 210 igured by Sacco (1891, pl. 1, ig. 57), MGPT BS.029.07.003, Colli Torinesi. Sigaretus (Sigaretotrema) michaudi var. pseudoaquensis Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 1, ig. 58), MGPT BS.029.07.004, Colli Torinesi. Material erroneously referred to as Sigaretus (Sigaretus) aquensis var. praecedens Sacco, 1890 in MGPT: Dego: 1 syntype, MGPT BS.029.08.001/01. Other material examined: Colli Torinesi: 7 spms., MGPT BS.029.07.001/01; Termofourà: 1 spm., MGPT-PU 23519 (Pl. 15, ig. 1); Valle Ceppi: 1 spm., MGPT-PU 25268 (specimen of Zunino 2007), 2 spms., PG 32. Description. Protoconch small, low-turbiniform, of 2.55 slightly convex whorls, tip very small. Teleoconch globose, rather thick. Spire low-conical, moderately depressed, whorls slightly convex. Suture adpressed. Last whorl globose, moderately depressed, slightly expanded toward aperture; subsutural shelf indistinct; periphery about at midline. Aperture ovate- to roundly-quadrangular in slightly prosocline plane, length about 1.5 times width. Abapical part of inner lip thick, almost straight. Parietal callus rather thin, with concave abapertural outline, ending with very slight lobe at level of umbilical border. Umbilicus deep, narrow; umbilical border broadly rounded; umbilical wall steep, with ine spiral threads. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture of rather coarse spiral cords, nine-ten on irst quarter of irst teleoconch whorl, soon increasing in number and with one-two threads or one cordlet in interspaces; spirals crossed and made rough by dense growth markings. Dimensions (mm) DHW PD H D SH 0.063* 0.597* 14.53219.420 14.65221.848 1.9944.850 * 1 protoconch measurable 16.976 18.250 3.422 AH AW UW IS SA 12.01815.090 8.65214.032 2.5045.020 38°-58° 112°128° 13.554 11.342 3.762 48° 120° Remarks. Repeated inquiries about the location of Michelotti collection had negative results and that collection is to be considered lost (Giulio Pavia, personal communication 2013). Since Sacco (1891, p. 97), in the remarks to Sigaretus (Sigaretotrema) michaudi, explicitly afirmed to have examined Michelotti’s type specimens, his specimens in MGPT, collected from the same area (colline de Turin = Colli Torinesi), stand as the most relevant reference material for Sigaretotrema michaudi. Thus, it is advisable to designate the shell igured by Sacco (1891, pl. 1, ig. 55) as the neotype of S. michaudi, in order to clarify and ix the characters of Michelotti’s species (ICZN 1999, Article 75 of the Code). The Eocene species Nerita clathrata Gmelin, 1791 (for the examined material, see the remarks on Sigaretotrema clathratum deshayesi) is the closest relative of Sigaretotrema michaudi. Both species have similarly shaped teleoconchs and identical umbilical characters. However, S. michaudi is distinguished from Sigaretotrema clathratum in that it has: 1) a protoconch with signiicantly smaller diameter and diameter of the initial half-whorl, 2) a more inlated last whorl, 3) a more distinctly angular inner lip, 4) the teleoconch sculpture starting with nine-ten spiral cords (with 16-18 in S. clathratum), and 5) more numerous threads/cordlets in between the spiral cords. Gmelin’s species has a spirally sculptured larval shell; nothing can be said as regards S. michaudi, since all the examined specimens have the protoconch decorticated to a variable extent. Sigaretotrema levesquei (Deshayes, 1864) is another closely related species. We have examined the specimen igured by Cossmann & Pissarro (1907, pl. 10, ig. 62-2; MNHNF-J02186) and can note that S. levesquei is distinguished from S. michaudi in having the protoconch with signiicantly greater diameter and smaller diameter of the irst half-whorl, and the sculpture of rough, uneven and unevenly spaced spiral cordlets without threads in the interspaces. In the course of consultation of the syntypes of the Burdigalian Sigaretus subcanaliculatus d’Orbigny, 1852 in MNHN (5 specimens collectively numbered F-B27769), we noticed that one syntype (the smallest) belongs in Sigaretotrema because of its more globose teleoconch with well distinct umbilicus (the other four syntypes belong in Sinum Röding, 1798). This specimen likely represents an unnamed species of Sigaretorema that resembles Sigaretotrema michaudi, but differs from it at least by its sculptured protoconch with signiicantly greater diameter and diameter of the irst half-whorl. S. subcanaliculatus is currently regarded as a synonym of Sinum patulum (Grateloup, 1847). Sacco (1891, p. 97) proposed the variety exclathrata of Sigaretus (Sigaretotrema) michaudi on the basis of the shell igured by Hoernes (1856, pl. 46, ig. 28) and referred to as Sigaretus clathratus Recluz, 1843. We obtained outstanding photographs Eocene, Oligocene and Miocene naticid gastropods of Northern Italy of the specimen of Sigaretus clathratus (NHMW 2013/0053/0001) illustrated by Schaffer (1912), supposedly identical to that of Hoernes (Mathias Harzhauser, personal communication 2013). From the photographs, it appears that the Austrian shells differ from Sigaretotrema michaudi in having a twowhorled protoconch (that of Michelotti’s species exceeds three whorls), a less globose teleoconch with the last whorl more extended toward the aperture, and a narrower umbilicus. Accordingly, Hoernes’ and Schaffer’s specimens represent a separate species from S. michaudi and cannot be regarded as a variety of it. The Austrian shells also seem not to fully agree with the characters of Sigaretotrema clathratum. Should they prove to represent an unnamed Sigaretotrema species (a decision is beyond the purpose of this study), the name exclathrata remains available for it. Concerning the other Sacco’s varieties of Sigaretus (Sigaretotrema) michaudi (see the above synonymy), we have examined the respective original material in MGPT and can state that they cannot be distinguished from S. michaudi consistently. We assign to S. michaudi also one Oligocene shell in MGPT erroneously referred to as Sigaretus (Sigaretus) aquensis var. praecedens Sacco, 1890 by Sacco (1891). The Tortonian Bulgarian shell igured by Kojumdgieva & Strachimirov (1960) and referred to as Sinum (Sigaretotrema) michaudi (Michelotti, 1847) appears to differ markedly from Michelotti’s species in that it has an oval, more elongated teleoconch with taller spire, wider umbilical opening and nearly D-shaped aperture with length twice the width. According to the igure published by Kojumdgieva & Strachimirov, that shell likely represents an unnamed species of the genus Eunaticina Fischer, 1885. Stratigraphic occurrence. Sigaretotrema michaudi (Michelotti, 1847) was hitherto known from lower Oligocene and lower Miocene (Burdigalian) units of Piedmont. Records from the Tortonian of Montegibbio (Doderlein 1864) and those from Germany are doubtful and need to be conirmed. Sigaretotrema sp. Pl. 15, ig. 2 Material examined: Cava Rossi: 1 spm., MPUM 11364 (Pl. 15, ig. 2), 1 spm., private collection. Description. Protoconch small, low-turbiniform, of 2.5 moderately convex, apparently smooth 211 whorls, tip small. Teleoconch globose, nearly as wide as high, rather thick. Spire broadly conical, depressed, whorls almost lat-sided. Suture adpressed (almost lush). Last whorl globular, slightly expanded toward aperture; subsutural shelf indistinct, only scarcely deined behind outer lip; periphery above midline. Aperture teardrop-shaped in slightly prosocline plane, length 1.5 times width. Outer lip subangular adapically; middle part of inner lip nearly straight, increasingly thick upward. Parietal callus rather thin, with concave abapertural outline, ending with faint lobe slightly below umbilical border. Umbilicus deep, moderately wide; umbilical border rounded; umbilical wall steep, bounded by groove. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture of seven-eight rounded, closely set spiral cords on irst quarter of irst teleoconch whorl, then slightly undulating, lat-topped, uneven, wider than interspaces that bear one-two threads, iner over base; spirals crossed by dense growth markings. Dimensions (mm) DHW PD H D SH AH AW 0.125 0.707 18.470 18.611 2.370 16.101 11.900 IS SA 39° 126° Remarks. The globular teleoconch nearly as wide as high is the primary character that differentiates the present specimens from the other European species of Sigaretotrema Sacco, 1890. They are distinguished from Sigaretotrema checchii sp. n. (see above) also in having the protoconch of 0.5 more whorls, with signiicantly smaller diameter of the irst half-whorl. Compared to Sigaretotrema michaudi (Michelotti, 1847), besides the different shell shape, they have a protoconch with signiicantly greater diameter of the irst half-whorl. The two examined specimens certainly represent a previously undescribed species, but more, better preserved material is required in order to name it. Stratigraphic occurrence. Sigaretotrema sp. was recovered from upper Ypresian-lowermost Lutetian deposits of Cava Rossi (Veneto). Genus Sinum Röding, 1798 Sinum Röding, 1798, p. 14. Type species by subsequent designation (Dall 1915, p. 109): Helix haliotoidea Linnaeus, 1758, Recent, Indo-Paciic. Robba E., Pedriali L. & Quaggiotto E. 212 Sigaretus Lamarck, 1799, p. 77. Type species by monotypy: Helix haliotoidea Linnaeus, 1758. Sigaretarius Duméril, 1806, p. 164. Type species by subsequent designation (Kabat 1991, p. 436): Helix haliotoidea Linnaeus, 1758. Cryptostomus Blainville, 1818a, p. 120. The genus was based on two species: Cryptostomus leachii Blainville, 1818 and Cryptostomus breviculus Blainville, 1818. The type species was designated neither originally, nor subsequently. Cryptostomus is currently regarded as a synonym of Sinum (cf. Kabat 1991 and Torigoe & Inaba 2011). Cryptostoma Blainville, 1818b, p. 126. Incorrect subsequent spelling of Cryptostomus. Catinus Blainville, 1827 (reference to Lamarck, Klein and Martini), p. 105. Nomen nudum. Catinus H. & A. Adams, 1853, p. 212. Type species not designated. All the species listed (25) belong in Sinum (see also Kabat 1991 and Torigoe & Inaba 2011). Ectosinum Iredale, 1931, p. 216-217, 232. Type species by original designation: Ectosinum pauloconvexum Iredale, 1931. Pedriali & Robba (2009) listed the distinctive characters of Sinum that are slightly modiied here as follows: 1) depressed-turbiniform to planorbid protoconch of from less than 2 up to 3 whorls, with faint spiral striation, 2) variably lattened, auriculate shell, oval in some species, 3) moderately high to very depressed spire, 4) strongly prosocline aperture, broad in most species, with markedly arched inner lip, 5) umbilicus closed or a small chink, 6) outer surface spirally sculptured, or smooth in some species. missing from last quarter of whorl. Teleoconch auriculate, depressed, wider than high, rather thin. Spire broadly conical, very low (scarcely protruding from outline of last whorl), whorls faintly convex. Suture adpressed. Last whorl lenticular, depressed, quickly expanding toward aperture; subsutural shelf indistinct; narrowly rounded periphery nearly at midline. Aperture large, oval, oblique, in markedly prosocline plane, distinctly wider than high. Outer lip arched; basal lip semicircular; inner lip with thin, narrow callus, bent adapically, relected toward umbilical area, then merging into thin parietal callus well below border of umbilical area. Umbilicus a very narrow chink. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture starting with 10-11 even spiral cords becoming uneven and lat-topped by second whorl; interspaces narrower than cords, a few in adapical part with one thread; moderately wide subsutural band with ive-six narrower cords; base with increasingly iner cordlets; spirals crossed by dense growth markings. PLATE 15 Sinum borellense sp. n. Pl. 15, igs. 3-5 Derivation of name: The name refers to the hamlet of Borelli, which is the type locality. Holotype: Borelli: MGPT-PU 135289 (Pl. 15, ig. 3). Paratypes: Borelli: 1 spm., MGPT-PU 135290 (Pl. 15, ig. 4), 1 spm., MGPT-PU 135291 (Pl. 15, ig. 5), 1 spm., MGPT-PU 135292, 20 spms., MGPT-PU 135293, 1 spm., MPUM 11365. Other material examined: Borelli: 7 spms., PG 31, 1 spm., NP 9819. Preservation: Most specimens are fairly well preserved, a few are variously damaged. Type locality: Borelli (see appendix). Horizon: Gray, medium sand of Tortonian age. Diagnosis: Protoconch I apparently smooth; protoconch II with spiral cordlets. Shell auriculate, depressed, markedly low-spired, with very narrow umbilical chink. Funicle and umbilical callus absent. Aperture oval, markedly prosocline. Sculpture starting with 10-11 even, rather widely spaced spiral cords becoming uneven and lat-topped by second whorl; interspaces narrower than cords; lowermost base with increasingly iner cords. Description. Protoconch medium-sized, depressed-turbiniform, of 2.75-2.8 rather convex whorls, tip very small; protoconch I apparently smooth, protoconch II with even spiral cordlets Fig. 1 - Sigaretotrema michaudi (Michelotti, 1847). Termofourà. MGPTPU 23519; protoconch. Fig. 2 - Sigaretotrema sp. Cava Rossi. MPUM 11364; a, apertural side (height of shell 18.47 mm); b, abapertural side. Fig. 3 - Sinum borellense sp. n. Borelli. Holotype, MGPT-PU 135289; a, apertural side (height of shell 6.02 mm); b, abapertural side. Fig. 4 - Sinum borellense sp. n. Borelli. Paratype, MGPT-PU 135290; a, protoconch; b, detail of protoconch. Fig. 5 - Sinum borellense sp. n. Borelli. Paratype, MGPT-PU 135291; a, apertural side (height of shell 5.89 mm); b, abapertural side. Fig. 6 - Sinum cryptostomoides (Sacco, 1890). Colli Torinesi. Holotype of Sigaretus (Sigaretus) cryptostomoides var. colligens Sacco, 1890. MGPT BS.029.08.007; a, apertural side (scale bar 4 mm); b, protoconch. Fig. 7 - Sinum oligopolitum (Sacco, 1890). Sassello. Holotype of Sigaretus (Sigaretus) oligopolitus Sacco, 1890. MGPT BS.029.08.006; apertural side, × 2. From Ferrero Mortara et al. (1984). Fig. 8 - Sinum oligopolitum (Sacco, 1890). Mioglia. MZB 60000; protoconch. Fig. 9 - Sinum patulum (Grateloup, 1847). Colli Torinesi. Lectotype (here designated) of Sigaretus (Sigaretus) aquensis var. longotriangula Sacco, 1890. MGPT BS.029.08.004; apertural side (scale bar 4 mm). Fig. 10 - Sinum patulum (Grateloup, 1847). Colli Torinesi. Holotype of Sigaretus (Cryptostoma) sigaretoides Sacco, 1890. MGPT BS.029.09.001; apertural side (scale bar 4 mm). Fig. 11 - Sinum patulum (Grateloup, 1847). Léognan, Le Coquillat (France). MPUM 11367; a, apertural side (height of shell 13.58 mm); b, abapertural side. Fig. 12 - Sinum patulum (Grateloup, 1847). Saucats (France). MPUM 11370; protoconch. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 213 Robba E., Pedriali L. & Quaggiotto E. 214 Dimensions (mm) Sinum cryptostomoides (Sacco, 1890) DHW PD H D SH 0.0760.092 0.9451.237 2.9097.637 6.43712.857 0.6751.139 0.084 1.091 5.273 9.647 0.907 AH AW IS SA 2.1736.561 5.0109.030 13°-41° 130°-142° 4.367 7.020 27° 136° Remarks. An unidentiied Sinum species (one specimen, MZB 60017) collected from middle Miocene deposits near Orthez (France) appears to be the most closely similar to Sinum borellense sp. n. in shell shape, umbilical and apertural characters. It differs from S. borellense in having the protoconch (also spirally sculptured) with slightly, but signiicantly greater diameter of the irst half-whorl, the teleoconch somewhat more depressed with the spiral cords that are more even and more widely spaced. The cited differences could be not enough for a separation at species level, but additional material of the French taxon is required to settle a decision in this respect. Sinum paviai sp. n. (see below), co-occurring with Sinum borellense in the type locality of Borelli, displays similar values of the characteristic elements of the protoconch, but its protoconch I has irregular spiral rows of minute granular microprotuberances, whereas that of S. borellense has not. S. paviai differs also in having a rather globose teleoconch with more inlated last whorl and the spaces in between the spiral cords with one cordlet or one-three threads; moreover, it lacks the subsutural band with iner cords present in S. borellense. The Pliocene species Sinum perregulare (Sacco, 1891) is another similar species in terms of shell shape. It differs from S. borellense in that it has the differently sculptured protoconch with a signiicantly greater diameter of the irst half-whorl and the teleoconch with alternating spiral ribbons and threads. The Pliocene species Sinum subhaliotideum (d’Orbigny, 1852) is also similar in shell shape and sculpture. However, the two syn-types in MNHN (FB35752 and F-B36276) have a signiicantly smaller, smooth protoconch. D’Orbigny’s species was regarded as a synonym of Sinum striatum (de Serres, 1829) by some workers (Sacco 1891; Cossmann & Peyrot 1919). However, Sinum subhaliotideum differs from Sinum striatum at least by its smooth protoconch with signiicantly greater diameter. Stratigraphic occurrence. Sinum borellense sp. n. was recovered only from the type locality. Pl. 15, ig. 6 1890b Sigaretus (Sigaretus) cryptostomoides Sacco, p. 39. 1890b Sigaretus (Sigaretus) cryptostomoides var. colligens Sacco, p. 39. 1891 Sigaretus (Sigaretus) cryptostomoides - Sacco, p. 101, pl. 1, ig. 68. 1891 Sigaretus (Sigaretus) cryptostomoides var. colligens - Sacco, p. 101, pl. 1, ig. 69. 1984 Sigaretus criptostomoides (sic) - Ferrero Mortara et al., p. 38. 1984 Sigaretus criptostomoides var. colligens - Ferrero Mortara et al., p. 38, pl. 4, ig. 5. Type Material: Sigaretus (Sigaretus) cryptostomoides Sacco, original material not found in MGPT and likely lost. Sigaretus (Sigaretus) cryptostomoides var. colligens Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 1, ig. 69) and reigured herein (Pl. 15, ig. 6), MGPT BS.029.08.007, Colli Torinesi. Material erroneously referred to as Sigaretus (Sigaretus) aquensis var. deshayesi Michelotti, 1847 in MGPT: Colli Torinesi: 3 spms., MGPT BS.029.08.003. Description. Protoconch large, lowturbiniform, of three convex, apparently smooth whorls, tip very small. Teleoconch auriculate, moderately depressed, wider than high, rather thin. Spire low-conical, pointed, whorls faintly convex. Suture adpressed. Last whorl dome-shaped in abapertural (dorsal) view, depressed, remarkably extended toward aperture; subsutural shelf indistinct; narrowly rounded periphery above midline. Aperture elliptical, oblique, in markedly prosocline plane, width nearly two times length. Outer lip gently arched in midadapical part, more so abapically; inner lip relected adapically toward umbilical area, then merging into thin parietal callus well below border of umbilical area. Umbilicus a very narrow chink. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture of 19 even, closely set spiral ribbons on irst quarter of irst whorl; subsequently, ribbons slowly increase in number and may be slightly undulating behind outer lip; interspaces narrower than ribbons, a few in adapical part with one thread; base with spiral cords increasingly iner downward; spirals crossed by dense growth markings. Dimensions (mm) DHW PD H D SH AH AW 0.081 1.548 13.87 17.11 1.39 12.48 12.09 IS SA 52° 125° Remarks. Sigaretus (Sigaretus) cryptostomoides can be easily recognized on the basis of the descrip- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy tion and the igure published by Sacco (1891). The auriculate teleoconch, the dome-shaped last whorl remarkably extended toward the aperture, and the sculpture of broad spiral ribbons with scarce intervening threads are the most relevant distinguishing elements. The co-occurring variety colligens of S. (Sigaretus) cryptostomoides was proposed by Sacco (1890b) and said to differ from the nominal species in having “testa minus depressa; spira subelato-conica; supericies undulato-striolata”. These differences do not warrant separation even at the subspecies level. Consequently, we include the variety colligens in the synonymy of the present species. Sinum cryptostomoides is similar to Sinum patulum (Grateloup, 1847), but differs from it in that it has: 1) a protoconch with signiicantly greater diameter and smaller diameter of the irst half-whorl, 2) a more distinctly auriculate teleoconch, 3) a sculpture starting with 19 spiral ribbons (seven-ten cords in S. patulum), and 4) an occasional occurrence of one thread in the interspaces, whereas S. patulum has one-three threads or one cordlet throughout. Stratigraphic occurrence. Considering the matrix inilling, Sinum cryptostomoides (Sacco, 1890) appears to have been collected from the Termofourà Formation of Burdigalian age (cf. Zunino & Pavia 2009). Sinum oligopolitum (Sacco, 1890) Pl. 15, igs. 7, 8 1890b Sigaretus (Sigaretus) oligopolitus Sacco, p. 38. 1891 Sigaretus (Sigaretus) oligopolitus - Sacco, p. 100, pl. 1, ig. 66. ig. 3. 1937 Sigaretus oligopolitus - Venzo, p. 48, pl. 2, igs. 40-40b. 1984 Sigaretus oligopolitus - Ferrero Mortara et al., p. 38, pl. 4, ? 1990 Sinum oligopolitum - Baglioni Mavros, p. 241, pl. 1, igs. 10, 11. Type material: Sigaretus (Sigaretus) oligopolitus Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 1, ig. 66) and reigured herein (Pl. 15, ig. 7; reproduced from Ferrero Mortara et al. 1984), MGPT BS.029.08.006, Sassello. Other material examined: Mioglia: 1 spm., MPUM 11366, 1 spm., MZB 60000 (Pl. 15, ig. 8), 1 spm., PG 33, 1 spm., PG 34, 1 spm., PG 35. Description. Protoconch medium-sized, lowturbiniform, of two convex whorls, tip apparently small. Teleoconch conical-ovate, slightly higher than wide, thick. Spire conical, pointed, rather elevated, whorls gently convex. Suture adpressed. Last whorl 215 broadly oval, slightly expanded toward aperture; subsutural shelf indistinct; periphery shortly above midline. Aperture ovate-quadrangular in markedly prosocline plane, length 1.3 times width. Outer lip slightly arched in mid-adapical part, more so abapically; inner lip thick, subangular medially, merging into also thick parietal callus well below border of umbilical area. Umbilicus absent, or a chink largely covered by everted inner lip. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture of uneven, rather widely spaced spiral cords, slightly iner over base; interspaces with two-three threads; spirals crossed by dense growth markings. Dimensions (mm) DHW PD H D SH - 1.486* 8.28125.305 8.83122.855 1.8072.967 2.387 * 1 protoconch measurable 16.793 15.843 AH AW IS SA 8.45918.591 4.651-18.255 30°-42° 106°118° 13.525 11.453 36° 112° Remarks. Sacco (1890b) named the species on the basis of the smooth surface of the single shell he examined. This specimen (the holotype) is poorly preserved, with the surface badly abraded. Our material demonstrates that Sacco’s species is spirally sculptured as are most species of the genus. Sinum oligopolitum is characterized by its two-whorled protoconch (nothing can be said about eventual sculpture and diameter of the irst half-whorl because of abrasion), its conical-ovate teleoconch with pointed, rather elevated spire (unusually elevated for the genus), and its spiral cords with twothree threads in the interspaces. Sigaretus philippii, introduced by Speyer (1870, p. 81) on the basis of upper Oligocene material from Cassel area, appears to be the most closely related species. It has a two-whorled protoconch, a similarly shaped teleoconch with rather elevated, pointed spire, and an identical apertural outline. The unique difference seems to be the sculpture (detail in Speyer’s pl. 12, ig. 12 d), consisting in even spiral cords without threads in the interspaces. This difference is deemed not suficient for consistent separation at species level. A decision in this respect would require the direct examination of the original material of S. philippii in BGR. However, that material was not found there and is possibly lost (Angela Ehling, 216 Robba E., Pedriali L. & Quaggiotto E. personal communication 2013) and the problem remains pending. Should the two taxa prove to be synonyms, S. philippii has priority by seniority. The Recent East Paciic species Sinum cymba (Menke, 1828) is supericially similar in that has a somewhat elevated spire. However, it is readily distinguished from Sinum oligopolitum by its protoconch of 0.5 more whorls, its markedly cyrtoconoid teleoconch with nearly lat base, and its sculpture of wide spiral ribbons with no threads in the narrow interspaces. Baglioni Mavros (1990) listed Sinum oligopolitum from upper Oligocene deposits of Belluno area. The author did not provide any description and the poor illustrations published rise some doubt about the assignment of that material. Consequently, we consider this citation as an uncertain reference and include it in the synonymy of the present species with much reservation. Stratigraphic occurrence. Sinum oligopolitum (Sacco, 1890) is known deinitely from the early Oligocene of Liguria, and from the late Oligocene of Veneto. Sinum patulum (Grateloup, 1847) Pl. 15, igs. 9-12; Pl. 16, igs. 1, 2 1847 Sigaretus haliotideus - Grateloup, pl. 48, ig. 19. 1847 Sigaretus haliotideus var. B patula Grateloup, pl. 48, ig. 20. 1852 Sigaretus subcanaliculatus d’Orbigny, p. 39, n° 589. 1890b Sigaretus (Sigaretus) aquensis var. praecedens Sacco, p. 38. 1890b Sigaretus (Sigaretus) aquensis var. longotriangula Sacco, p. 38. 1890b Sigaretus (Cryptostoma) sigaretoides Sacco, p. 39 (new synonym). 1891 Sigaretus (Sigaretus) aquensis var. praecedens - Sacco, p. 98, pl. 1, ig. 59. 1891 Sigaretus aquensis var. longotriangula - Sacco, p. 99, pl. 1, ig. 63. 1891 Sigaretus aquensis var. taurinensis Sacco, p. 100, pl. 1, ig. 62. 1891 Sigaretus (Cryptostoma) sigaretoides - Sacco, p. 102, pl. 1, ig. 70. 1919 Sigaretus aquensis - Cossmann & Peyrot, p. 234, pl. 12, igs. 47, 48. 1925 Sigaretus (Sigaretus) aquensis - Cossmann, p. 143, pl. 4, igs. 23, 24. 1952b Sigaretus aquensis - Glibert, p. 262, pl. 3, ig. 2. 1984 Sigaretus aquensis var. praecedens - Ferrero Mortara et al., p. 38. 1984 Sigaretus aquensis var. longotriangula - Ferrero Mortara et al., p. 38. 1984 Sigaretus aquensis var. taurinensis - Ferrero Mortara et al., p. 38. 1984 Cryptostoma sigaretoides - Ferrero Mortara et al., p. 39, pl. 3, ig. 10. 2001 Sinum patulum - Lozouet et al., p. 45, pl. 18, ig. 6. 2007 Sinum aquense - Zunino, p. 127 (pars); not specimen PU23669 in pl. 1, ig. 14 = Sinum sp. 3 (see below); not specimen BS.029.08.007 = Sigaretus cryptostomoides Sacco, 1890. 2007 Sinum sigaretoides - Zunino, p. 127, pl. 1, ig. 15. ? 2007 Sinum patulum - Pister & Wegmüller, p. 103, pl. 19, igs. 8-21. Type material: Sigaretus haliotideus var. B patula Grateloup, type material not seen. Grateloup’s original material is housed in the University of Bordeaux 1 in Talence, France (Didier Merle, personal communication 2013), but we failed to have any answer from the curator there. Sigaretus subcanaliculatus d’Orbigny, syntypes MNHNF-B27769, localities around Bordeaux (5 specimens). One specimen (out of 5) belongs in Sigaretotrema Sacco, 1890 (see remarks below). Sigaretus (Sigaretus) aquensis var. longotriangula Sacco, lectotype (here designated): the shell igured by Sacco (1891, pl. 1, ig. 63) and reigured herein (Pl. 15, ig. 9), MGPT BS.029.08.004, Colli Torinesi; 1 paralectotype, MGPT BS.029.08.004/01, Colli Torinesi. Sigaretus aquensis var. taurinensis Sacco, syntype: the shell igured by Sacco (1891, pl. 1, ig. 62), MGPT BS.029.08.005, Colli Torinesi (see the remarks on Sinum perinlatum below). Sigaretus (Cryptostoma) sigaretoides Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 1, ig. 70) and reigured herein (Pl. 15, ig. 10), MGPT BS.029.09.001, Colli Torinesi. Material erroneously referred to as Sigaretus (Sigaretus) aquensis var. deshayesi Michelotti, 1847 in MGPT: Colli Torinesi: 5 spms., MGPT BS.029.08.003. Other material examined: Léognan, Le Coquillat (France): 2 spms., MZB 60116, 125 spms., NP 9877, 1 spm., MPUM 11367 (Pl. 15, ig. 11), 16 spms., private collection; Saucats (France): 1 spm., MNHN-F-J03842 (specimen igured by Cossmann 1925, pl. 4, igs. 23, 24), 1 spm., MPUM 11370 (Pl. 15, ig. 12); Valle Ceppi: 1 spm., MGPT-PU 108134, 1 spm., MGPT-PU 25270 (specimen referred to as Sinum sigaretoides by Zunino 2007), 1 spm., MZB 30796, 1 spm., PG 40 (Pl. 16, ig. 1), 1 spm., PG 41 (Pl. 16, ig. 2); Val Sanfrà: 1 spm., MGPT-PU 107640. Description. Protoconch small, low-turbiniform, of 2.5-2.6 convex whorls, with remnants of spiral sculpture in some specimens, tip small. Teleoconch conical-auriculate, moderately depressed, nearly as high as wide, rather thin. Spire cyrtoconoid, low, rather pointed, whorls faintly convex. Suture adpressed. Last whorl dome-shaped in abapertural (dorsal) view, depressed, noticeably extended toward aperture; subsutural shelf indistinct; narrowly rounded periphery about at midline. Aperture ovaterectangular, oblique, in markedly prosocline plane, length nearly 1.3 times width. Outer lip arched; basal lip gently to very gently arched; inner lip relected toward umbilical area, subangular adapically, then curving to form an angle with thin parietal callus below border of umbilical area. Umbilicus absent or a very narrow chink. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture starting with seven-ten even, closely set spiral cords; subsequently, cords slowly increase in number and change into ribbons, undulating on last whorl; in- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy terspaces of last whorl narrower than ribbons, with one-three threads or one cordlet; base with spiral cords increasingly iner downward; spirals crossed by dense growth markings. Dimensions (mm) DHW PD H D SH 0.0980.118 0.8481.060 4.19826.722 4.45129.407 0.7023.270 0.108 0.954 15.460 16.929 1.986 AH AW IS SA 3.17823.770 3.30521.273 44°-60° 116°-140° 13.474 12.289 52° 128° Remarks. The present species was currently cited with the name aquensis/aquense either attributed to Recluz in Deshayes, 1843 (Cossmann & Peyrot 1919), or to Recluz in Chenu, 1843 (Sacco 1891; Glibert 1963), or to Recluz, 1851 (Anderson 1960; Nordsieck 1972; Janssen 1984; Wienrich 2001). Cossmann & Peyrot (1919, p. 234) made reference to Deshayes’ “Traité élémentaire de conchyliologie”; in this account (1839-1853) neither the genus Sigaretus nor the name aquensis are cited. The genus Sigaretus is covered in Deshayes & Milne Edwards (1843, pp. 7-14), but also in this case the name aquensis is never mentioned. The genus Sigaretus was illustrated by Chenu (1843, pls. 1-4) in his monumental atlas (1842-1853); again, the name aquensis is not present. The name aquensis was used for the irst time by Recluz (1851) to rename the shell igured by Grateloup (1847) and referred to as “Sigaretus haliotideus Lam”. This being stated, we note that: 1) Grateloup (1847) validly introduced the var. patula of Sigaretus haliotideus, 2) the var. patula was based on triling differences that do not warrant the separation from Sigaretus haliotideus (see the above synonymy), and 3) the name patula Grateloup has priority over the name aquensis Recluz. Consequently, we concur with Lozouet et al. (2001) in considering patula Grateloup as the oldest available name for the present species, this also in agreement with ICZN (1999, Article 23 of the Code). After its introduction, Sigaretus subcanaliculatus d’Orbigny, 1852 was considered as a synonym of the present species. We have examined the syntypes of S. subcanaliculatus in MNHN (5 specimens collectively numbered F-B27769) and note that one specimen belongs in Sigaretotrema Sacco, 1890 (see the remarks on Sigaretus michaudi Michelotti, 1847); 217 the other four specimens fully conform to the concept of d’Orbigny’s species. These four syntypes are indistinguishable from S. patulum; thence, we conirm the placement of S. subcanaliculatus in the synonymy of S. patulum. Sigaretus (Cryptostoma) sigaretoides Sacco, 1890 (holotype in MGPT examined) has protoconch and teleoconch characters that fully agree with those of Sinum patulum and is herein regarded as a synonym of Grateloup’s species. Sinum aquense was recorded from lower and middle Miocene units of the North Sea Basin (Sorgenfrei 1958; Anderson 1960; Nordsieck 1972; Janssen 1984; Wienrich 2001). Examination of the illustrations published by the cited workers and of abundant Dutch material in RGM (225.947, 793.934, 793.935, 793.936) rises doubt on whether these specimens fully agree with the characters of S. aquense (= patulum). They differ somewhat from the French and Italian shells in that have a protoconch with greater diameter and with the spiral striation restricted to the last half-whorl, a teleoconch with wider umbilical chink, and a sculpture starting with more numerous (13-15) spiral cords. The Dutch, German and Danish specimens likely represent a northern subspecies of Sinum patulum. A note is to be made on the material of Nordsiek (1972): according to the quite poor illustration published by the author, his specimens seem to have a wide, open umbilicus and hardly belong in Sinus. Concerning Sacco’s varieties praecedens, longotriangula and taurinensis of Sigaretus (Sigaretus) aquensis (see the above synonymy), we have examined the respective original material in MGPT and can state that they cannot be distinguished from Sinum patulum consistently. The varieties conicolonga Sacco, 1890 and tauroinlata Sacco, 1890 were not found in MGPT (cf. Ferrero Mortara et al. 1984). According to the igures published by Sacco (1891), these varieties seem to differ somewhat from S. patulum. However, in the absence of respective original material, nothing can be stated deinitely about their relationships with S. patulum. The variety patula Grateloup was just cited by Sacco (1890b, p. 38) without description; also in this case the related material was not found in MGPT. The Burdigalian specimens from Switzerland referred to as Sinum patulum (Grateloup, 1847) by Pister & Wegmüller (2007) are quite poorly preserved and their assignment to the present species is doubtful. Robba E., Pedriali L. & Quaggiotto E. 218 Stratigraphic occurrence. Sinum patulum (Grateloup, 1847) is known deinitely from the early Oligocene of Italy (var. praecedens Sacco, 1890) and from lower Miocene units of France and Italy. Its quotation from the Burdigalian of Switzerland needs to be conirmed as does that of Sinum (Sinum) aquense from the Eggenburgian of Paratethys (Steininger et al. 1971). Sinum paviai sp. n. Pl. 16, igs. 3-5 Derivation of name: The species is named after Prof. Giulio Pavia, University of Torino, who provided lot of material relevant to the present study. Holotype: Borelli: MGPT-PU 135294 (Pl. 16, ig. 3). Paratypes. Borelli: 1 spm., MGPT-PU 135295 (Pl. 16, ig. 4), 1 spm., MGPT-PU:135353 (Pl. 16, ig. 5), 1 spm., MGPT-PU 135354, 10 spms., MGPT-PU 135355, 1 spm., MPUM 11368. Other material examined: Borelli: 6 spms., PG 30, 1 spm., NP 9818; Rio di Bocca d’Asino: 1 spm., PG 36, 1 spm., PG 37. Prservation: Except for slight abrasion of the surface in some shells, the preservation is fair. Type locality: Borelli (see appendix). Horizon: Gray, medium sand of Tortonian age. Diagnosis: Protoconch I with minute granular microprotuberances; protoconch II with spiral cordlets. Shell rather globose, low-spired, with very narrow umbilical chink. Funicle and umbilical callus absent. Aperture oval, markedly prosocline. Sculpture starting with 10-11 even, rather widely spaced spiral cords tending to become lat-topped, with one cordlet or one-three threads in interspaces; lowermost base with increasingly iner cords. cords; approximately by second whorl cords tend to became lat-topped and one cordlet or one-three threads occur in interspaces; lowermost base with increasingly iner cords; spirals crossed and made somewhat beaded by dense growth lines. Dimensions (mm) DHW PD H D SH 0.0870.103 1.2991.559 3.80010.004 7.50610.678 1.0542.186 0.095 1.429 6.902 9.092 1.620 AH AW IS SA 2.5588.006 4.8626.722 33°-53° 99°-143° 5.282 5.792 43° 121° Remarks. The Oligocene to lower Miocene species Sinum patulum (Grateloup, 1847) is somewhat similar, differing from Sinum paviai sp. n. in that it has a signiicantly smaller protoconch diameter, a conical-auriculate, more depressed teleoconch with the depressed last whorl distinctly extended toward the aperture, and the sculpture of undulating spiral ribbons. The Burdigalian species Sinum cryptostomoides (Sacco, 1890) is also similar, but is easily distinguished from S. paviai by its smooth protoconch, its auriculate teleoconch, its dome-shaped last whorl PLATE 16 Description. Protoconch medium-sized, low-turbiniform, of 2.75 convex whorls, tip very small; protoconch I with minute granular microprotuberances irregularly arranged into spiral rows and inal coarse growth markings; protoconch II with last whorl sculptured with even spiral cordlets. Teleoconch rather globose, nearly as high as wide, moderately thick. Spire low, broadly cyrtoconoid, whorls very slightly convex. Suture adpressed. Last whorl broadly oval, slightly expanded toward aperture; subsutural shelf indistinct; periphery well above midline. Aperture oval in markedly prosocline plane, length about 1.35 times width. Outer lip arched, basal lip less so; inner lip with thin callus relected over umbilical area, subangular adapically, merging into thin parietal callus well below border of umbilical area. Umbilicus a very narrow chink, absent in some specimens. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture starting with 10-11 even, rather widely spaced spiral Fig. 1 - Sinum patulum (Grateloup, 1847). Valle Ceppi. PG 40; a, apertural side (height of shell 11.81 mm); b, abapertural side; c, basal view. Fig. 2 - Sinum patulum (Grateloup, 1847). Valle Ceppi. PG 41; a, protoconch; b, detail of protoconch. Fig. 3 - Sinum paviai sp. n. Borelli. Holotype, MGPT-PU 135294; a, apertural side (height of shell 9.37 mm); b, abapertural side; c, basal view. Fig. 4 - Sinum paviai sp. n. Borelli. Paratype, MGPT-PU 135295; a, protoconch; b, detail of protoconch. Fig. 5 - Sinum paviai sp. n. Borelli. Paratype, MGPT-PU 135353; a, apertural side (height of shell 6.37 mm); b, basal view. Fig. 6 - Sinum perinlatum (Sacco, 1890). Sassello. Holotype of Sigaretus (Sigaretus) aquensis var. perinlata Sacco, 1890. MGPT BS.029.08.002; apertural side (scale bar 4 mm). Fig. 7 - Sinum sp. 1. Case Soghe. MPUM 11369; protoconch. Fig. 8 - Sinum sp. 1. Case Soghe. NP 9820; a, apertural side (height of shell 10.58 mm). Fig. 9 - Sinum sp. 2. Villa Maiolo. PG 38; a, apertural side (height of shell 3.60 mm); b, abapertural side; c, basal view; d, protoconch; e, detail of protoconch. Fig. 10 - Sinum sp. 3. Albugnano. PG 39; a, apertural side (height of shell 11.37 mm); b, abapertural side; c, basal view. Fig. 11 - Sinum sp. 3. Valle Vergnana. MZB 19184; a, apertural side (height of shell 10.85 mm); b, abapertural side; c, protoconch. Eocene, Oligocene and Miocene naticid gastropods of Northern Italy 219 Robba E., Pedriali L. & Quaggiotto E. 220 remarkably extended toward the aperture, and its sculpture of broad spiral ribbons with scarce intervening threads. Stratigraphic occurrence. Sinum paviai sp. n. was recovered from upper Miocene (Tortonian) deposits of Piedmont. Sinum perinlatum (Sacco, 1890) stat. n. Pl. 16, ig. 6 1890b Sigaretus (Sigaretus) aquensis var. perinlata Sacco, p. 38. 1891 Sigaretus (Sigaretus) aquensis var. perinlata - Sacco, p. 99, pl. 1, ig. 60. 1984 Sigaretus aquensis var. perinlata - Ferrero Mortara et al., p. 38. Type material: Sigaretus (Sigaretus) aquensis var. perinlata Sacco, holotype (by monotypy): the shell igured by Sacco (1891, pl. 1, ig. 60) and reigured herein (Pl. 16, ig. 6), MGPT BS.029.08.002, Sassello. Material erroneously referred to as Sigaretus (Sigaretus) aquensis var. taurinensis Sacco, 1891 in MGPT: Colli Torinesi: 1 syntype (of 2), MGPT BS.029.08.005. Description. Protoconch abraded. Teleoconch ovate-cylindrical, higher than wide, thick. Spire broadly cyrtoconoid, rather depressed, whorls very gently convex. Suture adpressed. Last whorl 90% of total height, globose, subcylindrical in abapertural (dorsal) view; subsutural shelf poorly deined, more distinct behind aperture; periphery about at midline. Aperture teardrop-shaped in markedly prosocline plane, length about twice width. Outer lip roundly angular adapically, slightly arched in mid-abapical part, more so basally; inner lip very thick, bent at level of umbilical area, merging into moderately thick parietal callus below border of umbilical area. Umbilicus, funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture of rather even, closely set spiral ribbons, narrower and round-topped over base; interspaces very narrow, sharply incised, made pitted by crossing of coarse and dense growth markings. Dimensions (mm) DHW PD H D SH AH AW 18.640 17.291 3.500 15.140 12.802 - - IS SA 40° 122° Remarks. Sacco (1890b) validly introduced the variety perinlata of Sigaretus (Sigaretus) aquen- sis Recluz, 1843. Examination of the holotype in MGPT showed that the variety perinlata is quite unlike Sigaretus aquensis because of its ovate-cylindrical, higher than wide teleoconch and its sculpture of spiral ribbons with very narrow interspaces devoid of threads. The teleoconch shape is the most relevant distinguishing character of Sinum perinlatum and readily separates it from the other Italian species of Sinum that invariably have more depressed shells. Sinum oligopolitum (Sacco, 1890) is somewhat similar, but differs in that it has: 1) a conical-ovate teleoconch, 2) a more elevated, conical and pointed spire, 3) a broadly oval last whorl, expanded toward the aperture, 4) an ovate-quadrangular aperture, and 5) a sculpture of uneven, rather widely spaced spiral cords, with two-three threads in the interspaces. Sacco (1891) proposed the var. taurinensis of Sigaretus (Sigaretus) aquensis on the basis of material from an unspeciied locality in the Turin Hills (Colli Torinesi); the age was said to be “Helvetian”. Sacco’s material in MGPT consists of two syntypes collectively numbered BS.029.08.005. The larger syntype displays the characters of Sinum perinlatum and is herein assigned to this species. The other syntype (the smaller) fully agrees with the characters of S. (Sigaretus) aquensis. Stratigraphic occurrence. The holotype of Sinum perinlatum (Sacco, 1890) was collected from lower Oligocene deposits of Liguria. The Miocene specimen originally referred to as Sigaretus (Sigaretus) aquensis var. taurinensis Sacco (see remarks above), on the basis of the matrix inilling, results to have been collected from the Termofourà Formation of Burdigalian age (cf. Zunino & Pavia 2009). Sinum sp. 1 Pl. 16, igs. 7, 8 Material examined: Case Soghe: 1 spm., MPUM 11369 (Pl. 16, ig. 7), 1 spm., NP 9820 (Pl. 16, ig. 8), 1 spm., MGP-PD 31530. Description. Protoconch small, depressedturbiniform, of 2.5 rather convex whorls with remnants of spiral cordlets, tip small. Teleoconch auriculate, depressed, wider than high, rather thin. Spire broadly conical, pointed, low but distinctly protruding from last whorl, whorls faintly convex. Suture thin, adpressed. Last whorl lenticular, rather depressed, quickly expanding toward aperture; sub- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy sutural shelf indistinct; narrowly rounded periphery slightly above midline. Aperture large, elliptical, oblique, in markedly prosocline plane, distinctly wider than high. Outer lip arched; basal lip semicircular; inner lip also arched, with thin, narrow callus curved adapically to merge into thin parietal callus well below border of umbilical area. Umbilicus a rather narrow chink. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture of uneven, round-topped, rather closely set spiral cords, more widely spaced subsuturally, narrower toward periphery of last whorl, tending to fade away on base; spirals crossed by coarse growth markings. 221 expanding toward aperture; subsutural shelf indistinct; narrowly rounded periphery slightly above midline. Aperture large, oval, oblique, in markedly prosocline plane, distinctly wider than high. Outer lip arched; basal lip semicircular; inner lip gently arched, subangular adapically, then merging into thin parietal callus well below border of umbilical area. Umbilicus a narrow chink largely covered by everted labial callus. Funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture of uneven and unevenly spaced spiral cords, narrower toward periphery of last whorl, changing into threads on upper base, obsolescent on lowermost base; spirals crossed by incised growth markings. Dimensions (mm) Dimensions (mm) DHW PD H D SH AH AW 0.129 0.936 7.810 12.630 1.760 6.050 8.020 DHW PD H D SH AH AW IS SA 0.139 1.048 3.600 10.550 0.930 2.671 7.953 45° 124° IS SA 36° 138° Remarks. Sinum sp. 1 appears to be closely similar to Sinum borellense sp. n. (see above) from which it differs in that it has: 1) a protoconch with a signiicantly greater diameter of the irst halfwhorl, 2) a teleoconch with a low but distinctly prominent spire and with a wider umbilical chink, and 3) a sculpture of round-topped spiral cords (in S. borellense the cords become lat-topped by the second whorl) that tend to fade away over the base. The present specimens almost certainly belong to a previously unrecorded species, but naming it must await that more, better preserved material be recovered. Stratigraphic occurrence. Sinum sp. 1 was recovered from lower Oligocene deposits of Veneto. Sinum sp. 2 Pl. 16, ig. 9 Material examined: Villa Maiolo: 1 spm., PG 38. Description. Protoconch medium-sized, planorbid, of 2.5 lat whorls with increasingly robust peripheral cord overridden by short, irregular collabral riblets; tip small. Teleoconch auriculate, depressed, wider than high, rather thin. Spire broadly conical, very low, quite scarcely protruding from last whorl, whorls faintly convex. Suture thin, adpressed. Last whorl lenticular, markedly depressed, quickly Remarks. Sinum sp. 2 is readily distinguished from the other Sinum species by its planorbid protoconch with peculiar sculpture. It somewhat resembles Sinum borellense sp. n. (see above) in terms of teleoconch shape, but the sculpture of uneven and unevenly spaced spiral cords is different from that of S. borellense (lat-topped cords). Stratigraphic occurrence. Sinum sp. 2 seems to be a rare species occurring in Tortonian deposits of Piedmont. Sinum sp. 3 Pl. 16, igs. 10, 11 Material erroneously referred to as Sinum aquense (Recluz, 1851) in MGPT: Val Sanfrà: 1 spm., MGPT-PU23669 (specimen of Zunino 2007, pl. 1, ig. 14). Other material examined: Albugnano: 1 spm., PG 39 (Pl. 16, ig. 10); Valle Vergnana: 1 spm., MZB 19184 (Pl. 16, ig. 11). Description. Protoconch medium-sized, low-turbiniform, of slightly more than three convex, apparently smooth whorls, tip very small. Teleoconch auriculate, depressed, markedly wider than high, rather thick. Spire broadly cyrtoconoid, very low, whorls faintly convex. Suture adpressed. Last whorl lenticular, depressed, only slightly expanding toward aperture; subsutural shelf absent on irst whorl, poorly deined on subsequent whorls; narrowly rounded periphery above midline. Aper- Robba E., Pedriali L. & Quaggiotto E. 222 ture ovate-rectangular, oblique, in markedly prosocline plane, length about 1.3 times width. Outer lip subangular adapically in larger specimens, then gently arched; basal lip semicircular; inner lip with wide, lat and thick callus, subangular at adapical one-third, curving to form a sharp angle with thick parietal callus below border of umbilical area. Umbilicus, funicle and umbilical callus absent. Basal fasciole indistinct. Sculpture of rather even, closely set, slightly undulating spiral cords, iner adapically and much so over base; unsculptured subsutural band tends to develop in larger specimens; spirals crossed by dense growth markings. Dimensions (mm) DHW PD H D SH 0.068* 1.145* 6.20813.832 7.75717.637 0.7992.235 1.517 * 1 protoconch measurable 10.020 12.697 AH AW IS SA 4.08512.921 4.151-14.515 44°-68° 110°-146° 8.503 9.333 56° 128° Remarks. Sinum sp. 3 somewhat resembles Sinum aquense = Sinum patulum (Grateloup, 1847) and has been mistaken for it. The present form is readily distinguished from Grateloup’s species in that has: 1) a protoconch of 0.5 more whorls, with a signiicantly smaller diameter of the irst half-whorl, 2) an auriculate, more depressed teleoconch with the last whorl scarcely extended toward the aperture (the shell has a subcircular outline in dorsal view, whereas the outline is oval in S. patulum), 3) an outer lip subangular adapically instead of regularly arched, and 4) a sculpture of closely set spiral cords with no threads in the interspaces (S. patulum has spiral ribbons with intervening threads or cordlets). Sinum perinlatum (Sacco, 1890) also exhibits some resemblance to Sinum sp. 3 in that has a similar outline in dorsal view, but its teleoconch is more inlated and differently shaped, its aperture length is nearly twice the width, and its sculpture consists of spiral ribbons separated by very narrow, sharply incised grooves. The three examined specimens certainly belong to a previously undescribed species, but more material is required in order to name it. Stratigraphic occurrence. Sinum sp. 3 was collected from the Termofourà Formation of Burdigalian age (Torino Hills) and from Serravallian deposits near Albugnano. Acknowledgements. This paper has beneited from reviewing by Dr. A. Beu (Institute of Geological and Nuclear Sciences, Lower Hutt) and by Dr. B. Landau (Naturalis Biodiversity Center, Leiden). The following curators provided access to the collections on their care and/or kindly loaned specimens: Dr. A. Bonitto (Museo di Zoologia, Bologna); Dr. M. Fornasiero (Museo di Geologia e Paleontologia dell’Università, Padova); Dr. C. Francou (Museo Geologico G. Cortesi, Castell’Arquato); Dr. V. Frisone (Museo di Archeologia e Scienze Naturali “G. Zannato”, Montecchio Maggiore); Mr. Y. Gilly (Réserve Naturelle Géologique de Saucats, Muséum de Saucats-La Brède); Dr. L. Mazzini (Museo G. Scarabelli, Imola); Dr. D. Merle (Muséum National d’Histoire Naturelle, Paris); Dr. D. Ormezzano (Museo Regionale di Scienze Naturali, Torino); Dr. B. Pallozzi (Museo Civico D. Dal Lago, Valdagno); Dr. R. Pancaldi (Museo Paleontologico e Preistorico “P. Leonardi” dell’Università, Ferrara); Dr. C. Sendino (Natural History Museum, London); Dr. P. Serventi (Dipartimento di Scienze Chimiche e Geologiche, Università di Modena e Reggio Emilia); Dr. M. Taviani (ISMAR-CNR, Bologna); Drs. F. Wesselingh and R. Pouwer (Nationaal Naturhistorisch Museum, Naturalis, Leiden). Mr. H. Boerman (Rotterdam) and Dr. R. G. Moolenbeek (Instituut voor Taxonomische Zoölogie, Amsterdam) donated naticids from Winterswijk Miste (Hemmorian), Dr. A. Morton (Penrhyncoch, Aberystwyth) donated much reference material from the London Clay; Dr. S. Tracey (Natural History Museum, London) donated two shells and two opercula of Sigatica clarkeana (Aldrich, 1887). Drs. M. Aberhan and H. Götz (Museum für Naturkunde, Berlin), Drs. L. Cavin and A. Piuz (Museum d’Histoire Naturelle, Geneva), Dr. M. Harzhauser (Naturhistorisches Museum, Wien), Drs. A. Prieur and E. Robert (Université de Lyon 1), and Dr. C. Sendino (Natural History Museum, London) provided excellent digital photographs of syntypes or relevant specimens. Mr. J.-F. Lesport (Sainte-Hélène, France) showed his vast collection of naticids from the Miocene of Bordeaux area and provided outstanding digital photographs of shells and opercula. Mr. J.-P. Dupuy (Cadillac, France) helped during ield-work in Aquitaine. Dr. M. Aberhan (Museum für Naturkunde, Berlin), Drs. D. Merle and J.-M. Pacaud (Muséum National d’Histoire Naturelle, Paris) helped in trying to locate relevant collections including the original material of species (Speyer collection, Bayan and Grateloup collections). Dr. A. Ehling (Bundesanstalt für Geowissenschaften und Rohstoffe, Berlin), Dr. E. Robert (Université de Lyon 1) and Dr. L. Cavin (Museum d’Histoire Naturelle, Geneva) provided information about species presently available respectively in Speyer collection, Bayan collection and Defrance collection. Mr. J.-C. Hourcade (Villenave-d’Ornon) helped in going through the collections of the Muséum de Saucats-La Brède; Mr. C. Tabanelli (Cotignola) helped in locating Miocene naticids in the Museo G. Scarabelli, Imola. We are indebted to Prof. G. Pavia and Dr. M. Zunino (University of Torino) who informed on the most up-to-date cronostratigraphic framing of several naticid-bearing units of Piedmont and loaned a wealth of material relevant to the present study, to Mr. A. De Angeli (Vicenza) for information on the outcrops of Chiuppano and Sangonini, to Mr. M. Boscardin (Vicenza) for information on the locality Monte Gloso, to Mr. M. Gambillara (Treviso) for information on the hystorical localities near Possagno, and to Mr. Marco Vicariotto (Vicenza) for information on the section of Chiuppano. Prof. Maria Triantaphyllou (University of Atherns) provided calcareous nannoplankton evidence of the age of some shells. Prof. K. Bandel (Buchholz, Germany), Mr. J.-P. Dupuy (Cadillac, France), Dr. T. Huel- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy sken (University of Queensland, Brisbane), Dr. P. Rocher (Muséum d’Histoire Naturelle, Bordeaux), Dr. D. Scarponi (University of Bologna) and Dr. S. Tracey (Natural History Museum, London) assisted with some essential references. Mr. H. Boerman (Rotterdam), Mr. A. Bottazzi (Valdagno), Mr. E. Stadelmann (Hersbruck) and Mr. Z. Zordan (Valdagno) helped with translations from Dutch and German. The following private collectors generously donated or loaned much valuable material: Mr. R. Alberti (Bassano del Grappa); Mr. E. Bartoli (Lugo di Romagna); Mr. D. Battilani (Modena); Mr. R. Bertamini (Cesena); Mr. L. Bertolaso (Reggio Emilia); Mr. B. Bizzotto (Treviso); Mr. S. Boschele (Borgo Valsugana); Mr. R. Bourgeais (Paris); Mr. M. Brunetti (Rioveggio); Mr. A. Checchi (Montecchio Maggiore); Mr. G. Cracco (Arzignano); Mr. M. Cresti (San Casciano in Val di Pesa); G. Della Bella (Monterenzio); Mr. B. Dell’Angelo (Genova); Mr. J.-P. Dupuy (Cadillac); Mr. M. Forli (Prato); Mr. P. Frediani (Casteliorentino); Mr. P. Fremont (Bordeaux); Mr. G. Gariani (Milano); Mr. P. Giuntelli (Nole, Torino); Mr. S. Granelli (Noceto); Mr. L. Landini (San Donato, Parma); Mr. A. Maccaferri (Poggio Renatico, Ferrara); Mr. P. Magenes (Milano); Mr. S. Marsigli (Bazzano); Mr. S. Mezzalira (Bassano del Grappa); Mrs. P. Orru (Basiglio); Mr. P. Petracci (Cesena); Mr. M. Salmoiraghi (Castellanza); Mr. M. 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(2012) - Le campagne di scavo del Museo Paleontologico di Roncà in località Monte Duello (Comune di Montecchia di Crosara) e Valle della Chiesa (Comune di Roncà), nei Monti Lessini veronesi (Italia settentrionale): primi risultati. Boll. Mus. Civ. St. Nat. Verona, 36: 53-62. Zunino M. (2007) - Associazioni a molluschi del Miocene inferiore e medio della Collina di Torino: analisi tassonomica, tafonomica e paleobiogeograica. PhD Thesis, Università di Torino, 293 pp. (unpublished). Zunino M. & Pavia G. (2009) - Lower to Middle Miocene mollusc assemblages from the Torino Hills (NW Italy): synthesis of new data and chronostratigraphical arrangement. Riv. It. Paleont. Strat., 115: 349-370. APPENDIX (Locality data) PIEDMONT 1. Albugnano (Asti Province). There are no more precise locality data concerning the material so labeled. Ferrero Mortara et al. (1981, p. 14) considered the fossils from Albugnano as of Serravallian age. According to the Geological Map of Italy (scale 1:50,000, Sheet 156, Torino Est), the Serravallian deposits in Albugnano area are represented by the formation named Marne di Mincengo that crops out extensively south and southwest of the village. 2. Borelli, Moncucco Torinese (Torino Province). A small outcrop, approximately 0.15 km northeast of the hamlet named Borelli (formerly Tetti Borelli), exposes medium to coarse sand that was currently assigned a Messinian age. Recent research demonstrated a Tortonian age on the basis of nannoplankton and holoplanktonic mollusks. For additional information, reference to Pavia (1991) and Janssen (1999, 2010). 3. Cascina Pianiorito, Albugnano (Asti Province). Excavations near Cascina Pianiorito, approximately one km southeast of Albugnano, have unearthed sandstone and gray pebbly clay presently concealed by the cultivation of a vineyard. According to the Geological Map of Italy (scale 1:50,000, Sheet 156, Torino Est), the Langhian Baldissero Formation crops out in the area of Cascina Pianiorito. 4. Cassinelle (Alessandria Province). Graywacke and gray silt exposed near Cascina Fogli, approximately 1.7 km southsouthwest of the village of Cassinelle. According to the Geological Map of Italy (scale 1:50,000, Sheet 194, Acqui Terme) these lithotypes form the lowermost part of the formation named Marne di Rigoroso; the age is late Rupelian. For additional infor- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy mation, reference to Lorenz (1969). 5. Colli Torinesi (Torino Province). There are no more precise locality data concerning the material so labeled. According to Zunino & Pavia (2009), the bulk of the fossils was collected from the mid to upper Burdigalian Termofourà Formation and, less abundantly, from the Langhian Baldissero Formation. 6. Colli Tortonesi or Tortona (Alessandria Province). Again, no detail on the localities is given on the labels. Most likely the fossils are from Tortonian marly deposits of the Sant’Agata Fossili Formation that crops out in the hill area south of the town of Tortona. 7. Grazzano Badoglio (Asti Province). Outcrop north of the village of Grazzano Badoglio, close to the former shooting gallery (Tiro a volo). Gray marl belonging to the formation named Areniti di Tonengo of Langhian age. For additional information, reference to Arpa (2004). 8. Lerma (Alessandria Province). Exposures in the bed of Torrente Piota, upstream of the bridge located southwest of the village of Lerma. Coarse, gray sandstone with marly intercalations of the uppermost part of the Molare Formation; the location close to the upper boundary of the formation indicates a late Rupelian age. For additional information, reference to Lorenz (1969) and Gnaccolini (1978). 9. Madonna delle Rocche, Molare (Alessandria Province). Outcrops on both banks of Torrente Orba, approximately 2.8 km south of the village of Molare and 1.4 km southeast of the locality named Madonna delle Rocche. Coarse, gray sandstone with marly intercalations of the upper part of the Molare Formation; the age is late Rupelian. For additional information, reference to Lorenz (1969). 10. Molare (Alessandria Province). Cliffy bank on the right of Torrente Orba southeast of the village of Molare, upstream of the road connecting Molare to Ovada. Coarse, gray sandstone of the upper part of the Molare Formation; the age is late Rupelian. For additional information, reference to Lorenz (1969). 11. Moncucco Torinese (Asti Province). Outcrops on the southwestern slope of the hill on top of which the village of Moncucco Torinese is located. Medium to coarse sand belonging to the formation named Marne di S. Agata Fossili; the age is Tortonian. For additional information, reference to Arpa (2009). 12. Monte dei Cappuccini, Torino (Torino Province). Located on the northwestern slope of the Torino Hills facing the Po River. The fossils of this locality were collected from siltstones, sands and conglomerates forming the upper part of the Baldissero Formation of late Langhian age. For additional information, reference to Zunino & Pavia (2009). 13. Pareto (Alessandria Province). Outcrops in the locality named Sbarnera, three km east of the village of Pareto. Gray sandstone and sand belonging to the upper part of the Molare Formation; the age is late Rupelian. For additional information, reference to Lorenz (1969). 14. Rio di Bocca d’Asino, Stazzano (Alessandria Province). Exposures on both banks of the brook named Rio di Bocca d’Asino, upstream of the road connecting Stazzano to Sardigliano, about 1.9 km northeast of the former village. Resedimented sandstones and conglomerates forming lenticular bodies into the lower member of the S. Agata Fossili Formation; the age is Tortonian. For additional information, reference to Bongo (1914) and Ghibaudo et al. (1985). 15. Sant’Agata Fossili (Alessandria Province). Gully about 231 0.5 km east-southeast of the village of Sant’Agata Fossili, coinciding with the collecting site 10 of Robba (1968). Gray-bluish clayey marl belonging to the upper member of the S. Agata Fossili Formation; the age is late Tortonian. For additional information, reference to Robba (1968). 16. Squaneto, Spigno Monferrato (Alessandria Province). Outcrop along the road connecting Spigno Monferrato to Squaneto, approximately one km northwest of Squaneto. Gray sandstone and marl forming the uppermost part of the Molare Formation. The age is late Rupelian. For additional information, reference to Lorenz (1969). 17. Stazzano (Alessandria Province). There are no more precise locality data concerning the material so labeled. Some specimens might have been recovered at locations around Stazzano where the lower member of the S. Agata Fossili Formation (Tortonian) crops out extensively. Most of the material likely comes from Rio di Bocca d’Asino (see above). 18. Termofourà (Torino Province). Outcrop near Fontana dei Francesi, along the road leading to the locality named l’Eremo. Conglomerate made of small pebbles into a sandy-pelitic matrix pertaining to the topmost part of the Termofourà Formation; the age is late Burdigalian. 19. Tetti Civera, Pino Torinese (Torino Province). Outcrops close to the hamlet named Tetti Civera, approximately 1.1 km northnorthwest of the village of Valle Ceppi. Coarse sand of the Termofourà Formation; the age is mid-Burdigalian. Likely the outcrops are those referred to as Valle Ceppi (see ig. 3 of Zunino & Pavia 2009). 20. Val Sanfrà, Baldissero Torinese (Torino Province). Outcrops on right bank of Rio Baldissero, about 0,8 km north-northwest of the village of Baldissero, expose alternating pelites and conglomeratic sandstones (known as “Val Sanfrà Strata”) belonging to the Termofourà Formation; the age is mid-Burdigalian. For additional information, reference to Zunino & Pavia (2009). 21. Valle Ceppi, Pino Torinese (Torino Province). Outcrops along the small stream named Rio Civera, 1.5 km north-northwest of the village of Valle Ceppi and about three km east of Pino Torinese. Coarse sand belonging to the Termofourà Formation; the age is mid-Burdigalian. For additional information, reference to Zunino & Pavia (2009). 22. Valle Vergnana, Baldissero Torinese (Torino Province). Outcrop on right bank of the stream named Rio di Vergnana, about 0.84 km west-southwest of Baldissero Torinese, coinciding with the site UCLA 4927 of Hall (1964). Cemented sandy conglomerates belonging to the Termofourà Formation; the age is mid-Burdigalian, nearly coeval to Val Sanfrà. For additional information, reference to Hall (1964) and Arpa (2010). 23. Villa Allason, Torino (Torino Province). Outcrop along Val Salice, about 0.7 km west of Eremo (a district of Torino). The deposit, made of sandy pelites and poorly cemented sandstones, belongs to the lower part of the Baldissero Formation; the age is early Langhian. For additional information, reference to Zunino & Pavia (2009). 24. Villa Bertini, Torino (Torino Province). Outcrop about 0.8 km northeast of Villa Allason, close to the road connecting Eremo to the city of Torino. Conglomeratic layer made of small pebbles into a sandy-pelitic matrix pertaining to the topmost part of the Termofourà Formation; the age is late Burdigalian. For additional information, reference to Zunino & Pavia (2009). Robba E., Pedriali L. & Quaggiotto E. 232 25. Villa Maiolo, Marentino (Torino Province). Diggings nearby Villa Maiolo (also Castello Maiolo), approximately one km north of the village of Marentino, have unearthed coarse clayey sand belonging to the formation named Marne di S. Agata Fossili; the age is Tortonian. For additional information, reference to Arpa (2009). LIGURIA 26. Carpenaro, Mioglia (Savona Province). Outcrop approximately two km southeast of the village of Mioglia and 0.3 km east-southeast of the hamlet named Carpenaro. Gray sandstone belonging to the uppermost part of the Molare Formation; the age is late Rupelian. For additional information, reference to Lorenz (1969). 27. Colle del Giovo Ligure, Pontinvrea (Savona Province). Outcrop about 0.65 km northwest of the Colle del Giovo Pass (road 334). Marly sandstone belonging to the Molare Formation; the age is Rupelian. For additional information, reference to Lorenz (1969; ig. 123 and p. 618). 28. Dego (Savona Province). Outcrop at Case Ciappeiroli, about one km southeast of the village of Dego. Gray sandstone belonging to the Molare Formation; the age is Rupelian. For additional information, reference to Lorenz (1969) and Arpa (2010). 29. Mioglia (Savona Province). Outcrop 0.25 km east of Case Sciriti, approximately 1.5 km east of the village of Mioglia. Coarse, gray sandstone belonging to the upper part of the Molare Formation; the age is late Rupelian. For additional information, reference to Lorenz (1969) and Arpa (2010). 30. Sassello (Savona Province). Outcrop about 1.7 km northeast of Sassello, 0.2 km left of the road connecting Sassello to Palo. Gray marl (Case Capè Marl) belonging to the Molare Formation; the age is latest Rupelian. For additional information, reference to Lorenz (1969). EMILIA-ROMAGNA 31. Montegibbio, Sassuolo (Modena Province). Gully on the right side of the stream named Rio delle Bagole and small exposure about 0.4 km south of Cà del Chierico. Gray clayey marl belonging to the Formazione del Termina (Termina Formation); the age is Tortonian. For additional information, reference to Davoli (1972). 32. Passo dei Meloni, Sogliano al Rubicone (Forlì-Cesena Province). Small outcrop in the uppermost part of a gully close to the source of Rubicone River. Gray clay with abundant Amalda glandiformis (Lamarck, 1810) and Amalda obsoleta (Brocchi, 1814). The inferred age is Tortonian. 33. Pietracuta, San Leo (Rimini Province). Outcrop on the left bank of Marecchia River, approximately 1.5 km west of the village of Pietracuta. Blue-gray clay belonging to the formation named Argille di Montebello. According to the Geological Map of Italy (scale 1:50,000, Sheet 267, San Marino), the age is early Tortonian. 34. Scipione Ponte, Salsomaggiore Terme (Parma Province). Outcrops on the left of Torrente Stirone, approximately 0.9 km north-northeast of the hamlet named Scipione Ponte. Gray clay of Tortonian age. For additional information, reference to Marasti (1973). 35. Sogliano al Rubicone (Forlì-Cesena Province). Outcrop on the left of road 11 connecting Sogliano al Rubicone to Savignano, approximately 0.2 km before the junction with the road to San Giovanni in Galilea. Gray silty-clay belonging to the unit named Argille di Casa i Gessi; the age is early Messinian. For additional information, reference to Ruggieri & Davoli (1984). 36. Vigoleno (Piacenza Province). Val San Martino Section, slope on the right of Torrente Ongina near the village of Vigoleno. Gray clayey sand belonging to the Vigoleno Member of the Termina Formation. According to the Geological Map of Italy (scale 1:50,000, Sheet 180, Salsomaggiore Terme), the age of the Vigoleno Member is early Messinian. For additional information on the Val San Martino Section, reference to Venzo & Pelosio (1963). VENETO 37. Altavilla Vicentina (Vicenza Province). Decommissioned basalt quarry north of Colle della Chiesa. Excavations for basalt have exposed a sedimentary sequence of various age, from mid-late Eocene to early Miocene. The naticids were collected from a gray-brown marly level rich in turritellids; the mollusk assemblage was said to belong to the “Sangonini Horizon” of Rupelian age. For further information, reference to Fabiani (1915), Mietto (1988), Mellini & Quaggiotto (1990) and Beschin & De Angeli (2012). 38. Cà Sella, Salcedo (Vicenza Province). Outcrop along the road connecting Cà Sella to Cà Gnata, southwest of the locality named Laverda. Resedimented, reddish-brown volcanoclastic bed that yielded an assemblage belonging to the “Sangonini Horizon”. The age is Rupelian. For additional information, reference to Molon (1867) and Fuchs (1870). 39. Case Soghe, Barbarano Vicentino (Vicenza Province). A section near the locality named Case Soghe, approximately 2.4 km north-northeast of the locality named San Giovanni in Monte, exposes a 3.7 m thick bed of gray to yellowish-brown clayey silt with small pebbles that yielded the naticids. The age is Rupelian. For additional information, reference to Accorsi Benini (1971). 40. Cava Albanello, Nogarole Vicentino (Vicenza Province). Decommissioned quarry on the left side of Torrente Chiampo, about 1.5 km south of the village of Nogarole Vicentino. The naticids were collected from rather coarse, gray tuff forming the lower part of a thick, ining upward volcanoclastic bed underlying a limestone level (“San Giovanni Ilarione Horizon”). The age is early Lutetian. For additional information, reference to Beschin et al. (1991), Beschin et al. (1996) and De Angeli et al. (2010). 41. Cava Boschetto, Nogarole Vicentino (Vicenza Province). Quarry on the left side of Torrente Chiampo, 1.2 km southsoutheast of the village of Nogarole Vicentino. The naticids were collected from the lower part of a 16.5 m thick level of bedded, greenish-gray tuff (unit c of Beschin et al. 1991). The age is early Lutetian. For further information, reference to Fornasiero & Vicariotto (1997) and Beschin et al. (1991). 42. Cava Grola, Cornedo Vicentino (Vicenza Province). Active quarry on the right side of Torrente Agno, 1.6 km west of the village of Cornedo Vicentino and about 0.6 km southwest of road 246. The naticids were collected from greenish-gray volcanoclastic sandstone layers alternating with limestone beds; the fossil assemblage is said to have a close similarity to that of the “San Giovanni Ilarione Horizon”. The age is middle Lutetian. For additional information, reference to Mietto (1975), Beccaro & De Angeli (2001), Beschin et al. (2005) and De Angeli & Lovato (2011). 43. Cava Main, Arzignano (Vicenza Province). Decommis- Eocene, Oligocene and Miocene naticid gastropods of Northern Italy sioned quarry on the left side of Torrente Chiampo, 0.4 km northwest of the village of Arzignano. The naticids were collected from the uppermost level of gray volcanic sandstone reported to be of middle Lutetian age (cf. Beschin et al. 1996; Beschin et al. 2002). 44. Cava Rossi, Priabona (Vicenza Province). Active quarry about two km northeast of Priabona and approximately 1.9 km southeast of the hamlet of Monte di Malo, on the eastern margin of the Agno-Chiampo graben. Volcanoclastic layer in the lower part of the section. Research in progress demonstrates that the age is late Ypresian-earliest Lutetian. For further information, reference to Beschin et al. (1996), De Angeli et al. (2009), Checchi et al. (2012) and Zamberlan & Checchi (2014). 45. Chiuppano (Vicenza Province). Section exposed along Torrente Astico, approximately 1.4 km east of Chiuppano. The naticids were collected from grayish-brown marly sandstone layers of Rupelian age. For additional information, reference to Patrini (1902) and Fabiani (1912). 46. Ciupio, San Giovanni Ilarione (Verona Province). Outcrop on the left side of the brook adjacent to Case Ciupio, approximately two km east of San Giovanni Ilarione, exposes four to six m of a volcanoclastic deposit, greenish-gray basally, yellowish in the upper part; the age is early Lutetian. For additional information, reference to De Gregorio (1880) and Fabiani (1915). 47. Le Coe, Cavaso del Tomba (Treviso Province). Decommisioned quarry (cava Dalla Favera; site a of Bizzotto 2005) in the locality named Le Coe, approximately 1.2 km south-southeast of the village of Possagno. Blue-gray marl forming the uppermost part of the formation known as Marna di Possagno; the age is Priabonian. For additional information, reference to Fabiani (1915), Braga (1970), Reato (1983) and Bizzotto (2005). 48. Monte Gloso, Marostica (Vicenza Province). Decommissioned basalt quarry on Monte Gloso, near the locality of Marsan. Excavations for basalt have exposed intercalations of yellowish-gray, coarse-grained marly sandstone of Rupelian age. The mollusk assemblage is said to be a mixed assortment of species of the “Sangonini Horizon” and of the “Castelgomberto Horizon”. 49. Monte Merlo, San Giovanni Ilarione (Verona Province). Outcrop in the locality named Croce Grande, approximately 1.5 km northeast of San Giovanni Ilarione. Yellowish volcanoclastic deposit forming a single bed about ten m thick; the age is Lutetian. For further information, reference to De Gregorio (1880). 50. Roncà (Verona Province). Outcrop on the left side of Valle della Chiesa (formerly Val Nera), approximately one km upstream of Roncà and 0.15 km south of Casa Tessari. Volcanoclastic layer slightly more than one m thick (black tuff with cerithiids of Fabiani 1915); the age is middle Eocene (Bartonian). For further information, reference to Fabiani (1915) and Zorzin et al. (2012). 51. Sangonini (presently Santa Maria di Lugo), Lugo (Vicenza Province). Section approximately 0.3 km northwest of the hamlet named Santa Maria. The naticids were collected from a blackish volcanoclastic level and from a yellowish-gray marly sandstone level, both of Rupelian age. For additional information, reference to Fabiani (1915). 52. Soprasalmo, Borgo Valsugana (Trento Province). Outcrops on the left side of the valley of Torrente Fumola (Pissavacca), near the locality named Soprasalmo. Sandstone level of mid-Tortonian age. For further information, reference to Venzo (1934). 53. Valle Organa, Castelcucco (Treviso Province). Outcrops 233 close to the road connecting Castelcucco to Possagno, approximately 0.4 km north of the village of Castelcucco. Blue-gray marl belonging to the formation named Marna di Possagno; the age is Priabonian. For additional information, reference to Braga (1970).

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