Gastropoda, Pulmonata, Gastrodontidae) from the Zanclean (Early Pliocene) of Tuscany (Central Italy
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Bollettino della Società Paleontologica Italiana, 50 (3), 2011, 165-173. Modena, 30 dicembre 2011165 A new Janulus species (Gastropoda, Pulmonata, Gastrodontidae) from the Zanclean (early Pliocene) of Tuscany (central Italy) Giuseppe MANGANELLI, IVAN MARTINI & Andrea BENOCCI G. Manganelli, Dipartimento di Scienze Ambientali, Università di Siena, Via Mattioli 4, I-53100 Siena; [email protected] I. Martini, Dipartimento di Scienze della Terra, Università di Siena, Via Laterina 8, I-53100 Siena; [email protected] A. Benocci, Dipartimento di Scienze Ambientali, Università di Siena, Via Mattioli 4, I-53100 Siena; [email protected] KEY WORDS - Gastropods, Pliocene, Zanclean, Tuscany, Janulus spadinii n. sp. ABSTRACT - Janulus spadinii n. sp. (Gastropoda, Pulmonata, Gastrodontidae) is described herein from the Zanclean (early Pliocene) of Balze di Caspreno (Tuscany). It belongs to a Macaronesian genus occasionally reported from the Tertiary of Europe. The new species is very similar to the Recent Madeiran Janulus stephanophorus by virtue of its shape, dorsal sculpture and radial rows of palatal teeth. It is also apparently similar to some fossil European species, however, due to insufficient information regarding the latter, no well supported distinction may be proposed at present. A brief survey of all the nominal taxa of the species-group assigned to Janulus is also provided. RIASSUNTO - [Una nuova specie di Janulus (Gastropoda, Pulmonata, Gastrodontidae) dal Pliocene inferiore della Toscana (Italia centrale)] - Janulus spadinii n. sp. è descritto sulla base di materiale raccolti nelle argille continentali del Pliocene inferiore delle Balze di Caspreno (Toscana). La nuova specie appartiene ad un genere attualmente esclusivo della Macaronesia, ma conosciuto per il Terziario europeo. Janulus spadinii n. sp. è molto simile a Janulus stephanophorus, una specie vivente a Madeira (le due specie condividono: forma, scultura dorsale e file radiali di denti palatali) ma ricorda anche alcune entità descritte per il Terziario europeo. Una differenziazione adeguata rispetto a questi taxa è al momento impossibile, mancando del tutto informazioni sulla struttura dei loro denti palatali. Conclude il lavoro l’elenco di tutti i taxa nominali assegnati a Janulus, inclusivo di una breve discussione sul loro status. INTRODUCTION lacking. Moreover, this duct may not be homologous: it is absent in Pilsbryna and connects penial dart-sac and duct The genus Janulus Lowe, 1852 is regarded as a of bursa copulatrix in Gastrodonta, penis and free oviduct European palaeoendemite of the Tertiary warm climatic in Janulus, and penis and duct of bursa copulatrix in all the phases, associated with moist forest biotopes (Esu, 1999). others (however, in Zonitoides, a second accessory duct It survives in the Macaronesian refuges (Walden, 1983, joins duct of bursa copulatrix and free oviduct according 1984; Cameron et al., 2007) and includes only three Recent to Pilsbry, 1946; Riedel, 1980; Gittenberger et al., 1984). species: J. bifrons (Lowe, 1831) and J. stephanophorus Consequently the monophyly of the family, as currently (Deshayes, 1850) from Madeira and the probably recently conceived, is not certain, like the number of genera/ extinct J. pompilius (Shuttleworth, 1852) from La Palma, subgenera it includes (see different accounts by Hausdorf, Canary Islands (Riedel, 1980; Fontaine et al., 2007). 1998; Riedel, 1998; Schileyko, 2003; Slapcinsky & Cole, Another Madeiran species, J. calathoides (Lowe, 1863), 2004). tentatively added by Riedel (1998), is actually a species Recent Janulus species have a discoidal to trochiform of Discus Fitzinger, 1833 (Bank et al., 2002). However, shell, angled or shouldered at periphery, with small there is no definitive evidence that the three species really umbilicus, narrow ribbing on dorsal surface of teleoconch belong to the same genus, because only one of them (J. whorls and peristome simple and not reflexed. One bifrons) is known anatomically. Janulus has a holopod species (J. bifrons) is larger, with less tightly coiled and foot and distal genitalia with a penial dart-sac and an less narrowly ribbed whorls and peristome internally accessory duct connecting penis and free oviduct (Pilsbry, thickened (it strongly recalls the shells of hygromiids 1947). Holopod snails with apertural palatal teeth, a such as Trochulus Chemnitz, 1786, Monachoides Gude penial dart-sac and an accessory duct connecting penial & Woodward, 1921, etc.). Finally, at least one species (J. complex and female genitalia (duct of bursa copulatrix or stephanophorus) has radial rows of palatal teeth at 120 free oviduct) are assigned to the gastrodontids, a mainly degrees to each other, each row consisting of three-four Nearctic family including a few other genera besides teeth: one adaxial, very small and knob-like, one medial, Janulus [Gastrodonta Albers, 1850, Nastia Riedel, 1989, short and pleat-like and one abaxial, larger and inverted Poecilozonites Boettger, 1884, Pilsbryna Baker, 1929, comma-like or sometimes V-like, split into a longer outer Striatura Morse, 1864 and Zonitoides Lehmann, 1862 branch and a shorter inner branch (Pl. 1, fig. 9). (Riedel, 1998; Slapcinsky & Cole, 2004)]. Although Although the fossil record is much poorer than it the accessory duct joining penial complex and female was formerly thought (see for example Wenz, 1923), genitalia has been regarded as a unique autapomorphy sure reports spanning from the Oligocene (e.g. Janulus supporting the monophyly of the gastrodontids (Hausdorf, schottleri Wenz, 1922) to the end of the Pliocene and 1998), a robust phylogeny of these land snails is still ecology of Recent species (Seddon, 2008) are in line with ISSN 0375-7633 doi:10.4435/BSPI.2011.15 166 Bollettino della Società Paleontologica Italiana, 50 (3), 2011 Fig. 1 - Location map of the study area. a) Structural setting of southern Tuscany. b) Geological sketch of the Montaperti-Pianella area. Numbers (1-6) indicate sites sampled during the 1990s. Most of these outcrops are now covered by recent alluvial deposits, vegetation or dumped rubble. the palaeoecological meaning attributed to the genus in part of an elongated depression (Fig. 1a) extending for the literature. about 200 km in a NNW-SSE direction. The tectonic In this paper we describe a new Janulus species from setting responsible for the origin and evolution of this the Zanclean of Balze di Caspreno (Tuscany), already basin is still debated, and has been explained in different reported as Janulus suttonensis (Wood, 1872) by De ways: according to many authors (Costantini et al., 1982; Stefani (1880). We also shortly survey all the nominal Martini & Sagri, 1993; Pascucci et al., 1999; Carmignani taxa of the species-group assigned to this genus, many of et al., 2001; Brogi, 2011) its origin is connected with an which may belong to other groups of land snails. extensional regime acting until the late Miocene, while other researchers postulate a dominantly compressional tectonic setting (Boccaletti & Sani, 1998; Finetti et al., GEOLOGICAL SETTING 2001; Bonini & Sani, 2002). Regardless of its structural origin, the Siena Basin The Siena Basin is a post-collisional basin of the inner was interested by a continental and marine deposition Northern Apennines, a collisional belt formed during during the Neogene, and to limited and discontinuous the Cenozoic in response to the interaction between the sedimentation of alluvial deposits (Aldinucci et al., 2007) Adria and the Corso-Sardinian microplates (Carmignani and travertine (Brogi et al., 2010) during the Quaternary. et al., 2001). It corresponds to approximately the middle Its Neogene basin-fill history dates back to the latest G. Manganelli et alii - A new species of Janulus from Zanclean of Tuscany 167 Miocene (Messinian) with the deposition of a thick Fm. exposed in the northern Casino Basin (Bossio et fluvio-lacustrine succession that unconformably overlies al., 1998). However, micropalaeontological analysis pre-Neogene bedrock. Most of these Miocene deposits carried out in fine floodplain sediments (Bambini, pers. are buried beneath Pliocene sediments and crop out in comm.) shows a lack of recycled Serravallian calcareous limited western marginal areas of the basin (Lazzarotto nannofossils, while bedrock-derived nannofossils are & Sandrelli, 1977; Bossio et al., 2000). Marine conditions common. These findings suggest that the deposits date in the Siena Basin began at the base of the Zanclean back to the Pliocene. According to Martini et al. (2011), (Costantini et al., 1982; Bossio et al., 1993), with the integrated palaeontological and stratigraphical analysis deposition of a thick sedimentary succession, ranging of the continental and marine succession exposed in the from nearshore (sands and conglomerates) to offshore surrounding area also enables assignment of these deposits (mud) environments (Costantini et al., 1982; Manganelli to the MPl3 Zone of planktonic foraminifera zonation et al., 2010; Martini et al., 2011). Marine sedimentation (zonal scheme of Foresi et al., 2001). persisted until the late Piacenzian, when uplifting of southern Tuscany caused emergence of the Siena Basin (Costantini et al., 1982; Bossio et al., 1993). Local SYSTEMATIC PALAEONTOLOGY episodes of continental sedimentation have been reported within the marine deposits in the southwestern sector of Janulus spadinii n. sp. the basin (Bossio et al., 1992, 1993). (Pl. 1, figs 2-7) Janulus material was collected along the Arbia River