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Home , Bela alma, Clathurellinae, Haedropleura, Mangelia, Mangeliidae, Neogastropoda

Bollettino della Società Paleontologica Italiana, 52 (2), 2013, 71-79. Modena

Bela pseudoappeliusi n. sp. (: ) from the Plio-Pleistocene of Italy

Francesco Naldi, Giano Della Bella & Daniele Scarponi

F. Naldi, Via Corone di Berti 10, I-40137 Bologna, Italy; [email protected] G. Della Bella, Via dei Cedri 91, I-40050 Bologna, Italy; D. Scarponi, Dipartimento di Scienze Biologiche, Geologiche e Ambientali, Alma Mater Studiorum, Università di Bologna, Via Zamboni 67, I-40127 Bologna, Italy; [email protected]; corresponding author

KEY WORDS - , , Mangeliidae, , Plio-Pleistocene, Italy.

ABSTRACT - A new species of mangeliid, Bela pseudoappeliusi (Neogastropoda, Conoidea), is described from the Plio-Pleistocene of Italy. The systematic description is based on a series of eleven specimens selected from the numerous findings in northern and central Italy (Piedmont, Emilia-Romagna, Tuscany and Lazio). The new species shows affinities to Bela (s.l.) appeliusi (Bellardi, 1877) with regard to particular features such as shell dimension, teleoconch and morphology. The study includes a detailed survey of the type locality as well as a brief account of occurrences at other localities.

RIASSUNTO - [Descrizione di Bela pseudoappeliusi n. sp. (Neogastropoda, Mangeliidae) dal Plio-Pleistocene dell’Italia] - Sulla base di undici esemplari, riferibili al Plio-Pleistocene e selezionati fra quelli provenienti da varie zone dell’Italia settentrionale e centrale (Piemonte, Emilia-Romagna, Toscana e Lazio), è descritta la specie Bela pseudoappeliusi n. sp. (Conoidea, Mangeliidae). La nuova specie è confrontata con l’affineBela (s.l.) appeliusi (Bellardi, 1877), tramite un’accurata analisi morfometrica dei rispettivi olotipi unitamente ad altri esemplari provenienti da varie località fossilifere italiane. Bela pseudoappeliusi n. sp. condivide con B. (s.l.) appeliusi vari caratteri della teleoconca quali: dimensioni, tipo di ornamentazione, morfologia dell’apertura e seno esalante. Si distingue da B. (s.l.) appeliusi per la scultura della teleoconca più densa, la spalla non angolata e per le dimensioni e l’ornamentazione della protoconca. In ultimo è fornita una descrizione dettagliata della località tipo ed un elenco di altre località in cui è stata rinvenuta la nuova specie.

INTRODUCTION In this working classification the family Mangeliidae is characterized by small to medium shells (length < 30 mm), The family Mangeliidae was established by Fischer with prominent axial and/or spiral teleoconch sculpture; as the Mangiliinae in the second half of the 19th century the latter often being represented by spirally aligned (Fischer, 1883). However, its taxonomic placement has granules that are especially evident on the subsutural been highly debated and a general consensus regarding ramp. Furthermore, the anal sinus is subsutural, the the latter has only been agreed on recently (Taylor et shoulder is often angulated and the is usually al., 1993; Rosenberg, 1998 and others). Fischer (1883) multispiral with the nucleus being smooth or spirally lirate; included the Mangiliinae within the ; it was the remaining whorls often bear axial ribs sometimes subsequently assigned to the family (e.g., Hedley, overlapped by spiral elements. The consists of only 1922; Powell, 1966) and the Conidae (e.g., Taylor et al., marginal teeth, being variously shaped (from semi-enrolled 1993). Disagreement regarding the higher level taxonomic to true hypodermic; see Bouchet et al., 2011). classification persisted at least until the beginning of the Among the more than fifty extant genera of this family 21st century and may have been due to the differential the Bela Gray, 1847 is one of the most emblematic emphasis that taxonomists placed on the variety of of the ‘troublesome’ Mangeliidae. Bela was established by distinguishing characters used for systematic description Gray (1847) based on nebula Montagu, 1803. The (teleoconch-protoconch features, radula morphology, etc.). precise identification of the latter is unclear and a more In addition, factors related to convergence and homoplasy detailed analysis of this taxon is deemed necessary in order have rendered shell characters a much less reliable tool in to avoid erroneous assignments to the genus (Mariottini conoidean than was formerly assumed (e.g., et al., 2009). Indeed, since its establishment, Bela has Bouchet & Sysoev, 2001). This has led to contradictory often been used as a ‘wastebasket taxon’ for a vast array classifications and unsatisfactory consensus among the of unrelated eastern Atlantic-Mediterranean Cenozoic scientific community (Bouchet & Rocroi, 2005; Garilli & s.l. (see Powell, 1966). Hence, the numerous fossil Galletti, 2007; Figueira & Absalão, 2010 among others). and Recent species assigned to Bela require thorough A new and hopefully more stable classification of the revision. Among the hundreds of species documented as Conoidea has been outlined (Bouchet et al., 2011) based belonging to Bela (see Tucker, 2004), the World Register on recent molecular phylogenetic analyses (Puillandre et of Marine Species (http://www.marinespecies.org), al., 2008, 2011) and their relatively high congruence with estimated that approximately 28 can be stated as being a comprehensive anatomical classification of hard (e.g., Recent representatives, and Tucker (2004) reported only protoconch), and soft tissues (see Taylor et al., 1993). 44 as being valid Cenozoic fossil species.

ISSN 0375-7633 doi:10.4435/BSPI.2013.1 72 Bollettino della Società Paleontologica Italiana, 52 (2), 2013

In the present study research has been focused on a definition of Bela, the authors follow Mariottini et al. number of shells that preserve an original combination (2009) and include only species morphologically similar of features that, as far as the authors are aware, have to Bela zonata (Locard, 1891) within the genus. As for not been recognized to date in other mangeliids. Bela other representatives currently placed within this genus pseudoappeliusi is described herein as a new species for (such as Bela appeliusi), attribution to Bela (s.l.) pending the Plio-Pleistocene of Italy and is compared with the future revision has been adopted herein. closely related mangeliid, Bela (s.l.) appeliusi (Bellardi). This paper forms part of a series of contributions on the taxonomy and biodiversity of fossil conoideans from the Bela pseudoappeliusi n. sp. Plio-Pleistocene of the Mediterranean Basin (Scarponi & (Fig. 1a-c; Fig. 2a-c; Appendix I) Della Bella, 2004, 2010; Ceregato et al., 2006; Della Bella & Scarponi, 2007; Garilli & Galletti, 2007; Mariottini 1997 appeliusi Bellardi - Chirli, p. 62, Pl. 17, fig. 17; Pl. et al., 2009; Spada & Della Bella, 2010; Scarponi et al., 18, figs 1-2. 2011a, b among others). Etymology - After the Greek pseudo (false) and the word appeliusi (dedicated to Appelius), referring to the MATERIALS AND METHODS comparative species Bela (s.l.) appeliusi (Bellardi, 1877).

The material examined in the present study consists Type material - In total 11 specimens: Holotype of twenty specimens that have been ascertained herein as (MGGC-23470), and four paratypes (MGGC-23471 to belonging to Bela (s.l.) appeliusi and forty-three specimens -23474), from the Terre Rosse locality (Siena Basin; that have been attributed to Bela pseudoappeliusi n. sp. Pliocene sands of the S. Vivaldo Formation); three In order to better evaluate the variability of the latter and paratypes (MGGC-23475 to -23577), from Montelibretti to compare quantitatively the morphologic variations (Roma); two paratypes (MGGC-23478 to -23579), from between the two taxa, a batch of eleven specimens for Morrona (Terricciola, Pisa); one paratype (MGGC- each species was retained sufficient. Hence, a range of 23480), from Montopoli in Val d’Arno (Pisa). shell characters were carefully measured both on the holotype as well as on ten randomly selected shells for Additional material - Thirty-two specimens housed each species (see Appendixes I-II and Tab. 1). However, within the collections of the MGGC (inventory number it should be noted that both juvenile shells (teleoconch < MGGC-23482), even though attributed to the new species, 2.5 whorls) and excessively damaged shells were excluded are not considered as belonging to the type series (see (i.e., seven and 20 respectively) from the original batch provision 72.4.6 of International Code of Zoological of specimens prior to the selection process (see above). Nomenclature, 1999). The material includes: three shells Consequently, the description of the new species is based from Villalvernia (Alessandria); one shell from Vignola on five shells collected from the Terre Rosse outcrop (Modena); 15 shells from Terre Rosse (Siena); eight shells (Siena Basin, Italy) and six shells from localities in from Montelibretti (Roma), two shells from Morrona and Latium and Tuscany (Appendix I). All the studied material three shells from San Pietro quarry (Siena). is housed in the Museo Geologico Giovanni Capellini (MGGC) in Bologna, except for the holotype of B. (s.l.) Type locality - Terre Rosse (lat. 43°19’51’’N; long. appeliusi, housed in the Bellardi-Sacco (BS) collection in 11°35’11’’E, geographic coordinate on the World Geodetic the Museo di Scienze Naturali (Torino, Italy). System of 1984 [WGS84]), Castelnuovo Berardenga (Siena, Italy). Type material described herein is from the uppermost two meters of a 5.5 m thick, richly fossiliferous, SYSTEMATICS coarse to medium sand body (see below).

Class Gastropoda Cuvier, 1795 Diagnosis - Shell small-sized, protoconch of 3.1 Order Neogastropoda Thiele, 1929 whorls (on average), nucleus spirally lirate and immersed, Superfamily Conoidea Fleming, 1822 last protoconch with low granulose axial riblets on Family Mangeliidae Fischer, 1883 the final 0.5 portion, axial sculpture strong, consisting of suture-to-suture rounded ribs, spiral sculpture of thin, Genus Bela Gray, 1847 heterogeneous, granulose threads, rounded shoulder, anal sinus rounded, subsutural. Type species - ?Murex nebula Montagu, 1803. Description - Shell small (average L = 4.5 mm, σ = 0.8 Remarks - According to Mariottini et al. (2009), mm; D = 2.1 mm, σ = 0.4 mm) (L = length; D = diameter; σ as identification of the type species of this genus is = standard deviation; Tab. 1), biconical, turriculate, solid, still uncertain, a neotype should be proposed in order with conical and rounded body whorl. Protoconch to stabilize the genus name usage. Furthermore, the (Fig. 1a-c), dome-shaped of - on average - 3.1 convex opinion expressed by Mariottini et al. (2009) regarding whorls (average: L = 0.63 mm, σ = 0.06 mm; D = 0.65 the doubtful generic attribution to Bela of Pleurotoma mm, σ = 0.05 mm; Tab. 1). Apical whorl small, rather brachystomum Philippi, 1844 and its related species is immersed, spirally lirate, otherwise smooth except for the supported by the authors herein. Meanwhile, pending final half, where low and evenly curved axial riblets are molecular analysis in order to arrive at a better generic overrun by 3-4 rows of globose tubercles. Protoconch- F. Naldi et alii - Plio-Pleistocene Conoidea: Bela pseudoappeliusi n. sp. 73

σ = 0.8 mm; Tab. 1), running from suture to suture and thinner at the adapical suture. Spiral sculpture overrides the axial ribs, consisting on the subsutural ramp of several thin and uniform threads separated by incised lines; on the remaining whorl threads are thicker and irregularly alternated by thinner elements; interspaces of variable size. Each thread is densely indented by growing stage lirae and, especially on the subsutural ramp, it resembles a string of densely packed granules. Aperture rather narrow, elongated, sub-rectangular; very short, rather broad, left-deviated (aperture-canal average length = 2.0 mm, σ = 0.2 mm; Tab. 1). Outer without , thin, smooth within, rounded in profile; anal sinus subsutural, relatively broad and rounded. Inner lip straight with a thin parietal callus.

Remarks - The new species (Figs 1-2; Tab. 1 and Appendix I), shows a combination of features (i.e., rounded shoulder, marginated sutures, protoconch sculptural pattern, morphology of the aperture and sinus), suggesting assignment to the genus Bela Gray, 1847 and, more specifically, to theB. zonata group. Bela pseudoappeliusi n. sp. shows close affinities to the Mediterranean Pliocene mangeliid Bela (s.l.) appeliusi (Bellardi, 1877) in particular with regard to shell dimension and teleoconch sculpture (i.e., evident axial ribs overrun by densely indented spirals) (Fig. 2). In addition, certain features of B. (s.l.) appeliusi such as the sub-rectangular aperture, the short siphonal canal as well as the rounded, subsutural anal sinus show great similarity to those of B. pseudoappeliusi. Indeed, no significant (at this sample size) difference in the average value of a variety of teleoconch features (e.g., teleoconch length, diameter; see t-test in Tab. 1) was highlighted by basic morphometric analyses of selected specimens (Appendix I). However, when compared to B. pseudoappeliusi, it may be noted that Bela (s.l.) appeliusi shows flatter and shorter spire whorls with a strongly angulated shoulder as well as thinner and more numerous axial ribs that are overrun by less dense spiral threads (Fig. 2 and Tab. 1). Furthermore, the protoconch of B. (s.l.) appeliusi is notably larger in dimension, with a significantly higher protoconch embryonic whorl length/number ratio (Tab. 1). Lastly, the protoconch sculpture of B. (s.l.) appeliusi Fig. 1 - Scanning electron microscope photos of protoconch is clearly reticulate, being composed of marked axial and features. Bela pseudoappeliusi n. sp.: a) MGGC-23477-paratype, spiral elements (Fig. 1d-f). Montelibretti (Roma), Calabrian(?); b) MGGC-23482-spec. 1, In summary the new species differs from B. (s.l.) Villalvernia (Alessandria), Zanclean-Piacenzian; c) MGGC- 23482-spec. 2, Montopoli in Val d’Arno (Pisa), Piacenzian. Bela appeliusi in having a strongly convex whorl outline, a (s.l.) appeliusi (Bellardi): d-e) MGGC-23493 and -23494, Marano rounded shoulder, thicker and less numerous axial ribs, sul Panaro (Modena), Piacenzian; f) MGGC-23495, Monte Alto a shorter and more convex subsutural ramp, a denser (Piacenza), Piacenzian. Scale bars 0.1 mm; MGGC-23477 = shell spiral sculpture and a lower teleoconch whorl length/ number given in Appendix I; spec. = specimen. number ratio (see Fig. 2 and Tab. 1). At protoconch level, the main differences reside in the smaller dimension of B. pseudoappeliusi (Tab. 1) and in the sculpture teleoconch junction marked. Teleoconch (Fig. 2a-c) of the ultimate protoconch whorl (Fig. 1). Indeed, averaging 3.8 whorls (σ = 0.6; Tab. 1) with strong axial ornamentation in B. pseudoappeliusi is restricted to the and spiral sculptures. Spiral whorl convex, with rounded ultimate half whorl of the protoconch and is characterized shoulder and separated by undulating and marginated by granulose axial riblets. In contrast, ornamentation suture. Body whorl, relatively tall (average 2.9 mm, σ in B. (s.l.) appeliusi is reticulated, present all along the = 0.4 mm), about 2/3 of the shell length (Tab. 1). Axial ultimate whorl and shows more numerous and stronger sculpture of rounded, thick ribs (c. 9 on the second whorl, spiral elements. 74 Bollettino della Società Paleontologica Italiana, 52 (2), 2013

STRATIGRAPHICAL INFORMATION RELATED TO (alluvial to lacustrine) deposits have a scattered THE TYPE SERIES geographical distribution and depositional sequences are complex as a result of the intra-Messinian regional The greater part of the type series material (i.e., unconformity (Costantini et al., 2009). Conversely, holotype and four paratypes) was collected from a small Zanclean to Piacenzian successions outcrop extensively outcrop named Terre Rosse, located near Castelnuovo and are a few hundred meters in thickness. They overlay Berardenga (Siena Basin, Italy). both the upper Miocene deposits and the Ligurian units. The upper Miocene (Tortonian) to upper Pliocene Marine sedimentation persisted until the Piacenzian, when (Piacenzian) deposits of the Siena basin are among the a generalized uplift led to emersion of the area (see Bossio best-preserved of the richly fossiliferous successions et al., 1993 and references therein). Unfortunately, there occurring in Tuscany (Manganelli et al., 2010 and are relatively few detailed sedimentological studies of the references therein). However, Miocene continental area. According to Aldinucci et al. (2007) and Boscaini

Fig. 2 - Optical microscope photos of shell features. Bela pseudoappeliusi n. sp.: a) MGGC-23470-holotype, Terre Rosse (Siena), Zanclean- Piacenzian; b) MGGC-23480-paratype, Montopoli in Val d’Arno (Pisa), Piacenzian; c) MGGC-23475-paratype, Montelibretti (Roma), Calabrian(?). Bela (s.l.) appeliusi (Bellardi): d) BS.011.19.025-holotype, Colli Astesi (Asti), Piacenzian-Gelasian; e) MGGC-23491 Castell’Arquato (Piacenza), Piacenzian; f) MGGC-23487, Marano sul Panaro (Modena), Piacenzian. Scale bars 1mm; MGGC-23487 = shell number given in Appendix I. F. Naldi et alii - Plio-Pleistocene Conoidea: Bela pseudoappeliusi n. sp. 75

Fig. 3 - The Terre Rosse section showing stratigraphic information related to the sampled deposits and the distribution of Bela pseudoappeliusi n. sp. as well as other relevant mollusc species. Abbreviations: TAD = terraced alluvial deposits; Bk = bulk sample; MPMU = Mediterranean Pliocene Molluscan Unit.

(2011), the Pliocene deposits in the area of Castelnuovo and have been referred to the uppermost depositional unit Berardenga could be subdivided into two depositional of the latter by Boscaini (2011). units. The first unit is considered as being Zanclean in The section is ~ 9 m thick and consists of a variety age and is mainly represented in the study area by coarse of lithofacies that can be grouped into two main units of fluvio/deltaic deposits with subordinate sands. The second contrasting lithologies (Fig. 3). The lower to middle-upper unit is interpreted as being late Zanclean to Piacenzian in marine deposits of the section are richly fossiliferous and age and is represented mainly by massive to laminated are attributed to the San Vivaldo Formation (Fig. 3). The sands with minor tabular pebbly gravels that were succession includes 5.5 m of massive, fining-upward deposited in coastal to shallow marine settings. sandstone capped by 0.5 m of homogeneous silty-clays. The Terre Rosse section (Fig. 3) occurs along the bank The latter are in turn overlain by poorly laminated siltstone of a gully amid thick vegetation and the sedimentary to massive sandstone (about 1 m thick). The uppermost deposits crop out for some kilometers. According to the part of the section is characterized by barren alternations Geological Map of Italy (Costantini et al., 2009), the Terre of centimeter to decimeter tabular pebbles and plane- Rosse deposits form part of the San Vivaldo Formation paralleled bedded sands and clays that unconformably 76 Bollettino della Società Paleontologica Italiana, 52 (2), 2013

Teleoconch Protoconch Shell

length diameter # whorls on 2° whorl # L/# whorls L/D # of whorls length diameter L/# L/D length whorls

(mm) max aperture body whorl ribs threads

Bela pseudoappeliusi (n=11 specimens)

Mean 3.9 2.0 2.9 2.1 3.8 8.7 7.8 1.0 1.9 3.1 0.63 0.65 0.21 0.98 4.5

St. dev. 0.8 0.2 0.4 0.4 0.6 0.8 1.2 0.1 0.2 0.1 0.06 0.05 0.02 0.06 0.8

Mode 3.8 2.0 2.8 2.8 3.8 9.0 7.0 1.0 1.9 3.0 0.61 0.66 0.20 0.92 4.4

Max 5.8 2.6 3.8 2.8 4.9 10.0 10.0 1.2 2.1 3.2 0.74 0.71 0.24 1.07 6.4

Min 2.5 1.5 2.2 1.4 2.7 7.0 6.0 0.9 1.5 2.9 0.54 0.59 0.18 0.92 3.2

Bela (s.l.) appeliusi (n=11 specimens)

Mean 4.0 2.2 3.2 2.2 3.4 9.7 6.3 1.2 1.7 3.3 0.77 0.75 0.24 0.98 4.8

St. dev. 0.8 0.3 0.5 0.5 0.5 0.7 0.9 0.1 0.2 0.2 0.08 0.06 0.01 0.07 0.8

Mode 4.7 2.2 3.4 2.1 3.8 10.0 6.0 1.2 1.5 3.4 0.84 0.71 0.24 0.99 /

Max 4.8 2.6 3.8 3.1 4.0 11.0 8.0 1.4 2.0 3.5 0.89 0.84 0.26 1.15 5.7

Min 2.7 1.7 2.4 1.7 2.6 9.0 5.0 1.0 1.5 2.8 0.66 0.66 0.22 0.90 3.4

F test (p) 0.919 0.844 0.787 0.801 0.705 0.636 0.337 0.181 0.213 0.012 0.362 0.554 0.979 / 0.882

t test (p) 0.776 0.108 0.115 0.640 0.0946 0.005 0.007 <0.001 0.055 0.011* <0.001 <0.001 0.001 / 0.402

Tab. 1- Measures of central tendency and dispersion based on quantitative evaluation of fifteen shell parameters measured in eleven specimens for each species (see Appendix I). F test (p) indicates the probability that the two samples come from populations with equal variance; t test (p) indicates the probability that the two samples come from populations with equal mean. In bold, test results where the null hypothesis (given alpha < 0.05), can be rejected. Whenever the F test highlights unequal variance, the Welch test is performed instead of the classical t test (see asterisk). Abbreviations: D = diameter; p = probability of rejecting the null hypothesis; L = length; # = number; / = test not computed. overlay the marine deposits of the San Vivaldo Formation. 3. San Pietro quarry, Montopoli in Val d’Arno (Pisa), This uppermost part of the succession is interpreted as clay deposits attributed to the Piacenzian (lat. 43°37′24″N; being terraced alluvial deposits of Pleistocene-Recent age long. 10°52′54″E, WGS84). (i.e., TAD in Fig. 3). The holotype and four paratypes were collected from In the outcrops above, specimens of the newly described a level approximately 1 meter below the top of the thick species were found in association with infralittoral to fossiliferous sandstone (bulk sample 4 in Fig. 3). The use circalittoral molluscan assemblages. Unfortunately, a of traditional biomarkers to constrain the stratigraphic unit more detailed reference for the stratigraphic position of geochronologically proved unsuccessful due to the nature the occurrences and data related to paleoecology is not of the sediments (i.e., medium to coarse sands). However, available. the type material was found in association with several infralittoral molluscan species with subtropical affinities which have proved useful for dating the horizon (Fig. 3). ACKNOWLEDGEMENTS The concomitant occurrence of the following key species: Persististrombus coronatus (Defrance, 1827), Megaxinus This study was supported by funds from Ricerca Fondamentale ellipticus (Borson, 1825) and Callista italica (Defrance, Orientata 2011: project leader D. Scarponi. We acknowledge with 1818) clearly brackets this thick arenaceous deposit within thanks reviews by Bernard Landau and Marco Oliverio that have the Mediterranean Pliocene Molluscan Unit 1 (MPMU, notably improved the quality and clarity of this paper. We would like as defined in Raffi & Monegatti, 1993). The time-span of to thank Daniele Ormezzano (Museo Regionale di Scienze Naturali, the latter is comprised between the base of the Pliocene Torino) for help with type material under his care. and around 3.1Ma (Monegatti & Raffi, 2001), suggesting a Zanclean to early Piacenzian age. Integration with the above-mentioned stratigraphic scheme (see Boscaini, REFERENCES 2011), may allow a late Zanclean to early Piacenzian age to be assigned to the lowermost 5.5 meters of the Terre Aldinucci M., Ghinassi M. & Sandrelli F. (2007). Climatic and Rosse section (Fig. 3). tectonic signature in the fluvial infill of a late Pliocene valley The localities and horizons of the remaining type series (Siena basin, northern Apennines, Italy). Journal of Sedimentary specimens are as follows (Appendix I): Research, 77: 398-414. Bellardi L. (1877). I molluschi dei terreni terziarii del Piemonte e della Liguria. Parte II. Memorie della Reale Accademia delle 1. Montelibretti (Roma), silty clay deposits (lat. Scienze di Torino, serie 2, 29: 1-373. 42°08′0″N; long. 12°44′0″E, WGS84), attributed with Borson S. (1825). Continuazione del Saggio di orittografia doubt to the Calabrian. Piemontese. Memorie della Reale Accademia delle Scienze di 2. Morrona, Terricciola (Pisa), silty sand deposits Torino, serie 1, 29: 251-318. attributed to the Piacenzian (lat. 43°31′48″N; long. Boscaini N. (2011). I depositi plio-peistocenici di valle incisa del 10°39′49″E, WGS84). torrente Ambra (Toscana, Italia): interazioni tra tettonica e F. Naldi et alii - Plio-Pleistocene Conoidea: Bela pseudoappeliusi n. sp. 77

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Bollettino della Società Paleontologica Italiana, Revised manuscript accepted 19 March 2013 49 (3): 195-202. Zoobank registration number urn:lsid:zoobank.org:pub:46801EFB- Mariottini P., Smriglio C., Di Giulio A., & Oliverio M. (2009). A F4EB-47BF-86C3-C8878C745E32 new fossil conoidean from the Pliocene of Italy with comments Zoobank registration date 30 April 2013 on the Bela menkhorsti complex (Gastropoda: Conidae). Journal Published online 7 June 2013 of Conchology, 40: 5-14. Editor Fabio Massimo Petti 78 Bollettino della Società Paleontologica Italiana, 52 (2), 2013 Locality Morrona (Pisa) Morrona (Pisa) Colli Astesi (Asti) Terre Rosse (Siena) Terre Rosse (Siena) Terre Rosse (Siena) Terre Rosse (Siena) Terre Rosse (Siena) Terre Montelibretti (Roma) Montelibretti (Roma) Montelibretti (Roma) Monte Alto (Piacenza) Monte Alto (Piacenza) Monte Alto (Piacenza) Monte Alto (Piacenza) Monte Alto (Piacenza) Monte Alto (Piacenza) Marano sul P. (Modena) Marano sul P. (Modena) Marano sul P. (Modena) Marano sul P. (Modena) Marano sul P. San Pietro quarry (Pisa) 4.39 3.20 4.85 5.02 3.95 6.35 4.75 4.37 4.44 4.29 4.33 5.30 5.57 4.93 4.13 5.49 5.36 4.61 5.72 3.82 3.42 Shell (mm) >4.00 length (s.l.) ? D L / 0.9 B. 1.04 1.07 0.93 1.00 0.98 0.92 0.92 0.92 1.03 0.97 1.10 0.97 0.99 0.95 0.99 1.15 0.99 0.94 0.97 0.93 ? L / 0.21 0.24 0.21 0.22 0.20 0.18 0.19 0.20 0.20 0.22 0.20 0.24 0.22 0.25 0.24 0.22 0.24 0.26 0.25 0.23 0.24 # whorls n. sp. and 0.6 0.61 0.69 0.71 0.66 0.67 0.59 0.66 0.66 0.59 0.71 0.75 0.81 0.72 0.80 0.80 0.76 0.66 0.84 0.71 0.71 0.71 diameter Bela pseudoappeliusi 0.64 0.74 0.66 0.66 0.58 0.54 0.61 0.61 0.61 0.69 0.68 0.84 0.73 0.84 0.81 0.66 0.67 0.89 0.79 0.74 0.76 >0.61 length (mm, uncertainty ±0.02) Protoconch 3.0 3.1 3.1 3.0 2.9 3.0 3.2 3.0 3.1 3.1 3.4 3.5 3.4 3.4 3.4 3.0 2.8 3.4 3.2 3.2 3.2 >2.6 # whorls uncert. ±0.1 I ? ? ? ? D 1.9 L_t / 1.96 1.71 2.10 1.96 2.10 1.53 1.89 1.93 1.87 1.93 2.16 1.99 1.59 1.50 1.82 1.68 1.52 appeliusi ppe n dix (s.l.) A ? L_t / 1.00 0.91 1.05 1.08 1.01 1.18 0.96 0.99 1.02 0.93 1.03 1.10 1.24 1.17 1.18 1.23 1.17 1.16 1.38 1.03 1.02 Bela Bela pseudoappeliusi # whorls 7 8 8 9 7 9 7 6 7 ? 8 6 6 7 6 7 6 6 5 10 >4 >4 threads 9 9 9 9 8 9 7 9 9 8 9 9 9 9 9 on 2° whorl # 11 10 10 10 10 10 10 ribs 3.8 2.7 4.0 4.1 3.3 4.9 4.4 3.8 3.8 4.0 3.6 4.2 3.8 3.6 2.8 3.8 4.0 3.4 3.5 3.3 3.0 2.6 # whorls uncert. ±0.1 diameter; Hol. = holotype; L = length; Par. = paratype; # number; ? feature not quantifiable. = length; Par. diameter; Hol. = holotype; L 2.11 1.92 1.44 2.00 2.26 1.73 2.75 2.75 1.99 1.99 1.97 1.89 2.14 2.07 3.14 2.16 1.83 1.75 >2.40 >2.40 >2.30 >1.80 Teleoconch uncert. ±0.08 diameter (mm) (Bellardi, 1877). Ten randomly sampled shells as well as the holotype were examined for each species. Abbreviations: D = randomly sampled shells as well the holotype were examined for each species. Ten (Bellardi, 1877). 2.85 2.23 3.06 3.18 2.70 3.80 3.20 2.82 2.86 2.79 2.81 3.44 3.50 3.24 3.10 3.60 3.42 3.18 3.82 2.62 2.39 >2.50 body whorl appeliusi Quantitative evaluation of eleven shell features the twenty -two studied shells belonging to 2.11 1.99 1.53 2.23 1.96 2.55 2.15 1.97 2.02 1.92 1.99 2.42 2.41 2.31 2.23 2.65 2.39 2.23 2.31 1.83 1.75 >1.60 aperture ? 3.75 2.46 4.19 4.41 3.29 5.77 4.21 3.76 3.83 3.68 3.65 4.62 4.73 4.20 3.29 4.67 4.70 3.94 4.84 3.08 2.66 length (mm, uncertainty ±0.08) teleoc. Specimen Hol. MGGC-23470 MGGC-23471 Par. MGGC-23472 Par. MGGC-23473 Par. MGGC-23474 Par. MGGC-23475 Par. MGGC-23476 Par. MGGC-23477 Par. MGGC-23478 Par. MGGC-23479 Par. MGGC-23480 Par. Hol. BS.011.19.025 MGGC-23483 MGGC-23484 MGGC-23485 MGGC-23486 MGGC-23487 MGGC-23488 MGGC-23489 MGGC-23490 MGGC-23491 MGGC-23492 F. Naldi et alii - Plio-Pleistocene Conoidea: Bela pseudoappeliusi n. sp. 79

Appendix II Visual depiction of the morphological characters measured on the studied shells. Abbreviations: AL = aperture length; BWL = body whorl length; D = diameter; PD = protoconch diameter; SL = shell length; 2°wh = second whorl.