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Accepted Manuscript Palaeocological anatomy of shallow-water Plio-Pleistocene biocalcarenites (Northern Apennines, Italy)

Simone Cau, Alex Laini, Paola Monegatti, Marco Roveri, Daniele Scarponi, Marco Taviani PII: DOI: Reference:

S0031-0182(17)31253-1 doi:10.1016/j.palaeo.2018.08.011 PALAEO 8897

To appear in:

Palaeogeography, Palaeoclimatology, Palaeoecology

Received date: Revised date: Accepted date:

19 December 2017 7 August 2018 20 August 2018

Please cite this article as: Simone Cau, Alex Laini, Paola Monegatti, Marco Roveri, Daniele Scarponi, Marco Taviani , Palaeocological anatomy of shallow-water PlioPleistocene biocalcarenites (Northern Apennines, Italy). Palaeo (2018), doi:10.1016/ j.palaeo.2018.08.011

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ACCEPTED MANUSCRIPT Palaeocological anatomy of shallow-water Plio-Pleistocene biocalcarenites (Northern Apennines, Italy)

Simone Cau 1*, Alex Laini 1, Paola Monegatti 1, Marco Roveri 1, Daniele Scarponi 2, Marco

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Taviani 3,4,5

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1) Dipartimento di Scienze Chimiche, della Vita e della Sostenibilità Ambientale, Università

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degli Studi di Parma, Parco Area delle Scienze 157/A, 43124, Parma, Italy. 2) Dipartimento di Scienze Biologiche, Geologiche e Ambientali, University of Bologna,

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Via Selmi 3, I-40126 Bologna, Italy.

3) Istituto di Scienze Marine (ISMAR-CNR), Via Gobetti 101, 40129 Bologna, Italy.

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4) Biology Department Woods Hole Oceanographic Institution, 266 Woods Hole Road, Woods Hole, MA 02543, USA.

*Corresponding author

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5) Stazione Zoologica Anton Dohrn, Villa Comunale, 80121 Naples, Italy.

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Abstract

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Email: [email protected]

Shell-rich biodetrital carbonate lithosomes punctuate the Plio-Pleistocene marine shallowwater successions of the Mediterranean basins. These m-thick sedimentary packages often display large-scale clinostratified geometry and show a remarkable rhythmic alternation with mudstone lithosomes. Our palaeoecological analysis defines the depositional settings and hints to the factors involved in the development of these peculiar bio-detrital carbonate units. Six distinct biofacies, have been identified through a two-way cluster analysis based on the macrofossil content, mostly molluscs, and matched with sedimentary facies. These are

ACCEPTED MANUSCRIPT Aequipecten gr. opercularis and Bittium reticulatum biofacies, Anomia ephippium biofacies, Ditrupa arietina and Tritia semistriata biofacies, Corbula gibba and Saccella commutata biofacies, Timoclea ovata and Corbula gibba biofacies, Timoclea ovata and Anomia ephippium biofacies. Quantitative analyses suggest that these bio-detrital deposits have a multiphase history, and that, along with their bracketing marine mudstones, are not

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nearshore or shoreface deposits but developed in mid to outer shelf settings. Our results

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suggest that sea-level change was neither the only nor the main factor controlling the

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internal stacking pattern of individual biocalcarenites. They provide instead arguments in favour of periodic, high-amplitude climatic/oceanographic oscillations determining

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significant modifications of the source-to-sink dynamics of the basin and affecting the trophic structure, the supply of terrigenous sediments, as well as the energy and pattern of

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bottom currents of shelfal areas.

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Keywords: Stratigraphic palaeoecology, Mollusc, Heterozoan carbonates, Palaeoclimatology

1. Introduction

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Skeletal accumulations in the fossil record are the product of many taphonomic, environmental and ecologic filters intervening from the original carbonate factories to their

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ultimate stratal entombment (e.g., Carthew and Bosence, 1986; Hendy et al., 2006; Cherns et al., 2008; Kidwell, 2013; Brady et al., 2016; El Quot et al., 2017). In the marine realm, such fossil-rich beds are widespread in the Cenozoic record, resulting from the concentration of skeletal remains in coastal to outer shelf environments (e.g., Kidwell, 1982, 1991; Kidwell et al., 1986; Abbott, 1998; Nakashima and Majima, 2000; Del Rio et al, 2001; Hendy et al., 2006; Massari and D’Alessandro, 2012; Nalin et al., 2016). The origin of these bio-detritic accumulations is of great interest due to their important sedimentologic, stratigraphic, palaeoclimatic and tectonic implications (Kidwell, 1985; Beckvar and Kidwell, 1988;

ACCEPTED MANUSCRIPT Meldhal, 1993; Abbott, 1997; Avila et al., 2015; Negri et al., 2015; Scarponi et al., 2016; 2017a; Tomašovỳch et al., 2017; von Leesen et al., 2017).In the Mediterranean Basin, peculiar skeletal-rich units are clustered at specific time intervals within the Piacenzian and Gelasian successions of the Apennine-Maghrebian foreland basin system (Pomar and Tropeano, 2001; Di Bella et al., 2005; Massari and Chiocci, 2006; Chiarella and Longhitano,

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2012; Nalin et al., 2016). In the Northern Apennines, these macrofossil rich, mollusc-

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dominated beds are well developed in the Castell’Arquato Basin (CAB), where the bio-

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detrital carbonate rocks display a cyclical stacking pattern highlighted by the rhythmic alternation with mudstones. The integration of facies analysis, seismic, and bio-

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magnetostratigraphic data from boreholes and outcrops provided a robust chronostratigraphic framework for the CAB biodetrital-mudstone units (Roveri et al., 1998;

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Monegatti et al., 2001). Indeed, within the CAB, shell beds appear to be thicker and more carbonate-rich starting from 3.1 Ma, forming distinct clusters of major biocalcarenite-

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mudstone couplets with a time interval periodicity of 400-kyr, corresponding to the

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eccentricity maxima (see Roveri and Taviani, 2003 and references therein). The Castell’Arquato Basin is an ideal venue for Plio-Pleistocene stratigraphic and

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palaeontologic investigations, owing to its rich marine fossil record framed in a wellestablished chronostratigraphy (e.g., Barbieri, 1967; Iaccarino, 1967; Rio et al., 1988; Roveri

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et al., 1998; Monegatti et al., 2001, 2002; Roveri and Taviani, 2003; Ceregato et al., 2007; Crippa, 2013; Crippa et al., 2016; 2018). However, beside a general appreciation of their richness in fossils, a detailed palaeoecological analysis of the CAB biocalcarenitesmudstone couplets is missing so far. In addition, a direct control of relative base-level changes in the development of the biocalcarenite-mudstone couplets is not obvious and palaeontological analysis might prove fundamental for their correct interpretation. This research aims to provide a first in-depth qualitative and quantitative analysis of the macropalaeontological content of the CAB biocalcarenites and associated mudstone units, in order

ACCEPTED MANUSCRIPT to reconstruct their relevant environmental and depositional aspects and to disclose their architectural framework. As shown in other geological settings, a stratigraphic palaeobiology approach to sedimentary successions represents a powerful tool to shed light on the anatomy of complex, multifaceted units (e.g., Dominici, 2001; Scarponi and Kowalewski, 2007; Holland and Patwkowski, 2012; Scarponi et al., 2014; Capraro et al., 2015; Moisette et al.,

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2016; Danise and Holland, 2017).

2. Geological setting

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The Castell’Arquato Basin (CAB) is located between the Arda and Chiavenna river

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valleys in the foothills of the Northern Apennines (Fig. 1a), a fold-thrust belt made up of tectonic units emplaced since the Oligocene (Castellarin et al., 1992). Seismic and borehole

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data show that the CAB is a wedge-top basin developed above the Northern Apennines orogenic wedge and filled up by a Plio-Quaternary sedimentary succession (Roveri et al.,

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1998). The CAB developed in the innermost part of the Po Basin foreland basin system

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(sensu De Celles and Giles, 1996) whose foredeep depozone with turbidite deposits is now buried under the Po Plain (see Ghielmi et al., 2013 and references therein).

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The Plio-Pleistocene succession of the CAB forms a N-NE-dipping monocline and records a large-scale (3rd order) depositional sequence bounded by a regional unconformity

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(MES; Fig. 1a) related to an intra-Messinian tectonic phase (Roveri et al., 1998; 2003; 2014). The 3rd order sequence of the CAB can be further subdivided into four unconformitybounded units (sensu Mutti et al., 1994), labelled from the bottom UM, LP, MP and UP (Fig. 1). Their bounding surfaces are dated at ~5.6, 4.2, 2.7 and 2.1 Ma, respectively (Figs. 1 and 2; Monegatti et al., 2001; Roveri et al., 2003; Roveri and Taviani, 2003), and related to Pliocene-Quaternary phases of tectonic uplift of the Apennine fold-thrust belt. The LP depositional sequence in turn has been subdivided into three smaller scale sequences (LP1, LP2, LP3; Figs. 1 and 2). Stratigraphic and sedimentologic analyses allowed to reconstruct

ACCEPTED MANUSCRIPT the overall depositional profile of the basin. Coastal settings of sequence UM are not preserved and only epibathyal deposits are represented in the study area (Barbieri, 1967; Iaccarino, 1967; Rio et al.,1988; Channell et al., 1994). Coastal and inner shelf deposits of sequences LP, MP and UP occur in the northwestern sector of the CAB, whereas mid to outer shelf and bathyal deposits are observed respectively in the central and southeastern

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ones (Fig. 1b). Our study is focused in the central sector of the CAB (Figs. 1 and 2); here the

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sedimentary succession is characterized by a rhythmic alternation of m-thick biocalcarenites

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and mudstones, interpreted as deposited in mid- to outer shelf settings characterized by settling of fine-grained siliciclastic sediments and by biological activity resulting in mollusc

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shell accumulation. More proximal environments (i.e. nearshore/shoreface above the fairweather wave-base) can be ruled out for the absence of wave-related sedimentary structures;

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the absence of massive to graded sandstone or hybrid siliclastic-carbonate beds with erosional or sharp base and hummocky cross-stratification, which commonly occur in inner

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shelf settings, point to deposition in shelfal areas beyond the influence of fluvial floods and

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storms. The alignment and imbrication of shells, the planar and/or the small to large-scale cross-bedding often observed in biocalcarenites suggest the action of unidirectional traction

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currents. These could be interpreted as by-pass deposits of catastrophic fluvial floods or storm-related gravity flows depositing the bulk of their sediment load in more distal

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positions; however, field and subsurface data do not indicate the occurrence of sand-rich deposits downbasin with respect to biocalcarenites. On the other hand, micro- and macropalaeontological data exclude a deeper (e.g. epybathyal) deposits for sequences LP and MP in the studied sector of the CAB, which are found in the basin depocenter (e.g. Stirone Section, Figs. 1 and 2). In the central sector of the CAB, major biocalcarenite bodies form three distinctive clusters separated by mudstone horizons. According to the proposed sequence-stratigraphic reconstructions (Roveri et al., 1998; Roveri and Taviani, 2003), the lowermost cluster

ACCEPTED MANUSCRIPT belongs to the transgressive systems tract (TST) of sequence LP3, whereas the middle and upper ones are assigned to the TSTs of sequences MP and UP, respectively. The individual biocalcarenite bodies are always associated with the flooding surfaces separating the minor units (parasequences and/or elementary depositional sequences - EDS, sensu Mutti et al., 1996, 2000) which form the systems tracts (Fig. 1B).

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Each cluster is characterised by individual cycles formed by biocalcarenite-mudstone

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couplets (Figs. 1 and 2): the Monte Giogo cluster (MTGc, four couplets), the Monte Padova

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cluster (MTPc; five couplets), and the San Nicomede cluster (SNc; six couplets). Minor bioclastic-rich sandstone lithosomes (i.e., less developed in terms of thickness, lateral

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continuity and carbonate component) and their mudstone counterparts occur in sequence LP2, along the southern margin of the CAB, and are well exposed in the Lugagnano-Monte

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Giogo section (Figs. 1 and 3).

Bio- and magnetostratigraphic data (Monegatti et al., 2001; Roveri and Taviani, 2003

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and references therein) document that the clusters of the major biocalcarenitic bodies are

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coeval with the major Mediterranean sapropel clusters of late Piacenzian to Gelasian age (Hilgen, 1991a,b; 1999; Lourens et al., 1996; Roveri and Taviani, 2003; Fig. 2), which

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developed at 400 kyr eccentricity maxima in deep-water settings. The age correspondence and the similar stacking pattern of biocalcarenitic bodies, observed in the same time window

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in other Mediterranean basins (Sicily, Tuscany), prompted Roveri and Taviani (2003) to suggest common forcing factors for the two deposits, possibly related to astronomicallydriven climate and oceanographic changes at obliquity or precessional scale.

3. Methodology

3.1. Fieldwork Sampling intervals were established following the reconstructed stratigraphic

ACCEPTED MANUSCRIPT framework (biocalcarenite-mudstone couplets and embedding mudstones; Monegatti et al., 2001; Supplementary Data Table 1). Sampling volume was influenced by the abundance of fossils as seen in the field. Namely, the sample volume was limited in bioclastic deposits (0.5-1.0 litres) and larger in fossil-poor mudstones (25-32 litres). Though the sampling volume differed among lithologies, our goal was to collect samples with comparable

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abundances across the whole investigated succession. Two to four subsamples (especially in

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mudstone units) were picked from the same interval at a horizontal distance of 1 to 2 m. in

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order to buffer the effects of fossil patchiness. Following Wittmer et al.’s (2014) procedure, samples were dried (24 h at 40 °C), soaked in 3% H2O2 solution depending on lithology,

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and wet-sieved over a 1.0 mm screen. For each sample, fossils were separated from the residual material and the most abundant fossil fraction (i.e., molluscs, brachiopods and

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serpulids) were identified, when possible, to the species level and counted. Less common fossil remains (e.g., bryozoans, corals and echinoderm fragments) were not counted. The

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term specimen here refers to a complete fossil or a unique fragment (e.g., apex for

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gastropods and umbo for bivalves). In order to obtain a conservative estimate for the number of bivalve individuals, 0.5 for further analysis multiplied each valve. The number of serpulid

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specimens in a sample was estimated from the total weight of tube fragments in that sample

2012).

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divided by the estimated weight of an average complete specimen (see Martinell et al.,

3.2. Quantitative data analysis The data matrix comprises 35 samples, representing 333 species, and 109,455 specimens. Some of the samples utilised in this study were previously published by Monegatti and Raineri (1987) (MR871 to MR874, Rio Stramonte Section), Monegatti et al. (1997) (MR971 to MR978, Monte Padova Section; RR1-RR2, Riorzo Section) (see Fig. 2). All specimens identified at the genus level or higher were excluded from quantitative

ACCEPTED MANUSCRIPT analyses (i.e., 15 taxa and 982 individuals; Acantocardia gr echinata includes A. echinata and A. spinosa; Aequipecten gr. opercularis includes A. opercularis and A. scabrella; Euspira gr. helicina includes E. helicina, E. catena, E. pulchella and E. macilenta). Small samples (i.e., n < 50 specimens and < 10 species) along with rare species (i.e., present in only one sample) were also removed. The final matrix counts 33 sample and 103 species. R

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Statistics software (R 3.2.2 GUI; R Core Team, 2015) was used for multivariate analyses.

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3.3. Multivariate analyses

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Multivariate analyses have proven useful in identifying and dissecting complex relationships in palaeoenvrionmental studies (e.g., Dominici, 2001; Yesares-Garcìa and

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Aguirre, 2004; Tomašových and Kidwell, 2011; Holland and Patzkowsky, 2012; Amorosi et al., 2014; Kowalewski et al., 2015; Scarponi et al., 2017b; Azzarone et al., 2018; Rossi et al.,

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2018). One of the most beneficial advantages of these techniques is the ability to reduce

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large masses of data into a usable form while emphasizing the significance of their similarities and differences (Burma, 1949). However, palaeontological datasets usually

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suffer from a variety of sampling biases (e.g., poor/uneven sampling, varying sample sizes, etc.) that could make the interpretation of multivariate outputs not so straightforward. Alroy

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(2015), by means of simulations based on uneven sampling size, demonstrated that a series of well-known and widely used distance metrics (e.g., Dice or Euclidean) in cluster analyses tend to create clusters that reflect sample size. Whereas others (such as Simpson) tend to overemphasise differences within and between clusters. For these reasons and for the peculiar structure of our dataset, characterised by unequal sampling and strong differences in sample size, we employed the conservative binary coefficient-Forbes revised (Alroy, 2015) that seems to behave better in such circumstances and creates groups based strictly on faunal co-occurrence. In addition, by

ACCEPTED MANUSCRIPT employing a two-way cluster analysis (e.g., Patzkowsky and Holland, 2012), the matrix reporting species abundances will aid the taphonomic/ecological interpretation of Forbes binary-derived dendrograms. Samples (Q-mode) and species (R-mode) were clustered by the unweighted pair group method with arithmetic averaging (UPGMA). The two-way cluster analysis was conducted with default settings.

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Biofacies units were identified by Q-mode clusters, or singletons if formed by a

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single sample, and their ecological insights derived only by those R-mode clusters (or

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singleton) well represented in the examined Q-cluster. That is, at least 50% of the species forming an R-cluster have to be present in more than 50% of samples of one of the resulting

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Q-clusters. As for small R- clusters (i.e., n taxa ≤ 5), these are considered only if at least one species is present in all samples of a Q-unit with relatively high mean abundance (i.e.,

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average n > 30 specimens). Furthermore, if the totality (or a great part) of the taxa (n >80%) of large R-units are well represented in only a portion of a Q-cluster, these R-units are

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mentioned as sub-biofacies. Hence, Q-clusters recognize suites of biofacies and are usually

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named after the most abundant species therein retrieved. Finally, non-Metric Multidimensional Scaling (NMDS) on the same matrix employed for the two-way cluster

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analysis was performed as a cross-check on the robustness of the outpus. The NMDS output yield consistent even if less effective, output. Here we report two-way results as for

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ordination output (see Supplementary Data: Figure 1 and Table 2)

3.4. Bathymetric calibration of CAB biocalcarenite-mud couplets

Multiple strategies can be employed to develop a bathymetric model suitable for interpreting different facies tracts in a stratigraphic perspective (e.g., Scarponi and Kowalewski, 2004; Hendy and Kamp, 2007; Dominici et al., 2008; Wittmer et al., 2014; Capraro et al., 2017). In this research, preferred bathymetry attributes of extant species are

ACCEPTED MANUSCRIPT employed to develop sample-level water depth estimates on CAB samples, following a weighted averaging approach (see also Supplementary Data Table 3). This approach assumes that the preferred bathymetry of the most common and extant species recovered was approximately the same in the studied interval as today. Given the uncertainty related to the comparison of modern and fossil taxa, the bathymetric estimates obtained should be

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considered with prudence. However, these values may provide insight on bathymetric trends

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at the scale of the basin infill, and a robust ground for the recognition of the same trends

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within the biocalcarenite-mudstone couplets, which is the most relevant information for the aim of this work.

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We obtained present-day water depth estimates of the extant species retrieved in CAB samples from the New Technologies Energy and Environment Agency (ENEA) Italian

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Mollusc Census database

(http://www.santateresa.enea.it/wwwste/banchedati/bd_ambmar.html).

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In addition to water depth and to the abundance of retrieved individuals, the ENEA database

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includes substrate data (e.g., coralligenous, sandy, muddy, silty-mud, etc.). Hence, ENEA species entries were parsed for lithology and only those occurrences reporting an agreement

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in sediment composition between our samples and ENEA samples were utilised to acquire the average water (i.e., preferred) depth for each species representing at least 0.2% (or at

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least 20 specimens) of the total abundance of a CAB sample. However, for oligotypic samples (where a species >60% of total abundance) and represented by at least 20 taxa, the above-mentioned cut off value was lowered to five specimens. In the case of samples represented by less than 20 taxa and characterised by a dominant species (i.e., monotaxic sample), all taxa have been used. The sample water depth along with the associated standard error were then calculated for each sample by means of weighted (by species occurrences in ENEA dataset) average of species preferred depth (see Supplementary Data Table 4).

ACCEPTED MANUSCRIPT 4. Results

4.1. CAB biocalcarenite-mudstone units: field observations Qualitative field-based palaeoecological analyses identified clear differences between minor and major biocalcarenites, as well as between biocalcarenite and mudstone

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deposits.

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Minor biocalcarenites are 1-3 meter-thick, tabular bodies (Fig. 3), consisting of

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densely packed, comminute, variably abraded and bioeroded shells in a muddy matrix with subordinate coarse- to medium-grained sand. These bioclastic-rich units are characterised by

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oligotypical/low diversity assemblages dominated by epifaunal and sessile suspension feeders (mostly Anomia ephippium, Aequipecten gr. opercularis, A. scabrella, Glycymeris

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spp., Moerella donacina and rare Atlantella pulchella), along with herbivorous gastropods (e.g., Bittium, Pusillina, Alvania). The fabric is generally chaotic and internal bedding is not

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recognisable, as well as mechanical sedimentary structures; however, at the top of these

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bodies horizontal/imbricate valves of A. opercularis and A. scabrella, suggesting weak and/or sporadic winnowing by unidirectional currents, can be locally recognised.

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The major biocalcarenitic bodies are up to 10 meters thick (exceptionally 25 m in MTPc; Fig. 2) and are often characterized by well-developed progradational geometries

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showing a topset-foreset-bottomset facies tripartition (Figs. 3 and 4). The bottomset facies mainly consists of pectinid-dominated shell pavements (Flabellipecten flabelliformis, A. opercularis and A. scabrella), mostly showing convex-up orientation in a coarse- to medium-grained sandy matrix with planar lamination. Centimeter-scale bioclastic horizons dominated by Anomia ephippium are also commonly observed. The foreset facies is often characterised by weakly to well cemented clinoform beds showing oligotypical, poorly preserved (typically mouldic preservation) horizons of mainly imbricate A. ephippium, pectinids and ostreids valves associated with widespread Ophiomorpha and Thalassinoides

ACCEPTED MANUSCRIPT ichnotaxa. The topset facies is made up of sandstone containing disarticulated skeletal remains without preferential organization or locally forming nesting geometry (sensu Kidwell et al., 1986). These deposits enclose rich macrofaunal associations, in which herbivorous Tricolia pullus, Diodora italica and Emarginula spp and abundant epifaunal detritivorous gastropods (in particular Alvania spp.) are key-features. The latter are abundant

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also in bottomset facies.

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Macrobenthic assemblages retrieved in mudstone and sandy-mudstone associated to

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bioclastic-rich units are generally characterised by scattered and low diversity fossils content. However, thin, oligotypic veneer of shells or clumps of fossils are occasionally

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retrieved in these muddy units. Epifaunal detritivores or deposit-feeders are scant, whereas infaunal detritivores show a marked increase respect to coarser-grained bioclastic-rich units.

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In general, infaunal and semi-infaunal detritivores and suspension feeders (i.e., Turritella tricarinata, Saccella communtata, Abra nitida, Corbula gibba and Ditrupa arietina) are

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dominant or at least represent a substantial proportion of the fauna.

4.2. Two-way cluster analysis

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The top dendrogram (Q-mode analysis) shows three super-clusters (Fig. 5). Qsupercluster 1 groups mainly samples from biocalcarenite bodies (cluster 1a) along with a

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minority of coarse-to-medium-sandy samples (cluster 1b). Q-supercluster 2 shows chaining and heterogeneous sedimentological features: from biogenic-rich sediments (i.e. assemblages of the MTG1 biocalcarenites) to silty-muds. Whereas, in Q-supercluster 3 samples are all siliciclastic (mudstone to medium-grained sandstone) and chaining is still conspicuous. Hence, a better understanding of the depositional environment and processes along studied sections require analysis of single clusters defining biofacies. The Q-mode cluster analysis allowed the recognition and characterization of six biofacies and one sub-biofacies (i.e., alphanumeric codes in Fig. 5) along with two

ACCEPTED MANUSCRIPT singletons. In the R- and Q-mode dendrograms, the cut-off similarity value for the recognition of examined R- and Q-clusters was chosen in function of multiple outputs obtained varying analytical options (i.e., taxa and/or sample values, correlation coefficients and data transformations).

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4.2.1. Cluster 1a: Aequipecten gr. opercularis and Bittium reticulatum biofacies

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This biofacies is identified by Q-mode cluster 1a and is characterised by R-clusters

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A, B, E and R-singletons a-b (Fig. 5). R-cluster A is mainly represented by shell-gravel dwellers (i.e., Thylacoides arenaria, Limaria loscombi, Ostrea sp., Barbatia candida,

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Paracratis minuta, Clausinella punctigera and Astarte sulcata), dominated by A. gr. opercularis, a taxon indicative of sandy to shell-gravel bottoms from inner-to-middle shelf

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depths (Kamenos et al., 2004). This unit also contains gastropods sourced from mats or vegetated substrates, such as Bittium reticulatum, B. deshayesi, Jujubinus striatus, and

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Calliostoma conulus. R-cluster A shows also a residual stock of species characteristic of low

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energy muddy substrates (i.e., Venus nux, the extinct Turritella tricarinata and Dentalium sexangulum), commonly reported in muddy shelf settings subject to high rates of terrigenous

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input (Taviani et al., 1997; Amorosi et al., 2002). R-cluster B is characterised by eurytopic serpulid Ditrupa arietina, preferential of coarse to mixed unstable shelf substrates

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(Cosentino and Giacobbe, 2006) and the epifaunal Calyptrea chinensis, a sessile suspension feeder that exploits any suitable hard substrate, including cobbles and shells (Minchin and Nunn, 2006). The filter feeder bivalve Timoclea ovata even if not included in any of the selected R-units, is ubiquitous in Q-cluster 1a. T. ovata, as A. gr opercularis, reaches high abundances on sandy and shell-gravel substrata of inner to mid shelf settings subject to bottom currents (e.g., Kamenos et al., 2004). R-cluster E is well represented by Nucula nucleus, an infaunal bivalve common in sandy to muddy-gravelly oligophotic bottoms of the inner and middle-shelf (La Perna, 2007). Lastly, both Anomia ephippium and Corbula gibba

ACCEPTED MANUSCRIPT singletons are also well represented. A. ephippium is a eurybathic, sessile bivalve thriving on hard substrates (Poirier et al., 2010). Conversely, the infaunal suspension/deposit feeder C. gibba favours soft substrates and is retained an indicator of stressed settings caused by pollution, low oxygen content or increased turbidity (Hrs-Brenko, 2006). The multiple R-units charachterizing Aequipecten gr. opercularis and Bittium

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reticulatum biofacies indicate a complex eco-taphonomic setting, represented by the co-

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occurrence of taxa with ecologically non-overlapping distributions (e.g., R-singletons a vs.

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b, or B. reticulatum and D. aretina). Indeed, all key-taxa here recognized are nowadays recovered in inner to mid shelf settings but show different tolerance to water turbidity and

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prefer different substrates: coarse (R-units A and a), vegetated (R-units A and B), sandymuddy (R-units A, E, D, and a). Hence, the co-occurrence in this biofacies (Q-cluster 1a), of

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taxa with different ecological requirements is here interpreted as the result of intermittent periods of intensified winnowing of fines in heterogenic shelf bottoms subject to variable

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sedimentation rates. Therefore, the removal of fines allowed taxa with different ecological

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requirements to be packed within the same stratigraphic horizon.

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4.2.2. Cluster 1b: Anomia ephippium biofacies This biofacies is identified in the two-way dendrogram at the intersection of Q-

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cluster 1b and R-singletons a-b (i.e., A. ephippium and C. gibba singleton in Fig. 5). The peculiar trait of this biofacies is the dominance of A. ephippium in all the samples (average abundance 44.3%) coupled with the lowest species richness of the dataset. As mentioned in the previous biofacies, A. ephippium is a shelf eurybathic and epifaunal bivalve that favours hard to coarse substrates. Q-cluster 1b is also represented by a relevant presence of Aequipecten gr. opercularis not included in the relevant R-cluster. The ecological traits of this biofacies indicate a characteristic and ecologically compatible suite of taxa thriving primarily in shelf settings characterised by coarse-grained

ACCEPTED MANUSCRIPT sediments subject to bottom currents (A. gr. opercularis), which are not a limiting factor for A. ephippium (Maughan, 2001). C. gibba, which thrives in ecologically stressed environments along the shelf (see 4.2.1), is poorly represented in this biofacies, thus reinforcing such an interpretation.

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4.2.3. Cluster 2a: Saccella commutata and Tritia semistriata biofacies

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This biofacies, defined by R-clusters C, F plus A. ephippium and C. gibba singletons,

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groups samples with different sedimentological features and chaining is noticeable within this unit (Fig. 5). The bulk of this biofacies is made from three samples (MG1B, MG1T and

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CLB2) that show a high affinity with biofacies 1a (section 4.2.1) in terms of lithological (coarse-grained) and biological (sharing R-cluster A) features. The three samples record high

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abundance (within this Q-unit) of four key taxa: A. ephippium, C. gibba, D. arietina, and Saccella commutata, characterising respectively, R-singletons a-b and -clusters A, C. As for

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the environmental context of A. ephippium and its coexistence with C. gibba we refer to

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sections 4.2.1-2. Suspension-feeder D. arietina is indicative of turbidity/high sedimentation in mixed substrates, sensitive to increasing concentrations of fines, and rather tolerant to the

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presence of coarse sediments (Cosentino and Giacobbe, 2006; Hartley, 2014). S. commutata is an eurybathic infaunal bivalve that preferably thrives in interstices of shell-gravel bottoms

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along the shelf and is capable of withstanding increased sedimentation rates (Gianolla et al., 2010). R-cluster F is represented by Tritia semistriata (= Nassarius semistriatus), an extinct gastropod indicative of mid-outer shelf muddy bottoms and commonly reported in Pliocene siliciclastic successions (e.g., Dominici et al., 2018). An important stock of filter feeders is represented by Amusium cristatum, Limopsis aurita, Phaxas adriaticus and Anadara diluvii that thrive in mid-shelf muddy settings (Aguirre et al., 1996; Monegatti et al., 2001; Ceregato et al., 2007). The core of Q-cluster 2a grows by the addition of four more samples, showing a decrease in the richness of species indicative of coarse-to-mixed substrata (e.g.,

ACCEPTED MANUSCRIPT D. arietina, Ostrea sp. and G. minima for sub-biofacies A; S. commutata for r-unit C). Following the ecological requirements of the species characterizing Q-cluster 2a, the bulk of this biofacies (as in A. gr. opercularis and B. reticulatum unit, see section 4.2.1) groups ecologically mixed assemblages pointing to heterogenic shelf bottoms subject to periodic winnowing. However, respect to A. gr. opercularis and B. reticulatum biofacies, this

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unit reflects a progressively increase in sedimentation rates or decreasing of

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bypassing/winnowing events responsible for the relative increase of mud-loving taxa. This

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interpretation gets further support by the decrease of the sessile, epifaunal flat-bivalve A. ephippium (suggestive, when abundant of current-swept setting; Broekman, 1974), and

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variable abundance of the opportunistic C. gibba, which peaks in settings characterised by subtantial and aperiodical oversilting and/or hypoxia events (Di Geronimo et al., 1987; Hrs-

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Brenko, 2006).

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4.2.4. Cluster 2b: Corbula gibba and Saccella commutata biofacies

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The Q-cluster 2b represents a further step toward lower values of similarity in the highly chained supercluster 2 and is defined by a sub-set of species that characterise the

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previous biofacies, namely R-cluster C and C. gibba singleton (Fig. 5). Based on the considerations reported for the previous unit (section 4.2.3), this biofacies groups low

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diversity assemblages of mixed shelf bottoms under low to moderate sedimentation rates.

4.2.5. Cluster 3a: Timoclea ovata and Corbula gibba biofacies This biofacies is identified by Q-cluster 3a and characterised by R-clusters D-E and singletons a-b (Fig. 5). The R-clusters D-E record mainly suspension-feeders, the most common being: T. ovata, Parvicardium minimum, Pitar rudis (cluster D), Capulus hungaricus and Striarca lactea (cluster E). In modern marine settings, these molluscs are primarily found in offshore transition-to-middle shelf mixed and coarse bottoms (Zenetos,

ACCEPTED MANUSCRIPT 1996; Mastrotauro et al., 2008). In addition, R-clusters D and E show a subordinate number of deposit-feeders: Lembulus pella, Atlantella pulchella, Moerella donacina and the extinct gastropod Archimidiella spirata. R-cluster D also presents a stock of extinct carnivorous gastropods (i.e., Crassopleura sigmoidea, Roxania utriculus and Nassarius serraticosta) commonly found in mixed (fine- and coarse-grained) shelf deposits (Massari et al., 1999;

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Ceregato et al., 2007; Moisette et al., 2016).

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The relatively well-diversified structure of this biofacies, characterized by the

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presence of important stocks of filter-, deposit-feeders and carnivorous taxa, along with the reduced abundance of A. ephippium and C. gibba, that commonly peak in presence of

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ecosystem stressors, suggests a shelf setting characterised by mixed-substrates subject to

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low-moderate sedimentation rates.

4.2.6. Cluster 3b: Timoclea ovata and Anomia ephippium biofacies

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This biofacies is identified by Q-cluster 3b and is characterised by R-cluster D, and -

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singletons a-b (Fig. 5). The key species of this biofacies are already examined in previous sections to which we refer (see sections 4.2.2 and 4.2.5).

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The ecological requirements of the key-taxa belonging to R-cluster E and -singleton a point towards mixed grained, bottoms in offshore transition to middle shelf settings (see

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also biofacies 3a). The noticeable and concomitant presence of A. ephippium and C. gibba point to heterogenic shelf bottoms subject to periodic winnowing or high sedimentation rates.

4.2.7. Singleton 1c and 3c Two samples MF1 and RR1 are not included in any Q-mode cluster, as they join their respective Q-superclusters above the cut-off value (Fig. 5). Singleton 1c (= sample MF1) represents the extreme chaining of Q-supercluster 1 and is characterized by a low diversity

ACCEPTED MANUSCRIPT assemblage, mainly represented by R-singleton b (A. ephippium) and A. gr. opercularis and S. commutata species. Similarly, singleton 3c (= sample RR1) represents the extreme chaining of Q-supercluster 3 and is defined by R-cluster B, indicative of vegetated substrates (section 4.2.1) along with R-singletons a-b (both however, showing reduced abundance values; Fig. 5). Besides, it shows a certain similarity in composition with the T. ovata and

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A.ephippium biofacies (Q-cluster 3b in Fig. 5).

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As for Q-singleton 1c, the presence of a scant stock of species dominated by A.

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ephippium, A. gr. opercularis and N. commutata suggests a possible shelf setting characterized by mixed grained bottoms subject to currents (see also section 4.2.2). Whereas

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cluster 3c, following considerations expressed in the previous units (sections 4.2.1 and 4.2.6), is retained indicative of sandy bottoms in mid shelf settings characterised by a

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vegetated cover.

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4.3. Palaeobathymetric trends

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Our results, based on the preferred bathymetry of extant species retrieved in the CAB samples, support and refine previous qualitative estimates (Monegatti et al., 1997; 2001;

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2002). We conducted bathymetric analysis on a set of samples representative of the CAB stratigraphic architecture (Figs. 2 and 6). Only samples from Rio Stramonte section were not

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considered, as they overlap stratigraphically with those collected along the adjacent Lugagnano-Monte Giogo section (see Fig. 2). The palaeobathymetric curve reconstructed for the studied portion of the basin fill suggests an overall shallowing upward trend. At the scale of individual depositional sequences, a substantially invariant bathymetric trend characterizes sequence LP2. An overall shallowing upward trend characterizes sequence LP3, with deepest estimates attained by bioclastic-rich basal deposits, while the shallowest water depths are recorded within the uppermost, muddy deposits (Fig. 6). This is in good agreement with sedimentological

ACCEPTED MANUSCRIPT evidence and progradational geometries observed in seismic profiles suggesting the basinward migration of coastal and inner shelf depositional systems during the latest Piacenzian. Similarly, biocalcarenite samples from the base of the overlying MP sequence marks a phase of water-depth increase, followed by an overall shallowing-upward trend (Fig. 6). The two sandy samples of the UP sequence returned shallow-water settings. The

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fact that biocalcarenites clusters attain deeper palaeobathymetric estimates than their

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embedding mudstone successions (Figs. 2 and 6) is in good agreement with the

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reconstruction of the sequence-stratigraphic architecture of the CAB basin proposed by Roveri et al. (1998) and Roveri and Taviani (2003). Indeed, these authors considered the

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biocalcarenite clusters as developed in the transgressive systems tracts of large-scale composite depositional sequences related to the tectonic evolution of the basin.

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On a smaller scale, i.e. the biocalcarenite-mudstone couplets, palaeobathymetric trends suggest a similar scenario. Indeed, in most of the examined biocalcarenite-mudstone

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couplets, deeper water values are invariably attained in bioclastic lithosomes (Fig. 6).

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However, the estimated palaeobathymetry of samples collected from major biocalcarenites units, due to the uncertainties associated to the water-depth estimates, does not allow

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discriminating between the recognized facies tripartition (i.e., topset-foreset-bottomset). The overlying mudstone lithosomes are commonly characterised by shallower depth

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estimates and shallowing upward trends, attaining water depths as shallow as -20 m (see LP2 and MP sequences; Fig. 6); these deposits may have a modern equivalent in the highstand muddy wedges of the Late Quaternary depositional sequence of the Adriatic basin (Amorosi et al., 2016). The reconstructed palaeobathymetric trends suggest that the biocalcarenite-mudstone couplets may record transgressive-regressive cycles developed within a mid- to outer shelf setting. In sequence-stratigraphic terms, thus considering relative sea-level changes as the main factor controlling the stratigraphic architectures, these cycles may represent either parasequences or elementary depositional sequences forming the TST

ACCEPTED MANUSCRIPT and HST systems tracts of LP and MP sequences. According to this interpretation, the biocalcarenites would correspond to the transgressive phase of the cycle, characterized by a substantially reduced input of terrigenous sediments, while the mudstones could be related to the progradation of coastal muddy wedges during the regressive phase of the cycle. However, the amplitude of such inferred palaeobathymetric oscillations appears very small

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(less than 15 m), and not sufficient to justify the striking facies differences characterizing

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deposits of the two hemicycles (biocalcarenite vs. mudstone), thus suggesting the need for

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additional forcing factors.

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5. Discussion

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5.1. Palaeoecology of CAB biocalcarenites-mudstone couplets Our quantitative data and field observations on biocalcarenitic bodies of the CAB

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reveal a complex internal architecture resulting from a polyphasic development. Two-way

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cluster output documents an overall difference between assemblages characterizing bioclastic-rich strata (Q-supercluster 1, biofacies 1a-c) and assemblages associated to

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terrigenous-rich deposits (Q-superclusters 3, biofacies 3a-c; and Q-superclaster 2, biofacies 2b in Fig. 5). A biofacies showing transitional features between the two assemblage groups

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has been identified (i.e., Q-supercluster 2; biofacies 2a; Fig. 5). However, the cluster analysis was not able to identify specific topset-foreset-bottomset biofacies, nor to separate major from minor biocalcarenitic bodies. Indeed, the multivariate output groups all biocalcarenite samples under the A. gr. opercularis and B. reticulatum and S. commutata and T. semistriata biofacies (see section 5.2.1). These biofacies are characterised by rich macrobenthic assemblages of inner shelf and partially represented by non-ecologically overlapping species. Hence, the skeletal concentration that typifies biocalcarenites (biofacies 1a and part of biofacies 2a; Fig. 5), are considered the result of amalgamation due to intense

ACCEPTED MANUSCRIPT and different hydrodynamic actions. Specifically, our paleoecological insights suggest the presence of bottom currents (presumably generated by storm events) and affecting the shelf sea-bottom between 50 and 30 m water depth (Fig. 6). These currents are responsible for winnowing of muddy deposits accumulated during previous periods of decrease in the intensity of bottom currents. This

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hypothesis is supported by the presence of planar to high-angle cross-laminations,

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imbricated pectinid pavements and the bathymetric (Fig. 6) and ecological preferences of

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dominant taxa (i.e., thriving in current-swept setting) retrieved in studied biocalcarenites. Quantitative multivariate tools were not able to resolve internal organization of investigated

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bio-detrital lithosomes. Field-derived paleoecological inferences allowed getting insights on the not well-resolved tripartite structure of major biocalcarenites. Topset facies tend to show

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higher richness in herbivorous taxa respect to the otherwise indistinguishable bottomset facies. Whereas, foreset facies are commonly represented by oligotypic, imbricated

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assemblages (Anomia ephippium).

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As for mudstone units, two-way cluster output revealed a series of siliciclastic related biofacies representing a wider spectrum of environments ranging from relatively stable to

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highly stressed settings. The S. commutata and T. semistriata or the S. commutata and C. gibba biofacies (Fig. 2, Q-clusters 2a-b in Fig. 5) are commonly retrieved at the boundaries

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with the biocalcarenite units (or within the latters). These two biofacies, given the similar ecological features, have a vicariant distribution, and usually develop when a strong reduction of winnowing occurs on biodetrital shelf bottoms (~40-30 m water depth; Fig. 6) with consequent shift from detritic to muddy-detritic substrata. These biofacies have been also retrieved (samples MG2 and CLP2 Figs. 2 and 5) in the lowermost mudstone separating biocalcarenite clusters of LP3 sequence and has been interpreted as the result of a geologically short-lived phase of winnowing on a muddy bottom. Similarly, the C. gibba and S. commutata biofacies is considered vicariant of the previous mentioned paleoecological

ACCEPTED MANUSCRIPT unit given the overlapping ecological features and occurrence with the same position. A different setting is suggested by Q-cluster 3a (i.e., T. ovata and C. gibba biofacies, section 5.2.5), grouping samples of the central part of the decametric thick sandy-mudstone successions separating major biocalcarenites of the LP3 sequence. The palaeoenvironmental and bathymetric signatures of this unit suggest the progradation of coastal mudstone wedges

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on mid-shelf settings during phases of sea level still stands.

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5.2. Genetic model for the major biocalcarenite bodies of the Castell’Arquato Basin The analysis of CAB macrobenthic assemblages coupled with published bio-

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geochronological and cyclostratigraphic data (Monegatti et al., 2001; Roveri and Taviani, 2003) allow to reconstruct the depositional environments of the CAB succession and their

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dynamics. Our stratigraphic and palaeobiological inferences are consistent with currently accepted models for offshore biocalcarenite deposits (e.g., Massari and Chiocci 2006;

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Massari and D'Alessandro, 2012; Nalin et al., 2016; Puga-Bernabéu and Aguirre, 2017). In

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addition to previous models, our sedimentological and palaeontological insights indicate for biocalcarenite deposit and their biological component a taphonomic imprint (see section 6.1)

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related to a constant sea-bottom winnowing, possibly originated by wind-driven shelf currents (see Massari and Chiocci 2006) during periods of reduced fluvial runoff and silting

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(Roveri and Taviani, 2003). We suppose that the minor siliciclastic component may result from rare events of catastrophic fluvial floods and/or of exceptional storms able to carry sand-sized sediments in mid-shelf areas. By combining sedimentological and paleoecological insights derived from qualitative and quantitative analyses, it is possible to sketch a model for the formation of major biocalcarenites. In particular, four different evolutionary steps recording the depositional history of a complete biocalcarenite-mudstone cycle can be defined (Fig. 7): 1) low-energy inner shelf environment dominated by deposition of fine-grained siliciclastic

ACCEPTED MANUSCRIPT sediments with sparse molluscs tolerant to environmental stressors (Fig. 7A); 2) sharp-based, flat-bedded, shell-rich hybrid deposits mark the initial development of a biocalcarenite ramp over the mudstone substrate of step 1. This phase is characterised by deposition of coarse sandy sediment carried by high-density flows, which also originated ecologically mixed (sand-loving species along with those reworked from underlying muddy

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deposits) and imbricated shell-beds (Fig. 7B) forming the base and bottomset of overlying

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deposits;

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3) clinostratified (foreset) deposits recording peak energy conditions (e.g. high-frequency storm events and/or winnowing by steady unidirectional currents) and indicating the full

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development of sandwave/dune fields in shelfal settings, as suggested by the presence of low diversity assemblages embedded in a matrix of abraded shells and shell debris (Fig. 7C);

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4) flat-bedded deposits (topset) showing a higher fossil diversity respect to foreset facies, suggesting a lower hydrodynamic disturbance, oligotrophic and well-oxygenated bottom

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conditions (active winnowing) and presence of vegetation (Fig. 7D).

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The biocalcarenite/sandstone-mudstone couplets form the basic elements of the stratigraphic architecture of the studied interval. Previous field studies suggested that no

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lateral transitions exist between the biocalcarenitic and the muddy deposits forming the two hemicycles (Roveri et al., 1998; Roveri and Taviani, 2003). This means that they do not

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represent different depositional systems (or elements) developing synchronously along the depositional profile. Accordingly, their vertical superposition would not be related to the simple landward or seaward shift of depositional systems during a complete base-level change cycle. Our results seem to confirm this view. Biocalcarenites and mudstones likely reflect two distinct sets of palaeoenvironmental conditions and processes experienced by the same shelfal depositional setting within a complete cycle. The most evident sedimentologic and palaeontologic signals document regularly alternating phases characterised by the activation

ACCEPTED MANUSCRIPT on the shelf of powerful bottom-currents (e.g., geostrophic, thermohaline) reworking skeletal sediments followed by lower-energy conditions with higher silting and water turbidity. These phases probably record periodic changes of the whole source-to sink-system of the basin, i.e. of the factors controlling the type and volume of sediments produced, the processes operating their transfer and distribution along the depositional profile, thus

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pointing to a mainly climatic control (Amorosi et al., 2016). The eustatic forcing, albeit

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surely accompanying its development, does not appear to be the only and surely not the

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main factor responsible for the origin of cycles. The full exploration of the inferred climatic control on the observed cyclicity is beyond the scope of this work and needs further work;

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however, our results seem to indicate that high-frequency, probably orbitally-forced (Roveri and Taviani, 2003), climatic oscillations had a prominent role in shaping the stratigraphic

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architecture of the studied interval, and particularly during specific time intervals coincident

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with eccentricity maxima.

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6. Conclusions

In this work, the relationships between depositional settings and composition of the

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benthic assemblages of Plio-Pleistocene bioclastic-rich lithosomes of the Castell’Arquato Basin have been reconstructed at different time scales, through sedimentologic, qualitative

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and quantitative palaeontologic analyses. Biodetrital units are rhytmically alternated to mudstone horizons forming small-scale sedimentary cycles which record periodic changes in the dynamics of offshore settings. Bioclastic units show vertical facies sequences which record phases characterized by a high energy hydrodynamic regime, likely due to the activation of powerful unidirectional currents on the shelf. Multivariate analyses evidenced a strong taphonomic imprint/homogenization on all biodetrital samples; coupled quantitative and qualitative insights suggest that the taxonomic assemblage of the biocalcarenite bodies differs substantially from those

ACCEPTED MANUSCRIPT characterizing the mudstones. The palaeobathymetric curves reconstructed at the scale of individual biocalcarenites-mudstone sequences show a substantially invariant bathymetric trend. This evidence indicates that the eustatic forcing, albeit contributing the development of the observed cycles, does not appear to be the major player for origin of the

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biocalcarenite-mudstone couplets as well as for the individual biocalcarenitic architecture.

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Acknowledgements

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Luigi Cantelli is gratefully acknowledged for the drone aerial documentation of the Lugagnano-Mt Giogo section. We thank Alessandro Freschi, Gianluca Raineri (Riserva

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Geologica del Piacenziano e dello Stiorne) and Carlo Francou (Museo Geopaleontologico ‘G. Cortesi,’ Castell’Arquato) for their assistance with sample collection and Andrea Cau for

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his help during the editorial process and for constructive comments. This is ISMAR CNR, Bologna, scientific contribution n.1950. The drone aerial documentation was supported by

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Consorzio Parchi del Ducato.

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Figure captions

Fig. 1. A) Simplified geological map of the Castell'Arquato Basin (CAB) and location of the studied stratigraphic sections. The geological map shows the main allostratigraphic units recognised within

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the Plio-Pleistocene succession outcropping in the CAB area (modified by Monegatti et al., 2001).

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B) Stratigraphic architecture of the Plio-Pleistocene infill of the CAB, based on field and subsurface

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data. The model represents an idealised west-east section from the basin margin towards its

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depocenter (Stirone section) (modified by Monegatti et al., 2001).

Fig. 2. Columnar logs, bulk sample position (red arrows) and correlation of studied sections within

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the Castell’Arquato Basin. Stratigraphic correlation of CAB sections is based on bio- and chronostratigraphic references (foraminifera and calcareous nannoplankton biozone and bioevents,

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magnetostratigraphic data, time distribution of sapropel clusters marine isotope stages and

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astronomic parameters; see Supplementary Data Table 1). Sections numbering: 1) Rio Stramonte; 2) Lugagnano-Monte Giogo section; 3) Monte Padova section; 4) Monte Falcone section; 5) Madonna

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dell’Arda section; 6) Rio Riorzo section; 7) Arda section; 8) Castell’Arquato 2 borehole; 9) Stirone section. Biostratigraphical events: G.”small”: Gephyrocapsa