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Upper Miocene molluscs of Monti Livornesi (Tuscany, Italy): Biotic changes across environmental gradients

Article in Palaeogeography Palaeoclimatology Palaeoecology · April 2019 DOI: 10.1016/j.palaeo.2019.04.024

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Upper Miocene molluscs of Monti Livornesi (Tuscany, Italy): Biotic changes across environmental gradients T ⁎ S. Dominicia, , M. Benvenutib, M. Forlic, C. Bogid, A. Guerrinie a Museo di Storia Naturale, Università di Firenze, Via La Pira 4, 50121 Firenze, Italy b Dipartimento di Scienze della Terra, Università di Firenze, Via La Pira 4, 50121 Firenze, Italy c Via Grocco 16, 59100 Prato, Italy d Via Gino Romiti 37, 57124 Livorno, Italy e Via Piave 28, 57123 Livorno, Italy

ARTICLE INFO ABSTRACT

Keywords: The upper Miocene mollusc assemblages of Monti Livornesi, used as a means to study the nature of Messinian salinity crisis Mediterranean benthic communities at the edge of the Messinian salinity crisis, are framed in a high-resolution Mediterranean benthos stratigraphic scheme and quantitatively approached by the study of historical museum collections and modern Facies bias samples. Instead of a single assemblage from either Tortonian or Messinian age, as previously thought, this fauna Sedimentary bias comes from three consecutive shallowing-upward depositional sequences bounded by regional unconformities, Shell bed taphonomy the Luppiano (upper Tortonian-lowermost Messinian), Rosignano and Raquese units (early Messinian). Facies Water depth gradient analysis, taphonomy and quantitative paleoecology show that the Luppiano assemblage is characterised by aragonitic species from a eutrophic brackish-water shallow marine muddy bottom, the Rosignano assemblage by calcitic species from an oligotrophic coarse-grained seafloor close to a coral reef and the Raquese assemblage from an open-marine muddy bottom. Published comparisons between Miocene and Pliocene faunal lists should be considered only crude estimates of faunal change until more is known about the distribution of species along paleoenvironmental gradients. The analysis of Miocene and Pliocene abundance data allows us to frame the Monti Livornesi molluscs along carbonate-siliciclastic and water depth gradients and to reach a better under- standing of the effects of salinity crisis on the Mediterranean biota.

1. Introduction et al., 2013), Piacenzian-Calabrian interval (Monegatti and Raffi, 2007), Calabrian-Upper Pleistocene interval (Garilli, 2011). Species The Mediterranean area includes most modern Global Boundary counts and comparisons between fossil assemblages collected in dif- Stratotype Sections and Points (GSSP) of the Neogene (Hilgen et al., ferent stratigraphic intervals would however prove elusive unless con- 2012). The abundance of shelly beds within thick shallow marine sidering incompleteness and bias of the sedimentary record: occurrence successions that cover times of crucial paleogeographic and climatic of fossils is erratic and controlled by facies and sedimentary successions changes shaping the modern biota, has long hinted at the use of fossil are riddled with discontinuities (Holland, 2000). molluscs as a stratigraphic proxy. A general decrease in diversity par- Several studies on global changes in biodiversity through geological allels the climatic deterioration registered during the upper Neogene time have attempted to correct for these biases (e.g., Crampton et al., and Quaternary, suggesting that many species went extinct or migrated 2003; Smith, 2007; Foote et al., 2015). Others have focused on en- (Monegatti and Raffi, 2001; Garilli, 2011). Attempts at using mollusc vironmental bias on the distribution of benthic species at a regional assemblages as chronostratigraphic tools have targeted all stages of the scale and during shorter time intervals (Dominici and Kowalke, 2007; upper Neogene and Quaternary. Taking into account some recent stu- Scarponi and Kowalewski, 2007; Zuschin et al., 2011), posing similar dies, these have involved faunas of the Tortonian (Lauriat-Rage et al., problems on a smaller scale, as in the case study presented here. Upper 1999; Satour et al., 2011), Messinian (Ben Moussa et al., 1987; Miocene molluscs collected during more than one century in the hills Monegatti and Raffi, 2010; Harzhauser et al., 2013), Zanclean (Lauriat- south of Livorno (Monti Livornesi, Tuscany) have been compared to Rage, 1981; Rouchy et al., 2003; Monegatti and Raffi, 2007; Satour younger and older fossil assemblages to understand the effects of the

⁎ Corresponding author. E-mail address: stefano.dominici@unifi.it (S. Dominici). https://doi.org/10.1016/j.palaeo.2019.04.024 Received 2 February 2019; Received in revised form 17 April 2019; Accepted 24 April 2019 Available online 30 April 2019 0031-0182/ © 2019 Elsevier B.V. All rights reserved. S. Dominici, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 527 (2019) 103–117

Messinian salinity crisis on the composition of the Mediterranean is made of poorly exposed greyish mudstone (subunit LUPb, Fig. 3 B) benthos (Monegatti and Raffi, 2010), but the facies distribution of with an abundant molluscan fauna, up to 30 and up to 10 m thick in the mollusc species richness and abundance and its relationship with stra- Quarata and Popogna areas, respectively (Fig. 2). tigraphic architecture has not been addressed yet. Moreover, the age of For its lithological features LUPb is equivalent to the Argille del the fauna has been the subject of controversy, some authors considering Torrente Fosci Formation, a muddy interval not recognized before in it Tortonian (Trentanove, 1901, 1911; Harzhauser et al., 2013) others the Livorno Hills (e.g., Polino et al., 2015), but mentioned at Quarata by Messinian (Ruggieri, 1956; Monegatti and Raffi, 2010). We have un- both Sacco (1895) and Trentanove (1911) for its peculiar mollusc as- dertaken a stratigraphic and sedimentary analysis of the succession and sociation. This facies can be also equated to the “dark grey mudstone a taphonomic and paleoecologic study of the fauna, reconsidering its with Ostrea lamellosa” described along the Morra creek, a few km age and the value of its comparison with younger and older Medi- northeast of the study area, and containing a distinct microfauna with terranean shell beds considering their distribution across paleoenvir- miliolids and fragments of echinoids (Giannini, 1960). onmental gradients. Our final aim is to understand how overall mollusc The above features, albeit observed for a limited extent, suggest a species richness and abundance are distributed among the different depositional setting characterised by torrential streams evolving into a facies, how this particular case relates to the general patterns of bio- subtidal, shallow marine environment, in an overall transgressive trend. diversity during the Neogene and Quaternary and what were the effects At Quarata the unconformable contact between LUPa and the bedrock of the Messinian salinity crisis on the marine biota. suggests deposition in a paleovalley sloping to NE in the Botro della Casina area, turning downstream toward SE, i.e., funnelled along the 2. Geological background axis of the syncline. Similarly, in the Popogna area the limited occur- rence of this unit toward the NW sector suggests an original confine- The upper Miocene continental, restricted and fully marine suc- ment of the depositional area in a valley opening eastward and occu- cession here under study is exposed in limited patches upon reliefs pying a structural low. Wedging of strata suggests syn-depositional made of a complex structural stack of strongly deformed Upper growth of the Popogna syncline. Cretaceous-Paleogene ocean-related limestone, mudstone and sand- stone, including blocks of Upper Jurassic ophiolites (Fig. 1A; Ligurian 2.1.2. Rosignano unit Units; Lazzarotto et al., 1990; Nirta et al., 2007). Following repeated Two lithological divisions also characterise the overlying Rosignano stages of folding and thrusting related to the Europe-Africa plates unit (ROS) in the two examined areas, resting through an high-relief convergence and collision, this tectono-sedimentary complex piled up erosional surface above unit LUP (Fig. 2), or directly on the bedrock. onto Upper Oligocene-Lower Miocene turbiditic sandstone, forms the These divisions coincide respectively with the Villa Mirabella Con- youngest unit in a sedimentary succession accumulated since the glomerates and the Castelnuovo Limestone of previous studies. The

Triassic upon the basement of Adria, a promontory of the Africa plate. basal deposits (subunit ROSa) show facies variation in the two areas. In The Upper Miocene of the Monti Livornesi, resting unconformably over the Quarata they are pebble-sized conglomerate, visible for few meters this bedrock complex (Bossio et al., 1999; Lazzarotto et al., 1990), around the Podere del Gorgo. Clasts are well-rounded, framed in a clast- occurs in the distinct areas of Quarata, Popogna and Poggio Cafaggio supported texture with abundant interstitial sandy matrix, within (Fig. 1A), as recognized by early authors (Capellini, 1878; Sacco, 1895; amalgamated massive or graded beds. In the Popogna area this subunit, Trentanove, 1901). These are narrow elongated depressions trending about 20 m thick, is exposed in the Popogna Creek, starting with a WNW-ESE, a direction that fits the orientation of the axes of folds in the breccia made of blocks of the adjacent Ligurid substratum followed Ligurian bedrock, resulting from a shortening pulse that occurred at the upwards by matrix-rich massive cobbly-pebbly conglomerates (Fig. 3 Eocene-Oligocene transition (D3 stage in Nirta et al., 2007). The B–C). These are arranged in decimeter-thick beds: clasts are sub-angular synclinal attitude of the Upper Miocene successions in the three areas forming a framework characterised by abundant muddy interstitial outlined in existing maps (Malatesta, 1954; Lazzarotto et al., 1990; matrix and containing fragments of the molluscs associated with the

Polino et al., 2015) is coherent with these folds in the bedrock, con- LUPb mudstone (Fig. 4 A). Upward, the gravelly facies is similar to that sistent with a reactivation of compression during their deposition. observed at Quarata, including also channelized pebbly sandstones These successions, some tens of meters thick, are in large part lithos- suggesting confined paleoflows directed to the ENE. A peculiar feature, tratigraphically equivalent to the thicker Upper Miocene infill of the not observed at Quarata, is the change of the inclination of bedding, adjacent wider Fine Basin (Bossio et al., 1999), separated by a threshold dipping to ESE of about 45° in the lower, coarser strata to about 30° in of bedrock. The successions outcropping at Quarata and Popogna, the upper strata (Fig. 3 B). where some of the shell beds were newly sampled, are discussed for The upper portion (subunit ROSb) marks a drastic lithological their depositional origin and correlation with the regional Late Miocene change from terrigenous to hybrid‑carbonate deposition, being re- stratigraphy. Three main unconformity-bounded units are recognized in presented by biohermal limestone, marl and shelly sandstone, the latter both areas (Fig. 1 B–C), matching main litostratigraphic units re- particularly exposed in the Popogna area (Figs. 2; 3 C–D). Subunit ROSb cognized by Trentanove (1901, 1911). is associated with a rich fossil content characterised by the hermatipic corals Siderastrea crenulata and Porites lobatosepta (Trentanove, 1911; 2.1. Facies succession and stratigraphic correlation Bossio et al., 1996), bryozoans and red algae. In the sandstone facies, molluscs and echinoderms are found in life position (Fig. 4 B–C; see also 2.1.1. Luppiano Unit Bossio et al., 1996). Corals and bryozoans are also reported by authors The lowermost unit of the upper Miocene succession, called who have described this unit elsewhere (Giannini, 1960; Esteban, 1979;

Luppiano Unit (LUP), includes two lithological portions stacked in a Bossio et al., 1996; Polino et al., 2015). ROSb is topped by a mono- fining-upward succession (Fig. 2). The lower portion is formed by specific Crassostrea gryphoides shell bed (Trentanove, 1911). reddish cobble-pebble sized massive conglomerate alternating with Unit ROS developed after the deep incision of the Luppiano Unit, sandstone and mudstone (subunit LUPa). Albeit scanty, the above ob- thus following a regressive stage. The early deposition occurred in a servable features allow us to identify it with the Conglomerati di Podere fluvio-deltaic setting where ROSa accumulated as the infill of a valleys Luppiano Formation (Lazzarotto et al., 2002; Polino et al., 2015). These generally trending to the ESE. At Quarata, this conclusion is supported deposits rest with angular unconformity on the bedrock, ranging in by the overall geometry of ROSa as resulting from a revision of existing thickness from about 25 m south of the Quarata creek (Fig. 1 B) to 4 m maps, showing a NE-dipping contact on LUPb to the south and a SW- at the north-western end of the Popogna area, where the poorly-bedded dipping contact on the bedrock to the north, again outlining the conglomerate show wedging of strata (Figs. 1 C; 3 A). The upper portion synclinal shape of the succession. The wedging of ROSa beds

104 .Dmnc,e al. et Dominici, S. 105 Palaeogeography, Palaeoclimatology,Palaeoecology527(2019)103–117

Fig. 1. Geology of the upper Miocene of the Monti Livornesi. A) Schematic structural map of the Monti Livornesi: Q: Quarata area; C: Poggio Castello area; P: Popogna area; FB: Fine Basin; TMS: Torrente Morra section; Q: area detailed in B); P: area detailed in C. B) Geological map of the Quarata area, with position of three sampled mollusc assemblages and lateral cross section displaying the syncline structure. C) Geological map of the Popogna area, with position of two sampled mollusc assemblages and lateral cross section displaying the syncline structure (geological maps revised after Polino et al., 2015). S. Dominici, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 527 (2019) 103–117

Fig. 2. Stratigraphic scheme of the Monti Livornesi upper Miocene in the Quarata and Popogna areas. documented in the Popogna outcrop (Fig. 3 B) testifies to a continued western end of the Popogna area. An almost continuous outcrop along syn-depositional deformation related to pulses of crustal shortening. the creek offers a natural section across the Popogna syncline, showing

This limited outcrop suggests a depositional transition from a slope the RAQb for a partial thickness of about 30 m. At least four lithological setting dominated by denudation of the bedrock and of the overlying intervals have been observed (Figs. 2; 5): the lowermost deposits occur

LUPb, mudstone, recorded by the basal matrix-rich conglomerates, into on the SW limb of the syncline and consist of massive greyish mud- a torrential stream flowing to the ENE. stones resting on top of the RAQa calcarenites. The mollusc association Unit ROSb attests to a significant reduction of clastic supply under is characterised by abundant specimens of Turritella tricarinata asso- raised relative sea level, promoting carbonate production and favouring ciated with Tritia semistriata, Fissidentalium inaequale, Aporrhais uttin- the development of patch reefs in nearby waters. The overall macro- geriana. Abundant specimens of Microloripes dentatus and Corbula gibba palentological association indicates that ROSb was deposited in a are found articulated, in loosely-packed shell beds (Fig. 4 D). All shells marginal-marine environment (see also Bossio et al., 1996). Particularly are aragonitic and the overall taxonomic composition suggests an open- in the Popogna area, the development of limited carbonate factories is marine muddy paleobottom. These deposits pass upward to greyish testified by the shelly debris reworked in a reef-slope setting. mudstones and siltstones with occasional cm-thick beds of graded and laminated sandstones (interval 2 in Figs. 2; 5 A–B). Mollusc remains are almost absent, with the exception of a thin bed containing shell frag- 2.1.3. Raquese Unit ments. The overlying interval 3, about 1 m-thick, includes thin beds of The third and uppermost Raquese Unit (RAQ), also comprising two siliceous mudstone passing upward to mudstones and siltstone of fourth distinct lithologies, onlaps onto the erosional surface on ROS deposits b and uppermost interval (intervals 3 and 4 in Figs. 2; 5 C–D). At the NE (Fig. 2). Unit RAQ is equivalent to the Formazione del Torrente Raquese limb of the syncline the basal mollusc-bearing mudstones are missing or “Pycnodonta Clays” of previous authors (Mazzanti, 1993). The low- and interval 2 rests over a thin shelly sandstone sampled for its mollusc ermost portion observed only in the Quarata area is a thin conglomerate assemblage (BR in Fig. 1 C) and referred to subunit ROSb. level (subunit RAQ ), poorly exposed north of Podere del Gorgo. The a The depositional signature of RAQ suggests a renewed aggradation upper and thicker portion is made of fine-grained sediments (subunit in a transgressive trend, following a new relative sea-level fall. RAQa RAQ ), bearing a distinctive association of marine bivalves and gas- b deposits in the Quarata area mark deposition in a fluvio-deltaic system tropods (Fig. 4 D). These deposits are poorly exposed at Quarata, fed by catchments almost coinciding with those active during the early whereas they can be observed in the Botro Rosso creek, at the north-

106 S. Dominici, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 527 (2019) 103–117

Fig. 3. Outcrops of the LUP and ROS units in the Popogna area. A) recent roadworks on the right bank of the Botro Rosso Creek showing the LUP unit: pebbly conglomerates are poorly bedded, dipping to ENE with inclination decreasing up-

ward. LUPb mudstones bear the mollusc assem- blage recognized in MQLI1-2 samples of the Quarata area (hammer for scale: 35 cm); B) pa- noramic view of the ROS unit from the left bank of the Popogna Creek looking upstream: the

variation of bed tilting from the basal ROSa to the ROSb strata is outlined; C) detail of the

transition between the ROSa upper gravelly-

sandy strata and the ROSb calcarenites: D) detail of the lowermost shelly sandstone showing a graded structure, with a 20 cm-thick tightly- packed shell bed at the bottom.

deposition of the Rosignano Unit. A full transgression is recorded by productivity related to upwelling of nutrients or to river runoff and sub-unit RAQb, deposited in an inner shelf setting under a varying re- expansion of opal-rich terrestrial biomes (Pellegrino et al., 2018). On gime of sediment supply as indicated by the record in the Popogna area. the whole, the latter unit is equated to the Raquese Unit. The bios- Specifically, the setting recorded by the basal interval 1 points to a tratigraphic calibration of upper Miocene deposits in the Fine Basin muddy bottom populated by an open-marine benthic fauna (Fig. 4 D). allows to refer the ROS- and RAQ-equivalent intervals to the early The overlying interval 2 suggests deposition under an higher sediment Messinian (Bossio et al., 1997; Polino et al., 2015), whereas the basal supply transferred by muddy and subordinate sandy hyperpycnal flows LUP-equivalent deposits, in absence of direct biostratigraphic evidence, (Fig. 5 A-B). This condition possibly prevented bottom colonization by are tentatively ascribed to the late Tortonian, based on lithostrati- microfaunal benthos. The siliceous interval 3 (Figs. 2; 5 C–D) can be graphic correlation with other areas. The near-equivalency of the possibly related to an episode of high organic productivity that affected Quarata-Popogna succession with the upper Miocene of the Fine Basin the adjacent Fine Basin during the early Messinian (see below). The implies a depositional connection during the late Tortonian-early uppermost interval 4 outcropping in the Popogna area (Figs. 2; 5 C) Messinian, expressed in LUPa time by a proximal fluvial catchment in indicates a return to depositional conditions similar to those of interval the Quarata-Popogna areas pointing toward the western margin of the

2. The RAQb outcrop in the Botro Rosso documents a possible north- Fine basin. During a relative lowstand of sea-level, fluvial valleys in- ward shift of the depocenter as a consequence of the syn-depositional cised in the Ligurian bedrock fed deltaic systems in the basin. A growth of the Popogna syncline. transgressive event in the inner portions of the catchment, funnelled by the drainage network, is attested to by the establishment of brackish- 2.2. Upper Miocene depositional dynamics and stratigraphic correlation water conditions, recorded by LUPb (Fig. 2). Lithological and deposi- tional analogies with the upper Miocene of the eastern Volterra Basin, The stratigraphic revision provides arguments for a correlation with about 50 km ESE of the study area (Sandrelli, 1996; Ielpi and the upper Miocene of nearby basins, extending the significance of the Cornamusini, 2016), support the idea of a generalised transgression and local depositional history over the limits of the study areas. A correla- the ensuing of a highstand of sea-level. Mudstones resting on top of tion is done with the succession, about 350 m thick, of the adjacent Fine conglomerates record here an environmental transition from a shallow Basin exposed along the Morra Creek (Giannini, 1960; Bossio et al., lake to a lagoonal sedimentary environment, as evidenced by ostracod 1997; Sarti et al., 1998) about 6 km north-east of the study areas. Here, faunas, referred to the Tortonian-Messinian transition (Sandrelli, 1996). fluvial conglomerates, sandstone and mudstone, unconformably over Further east, in the Casino Basin (Bossio et al., 1993) and south, in the the Ligurian bedrock, lithologically equivalent to the Luppiano Unit, Albegna Basin (Bossio et al., 2003-2004), the upper part of the Argille are overlain by limestone correlated to the Rosignano unit (subunit del Fosci formation records a transition from fresh-water to brackish- water conditions around the Tortonian-Messinian boundary. This ROSb). The overlying deposits are a thick sequence of shelfal mudstone including an interval of laminitic diatomite, hinting at episodes of high widespread environmental signal attests to a transgressive trend

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Fig. 4. Shell beds of the Monti Livornesi. A: Dispersed bioclastic fabric in the LUPb mudstone in the Quarata area (sample MQ1); B: Shelly sandstone of the ROSb subunit, with a large bivalve in life position (sample BR in Fig. 1C); C: Regular echinoderm from the ROSb subunit in the Popogna area; D: Dispersed bioclastic fabric in the RAQb mudstone in the Popogna area, with many articulated specimens of Corbula gibba and a specimen Omniglypta jani in the upper left part. All scales in cm. analogous to what was recorded in the studied areas at the transition 3. Composition of the mollusc fauna between LUPa and LUPb. The molluscan fauna partly described by Trentanove (1911), and quantitatively approached here (samples MQ1 3.1. Previous studies and MQ2 in Figs. 1; 2), can be thus referred to a highstand systems tract (HTS) dated around the Tortonian-Messinian boundary, based on se- The dating of the upper Miocene molluscs collected in the Monti quence-stratigraphic correlation with the upper Miocene of other Livornese and their affinity with older and younger faunas have been a

Tuscan basins. A second sea-level lowstand is recorded by ROSa, fol- subject of debate since the nineteenth century, until very recently. lowed by a transgressive systems tract represented by ROSb. Highstand Capellini (1878) carried out a first study of the area around Popogna deposits can be represented by the sandier facies of ROSb, or it can be and Cafaggio, drawing a geological map and publishing a list of 19 missing due to subsequent erosion. The coral biostrome dominated by species, some of which were found within a coral biohermal limestone Siderastrea crenulata, passing laterally to a coarse-grained sandstone outcropping near the town of Rosignano. A thorough study of the Po- facies (Fig. 2) with a fauna dominated by large suspension-feeding bi- pogna and Cafaggio fossil molluscs was carried out by Trentanove valves, both epifaunal and infaunal, testifies to oligotrophic shallow (1901), building also on the preliminary notes published by Fuchs marine environment during deposition of the TST (Esteban, 1979; (1878) and Sacco (1895). After studying a collection hosted in Florence Bossio et al., 1996, 1998) and the Crassostrea gryphoides shell bed to the and fossils collected during his own field work, Trentanove (1901) maximum flooding surface. A third depositional sequence is re- expanded the list to 16 species for the “limestone” and 29 species for presented by the two parts of RAQ, respectively conglomerate of the the overlying mudstone, attributing both assemblages to the Tortonian

LST-FSST (RAQa) and mudstone of the TST-HST (RAQb) with its distinct by comparison with the better-known fossil molluscs found at Mon- assemblage dominated by suspension-feeders such as Turritella tricar- tegibbio, Stazzano and Sant'Agata, in the Northern Apennines. Ten inata and Corbula gibba, open marine scavengers, such as Tritia semi- years later, he completed the analysis with the description of fossils striata, and at places by a chemiosymbiotic, Microloripes dentatus fauna collected at Quarata, a few km from Popogna, where he also described a (Fig. 4 D). Decapods and infaunal echinoderms are also present (Fig. 2). third, brackish-water assemblage underlying the coral biohermal A marked syn-tectonic control on this depositional dynamic is tes- limestone (Trentanove, 1911), only briefly outlined in previous litera- tified by the funneling of the clastic supply along the axis of the Quarata ture (Sacco, 1895; Trentanove, 1901). The Popogna and Quarata spe- and Popogna synclines during the regressive stages, by wedging of cimens collected by Capellini and hosted at the University of Bologna strata recorded in the Popogna area and depocenter migration inferred were later studied by Ruggieri (1956), who largely criticised the taxo- in both areas. nomic work of Trentanove and reinterpreted all fossils of the Livorno hills as Messinian. Possibly because of the “very bad outcrop

108 S. Dominici, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 527 (2019) 103–117

Fig. 5. Outcrops of the RAQb sub-unit in the Botro Rosso Creek. A) Panoramic view of the northern limb of the Popogna syncline (the rod in the rectangle box is 1 m long); B) detailed view of the bed stacking in the lithological interval 2 of RAQb: the deposition is referred to hyperpycnal flows carrying muddy to sandy suspensions and reworking shell material (rod for scale: 50 cm); C) panoramic view of the southern limb of the Popogna syncline (rod for scale: 100 cm); D) detail of the sub- horizontal siliceous mudstones of the lithological interval 3) in the axial zone of the Popogna syncline (hammer for scale: 35 cm).

109 S. Dominici, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 527 (2019) 103–117

Table 1 Per-collection and per-sample abundance distributions of upper Miocene molluscan species. The first columns report abundance data for the historical material, divided among the three fossil assemblages discussed in the text based the lithologic descriptions given by Trentanove (1901, 1911; columns 1–3), with a minor number of occurrences that could not be independently assigned to an assemblage (column 4). Columns 5–10 report abundance data for novel samples, also subdivided among the three upper Miocene assemblages (see text for further information).

Historical collections LUPb ROSb RAQb

LUPhist ROShist RAQhist Uncertain MQ1 MQ2 BR TR1 TR3 TR5

Gibbula dertosulcata (Sacco, 1896) 0 0 0 0 28 34 0 0 0 0 Gibbula monterosatoi (Sacco, 1896) 0 0 0 0 0 79 0 0 0 0 Oxystele rotellaris (Michelotti, 1847) 0 0 0 3 0 0 3 0 0 0 Turritella tricarinata (Brocchi, 1814) 0 0 2568 0 0 0 0 21 16 104 Tenagodus anguinus (Linnaeus, 1758) 0 0 0 9 0 0 0 0 0 0 Thericium europaeum (Mayer, 1878) 3 0 0 0 0 1 0 0 0 0 Thericium italicum (Mayer, 1878) 552 0 0 0 15 260 0 0 0 0 Bittium deshayesi Cerulli Irelli, 1912 0 0 0 0 82 0 0 0 0 0 Granulolabium bicinctum (Brocchi, 1814) 74 0 0 0 12 29 0 0 0 0 Granulolabium tuberculiferum (Cocconi, 1873) 469 0 0 0 45 45 0 0 0 0 Epitonium sp. 0 0 2 0 0 0 0 0 0 2 Cochlis sp. 0 54 0 0 0 0 2 0 0 0 Alvania cioppii Chirli, 2006 0 0 0 0 1 0 0 0 0 0 Alvania granosa Tabanelli et al., 2011 0 0 0 0 3 0 0 0 0 0 Pusillina sp. 0 0 0 0 43 0 0 0 0 0 Setia turriculata Monterosato, 1884 0 0 0 0 13 0 0 0 0 0 Hydrobia frauenfeldi Hörnes, 1856 0 0 0 0 400 0 0 0 0 0 Stenothyrella schwartzi (Frauenfeld in Hörnes, 1856) 0 0 0 0 12 0 0 0 0 0 Tornus subcarinatus (Montagu, 1803) 0 0 0 0 11 0 0 0 0 0 Xenophora sp. 0 0 0 0 0 0 1 0 0 0 Aporrhais uttingeriana (Risso, 1826) 0 0 61 0 0 0 0 9 0 0 Amalda glandiformis Lamarck 1810 0 2 0 0 0 0 1 0 0 0 Streptochetus ornatus (d'Orbigny, 1852) 0 0 0 2 0 0 0 0 0 0 Heteropurpura dertonensis (Mayer in Bellardi, 1873) 8 0 0 0 4 20 0 0 0 0 Hexaplex austriacus (Tournoüer, 1875) 0 0 0 1 0 1 0 0 0 0 Tritia agatensis (Bellardi 1882) 24 0 0 0 10 36 0 0 0 0 Tritia brugnonis (Bellardi, 1872) 0 0 38 0 0 0 0 0 0 0 Tritia striatulus (Eichwald, 1829) 0 0 560 0 0 0 0 21 27 47 Tritia coloratus (Eichwald, 1830) 4 0 0 0 60 6 0 0 0 0 Tritia semistriata (Brocchi, 1814) 0 0 61 0 0 0 0 0 0 3 Clavatula coppii Bellardi, 1878 28 0 0 0 39 16 0 0 0 0 Clavatula mystica (Doderlein in Manzoni, 1869) 41 0 0 0 9 5 0 0 0 0 Clavatula sotterii (Michelotti, 1847) 3 0 0 0 23 31 0 0 0 0 Odostomia cfr. O. carrozzai van Aartsen, 1987 0 0 0 0 2 0 0 0 0 0 Nisosyrnola concava (Boettger, 1907) 0 0 0 0 18 0 0 0 0 0 Parthenina indistincta (Montagu, 1808) 0 0 0 0 2 0 0 0 0 0 Parthenina terebellum (Philippi, 1844) 0 0 0 0 1 0 0 0 0 0 Ebala pointeli (de Folin, 1868) 0 0 0 0 31 0 0 0 0 0 Henrya wareni Landau et al., 2013 0 0 0 0 2 0 0 0 0 0 Acteocina lajonkaireana (Basterot, 1825) 0 0 0 0 6 0 0 0 0 0 Retusa truncatula (Bruguière, 1792) 0 0 0 0 2 0 0 0 0 0 Haminoea sp. 0 0 0 0 0 0 0 0 0 0 Ringicula minor (Grateloup, 1838) 0 0 0 0 8 0 0 0 0 0 Nucula sp. 0 0 0 0 0 0 0 0 1 0 Lembulus pella (Linnaeus, 1758) 0 0 25 0 0 6 0 0 0 0 Yoldia nitida (Brocchi, 1814) 0 0 2 0 0 0 0 0 0 0 Yoldia philippi Bellardi, 1875 0 0 8 0 0 0 0 0 0 0 Anadara diluvii (Lamarck, 1805) 0 0 652 0 0 0 0 0 20 0 Anadara firmata (Mayer, 1868) 15 0 0 0 0 31 0 0 0 0 Anadara fichteli (Deshayes, 1850) 0 5 0 0 0 0 3 0 0 0 Limaria hians (Gmelin, 1790) 0 0 0 1 0 0 0 0 0 0 Gibbomodiola adriatica (Lamarck, 1819) 0 1 0 0 0 0 0 0 0 0 Pecten aduncus Eichwald, 1830 0 34 0 0 0 0 2 0 0 0 Talochlamys multistriata (Poli, 1795) 0 1 0 0 0 0 0 0 0 0 Aequipecten malvinae (Dubois de Montpéreux 1831) 0 2 0 0 0 0 0 0 0 0 Anomia ephippium Linnaeus, 1758 7 0 0 0 0 1 0 0 0 0 Podedesmus patelliformis (Linnaeus, 1758) 1 0 0 0 0 0 0 0 0 0 Ostrea edulis Linnaeus, 1758 8 0 0 0 0 0 0 0 0 0 Crassostrea gryphoides (Schloteim, 1813) 0 0 0 5 0 0 0 0 0 0 Neopycnodonte navicularis (Brocchi, 1814) 0 0 1 0 0 0 0 0 0 0 Centrocardita aculeata (Poli, 1795) 0 0 2 0 0 0 0 0 0 0 Chama gryphoides Linnaeus, 1758 19 0 0 0 0 13 0 0 0 0 Microloripes dentatus (Defrance, 1823) 685 0 684 0 100 232 0 0 27 0 Myrtea spinifera (Montagu, 1803) 0 0 12 0 0 0 0 0 0 0 Myrtina meneghinii (De Stefani & Pantanelli, 1888) 0 0 4 0 0 0 0 0 0 0 Hemilepton nitidum (Turton, 1822) 0 0 0 0 1 0 0 0 0 0 Lepton squamosum (Montagu, 1803) 0 0 0 0 1 0 0 0 0 0 Montacuta phascolionis Dautzenberg & H. Fischer, 1925 0 0 0 0 1 0 0 0 0 0 Cerastoderma glaucum (Bruguière, 1789) 138 0 0 0 0 62 0 0 0 0 (continued on next page)

110 S. Dominici, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 527 (2019) 103–117

Table 1 (continued)

Historical collections LUPb ROSb RAQb

LUPhist ROShist RAQhist Uncertain MQ1 MQ2 BR TR1 TR3 TR5

Procardium cfr. P. danubianum (Mayer, 1866) 0 1 0 0 0 0 0 0 0 0 Acanthocardia echinata (Linnaeus, 1758) 0 8 0 0 0 0 1 0 0 0 Laevicardium oblongum (Chemnitz, 1791) 0 2 0 0 0 0 0 0 0 0 Papillicardium papillosum (Poli, 1791) 0 0 2 0 0 0 0 0 0 0 Parvicardium minimum (Philippi, 1836) 0 0 0 0 0 0 0 0 0 0 Gastrana fragilis (Linnaeus, 1758) 0 23 0 0 0 0 1 0 0 0 Peronaea planata (Linnaeus, 1758) 0 1 0 0 0 0 1 0 0 0 Thracia pubescens (Pultney, 1799) 0 2 0 0 0 0 0 0 0 0 Pelecyora islandicoides (Lamarck, 1818) 0 13 0 0 0 0 1 0 0 0 Clausinella fasciata (da Costa, 1778) 38 0 0 0 7 62 0 0 0 0 Clausinella sp. 0 9 0 0 0 0 0 0 0 0 Venus nux Gmelin, 1791 0 0 221 0 0 0 0 0 1 0 Pitar rudis (Poli, 1826) 0 0 16 0 0 0 0 0 0 0 Gouldia minima (Montagu, 1803) 0 0 0 0 25 0 0 0 0 0 Paphia vetula (Basterot, 1825) 0 15 0 0 0 0 0 0 0 0 Circomphalus subplicatus (d'Orbigny, 1852) 0 1 0 0 0 0 0 0 0 0 Venerupis basteroti (Mayer, 1857) 0 4 0 0 0 0 0 0 0 0 Dosinia exoleta (Linnaeus, 1758) 0 0 49 0 0 0 0 0 0 0 Timoclea ovata (Pennant, 1777) 0 0 1 0 0 0 0 0 0 0 Lutraria oblonga (Chemnitz, 1791) 0 1 0 0 0 0 0 0 0 0 Corbula birostrata Trentanove 1901 50 0 0 0 0 9 0 0 0 0 Corbula gibba (Olivi, 1792) 0 0 846 0 0 0 0 8 26 68 Clavagella bacillum (Brocchi, 1814) 0 0 0 1 0 0 0 0 0 0 Fissidentalium inaequale (Bronn, 1831) 0 0 293 0 0 0 0 0 1 16 Omniglypta jani (Hörnes, 1853) 0 0 0 0 0 0 0 0 6 0 S = 94 2167 179 6108 22 1017 979 16 59 125 240 conditions” experienced during his field survey, Ruggieri (1956) dis- collections, together with species-level data from the literature cussed the fauna as a whole, without distinguishing between assem- (Trentanove, 1901, 1911), allowed to construct a quantitative database. blages from different sedimentary facies. Giannini (1960) confirmed The historical collection amounted to 8476 individuals divided in 66 instead the existence of three distinct macrofaunal associations in the species. Newly-sampled molluscs belonged to 2436 additional in- upper Miocene of nearby areas. More than fifty years later, Ruggieri's dividuals and 28 additional species, all smaller than 1 cm. The total paper, together with other upper Miocene fossil lists, formed the basis abundance is 10,912 individuals belonging to 94 different species. of a quantitative comparison on the biological impact of the Messinian Based on lithologic descriptions given by Trentanove (1901, 1911), Salinity Crisis on the Mediterranean benthos (Monegatti and Raffi, most species found in historical collections could be assigned to one and

2010). The interest on the Livorno upper Miocene molluscs continued, only one of the three different fossiliferous units described above (LUPb, as some of the species introduced by Trentanove (1901, 1911) were ROSb or RAQb: Table 1). Exceptions to the rule are a very few species discussed in a study of an early Messinian fauna from Sicily and con- that were not ascribed to a lithology by Trentanove (1901, 1911) and sidered Tortonian (Harzhauser et al., 2013). The study of mollusc the bivalve Microloripes dentatus, abundant both in LUPb, and RAQb faunas has been accompanied by that of the Monti Livornesi corals, fossil assemblages. For the purpose of statistical analysis, we arbitrarily framed in the general evolution of the Mediterranean during the late decided to divide the Microloripes dentatus museum lot in two equal Miocene (Esteban, 1979; Bossio et al., 1996; Perrin and Bosellini, parts among the two above-mentioned assemblages. Amanda glandi-

2013). formis was assigned to ROSb assemblage based on the lithology of the associated matrix, which is very distinctive (a strongly-cemented sandstone). Museum samples and new field samples, pooled for each 3.2. Methods fossil assemblage, were confronted by means of Spearman correlation using the software PAST 3.23 (Hammer, 2019). A more detailed com- We revised the Popogna and Quarata collection hosted at the parison of historical and new samples is the subject of a separate mu- Museum of Natural History (MSN) of the University of Florence, con- seological and statistical study that also verifies the integrity of the taining all fossils described by Trentanove (1901, 1911). This collection MSN collection (Dominici et al., submitted). Abundances of genera formed during the years, grouping specimens mainly collected by Igino within collections were confronted with a dataset of analogue Miocene Cocchi and Giorgio Trentanove (late nineteenth-early twentieth cen- and Pliocene data. Miocene data were taken from the literature, tury: Dominici, 2011). The MSN material was catalogued and labelled. whereas most of the Pliocene dataset is largely formed by bulk-samples Species were discussed and specimens counted. Five samples of the treated and studied in the same way as bulk samples collected in the Monti Livornese fauna were picked from the surface in the two study Monti Livornesi (Supplementary Table 1). All samples with a total areas (Fig. 1 B,C), two in LUPb at Quarata (samples MQ1 and MQ2), one abundance smaller than 26 individuals (e.g., sample BR) were excluded in ROSb at Popogna (sample BR) and three in RAQb, both at Quarata from the quantitative comparison. (samples TR1, TR3) and Popogna (TR5: Figs. 1; 2; Table 1). In addition, Nonmetric multidimensional scaling (MDS) was used as an ordina- two bulk samples of about four liters each were collected in proximity tion method to detect ecological gradients, based on the Bray-Curtis of MQ1 and TR1, washed and sieved (1 mm mesh size) to look for small- similarity coefficient computed on square-transformed, standardised sized molluscs, the content being added to surface collections. During abundance data (Patzkowsky and Holland, 2012). The ordination was surface picking in the field, we collected all visible fossils within a performed with the computer software package PRIMER 6 (Clarke and limited area, allowing to connect each sample with one facies and Warwick, 1994). This and other statistical analyses were also important possibly an individual shell bed. Gastropods, bivalves and scaphopods to verify how abundance data of the Monti Livornesi museum were determined to the species-level and specimens were counted. New

111 S. Dominici, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 527 (2019) 103–117 collections compare to data from novel samples. 1485 individuals. The new collection is larger in abundance, with N = 1995, and more than double in richness, with S = 44, due to the 3.3. Paleoenvironmental gradients inclusion of small-sized molluscs, overlooked by previous authors (total

SLUP = 49). The ten most abundant species in each set show a com- Most species found in historical collections could be attributed to parable distribution of abundances, with the sole exception of the one and one only of the three vertically-stacked stratigraphic units LUP, small-sized Hydrobia frauenfeldi, absent in the historical collection, but ROS and RAQ. Exceptions include Microloripes dentatus, found both in dominant in new samples (about 40% frequency in sample MQ1: modern LUP and RAQ samples, and a few other species of unknown Table 1). The other three dominant species Microloripes dentatus, Ther- facies attribution (Table 1). The quantitative comparison of abundance icium italicum and Granulolabium tuberculiferum show similar dominance data of the Monti Livornesi fauna with abundance distribution of in the two quantitative sets, with similar ranks (Fig. 7 A). The fossil mollusc genera in 90 Miocene and Pliocene samples (Supplementary assemblage suggests an ecologically diversified benthic community, Table 1) allowed to better frame the relative significance of historical comprising among the infaunal bivalves chemosimbiotic forms of fa- and modern collections and to cast them within wider paleoenviron- mily Lucinidae (Microloripes dentatus), suspension-feeders (Cer- mental gradients. The MDS plot can be interpreted in terms of a astoderma glaucum) and, among the gastropods, herbivores (Thericium bathymetric gradient where samples are ordered along the main axis italicum, Gibbula spp., Bittium deshayesi), detritus-feeders (Hydrobia from onshore to offshore (from right to left in Fig. 6; see Dominici et al., frauenfeldi, Granulolabium tuberculiferum), scavengers (Tritia coloratus) 2018). In particular, most samples are ordered following an upper and active predators (Clavatula spp.). The distribution of abundances Miocene and a Pliocene gradient, where differences are relevant for (Fig. 7 A) suggests that the trophic nucleus was based on a large onshore communities becoming irrelevant going offshore, where Mio- quantity of organic matter deriving from the vegetal cover, like in cene and Pliocene samples intermingle. Samples collected in the Lup- modern seagrass meadows of the Northeastern Adriatic in very shallow piano unit (MQ1, MQ2 and LUP in Fig. 6) plot close to one another in subtidal settings (Weber and Zuschin, 2013). Very similar fossil asso- the onshore half of the diagram, whereas those from the Raquese unit ciations are known in the Burdigalian of the Paratethys (Agapilia pachii- (TR1, TR3, TR5 and RAQ in Fig. 6) occupy an offshore position along Granulolabium plicatum-Microloripes dentatus associations: Zuschin et al., the main gradient. On the offshore portion of the diagram, but sepa- 2004, 2014) and the Tortonian of the Mediterranean (Hydrobia rated from all the rest, are samples from coarse-grained shelly bottoms fraeunfeldi-Microloripes dentatus association: D'Amico et al., 2012). (carbonate or mixed carbonate-siliciclastic bottoms), as they have been Judging from the present-day environmental distribution of Hydrobia interpreted by previous authors (Merle et al., 1988; Bernasconi and (Wilke and Davis, 2000; Weber and Zuschin, 2013), the local abun-

Robba, 1993). The historical collection from the Rosignano unit plots in dance of Hydrobia frauenfeldi suggests that unit LUPb formed under this lower left corner of the ordination space (sample ROS in Fig. 6). brackish water conditions, with salinities either higher or lower than normal marine, in an outer flat setting. 3.4. Composition of the Monti Livornesi fauna The mollusc association found in calcirudites and calcarenites of

subunit ROSb (N = 245; total SROS = 26) is very distinctive, sharing no ff The molluscs associated with mudstones of subunit LUPb, char- species with the other Livornese assemblages and showing a di erent acterised by aragonitic shells, are dominated by a few species. The taphonomic signature, with aragonite shells either dissolved, and bi- historical collection attributable to this subunit contains 20 species with valves preserved as coupled inner and outer molds, or substituted from

Fig. 6. Non-Metrical Multidimensional Scaling (MDS) ordination of 93 Miocene and Pliocene samples, subdivided per stage, based on the distribution of abundances of mollusc genera. Monti Livornesi historical museum collections and modern samples shown in bold characters and larger triangles. The main axis of the ordination shows the presence of a bathymetric gradient, deepening from right to left. Samples from coarse-grained shelly bottoms occupy the lower left corner of the ordination. Samples from siliciclastic bottoms are ordered following an upper Miocene and a Pliocene gradient. Differences in abundance distributions are more relevant for onshore communities than for those offshore.

112 S. Dominici, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 527 (2019) 103–117

Fig. 7. Distribution of abundances of the most abundant mollusc species in historical and new col- lections, divided in the three assemblages. A) Ten

dominant species of unit LUPb (shallow-marine to brackish-water paleoenvironment, upper Tortonian- lower Messinian). B) Twelve dominant species of

unit ROSb (reef-slope, shallow marine paleoenvir- onment; lower Messinian). C) Ten dominant species

in unit RAQb. (open shelf marine environment, lower

Messinian). Spearman correlation (rs) suggests weak to moderate correlation between historical and new collections, increasing from older to younger fossil assemblages (samples MQ1-MQ2 and TR1-TR3-TR5 were pooled, to represent LUP and RAQ field col- lections, respectively); p-values ranging 0,05–0,02 suggest that no correlation is highly improbable.

calcite. It is dominated by large suspension-feeding bivalves (Fig. 7 B), both at reef-talus (Pecten aduncus, Talochlamys multistriata, Aequipecten both infaunal siphonate (Peronaea planata, Acanthocardia echinata, Pe- malvinae, Venerupis basteroti, Dosinia exoleta) or proximal slope (Acan- lecyora islandicoides, Dosinia exoleta, Venerupis basteroti) and epifaunal thocardia echinata, Lutraria oblonga) shallow subtidal settings (Níjar free-swimmers or byssate (Pecten aduncus, Talochlamys multistriata, reef: Jiménez and Braga, 1993; carbonate shelf with Porites colonies: Aequipecten malvinae). Gastropods include abundant carnivores (Cochlis Freneix et al., 1987a, 1987b; Lacour et al., 2002; Harzhauser et al., sp.) and large-sized (Xenophora sp.) and medium-sized herbivores 2013). Also the taphofacies is similar to that of other Messinian peri- (Oxystele rotellaris). Many of these taxa are known from other Messi- reefal settings (Freneix et al., 1987a, 1987b; Jiménez and Braga, 1993; nian, pre-evaporitic, Porites-dominated tropical paleoenvironments, Roveri et al., 2018). The lack of borers and Porites fragments and the

113 S. Dominici, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 527 (2019) 103–117 rather good sorting with respect to reef-slope facies (see Jiménez and tropical and subtropical species surviving in the Mediterranean during

Braga, 1993), suggests that subunit ROSb formed in a proximal slope the preglacial Pliocene and today either extinct or confined to tropical paleosetting. In Zanclean shallow marine successions of Southern Spain, and subtropical latitudes in the Atlantic (Monegatti and Raffi, 2007, a similar taphofacies, with Peronaea planata, is associated with a 2010). The uppermost assemblage from the Raquese Unit (RAQ) is widespread condensed section formed during a major marine flooding composed of geologically long-lived species presently thriving in open (Roveri et al., 2018). marine settings of the Modern Mediterranean and Northeastern Atlantic

The mollusc association found in mudstones of unit RAQb,is and of no chronostratigraphic significance in this context, such as dominated by different species with respect to underlying associations, Turritella tricarinata, Aporrhais uttingerianus, Tritia semistriatus, Anadara although some species are shared with the association of subunit LUPb. diluvii and Venus nux. However, RAQ also contains species previously The historical RAQ collection is the most abundant of the three, but unknown in successions younger than the Tortonian, such as Tritia with fewer species, with 5366 individuals belonging to 18 taxa. The brugnonis, Tritia striatulus, and Streptochetus ornatus, confirming that the new collection is smaller in size, with N = 402 and S = 12 (total use of benthic molluscs to the Raquese Unit is at least problematic.

SRAQ = 22). Dominant molluscs belong to the guilds of shallow infaunal Considering all evidence, we conclude that the Luppiano Unit deposited suspension-feeders (Turritella tricarinata, Corbula gibba, Clausinella sub- in the uppermost Tortonian or lowermost Messinian and that both the fasciata, Venus nux), deposit-feeders (Fissidentalium inaequale), selective Rosignano and Raquese units formed during the pre-evaporitic Messi- detritivores (Aporrhais uttingeriana) and scavengers (Tritia spp.; Fig. 7 nian, consistently with the correlation of the Quarata-Popogna succes- C). Microloripes dentatus attests to the availability of sulphide. The as- sion with better time-constrained, analogue sequence stratigraphic sociation is characterised by shallow-infaunal mud-dwellers with many units in other basins. species known in modern seas at shelf depths, close to the coast. Ana- logue associations are known in some classical localities of the Medi- 4.2. Facies relationships terranean area, of both Tortonian (Robba, 1968; Bernasconi and Robba, 1993) and Messinian age (Moroni, 1956; see Manzi et al., 2007). With respect to the species richness indicated for the whole Livornese fauna in previous systematic lists, we propose a slightly 4. Discussion higher figure than that of Ruggieri (1956, where S = 63). Our study adds to previous lists all the small-sized species found in quantitative 4.1. Age of the Livornese fauna samples, reaching a richness S = 93, almost doubling the value used by Monegatti and Raffi (2010, with S = 54). All small-sized molluscs have Previous authors have treated the Livornese upper Miocene assem- aragonite shells. They were found exclusively in LUP and RAQ, but blage as a whole, leaning toward either a Tortonian or a Messinian were missing altogether from ROS, probably due to their dissolution chronostratigraphic interpretation. Trentanove (1901, 1911) adopted a during fossil diagenesis. Taphonomy is here found to select species Tortonian interpretation, followed by Harzhauser et al. (2013). depending on grain size of the matrix (the coarse-grained matrix found

Ruggieri (1956), followed by Monegatti and Raffi (2010), opposed this in subunit ROSb facilitated the movement of pore water and the un- view and stated that if the fauna is Messinian, the “aberrant presence of dersaturation of carbonates), shell composition (calcitic shells of pec-

Tortonian limbs” in Tuscany would be removed. So where Trentanove tinids and ostreids, abundant in ROSb, were not dissolved) and thick- (1901) re-interpreted “Nassarius” semistriatus listed by Capellini (1878) ness (casts of larger aragonitic shells still survived taphonomic as “Nassarius” hoernesi, a form with middle Miocene distribution pathways and many ROSb species are still recognisable: Table 1). The (“Nassarius” striatulus: Harzhauser and Kowalke, 2004), Ruggieri (1956) Livornese example therefore suggests that there are facies-dependant reestablished “Nassarius” semistriatus, a species very common in the taphonomic factors that influence the measurement of biodiversity of Pliocene of the Mediterranean Sea. Where Trentanove listed “Nassarius” fossil assemblages. The systematic revision and the quantitative study brugnonis (Bellardi, 1882), of middle Miocene-Tortonian affinity, Rug- of the three Livornese fossil assemblages, coupled with upper Miocene gieri extended to the Messinian its distribution, relying on con- sequence stratigraphic interpretation, indicate three different deposi- temporaneous work carried out at San Marino (Moroni, 1956). Where tional environments formed at three consecutive transgressive events Trentanove (1911) split Clavatula in four different species, reflecting separated by erosional unconformities. The three distinct paleoecolo- the typical Tortonian diversity of the genus in the Northern Apennines gical units indicate, from bottom to top: 1) an outer tidal flat or very (e.g., Montanaro, 1937), Ruggieri (1956) saw only one, while re- shallow subtidal seagrass eutrophic setting, with brackish-water influ- interpreting Aequipecten malvinae, a pectinid common in the middle ence (HST of the Luppiano Unit); 2) a subtidal, slightly-deeper and Miocene (e.g., Mandic, 2004), as the living Aequipecten opercularis. oligotrophic carbonate shelf proximal to a Siderastrea-Porites reef (TST- Among eight new species introduced by Trentanove (1901, 1911), only HST of the Rosignano Unit); 3) an open-shelf muddy bottom with in- one survived in Ruggieri's revision. The Messinian interpretation was termediate nutrient content with respect to the previous two (TST of the inherited by modern authors in a comparison between upper Miocene Raquese Unit). These three distinct assemblages reflect the stacking and Pliocene mollusc species on the effects of the Messinian salinity pattern of the three units, subdivided in three distinct fining- and crisis (Monegatti and Raffi, 2010). The difference between taxonomic deepening-upward facies successions separated by erosional un- lists is also evident, the latter authors calculating percent of holdovers conformities, indicating that they were deposited during three distinct starting from a list of 54 species based on the work of Ruggieri (1956); cycles of relative sea-level variation. Each time the sea transgressed including 63 species according to the present revision, see Table 1), over the emerged land, a different marine setting ensued. Each time the instead of 73 species listed by Trentanove (1901, 1911). Our study sea regressed, an unknown part of the sedimentary record was eroded. suggests that we should make a distinction between three Livornese fossil assemblages, each corresponding to one of three stacked sequence 4.3. Comparing diversities across the Messinian salinity crisis stratigraphic units. The assemblage of the lowermost Luppiano Unit (LUP) contains species presently unknown in sediments younger than Because of the nature of the sedimentary rock record, estimates of the Tortonian, such as Hydrobia fraunefeldi, Clavatula coppii and Clava- diversity measured simply by summing up the numbers of taxa (species, tula mystica. Living species are rare, a case being Cerastoderma glaucum, genera, families) recorded from successive geological time intervals surviving in modern Atlantic and Mediterranean coasts (Malham et al., brings only to crude approximation that may be subject to a number of 2012; Weber and Zuschin, 2013). The assemblage of the intermediate biases (Smith and McGowan, 2011). Species appearing or disappearing Rosignano unit (ROS) shows instead many affinities with early Messi- at a boundary are often associated with shallow marine onshore facies, nian, pre-evaporitic assemblages of the Mediterranean area, with those where turnover is statistically more frequent (Smith et al., 2001;

114 S. Dominici, et al. Palaeogeography, Palaeoclimatology, Palaeoecology 527 (2019) 103–117

Smith, 2007; Zuschin et al., 2011; Patzkowsky and Holland, 2012). stratigraphic context and position along the depositional profile of the Many studies related to species counts across Neogene sequence basins represented in the dataset (see Holland and Patzkowsky, 2015, boundaries did not explicitly account for the effect of facies control on for numerical simulations, and Nawrot et al., 2018, for an empirical fossil assemblage composition (e.g., Ben Moussa et al., 1989; Lauriat- example in brackish and marine molluscs). The lower preservation Rage et al., 1999; Monegatti and Raffi, 2001, 2010; Mandic and potential of onshore facies due to preferential uplift and erosion of Steininger, 2003) and the resulting interpretation in terms of extinction proximal sectors of sedimentary basins, which is known to lead to more rate remain questionable. In one of these studies (Monegatti and Raffi, patchy record of these environments relative to offshore facies (Smith 2010), the composition of the whole Livornese fauna has been com- et al., 2001; Zuschin et al., 2011), might also have played a role. Some pared to, among others, a late Tortonian assemblage (from locality of the dominant species of intertidal upper Miocene assemblages, such Stirone, Northwestern Italy: Marasti, 1973), six early Messinian as- as Microloripes dentatus and Hydrobia fraunefeldi, are however unknown semblages (localities including Casa Gessi, Northeastern Italy: Moroni, in the Pliocene, possibly substituted in the ecological niche by the 1956; Manzi et al., 2007; Sorbas, Southern Spain: Lacour et al., 2002; modern confamiliar (or congeneric) species Loripes lacteus and Hydrobia Sais, Marocco: Lauriat-Rage et al., 1999; Barbieri and Ori, 2000) and six ulvae, so that what took place during the Messinian salinity crisis was a middle Messinian assemblages (including molluscs from Oranie, true evolutionary and ecological turnover in taxonomic composition of Northern Algeria: Freneix et al., 1987a, 1987b, 1988; Melilla, Northern intertidal communities. Marocco: Ben Moussa et al., 1987, 1988, 1989). Some of the confronted faunas were collected exclusively in a carbonate facies, with a differ- 5. Conclusions ential preservation selecting in favour of calcitic shells (e.g., Ben Moussa et al., 1989; Freneix et al., 1987a, 1987b, 1988; Lacour et al., Species of the upper Miocene fauna of the Monti Livornesi, in 2002). These assemblages are comparable to the ROS fauna, but not to Tuscany (Italy), have been used to document faunal change across the the LUP and RAQ faunas. Others came from open-marine marls and Messinian Mediterranean salinity crisis. Historical museum collections mudstones, with aragonite shells preserved (e.g., Marasti, 1973; proved to be efficient tools for paleoecologic analysis, abundance dis- Moroni, 1956), a facies directly comparable to the RAQ assemblage, but tributions being analogous to those measured from new quantitative not to LUP and ROS. Finally, none of the study cases included assem- samples. What has been considered a single assemblage of Tortonian or blages from onshore, outer flat paleoenvironments with brackish-water Messinian age, depending on the author, is formed by three faunas species, such as the LUP assemblage. Published comparisons need to be originated in distinct paleoenvironments. These have subsequently revised. Gradient analysis of abundance data shows that when faunal undergone different taphonomic pathways. The succession is char- change takes place during million-years time spans, offshore faunas are acterised by the stacking of three depositional sequences, the Luppiano, more similar to each other than onshore faunas (Dominici and Kowalke, Rosignano and Raquese units. Each unit is bounded by an unconformity 2007), particularly across Neogene climatic and oceanographic per- that can be tracked in the nearby Fine Basin and is internally char- turbations (Tomašových et al., 2014) and that, at times of disruption of acterised by a transgressive trend, from fluvial conglomerate to onshore-offshore faunal gradients, deep subtidal faunas might expand brackish-water or marine shelly sandstone or mudstone. The quantita- into shallow habitats (Holland and Patzkowsky, 2007). One way to tive study of shell beds based on museum collections and new field overcame misinterpretations of faunal change or stasis from patterns samples shows that the Luppiano mudstone contains a marine to generated by facies control, is to carry out multiple studies so as to brackish water mollusc fauna with aragonite shells, the Rosignano cover the whole range of facies that occur along gradients (Holland, sandstone a reef-slope assemblage deprived during diagenesis of the 1996) or to confi ne a study to a single facies available at multiple aragonitic component, and the Raquese mudstone an open-marine as- stratigraphic levels (Ivany et al., 2009). A carbonate-siliciclastic gra- semblage of aragonitic species. By correlation with adjacent Fine suc- dient and a water depth gradient are recognized at different strati- cessions and consistently with known distribution of upper Miocene graphic intervals of the Phanerozoic (e.g., Dominici and Kowalke, 2007; molluscs, the Luppiano fauna is dated to the late Tortonian-early Holland and Patzkowsky, 2007; Brett et al., 2016; Danise and Holland, Messinian, whereas the other two are early Messinian. A quantitative 2017). Similarly, the distribution of abundance data consistently places comparison with other Mediterranean Miocene and Pliocene mollusc the Monti Livornesi collections within upper Neogene Mediterranean assemblages allows to place the three faunas along a water depth gra- environmental gradients. Documenting the history of species depends dient and possibly also along a carbonate-siliciclastic gradient. The on knowledge of environmental gradients in geological time. Molluscs results of the ordination analysis suggests that in intertidal and onshore from a carbonate and mixed coarse-grained facies comparable to ROSb settings some true evolutionary and ecological turnover in taxonomic are known in the Messinian (Jiménez and Braga, 1993), Zanclean composition took place during the Messinian salinity crisis, while off- (Yesares-García and Aguirre, 2004), Piacenzian and Gelasian (Cau shore communities remained relatively unchanged. et al., 2018), whereas biofacies comparable to LUPb and RAQb, and Supplementary data to this article can be found online at https:// others that make up the whole depth environmental gradient are doi.org/10.1016/j.palaeo.2019.04.024. widespread in the Neogene (Fig. 6 and Dominici et al., 2018) and Quaternary (see for example Scarponi and Kowalewski, 2007), allowing Acknowledgements us to consistently detect the effect of major events shaping the com- position of the Mediterranean marine benthos. The ordination analysis We thank two anonymous reviewers for precious comments that of Miocene and Pliocene fossil assemblages suggests that some change have improved the final quality of the manuscript. Thanks to Stuart in taxonomic composition took place, higher in onshore with respect to Wallace for revising the English. offshore habitats consistently with previous works (e.g., Tomašových et al., 2014; but see Danise and Holland, 2017, for a counter example of References higher turnover in offshore settings during an interval of sea-level and climate change). The Pliocene dataset used in the present comparison Barbieri, R., Ori, G.G., 2000. Neogene paleoenvironmental evolution in the Atlantic side covers all facies along the depositional gradient from intertidal to upper of the Riftian Corridor (Morocco). Palaeogeogr. Palaeoclimatol. Palaeoecol. 163, 1–31. slope (Dominici et al., 2018), including the facies encountered in LUP Bellardi, L., 1882. 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