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Original Citation: Pliocene stratigraphic paleobiology in Tuscany and the fossil record of marine megafauna / Dominici, Stefano; Danise, Silvia; Benvenuti, Marco. - In: EARTH-SCIENCE REVIEWS. - ISSN 0012-8252. - ELETTRONICO. - 176(2018), pp. 277- 310. [10.1016/j.earscirev.2017.09.018]

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10 October 2021 Earth-Science Reviews 176 (2018) xxx–xxx

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Pliocene stratigraphic paleobiology in Tuscany and the fossil record of MARK marine megafauna

⁎ Stefano Dominicia, , Silvia Daniseb, Marco Benvenutic a Università degli Studi di Firenze, Museo di Storia Naturale, Firenze, Italy b Plymouth University, School of Geography, Earth and Environmental Sciences, Plymouth, United Kingdom c Università degli Studi di Firenze, Dipartimento di Scienze della Terra, Firenze, Italy

ABSTRACT

Tuscany has a rich Pliocene record of marine megafauna (MM), including mysticetes, odontocetes, sirenians and seals among the mammals, and six orders of sharks among the elasmobranchs. This is reviewed with respect to paleogeography and sequence-stratigraphy in six different basins. Conditions at the ancient seafloor are explored by means of sedimentary facies analysis, taphonomy and multivariate techniques applied to a large quantitative dataset of benthic molluscs. MM is rare or absent in most basins during the Zanclean, except in one basin, and most abundant in Piacenzian deposits in all six basins. MM occurs preferentially in fine-grained, shelfal high- stand-deposits of small-scale depositional sequences, or at condensed horizons of the maximum flooding in- terval. It is rare in shallow marine paleonvironments and nearly absent in bathyal paleosettings. Paleogeographic and paleoecological evidence and a comparison with modern patterns of marine upwelling suggest that a wedge of nutrient-rich waters sustained in the offshore during the Pliocene a high biomass of primary producers and a community of apex consumers and mesopredators, similarly to the modern northwestern Mediterranean Sea, with a species-richness higher than the modern and a more complex trophic structure. The highest MM diversity coincides with the mid-Piacenzian warm period, suggesting that facies control does not obscure a link between climate and diversity. We underline however that not all marine environments were suitable for marine mammal preservation. Buoyant carcasses were preferentially dismembered and destroyed in high-energy shallow waters, with the possible exception of delta front deposits, where sudden sediment input occasionally buried pristine carcasses. We hypothesise that carcasses sunken on the seafloor below the shelf break underwent destruction through the activity of a whale-fall biota of modern type, specialised in the consumption of decomposing tissues, both soft and mineralised. A taphonomic window was left between storm wave base and the shelf break. Here water pressure is high enough to prevent the formation of decomposing gases and the resurfacing of carcasses, while the lack of a specialised whale-fall biota slows down bone degradation with respect to deeper settings. Sedimentation rate was high enough to cover skeletal material before its complete destruction. An estimate of paleobathymetries based on multivariate techniques suggests that the preferential depth for the inclusion of MM in the fossil record was 30–300 m. The results are compared with major Mesozoic and Cenozoic MM records worldwide. Available evidence suggests that the late Neogene radiation of large whales, true ecosystem engi- neers, and their size increase, triggered the radiation of a bone-eating fauna that hampered, and hampers, MM preservation in the deep sea. Stratigraphic paleobiology and an ecosystem-level approach deliver useful insights to the nature of the fossil record.

1. Introduction macroevolutionary scale, predation pressure has shaped the evolution of marine preys, with feedbacks on predators, setting the stage for the The modern marine megafauna (MM) includes all marine mammals, Mesozoic marine revolution (Vermeij, 1977; Chen and Benton, 2012; seabirds, sea turtles and sharks, apex consumers that influence their Benton et al., 2013). The new ecosystem structure started in the Early associated ecosystems (Lewison et al., 2004), both pelagic and near- and Middle Triassic with several lineages of actinopterygian fishes shore, through top-down forcing and trophic cascades, and now se- (Chen and Benton, 2012), continuing with marine reptiles possessing verely affected by human impact (Estes et al., 1998, 2011, 2016). On a feeding styles (Fröbisch et al., 2013; Motani et al., 2015; but see also

⁎ Corresponding author. E-mail address: stefano.dominici@unifi.it (S. Dominici). http://dx.doi.org/10.1016/j.earscirev.2017.09.018 Received 23 May 2017; Received in revised form 21 September 2017; Accepted 22 September 2017 Available online 28 September 2017 0012-8252/ © 2017 Elsevier B.V. All rights reserved. S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx

Motani et al., 2013) and reproductive adaptations (Motani et al., 2014) holistic approach that includes functional diversity and embraces as of modern type. Triassic and Jurassic novelties underwent a prolonged many ecosystem components as possible (Dineen et al., 2014). By crisis during the Cretaceous, with the gradual extinction of plesio- analogy with ecologists who shift focus from models based on single saurians, mosasaurs (Benson et al., 2010) and ichthyosaurs (Fischer groups (e.g., Steeman et al., 2009) to an all-embracing vision of marine et al., 2016), and a diversity drop of sharks (Guinot et al., 2012). A life (Lawton, 1994; Sergio et al., 2014), connecting food web ecology marine megafauna of comparable size returned in the Paleogene, with with landscape ecology (Polis et al., 1997; Estes et al., 2011), strati- the new diversification of neoselachian elasmobranchs (Kriwet and graphic paleobiology can draw from the fossil record and offer multi- Benton, 2004) and the evolution of large marine mammals: Eocene dimensional insights on the complex geological history of modern archaeocetes (Uhen, 2008; Gingerich et al., 2009) and Oligocene marine ecosystems. odontocetes and mysticetes (Gingerich, 2005; Marx and Uhen, 2010; After revising fossil MM hosted in major museums of Tuscany, both Berta, 2012; Marx et al., 2016) empowered by high metabolic rates and isolated and articulated remains, we focus on all fossil bones that can be new anatomic features (Armfield et al., 2013). Among the largest ver- stratigraphically (e.g., Bianucci, 1995; Bianucci et al., 1998, 2001; tebrates of all times, after a dramatic size increase at the outset of Tinelli, 2013) and taphonomically framed (e.g., Dominici et al., 2009; glacial age (Marx et al., 2016; Bisconti et al., 2017; Slater et al., 2017), Bianucci et al., 2010; Danise and Dominici, 2014). MM lists for the baleen and sperm whales are among today's ocean's ecosystem en- Mediterranean Pliocene have been recently updated (marine mammals: gineers (Roman et al., 2014) with which to compare their Mesozoic Landini et al., 2005; Bianucci et al., 2009a; Sorbi et al., 2012; Bianucci analogues (Smith et al., 2016). Notwithstanding a crucial role in and Vomero, 2014; sharks: Marsili, 2006). Species-level ecological data ecology and evolution, the nature and distribution of the MM fossil are available on modern apex consumers and mesocarnivores (Pauly record has been less explored, compared to that of marine invertebrates et al., 1998; Cortés, 1999), with detailed information made available and terrestrial vertebrates. Available data suggest a strong correlation for Mediterranean species following conservation concerns (marine between taxic diversity and the number of marine fossiliferous forma- mammals: Notarbartolo di Sciara et al., 2016a; sharks: Cavanagh and tions, resulting in megabiases in the fossil record (e.g., Cretaceous: Gibson, 2007), allowing for a detailed paleoecological evaluation of the Benson et al., 2010). Within its vast history, studies on the geologically Tuscan fossil record. The actualistic approach is also viable for species recent marine megafauna offer important insights, considering our of benthic molluscs, about half of which are still extant in modern better knowledge of: (1) geological setting, in terms of outcrop extent Mediterranean bottoms (55% of extant species of Mediterranean and and high-resolution stratigraphy; (2) ecologic role played by individual North Sea bivalves, excluding strictly brackish and bathyal forms, i.e., species, whether extant or extinct, in terms of habitat, trophic role, life 202 out of 367 species, survive from the Zanclean: Raffi et al., 1985). histories and population structure, thanks to a comparison with extant The regional quantitative study of molluscan assemblages was the basis descendants, or close relatives; (3) MM taphonomy, based on actuo- for an independent assessment of paleoenvironments, paleoecology and paleontology. A recent global study revealed that MM extinction paleobathimetry. A revision of abundance distributions of marine peaked in the late Pliocene, between 3.8 and 2.4 Ma, linked to the molluscs, the largest contributors to Mediterranean Pliocene shell beds sudden drop in the extension of nearshore environments after a large and a key component of Mesozoic and Cenozoic marine ecosystems sea level regression (Pimiento et al., 2017), confirming that the fossil (Stanley, 1975; Vermeij, 1977), allowed to further explore the structure record offers important clues on the vulnerability of keystone marine and composition of Pliocene marine ecosystems, and reconstruct a pa- species to climate change. We contribute here to a better understanding leobathymetric gradient (e.g., Scarponi and Kowalewski, 2004) along of the Pliocene fossil record by reviewing the rich MM of Tuscany, in which to frame MM occurrence. The present work must necessarily start Italy. In particular, we consider all reports of Pliocene marine mammals with a review of the chronostratigraphy and physical stratigraphy of the and sharks and revise taphonomy and sedimentary facies associated Tuscan marine Pliocene. with all known findings, setting them within a sequence stratigraphic framework. We also reconstruct the paleoenvironmental context and 2. Geological setting review data on a part of the marine ecosystem through the paleoecology of fossil invertebrates on a regional basis, following a stratigraphic The Pliocene succession of Tuscany was deposited in a complex paleobiological approach that can be applied to both the recent and the setting characterised by continental collision related to the later evo- distant geological past (Patzkowsky and Holland, 2012). Published lution of the northern Apennines chain. According to a well-established studies that have taken this direction are still not many, examples hypothesis, the region, affected by shortening before the middle-late concerning Jurassic ichthyosaurs, plesiosaurs and pliosaurs (McMullen Miocene, accommodated by NE-verging thrust and fold systems, un- et al., 2014), Cretaceous turtles, plesiosaurs, bony fish and sharks derwent crustal extension during the late Neogene and the Quaternary (Schmeisser McKean and Gillette, 2015), Eocene archaeocetes, sea cows (DeCelles, 2012; Fig. 1). Crustal extension generated differential sub- and sharks (Peters et al., 2009), and Neogene marine mammals and sidence in a series of normal-fault controlled hinterland sedimentary sharks (Boessenecker et al., 2014). All of these papers record the co- basins, filled throughout by continental and shallow marine, mostly occurrence of shelly faunas, only one undertaking quantitative studies clastic successions (Martini and Sagri, 1993; Pascucci et al., 2006; of the distribution of fossil invertebrates (Jurassic of the Sundance Brogi, 2011). An alternative hypothesis places the late orogenic hin- Formation: McMullen et al., 2014, see also Danise and Holland, 2017). terland basins in a more complex tectonic setting characterised by the The benefits of an outcrop-scale sequence stratigraphic approach in- alternation of compressive, extensional and transcurrent stress fields clude: (1) an independent record of relative sea-level change to test (Benvenuti et al., 2014; Bonini et al., 2014). paleobiological hypotheses (see also Pyenson and Lindberg, 2011; The Neogene Tuscan basins considered in this work include, from Noakes et al., 2013); (2) a chronostratigraphic scheme for high-re- West to East, and from North to South, the Fine Basin (FB; Bossio et al., solution correlations; (3) a means to recognise minor and major breaks 1997), the Volterra-Era Basin (VEB; Bossio et al., 1994), the Elsa Basin of the record; (4) an ecological and sedimentary framework for ta- (EB; Benvenuti et al., 2014), the Ombrone-Orcia Basin (OOB; Bossio phofacies distribution (Patzkowsky and Holland, 2012); and (5) an et al., 1991; Nalin et al., 2010), the Siena-Radicofani Basin (SRB; independent control of onshore-offshore patterns of fossil assemblages Ghinassi and Lazzarotto, 2005; Martini et al., 2011, 2016), and the (e.g., Tomašových et al., 2014). Chiana Basin (CB; Fig. 1: Pesa Basin not considered here). With one Researchers that study the geologic history of marine ecosystems exception (OOB, see below), these basins show a shape conditioned by have focused on patterns of ecological restructuring based on the tax- the structural and physiographic features of the inner portion of the onomy of selected groups (e.g., Thorne et al., 2011; Benton et al., 2013; Northern Apennines. Their NW-SE general elongation reflects the trend Scheyer et al., 2014; Fischer et al., 2016), at the expenses of a more of the thrust-related anticline ridges developed during earlier

2 S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx

the structural history also justifies the preservation in OOB of Zanclean fluvial and shallow marine facies (Fig. 2).

2.1. Pliocene stratigraphy

The Neogene succession of Tuscany is up to 2000 m-thick, about half of which belongs to the uppermost Miocene-Pleistocene interval (Bossio et al., 1997; Benvenuti et al., 2014). The Pliocene has been traditionally subdivided into three main informal lithostratigraphic units: continental conglomerates and sandstones at the base, overlain by the “Blue Clay Formation” (Argille Azzurre: Zanclean-lower Pia- cenzian), marking the post-Messinian Mediterranean marine trans- gression and forming the thickest part of the basin infill (e.g., Bossio et al., 1994, 1997; Ghinassi and Lazzarotto, 2005), and the “Upper Sands” (Sabbie superiori: Piacenzian-Gelasian) and conglomerates, de- posited during the ensuing regression. Several finer lithostratigraphic units have been introduced to define the local stratigraphy, resulting in a complex and largely informal lithostratigraphic terminology which includes Zanclean lower “Blue Clays” and Zanclean-Piacenzian upper “Blue Clays” (Capezzuoli et al., 2005), the latter eventually further separated by the widespread occurrence of Piacenzian carbonates (Nalin et al., 2016). The lower Zanclean (OOB: Ghinassi, 2007; Nalin et al., 2010) and the Piacenzian, are characterised by the high-fre- quency alternation of coarse-grained and fine-grained facies, ranging from fluvial to marine shelf settings (Benvenuti et al., 1995a, 1995b, 2007, 2014; Martini et al., 2011, Fig. 2). The dynamics of the Pliocene infilling are better-understood in the EB, where six synthems have been defined, each up to > 200 m-thick, further subdivided in a number of elementary and composite depositional sequences and chronologically calibrated through marine biostratigraphy and continental vertebrate biochronology (Benvenuti and Del Conte, 2013; Benvenuti et al., 2014, with references).

3. Materials and methods Fig. 1. Location of sedimentary logs within the largest Pliocene basins of Tuscany. Fine Basin (FB): Pagliana (1), Pieve Vecchia (2) and Orciano Pisano (3). Volterra-Era Basin Stratigraphic sections were measured and described at several lo- (VEB): Parlascio (4), Lajatico (5), Fabbrica (6) and Volterra (7). Era Basin (EB): San calities (Fig. 2). Siliciclastic and carbonate facies were described, sub- Lorenzo (8), La Serra (9), Poggio al Lupo (10), San Maiano (11), Canneto (12), Casenuove divided into groups of facies based on lithology, sedimentary structures fi (13), Castel orentino (14), Fiano (15) and San Gimignano (16). Orcia-Ombrone Basin and ichnology, and interpreted in terms of process and depositional (OOB): Arcille (17) and Poggio alle Mura (18). Siena-Radicofani Basin (SRB): Siena (19), environment (Table 1). Each group represents a set of individual facies Monteaperto (20), Castelnuovo Berardenga (21), Radicofani (22) and Fastelli (23). Chiana Basin (CB): Sinalunga (24), Cetona (25) and Allerona (26). forming monogenic associations (in the sense of Mutti et al., 1994), i.e., the meter-scale stacking of facies which express the autocyclic beha- viour of specific depositional systems within a given accommodation collisional stages. These compressive structures have bounded most space (Benvenuti and Del Conte, 2013). Sequence stratigraphic con- basins through their infilling, only to be obliterated by younger parallel cepts have been applied to reconstruct the dynamics of basin infills at a normal fault systems, leaving an invariant stratigraphic onlap of the hierarchy of scales, advancing hypotheses on controlling factors. The Pliocene successions onto the basin margins. Despite a NW-SE dis- sequence stratigraphic nomenclature used in this paper, taken from tribution of the hinterland basins, the structural setting is responsible Catuneanu et al. (2009), includes concepts such as sequence boundary for a NE-trending physiographic and paleogeographic gradient, where (SB), transgressive surface (TS), maximum flooding surface (MFS), the FB is closest, and the CB furthest, from offshore settings throughout maximum flooding (MF) interval, falling-stage systems tract (FSST), the late orogenic phase, with important implications for the facies ar- lowstand systems tract (LST), transgressive systems tract (TST) and chitecture and the distribution of marine vertebrates and shell beds. highstand systems tract (HST). The chronostratigraphic subdivision of Differential active uplift of the basin shoulders during the Pliocene, Benvenuti et al. (2014), which divides the Pliocene into six synthems, coupled with important erosional phases, resulted in a different pre- S1–S6 from older to younger, was extended to all six Tuscan basins by servation of the original stratigraphic architecture. The infill during the referring to available biostratigraphic schemes (Fig. 2). The sequence Zanclean is generally characterised by relatively continuous open stratigraphic interpretation of S2 in OOB is based on Tinelli (2013). marine successions, the correlative fluvial-coastal systems missing due Other parts of the S1–S3 succession were drawn based on available li- to uplift and erosion of basin margins. On the other hand, the Pia- thostratigraphic literature (see below). Studies integrating sedimentary cenzian infill is characterised by genetically-connected, fluvial, coastal facies analysis, biostratigraphy and magnetostratigraphy (Nalin et al., and shallow marine facies tracts, particularly well-preserved in the EB, 2016) have helped correlating carbonate bodies typical of S4. The hinting to reduced uplift of the marginal areas. The modern physio- reader is referred to Benvenuti et al. (2007, 2014) for details on facies graphy of OOB, escaping the structurally-controlled geometry of the analysis and sequence stratigraphic interpretation of synthems S3–S6. other basins, mimics instead an original fluvial network developed Data on the geographic distribution of fossil marine mammals, large during the latest Messinian, flooded after the Salinity Crisis (Bossio sharks and sirenians were largely based on collections housed at the et al., 1991; Benvenuti et al., 2015), filled during the Pliocene, and Natural History museums of the University of Florence (UFMSN), finally disrupted by post-Pliocene uplift and erosion. This difference in University of Pisa (UPMSN), and Accademia de' Fisiocritici of Siena

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Fig. 2. Pliocene stratigraphy of Tuscany, see Fig. 1 for the location of numbered sections. All logs measured and described by the authors, except Volterra (log 7: Bianucci et al., 1998), Arcille (log 17: Tinelli et al., 2012; Tinelli, 2013), Siena (log 19: Bianucci et al., 2001), Monteaperto (log 20: Martini et al., 2011), Castelnuovo Berardenga (log 21: Martini et al., 2016) and Radicofani (log 22: Ghinassi and Lazzarotto, 2005). Chronostratigraphy of locality 26 (Allerona, Danise, 2010) is unknown. Correlation between sections is also shown as boundaries of planktonic foraminifera biozones (dashed lines), following the scheme of Sprovieri (1992), based on available biostratigraphic studies for each basin (see main text for relevant references; chrono- and magnetostratigraphy modified after Hilgen et al., 2012).

(AFMSN), the three largest collections of Tuscany, and at the Geological living in the North-Western Mediterranean Sea (NWMS). Museum Giovanni Capellini, Bologna University (MGGC). In particular, All known MM-bearing sedimentary facies are associated with a counts of cetaceans were based on UFMSN collections (Mysticeti and mollusc-dominated benthic fauna. The third step of the analysis con- Odontoceti, N = 142), large shark on the sum of UFMSN, UPMSN and cerned a quantitative study of shell beds, allowing: 1) to interpret the AFMSN collections (Carcharhiniformes and Lamniformes, N = 337, regional evolution from a perspective independent from the sedimen- data synthesised from Marsili, 2006), sirenians from all reports in tary facies, 2) to characterise MM paleoenvironmental and bathymetric Table 2 (N = 10). Each record is formed either by a single element distribution, and 3) to explore the structure of the benthic component of (e.g., whale bone, shark tooth), by a few elements of the same in- marine ecosystems and to identify underlying environmental controls. dividual, or by a whole, quasi-articulated skeletons. A large proportion 72 bulk samples were collected at major shell beds at bed resolution of this dataset lacks precise location, allowing only for some crude throughout the succession and sieved with 1 mm mesh size. Fossils of stratigraphic attribution (Fig. 3). bivalves, gastropods and scaphopods were identified to species level. On a second step, all fossil Tuscan Pliocene cetaceans, sirenians and The minimum number of individuals was calculated following standard pinnipeds that could be framed within the available high-resolution approaches (see Patzkowsky and Holland, 2012), resulting in a richness stratigraphic framework and associated with taphonomic data, were of 525 species (S) and a total abundance of 64,206 individuals (N). We selected. At this step, after excluding unidentified MM remains, a da- coded each fossil assemblage by synthem, tract of small-scale deposi- taset of 64 specimens (cetaceans N = 50; sirenians N = 10; pinnipeds tional sequence, and depositional environment. Most samples belonged N = 4) was assembled. Association with shark teeth is frequent (55% of to facies types F2–F5 (Table 1; see also Tomašových et al., 2014). Facies 25 cases according to Danise and Dominici, 2014 for the Italian Plio- F6 usually lacks macrofossils and allowed for the collection of only one cene; see also Bianucci et al., 2002, 2010). The majority of the 64 sample. No samples were collected in facies F1, lacking marine shells, specimens are included in the catalogue of UFMSN, UPMSN, AFMSN facies F7, mostly devoid of shells, and facies F8, which is richly fossi- and MGGC, whereas a few are stored in smaller collections of the mu- liferous, but lacks aragonite shells and is associated with specimens nicipalities of Montaione, Scandicci (Florence province), and Certaldo hardly extractable from the rock. The resulting quantitative dataset (Pisa province), one in a private property (Castello di Villa Banfi, near served for statistical analyses on the distribution of species-level Montalcino, Siena province), and one in the Museum National d'His- abundances on a siliciclastic shelf depositional system, performed with toire Naturelle in Paris (France). Whenever possible, large marine the software Primer 6.0 (Clarke and Gorley, 2006). Analyses included vertebrates were coded by synthem (N = 60) and depositional en- clustering and NMDS ordination techniques on a Bray-Curtis similarity vironment (N = 54). We analysed abundance distributions among matrix, of standardised, square-root transformed data (72 samples; marine mammals, and species richness of marine mammals and sharks. S = 333 and N = 63,518 after the exclusion of singletons). To test To infer Pliocene paleoecology, fossil taxa recognised in Tuscany were statistically whether there is a significant difference between two or compared with their closest descendants, focusing on the species today more groups of sampling units based on sedimentary facies, we

4 .Dmnc tal. et Dominici S.

Table 1 Sedimentary facies types.

Facies type Main lithology Description Main fossil content Process interpretation Paleoenvironment References

F1a Conglomerate Crudely bedded pebble-sized conglomerate and Sparse plant remains, very rare bones of Poorly-confined subaereal or Multi-storey fluvial channel; Benvenuti and Degli Innocenti, subordinate coarse sandstone; graded gravelly terrestrial vertebrates subacqueus flows; channelized fiumara-like river 2001; Benvenuti and Del Conte, sandstone tractional flows 2013; Bianchi et al., 2015 F1a Sandstone Tabular trough cross-bedded medium and coarse Sparse plant remains, very rare Dune- to bar-scale bedforms Braided to low-sinuosity Benvenuti and Degli Innocenti, sandstone; lens-shaped beds with internal inclined fragmented bones of terrestrial characterised by both frontal and river channel 2001; Benvenuti and Del Conte, lamination vertebrates lateral accretion 2013; Benvenuti et al., 2007 F1b Mudstone Massive sandy silt and greyish clayey silt in tabular Sparse plant remains, very rare shells of Settlement from suspension; Floodplain Benvenuti and Degli Innocenti, beds, with root traces and vegetal matter, terrestrial molluscs bioturbation and pedogenesis 2001; Benvenuti and Del Conte, occasionally caliches 2013; Benvenuti et al., 2007 F2 Mudstone Massive dark grey mudstone with vegetal matter and Abundant plant remains, common or Settlement from suspension; crevasse Coastal lagoon; tidal flat Benvenuti et al., 2007; Dominici subordinated sandy lenses; bivalves in life position abundant shells of brackish-water splay and some channelized flow; et al., 2008 molluscs biogenic reworking of sediments F3 Sandstone Massive or small-scale cross-laminated medium and Common shells of marine molluscs, very Crude settling in high-density Proximal delta front; upper Benvenuti et al., 2007; Dominici 5 fine-grained sandstone with occasional interspersed rare bones of large terrestrial and marine suspensions; some wave reworking shoreface et al., 2008 mud clasts and fine pebbles; sedimentological shell vertebrates beds F4 Sandstone Massive bioturbated silty fine-grained sandstone, Common or abundant shells of marine Biogenic reworking of sediments; Distal delta front; lower Benvenuti et al., 2007; Dominici with shell beds and a diverse association of trace molluscs, rare crustaceans and corals, rare sediment starvation shoreface et al., 2008 fossils; occasional bioeroded and encrusted pebbles bones of large terrestrial and marine vertebrates F5 Mudstone Massive grey mudstones with sparse shells and shell Common shells of marine molluscs, rare Settling of fine-grained suspended Prodelta and open shelf Bossio et al., 1994; Benvenuti et al., beds; some bioeroded and encrusted shells crustaceans and corals, rare articulated load in low-energy environment; 2007; Dominici et al., 2008; remains of large marine vertebrates biogenic reworking Bianucci et al., 2009b; Martini et al., 2016 F6 Claystone Massive grey mudstone and claystone; shells very rare very rare shells of marine molluscs Settling of fine-grained suspended Outer shelf and upper slope Bossio et al., 1994; Benvenuti et al., load in low-energy environment 2007; Dominici et al., 2008; Bianucci et al., 2009b F7a Conglomerate Embricated pebbles, with occasional bioerosion Absent ? ? Bossio et al., 1994 F7b Sandstone Graded medium- and fine-grained sandstone in Absent Turbidity current Outer shelf and upper slope Bossio et al., 1994, 1997; Ghinassi

tabular beds and Lazzarotto, 2005 Earth-Science Reviews176(2018)xxx–xxx F8 Carbonate Polymictic conglomerate; bioturbated packstone in Common or abundant shells of molluscs, Settling of fine-grained suspended Shallow shelf Nalin et al., 2016 massive, laminated or clinostratified bodies, with rare red algae, brachiopods and load in low-energy environment; locally abundant shell beds crustaceans, locally abundant tests of biogenic reworking foraminifera .Dmnc tal. et Dominici S.

Table 2 Stratigraphic, taphonomic and paleoenvironmental framework for Pliocene marine mammals recovered in Tuscany, with abundance data (N = 64). A: Articulated and quasi-articulated skeleton; C: Complete and quasi-complete skeleton; C + PC: Cranial and post-cranial remains.

Marine Hosting institution Order Family Species Description A C C + PC Minimum Stage mammal abundance

WFi4-1 Museo di Storia Naturale, Cetacea Balaenopteridae Indet. Articulated, highly 1 1 1 1 Piacenzian Firenze (IGF 9299V) complete skeleton. Cortical bone layer deeply consumed WFi4-2 Museo di Storia Naturale, Cetacea Balaenidae Balaenula forsythmajori Periotic 1 Piacenzian Firenze (IGF 1624V) WFi4-3 Museo di Storia Naturale e del Cetacea Balaenidae Balaeonotus sp. Tympanic bullae 1 Piacenzian Territorio dell'Università di Pisa (MSNUP I-15458) WFi4-4 Museo di Storia Naturale e del Cetacea Delphinidae Etruridelphis giulii Partial skeleton (rostrum, 1 1 1 Piacenzian Territorio dell'Università di Pisa fragmented mandible, teeth, (MSNUP I-15458) parts of the skull, 17 thoracic and lumbar vertebrae, fragments of costae). No information on articulation WFi4-5 Museo di Storia Naturale e del Cetacea Balaenidae Balaenula sp. Partial skeleton (tympnic 1 1 1 Piacenzian Territorio dell'Università di Pisa bullae, fragments of the 2 (MSNUP I-12558) mandibles and sternum). No information on articulation WFi4-6 Museo Geologico Giovanni Cetacea Balaenidae Balaenula balaenopsis Partial skeleton (mandible 1 1 1 Piacenzian Capellini (MGGC 8908, fragment, elements of limb, – 6 8942 8947) tympanic bulla). No information on articulation WFi4-7 Museo di Storia Naturale e del Cetacea Kogiidae Kogia sp. Three periotics, teeth 2 Piacenzian Territorio dell'Università di Pisa (MSNUP I-16827) WFi4-8 Museo di Storia Naturale e del Cetacea Balaenotperidae Cetotheriophanes cf. C. capellinii Left limb 1 Piacenzian Territorio dell'Università di Pisa (MSNUP I-12559) WFi4-9 Museo di Storia Naturale e del Cetacea Balaenidae Balaenula sp. Left mandible 1 Piacenzian Territorio dell'Università di Pisa (MSNUP I-16838) WFi4-10 Museo di Storia Naturale, Cetacea Balaenidae Balaenotus orcianensis Periotic 1 Piacenzian Firenze (IGF 1648V) WFi4-11 Museo di Storia Naturale, Cetacea Ziphiidae Indet. Radius 1 Piacenzian Firenze (IGF 9361V) DFi4-1 Museo Geologico Giovanni Cetacea Delphinidae “Hemisyntrachelus pisanus” Partial skull, periotic, 1 Piacenzian Capellini (MGGC 8560) tympanic bulla

DFi4-2 Museo Geologico Giovanni Cetacea Delphinidae Etruridelphis giulii An incomplete skull, several 1 1 Piacenzian Earth-Science Reviews176(2018)xxx–xxx Capellini (MGGC 8560) vertebrae and ribs, sternum and one limb DFi4-3 Museo di Storia Naturale, Cetacea Delphinidae Hemisyntrachelus cf. H. cortesii Fragments of mandibles, 12 1 Piacenzian Firenze (IGF 1546V, 1547V, teeth, ulna 1551V, 1553V) DFi4-4 Museo di Storia Naturale, Cetacea Delphinidae Hemisyntrachelus sp. Ulna fragment, tens of teeth 1 Piacenzian Firenze (IGF 1527V, 1553V, 1556V, 1562V, 1564V) DFi4-5 Cetacea Delphinidae Etruridelphis giulii Three rostra 3 Piacenzian (continued on next page) .Dmnc tal. et Dominici S.

Table 2 (continued)

Marine Hosting institution Order Family Species Description A C C + PC Minimum Stage mammal abundance

Museo di Storia Naturale, Firenze (IGF 1541V, 1542V, 1544V) MFi4-1 Museo di Storia Naturale, Sirenia Dugongidae Metaxytherium cf. M. Two teeth 1 Piacenzian Firenze (IGF 102766) subapenninum PFi4-1 Museo di Storia Naturale e del Carnivora Phocidae Pliophoca etrusca Partial skeleton (skull, 1 1 1 Piacenzian Territorio dell'Università di Pisa fragmentary cervical, (MSNUP I-13993) thoracic, lumbar, sacral, and caudal vertebrae, radii, ulnae, carpals, metacarpals and phalanges, femora, tibiae, fibulae, tarsals, metatarsals and phalanges). No information on articulation WOm2-1 Castello Banfi, Montalcino Cetacea Balaenidae Indet. Disarticulated skeleton, 1 1 1 Zanclean with many cranial and post- cranial elements MOm2-1 Museo di Storia Naturale e del Sirenia Dugongidae Metaxytherium subapenninum Disarticulated skeleton with 1 1 1 Zanclean Territorio dell'Università di Pisa several cranial and post- (MSNTUP I-15892) cranial bones MOm2-2 Gruppo geopaleontologico Sirenia Dugongidae Metaxytherium subapenninum Highly articulated, highly 1 1 1 1 Zanclean GAMPS, Scandicci (GAMPS complete skeleton

7 62M) (incomplete skull and mandible, some teeth, 20 vertebrae, costae and sternum) MOm2-3 Gruppo geopaleontologico Sirenia Dugongidae Metaxytherium subapenninum Partial skeleton (teeth, 1 1 Zanclean GAMPS, Scandicci (GAMPS costae and vertebrae) 63M) MOm2-4 Gruppo geopaleontologico Sirenia Dugongidae Metaxytherium subapenninum Partial skeleton (costae and 1 Zanclean GAMPS, Scandicci (GAMPS vertebrae) 64M) MOm2-5 Museo di Storia Naturale, Sirenia Dugongidae Metaxytherium subapenninum Humerus 1 Zanclean Firenze (IGF 8743V) WEr1-1 Museo di Storia Naturale, Cetacea Physeteridae Indet. Isolated tooth 1 Zanclean Firenze (IGF 1632V) WEr1-2 Museo di Storia Naturale, Cetacea Physeteridae Indet. Isolated tooth 1 Zanclean Firenze (IGF 1577V) PEr1-1 Museo di Storia Naturale, Carnivora Phocidae Pliophoca cf. P. etrusca Partial skeletons (right and 1 1 2 Zanclean

Firenze (IGF 1473V–1486V) left mandibles, mandible Earth-Science Reviews176(2018)xxx–xxx fragment, teeth, cranial elements, several limb elements). No information on articulation WEl3-1 Museo di Storia Naturale, Cetacea Balaenopteridae Indet. Highly articulated, highly 1 1 1 1 Piacenzian Firenze (IGF 8184V) complete skeleton DEl3-1 Certaldo Municipality Cetacea Delphinidae Hemisyntrachelus sp. Skull 1 Piacenzian DEl3-2 Florence Archaeological Cetacea Delphinidae Indet. Partially articulated 1 1 Piacenzian Superintendence vertebrae and costae WEr4-1 Museo di Storia Naturale, Cetacea Kogiidae Kogia pusilla Skull 1 Piacenzian Firenze (IGF1540V) WEr4-2 Museo di Storia Naturale, Cetacea Delphinidae Globicephala etruriae Incomplete mandible with 1 Piacenzian Firenze (IGF 1675V) 30 teeth

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Table 2 (continued)

Marine Hosting institution Order Family Species Description A C C + PC Minimum Stage mammal abundance

DEr4-1 Museo di Storia Naturale, Cetacea Delphinidae Hemisyntrachelus sp. Teeth 1 Piacenzian Firenze (IGF 1562V) DEr4-1 Museo di Storia Naturale, Cetacea Delphinidae Etruridelphis cf. E. giulii 7 and 2 thoracic vertebrae 1 Piacenzian Firenze (IGF 15489 WEl-1 Museo di Storia Naturale e del Cetacea Balaenidae Balaena montalionis Skull 1 − Territorio dell'Università di Pisa (MSNUP I-12357) WEl4-1 Museo di Storia Naturale, Cetacea Balaenopterida- Indet. Partially articulated 1 1 Piacenzian Firenze (IGF 2800V) e? skeleton, incomplete (a few vertebrae, intervertebral disks, two radii, ulna, humerus, many costae) WEl5-1 Museo di Storia Naturale e del Cetacea Balaenidae Indet. Disarticulated mandible, 1 1 Piacenzian Territorio dell'Università di Pisa maxilla, occipital shield, (MSNUP I-16839) premaxilla, costae, zygomatic process of the squamosal WEl5-2 Museo di Storia Naturale, Cetacea Balaenidae Indet. Highly articulated skeleton, 1 1 1 1 Piacenzian Firenze (IGF 2801V) fairly complete, lacking lumbar and caudal vertebrae WEl6-1 Museo di Storia Naturale, Cetacea Balaenidae Idiocetus guicciardinii Skull fragments, bulla, 1 1 1 Piacenzian Firenze (IGF 16996) mandibles, three vertebrae,

8 22 costae, scapula, intervertebral disks. No information on articulation WEl6-2 Museo Civico di Montaione Cetacea Balaenidae Eubalaena sp. Partial skull 1 Piacenzian (M_Pal_002-005) WSi2-1 Museo di Storia Naturale, Cetacea Ziphiidae Tusciziphius crispus Partial skull 1 Zanclean Firenze (IGF 1594V) WSi2-2 Museo di Storia Naturale, Cetacea Ziphiidae Indet. (=Mesoplodon sp.) Incomplete skull 1 Zanclean Firenze (IGF 8736V) DSi2-1 Museo di Storia Naturale, Cetacea Delphinidae Etruridelphis giulii Highly articulated, highly 1 1 1 1 Zanclean Firenze (IGF 8736V) complete skeleton MSi2-1 Museo dell'Accademia dei Sirenia Dugongidae Metaxytherium subapenninum Fragmentary skull, 1 1 Zanclean Fisiocritici at Siena (496) mandible, costae MSi2-2 Museo di Storia Naturale, Sirenia Dugongidae Metaxytherium subapenninum Incomplete skull 1 Zanclean Firenze (IGF 2968V) MSi2-3 Museo di Storia Naturale, Sirenia Dugongidae Metaxytherium subapenninum Complete skull 1 Zanclean Firenze (IGF 13747)

MSi2-4 Museum National d'Histoire Sirenia Dugongidae Metaxytherium subapenninum Incomplete cranium, 1 1 Zanclean Earth-Science Reviews176(2018)xxx–xxx Naturelle, Paris, France incomplete mandible, rib (MUSNAF 4960, 4962) fragment WSi5-1 Museo dell'Accademia dei Cetacea Ziphiidae Indet. (=Mesoplodon sp.) Rostrum fragment 1 Piacenzian Fisiocritici at Siena (IG 4940) WSi5-2 Museo dell'Accademia dei Cetacea Mystecete Indet. 5 vertebrae Piacenzian Fisiocritici at Siena (IG 4401, 4417,4518, 4805?, 4945?) DSi5-1 Museo dell'Accademia dei Cetacea Delphinidae Indet. Left limb 1 Piacenzian Fisiocritici at Siena DSi5-2 Museo dell'Accademia dei Cetacea Delphinidae Indet. Teeth 1 Piacenzian Fisiocritici at Siena (IG 4300) DSi5-3 Cetacea Delphinidae Tursiops osennae 1 1 Piacenzian (continued on next page) .Dmnc tal. et Dominici S.

Table 2 (continued)

Marine Hosting institution Order Family Species Description A C C + PC Minimum Stage mammal abundance

Museo Geologico Giovanni Highly complete skull, Capellini (MGGC 8561–8562) mandible fragment, periotic, tympanic bullae, four cervical vertebrae PSi5-1 Repository unknown Carnivora Phocidae Pliophoca etrusca “Fragmentary specimen”. 1 Piacenzian No information on articulation or completeness WSi-1 Museo ‘G. Capellini’ (MGGC Cetacea Balaenidae Balaenotus insignis Highly complete skeleton 1 1 1 Zanclean/ 8922, 8923) (mandibles, jaw, vertebrae, Piacenzian ribs, elements of limbs). No information on articulation WSi-1 Museo dell'Accademia dei Cetacea Balaenidae Indet. Three vertebrae, one 1 Piacenziano Fisiocritici at Siena (IG mandible fragment, 4402–4408, 4418, 4420–4422) fragments WSi-3 Museo dell'Accademia dei Cetacea Ziphiidae Indet. (=Mesoplodon sp.) Rostrum fragment 1 Zanclean? Fisiocritici at Siena (IG 4941?, 4796?) DCh3-1 Museo ‘G. Capellini’ (MGGC Cetacea Delphinidae Orcinus citonensis Highly complete skeleton 1 1 1 Piacenzian 8560, 8563–8565) (skull, mandible, vertebrae, costae, elements of limbs). No information on articulation WCh3-1 Museo ‘G. Capellini’ (MGGC Cetacea Balaenidae Indet. (=Balaena etrusca) 8 vertebrae 1 Piacenzian

9 9066) WCh-1 Museo dell'Accademia dei Cetacea Balaenidae Indet. (=Balaenula Tympanic bulla, periotic, 11− Fisiocritici at Siena praediolensis) mandible, costa, maxillar fragment DCh4-1 Museo dell'Accademia dei Cetacea Delphinidae Stenella sp. Almost complete skeleton. 1 1 1 1 Piacenzian Fisiocritici at Siena No information on articulation WCh6-1 Museo Civico, Cetona Cetacea Balaenopteridae Indet. Articulated skeleton, one 1 1 1 1 Piacenzian complete limb, most vertebrae, skull. Pristine cortical layer Abundance (N): 9 17 24 64 Frequency (%): 14 27 38

Marine Biozone Basin Synthem Locality Municipality Lithofacies Sequence Reference Shell bed Sample mammal stratigraphy Earth-Science Reviews176(2018)xxx–xxx WFi4-1 MPL5a Fine 4 Case Nuove Orciano Pisano Shelf mudstone HST Dominici et al., Archimedella bed OP1-13 2009 WFi4-2 MPL4b/MPL5a Fine 4 Pieve Santa Santa Luce Shelf mudstone HTS Cioppi, 2014 Lucinoma bed OP17 Luce WFi4-3 MPL4b/MPL5a Fine 4 Santa Luce Santa Luce −−Bianucci and −− Landini, 2005 WFi4-4 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano Shelf mudstone HST Ugolini, 1899; −− Bianucci et al., 2009b; Bianucci and Sorbini, 2014 WFi4-5 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano Shelf mudstone HST Bianucci and −− Sorbini, 2014

(continued on next page) .Dmnc tal. et Dominici S.

Table 2 (continued)

Marine Biozone Basin Synthem Locality Municipality Lithofacies Sequence Reference Shell bed Sample mammal stratigraphy

WFi4-6 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano Shelf mudstone HST Bianucci and −− Landini, 2005; Sarti and Lanzetti, 2014 WFi4-7 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano Shelf mudstone HST Bianucci, 1999, −− Bianucci and Sorbini, 2014 WFi4-8 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano Shelf mudstone HST Ugolini, 1907; −− Bianucci, 1999; Bianucci and Sorbini, 2014 WFi4-9 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano Shelf mudstone HST Bianucci, 1999; −− Bianucci and Sorbini, 2014 WFi4-10 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano Shelf mudstone HST Bianucci and −− Landini, 2005; Cioppi, 2014 WFi4-11 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano −−Higgs et al., −− 2012 DFi4-1 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano Shelf mudstone HST Sarti and −− Lanzetti, 2014 DFi4-2 MPL4b/MPL5a Fine 4 Greppoli Lorenzana Shelf mudstone HST Lawley, 1876; −− Bianucci et al.,

10 2009b; Sarti and Lanzetti, 2014 DFi4-3 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano Shelf mudstone HST Cioppi, 2014 −− DFi4-4 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano Shelf mudstone HST Cioppi, 2014 −− DFi4-5 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano Shelf mudstone HST Bianucci et al., −− 2009b; Cioppi, 2014 MFi4-1 MPL4b/MPL5a Fine 4 Orciano Pisano Orciano Pisano −−Previously −− unpublished PFi4-1 MPL5a Fine 4 Case Nuove Orciano Pisano Shelf mudstone HTS Ugolini, 1902; Archimedella bed OP1-13 Berta et al., 2015 WOm2-1 MPL2 Ombrone 2 Poggio alle Montalcino Shelf mudstone MFS Danise and Haustator bed MON1-2 Mura Dominici, 2014 MOm2-1 MPL2 Ombrone 2 Arcille Grosseto Shelf mudstone MFS Tinelli et al., Haustator bed − 2012 MOm2-2 MPL2 Ombrone 2 Arcille Grosseto Shelf mudstone MFS Sorbi et al., Haustator bed −

2012 Earth-Science Reviews176(2018)xxx–xxx MOm2-3 MPL2 Ombrone 2 Arcille Grosseto Shelf mudstone MFS Tinelli et al., Haustator bed − 2012 MOm2-4 MPL2 Ombrone 2 Arcille Grosseto Shelf mudstone MFS Tinelli et al., Haustator bed − 2012 MOm2-5 MPL2 Ombrone 2 Camigliano Montalcino Shelf mudstone MFS Sorbi et al., Haustator bed CAM1 2012 WEr1-1 MPL1 Era 1 Podere Volpaia Saline di Shelf mudstone − Pilleri, 1987 −− Volterra WEr1-2 MPL1 Era 1 Saline di Saline di Shelf mudstone − Pilleri, 1987 −− Volterra Volterra PEr1-1 MPL1 Era 1 Saline di Saline di Shelf mudstone TST Berta et al., −− Volterra Volterra 2015

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Table 2 (continued)

Marine Biozone Basin Synthem Locality Municipality Lithofacies Sequence Reference Shell bed Sample mammal stratigraphy

WEl3-1 MPL4b? Elsa 3-2 Fornace Silap Castelfiorentino Shelf mudstone HST Danise and Dendrophyllia CST3-4 Dominici, 2014 bed DEl3-1 MPL4b? Elsa 3-2 Canonica Certaldo Shelf mudstone HST Alderighi et al., Serpulorbis bed − 2011 DEl3-2 MPL4b? Elsa 3-2 San Giorsolè Certaldo Shelf mudstone HST Arbeid et al., Serpulorbis bed − 2015 WEr4-1 − Era 4 La Rocca Volterra Shelf mudstone HST Bianucci et al., −− 1998 WEr4-2 − Era 4 La Rocca Volterra Shelf mudstone HST Bianucci et al., −− 1998 DEr4-1 − Era 4 La Rocca Volterra Shelf mudstone HST Bianucci et al., −− 1998 DEr4-1 − Era 4 La Rocca Volterra Shelf mudstone HST Bianucci et al., −− 1998 WEl-1 − Elsa − Casina Montaione −−Bianucci and −− Sorbini, 2014 WEl4-1 MPL5a Elsa 4-2 Castel San Castel San −−Elisabetta − CCN1-2 Gimignano Gimignano Cioppi, pers. comm., 2017 WEl5-1 MPL5a Elsa 5-2 Casenuove Empoli Intertidal MFS Dominici et al., Bittium bed − mudstone 1995; Bianucci, 1995; Collareta et al., 2016

11 WEl5-2 MPL5a Elsa 5-4 Poggio Tagliato San Miniato Shelf mudstone HST Borselli and Glycymeris bed CP2 Cozzini, 1992; Bianucci et al., 2002; Danise and Dominici, 2014 WEl6-1 MPL5a Elsa 6 Villa Montopoli Shelf mudstone HST Capellini, 1905; Pteria bed SRR3 Guicciardini Cioppi, 2014; Steeman, 2010 WEl6-2 MPL5a Elsa 6 Rio Ricavo Palaia −−Bisconti, 2002; −− Sorbini et al., 2014 WSi2-1 MPL3/MPL4a Siena 2/3 Val di Pugna Siena Delta sandstone TST Bianucci et al., Panopea bed − 2001; Cioppi, 2014 WSi2-2 MPL3/MPL4a Siena 2/3 Val di Pugna Siena −−Bianucci, 1997b −− DSi2-1 MPL3 Siena 2 Lucciolabella Pienza Outer shelf or HST Bianucci et al., −−

upper slope 2009b Earth-Science Reviews176(2018)xxx–xxx mudstone MSi2-1 MPL3/MPL4a Siena 2/3 Val di Pugna Siena Delta sandstone TST Bianucci et al., Panopea bed − 2001; Sorbi et al., 2012 MSi2-2 MPL3/MPL4a Siena 2/3 Val di Pugna Siena Shelf mudstone HST Bianucci et al., −− 2001; Sorbi et al., 2012 MSi2-3 MPL3/MPL4a Siena 2/3 Val di Pugna Siena −−Sorbi et al., −− 2012 MSi2-4 MPL3/MPL4a Siena 2/3 Val di Pugna Siena −−Sorbi et al., −− 2012 WSi5-1 MPL4b/MPL5a Siena 5 Guistrigona Asciano Shelf mudstone HST − CGR1 (continued on next page) .Dmnc tal. et Dominici S.

Table 2 (continued)

Marine Biozone Basin Synthem Locality Municipality Lithofacies Sequence Reference Shell bed Sample mammal stratigraphy

Martini et al., 2011; Manganelli and Benocci, 2014 WSi5-2 MPL4b/MPL5a Siena 5 Guistrigona Asciano Shelf mudstone HST Martini et al., − CGR1 2011; Manganelli and Benocci, 2014 DSi5-1 MPL5a Siena 5 Cava i Sodi Asciano Shelf mudstone HST Martini et al., − SD1 2016; Manganelli and Benocci, 2014 DSi5-2 MPL5a? Siena 5? Troiola Siena Shelf mudstone HST Manganelli and −− Benocci, 2014 DSi5-3 MPL5a? Siena 5? San Quirico Siena Shelf mudstone HST Bianucci et al., −− d'Orcia 2009b; Sarti and Lanzetti, 2014 PSi5-1 MPL4b/MPL5a Siena 5 Castelnuovo Castelnuovo Shelf mudstone HST Martini et al., − CGR1 Berardenga Berardenga 2011, 2016; Bianucci et al., 2009a; Berta et al., 2015 WSi-1 MPL4 Siena 3 Poggiarone Castelnuovo Shelf mudstone HST Martini et al., −−

12 Berardenga 2011; Sarti and Lanzetti, 2014 WSi-1 MPL4b Siena 4 Siena (Vicolo di Siena Nearshore − Costantini et al., −− Tone) sandstone 2009; Manganelli and Benocci, 2014 WSi-3 − Siena 2/3 Fangonero? Siena −−Bianucci et al., −− 2001; Manganelli and Benocci, 2014 DCh3-1 − Chiana 4/5 Poltriciano Cetona −−Capellini, 1883; −− Sarti and Lanzetti, 2014 WCh3-1 − Chiana 4/5 Poggio di Chiusi −−Capellini, 1876; −− Pasqualone Sarti and Lanzetti, 2014 WCh-1 − Chiana − Palazzone San Casciano dei −−Manganelli and −−

Bagni Benocci, 2014 Earth-Science Reviews176(2018)xxx–xxx DCh4-1 − Chiana 4? Guazzino Sinalunga Shelf mudstone HST Manganelli and −− Benocci, 2014 WCh6-1 − Chiana 6 Sinalunga Sinalunga Delta sandstone FSST Cioppi, 2014 −− S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx

Fig. 3. Abundance of fossil MM records in major museums of Tuscany, distributed by locality of provenance. Each record ranges from a single fragment or tooth, to articulated, nearly complete skeleton. A: mysticetes and odontocetes; B: large sharks; C: sirenians — scuba diver for scale in each figure. Symbols for basins as in Fig. 1. performed an analysis of similarity (ANOSIM). To interpret the out- delta-front hyperpycnal sandstones and conglomerates. In a very broad comes of the quantitative study and the significance of clusters we used sense, the concept of S2 is extended to these basins, by assuming that species-level autoecologic information available for the most abundant erosional unconformities in EB pass into correlative conformities in species, based on the distribution of extant forms. This information, rapidly subsiding adjacent basins, where thick successions could be retrieved from the Marine Biodiversity and Ecosystem Functioning EU accommodated. Apart from exceptions, no shells were found in bathyal website (MARBEF: www.marbef.org), included the average life depth of mudstone or in deltaic sandstone. In OOB, the same time span is 23 modern species that in our dataset had an overall abundance > marked by a N-S facies gradient characterised by a single deepening- 0.15%. upward succession, from fluvio-deltaic sandstone to shelfal mudstone, replaced by a succession made of four distinct regressive-transgressive 4. Results units in the Orcia valley to the north (Ghinassi and Lazzarotto, 2005; Benvenuti et al., 2015). A laterally-continuous shell bed, with sharks 4.1. Unconformity-bounded units remains and skeletons and articulated bones of whales, sirenians and large teleost fishes (Danise, 2010; Sorbi et al., 2012; Tinelli, 2013), 4.1.1. Synthem S1: the early Zanclean transgression marks a major transgressive surface overlain by open shelf mudstones The Miocene-Pliocene transition, marking the return to marine ( Sorbi et al., 2012; Tinelli, 2013: biozone MPL2) that is hypothetically conditions after the Messinian salinity crisis (Krijgsman et al., 1999), is traced along a NE-SW profile (Figs. 4, 5). recorded in limited exposures of earliest Zanclean, open marine mud- stones resting both unconformably or conformably onto latest Messi- 4.1.3. Synthem S3: Zanclean-Piacenzian transition nian non-marine deposits (Lago-Mare), an isochronous boundary being Synthem S3 is subdivided in EB into a lower and an upper interval dated in the Mediterranean at 5.33 Ma (Roveri et al., 2014). Differences (Benvenuti et al., 2014). The lower division is represented by dee- depend on the specific structurally-controlled distribution of hinterland pening-upward, coarse-grained delta front system, overlain by an upper basins, where an uplifting chain determined the presence of thresholds mudstone division from an open shelf setting. The upper part is rich delaying the early Zanclean marine flooding from inner (EB, OOB: with shell beds, and occasional articulated whale skeletons, associated Benvenuti et al., 2015) to outer hinterland basins (FB, VEB). In inner with shark teeth, have been recovered (Danise and Dominici, 2014). basins such as EB, where S1 has been defined, continental deposition Similar shelf mudstones of the MPL4 biozone crop out in VEB and SRB continued into the earliest Zanclean, marine flooding occurring within (Bossio et al., 1993; Riforgiato et al., 2005) whereas biostrati- the MPL1 biozone (references in Benvenuti et al., 2015). A chronos- graphically equivalent mudstones in FB testify to an upper epibathyal tratigraphical equivalent of S1 is represented in FB and VEB by an open- paleoenvironment. shelf mudstone, conformably resting on latest Messinian Lago-Mare deposits (Roveri et al., 2014). An apparently analogous situation is 4.1.4. Synthem S4: early-middle Piacenzian warm climate and high sea- documented in OOB, where MPL1 shelfal mudstones of the early Zan- level clean (normal chron C3n) rest on a Messinian to basal Pliocene paleo- Synthem S4 (Benvenuti et al., 2014) has been recognised in FB and valley fill (Benvenuti et al., 2015). VEB by facies similarities and chronostratigraphic correlation. In EB, S4 comprises a lower interval dominated by richly fossiliferous, massive 4.1.2. Synthem S2: Zanclean differential preservation mudstone or very-fine-grained sandstone (prodelta-inner shelf), over- Synthem S2, as recognised in the EB (Benvenuti et al., 2014), is lain by bioclast-rich sandstones recording prograding mixed carbonate- represented by relatively thin fluvial conglomerates unconformably clastic ramp, outcropping in the southeastern part of the Elsa valley. resting on S1 (biozones MPL1-MPL2), capped by S3 (biozone MPL4a: Equivalent deposits, also comprised in biozone MPL4b, are patchily Bossio et al., 1993), comprising important stratigraphic gaps at its base distributed in FB, VEB, SRB, OOB and other basins of southern Tuscany and top. On the other hand, in FB (Bossio et al., 1997), VEB (Bossio (Ghinassi and Nalin, 2016). The upper interval of S4 is formed by a et al., 1994) and SRB (Ghinassi and Lazzarotto, 2005), the same succession of delta front sandstones, passing in EB eastern margin to a chronostratigraphic interval is recorded by monotonous epibathyal few tens of m-thick fluvial succession, hinting at an original deposi- mudstones several hundred meters thick, locally intercalated with tional gradient. S4 is apparently missing due to erosion north of San

13 S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx

Fig. 4. - Detailed sedimentary logs measured at three lo- calities, representing three different stratigraphic contexts for the large marine vertebrate fossil record of the Tuscan Pliocene. The succession at Orciano Pisano is included in synthems S3–S4 of FB, at Arcille-Poggio alle Mura- Camigliano in synthem S2 of OOB, at Sinalunga in synthem S6 of CB. See Figs. 1–2 for the location of the numbered localities and references in the main text for facies analysis and sequence stratigraphy of synthem S5. Arcille log from Tinelli et al. (2012), and Tinelli (2013).

Gimignano (EB), and around Lajatico (VEB). Biostratigraphic data grained sediments, to open shelf mudstones, sometimes topped by a allow to refer S4 to the upper part of biozone MPL4b and the lower part regressive shoreface or delta sandstone, other times directly overlain by of MPL5a, thus comprising the lower-middle part of the Piacenzian, the next sequence through a sharp contact. Fluvial, brackish-water, and globally characterised between 3.264 and 3.025 Ma by warm climate intertidal deposits mark the lower part of each composite sequence, and relatively high sea level (Raymo et al., 2009; Dowsett et al., 2013; usually topped by a laterally-continuous shell bed, from a few cm to a Prista et al., 2015). few dm-thick, representing a surface of transgression. Shell beds are particularly well-developed around the middle part of sequences, where they separate shoreface and delta sandstones from overlying open shelf 4.1.5. Synthem S5: mid-Piacenzian high-frequency sea level variation mudstones, marking the time of maximum flooding. Large marine Synthem S5, recognised in EB and VEB, is bounded below by an vertebrates, including articulated whale skeletons and large sharks erosional unconformity that cuts deeply into underlying units, bringing (Danise and Dominici, 2014), are often recovered both at deposits of – S5 directly on top of S3 (EB: log 15; VEB: logs 5 6inFig. 2). S5 is up to the MF interval and in overlying mudstone (Fig. 2). Towards the north- about 200 m in EB, where it has been subdivided into a hierarchy of eastern margin of EB, cyclothemic fluvial conglomerates, sandstones small-scale depositional sequences (Benvenuti et al., 2007; Dominici and mudstones replace coastal and fully marine deposits, testifying to et al., 2008: see following paragraphs). Each composite depositional an original facies gradient. In the central part of EB, composite se- sequence forms a tens-of-m-thick asymmetric sedimentary cycle, com- quences are stacked to form a deepening-upward succession, with a posed by a deepening-upward part, from fluvial or coastal coarse-

Fig. 5. Taphonomy of large marine vertebrates at Poggio alle Mura (Panels A–D) and Arcille (Panel E, see Fig. 4 for the sequence stratigraphic and sedimentary context; plan view of the sirenian skeleton is modified from Tinelli et al., 2012), synthem S2. A: Plan view of the Poggio alle Mura undetermined balaenid (WOm2-1 in Table 2). Parts of the skeleton are quasi-articulated, others are scattered, but not far from the original position. B: Quasi-articulated ver- tebrae of the same specimen, lying on top of a Haustator shell bed. C: Side view of a vertebra on top of the densely- packed shell bed. D. Detail of the shell bed, in top view. The turritelline gastropod Haustator vermicularis is visible in the upper left, a large fragment of wood in the lower right, with the inchnofossil Teredolithes produced by wood-dwelling teredinid bivalves, in the center of the photograph. E: Plan view of one of the Arcille sirenian specimens of Metax- itherium appenninicum (MOm2-1 in Table 2). Same scale as in Panel A, the arrow points to the North.

14 S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx topmost thick and laterally-continuous open shelf mudstone interval, 4.3.1. Zanclean EDS (synthems S1–S3) directly onlapping the S4-S5 basal unconformity on the eastern EB (log In most basins, deposition of synthems S1-S2-S3 takes place at 15, Fiano: Fig. 2). outer-shelf or bathyal depths, well below the point on a depositional profile where the rate of relative sea level change is zero (equilibrium point). Here the sediment supply is not sufficient to fill the available 4.1.6. Synthem S6: Piacenzian-Gelasian climate change and regression accommodation space and an aggradational style of deposition prevails, As S5, synthem S6 is also built through a hierarchy of small-scale with the result that in most Zanclean settings smaller cycles of sea level depositional sequences, better expressed in EB, but also documented in variations are not marked by a facies change. The sharp facies change fl fi SRB and CB. In EB, uvial coarse-grained sandstones ll a deep valley recorded where the monotonous muddy deposition is interrupted by – incised in S5 deposits (logs 8 10 in Fig. 1), resting on the basal un- turbidite sandstone and conglomerate, is connected with synthem conformity of S6. Intertidal or coastal lagoon deposits form the trans- boundaries and major tectonic phases of restructuring of the region. gressive systems tract of the composite depositional sequence. A lat- MM and shell beds are practically absent. This situation reverses in the fi erally-continuous shell bed testi es to the MF of S6, topped by Orcia-Ombrone basin, where depths of deposition are shallower and highstand shoreface and delta front sandstones. The Piacenzian-Gela- EDS are expressed. At Arcille deltaic sandy conglomerates and sand- sian boundary, corresponding to a major climatic transition from stones (facies F3) are overlain by a fluvial cross-bedded sandstone warmer-moister to colder-drier conditions (Benvenuti et al., 1995b, (facies F1a), separated by a TS from an overlying bioturbated shallow 2007), is marked by the Gauss-Matuyama reversal detected at Mon- marine sandstone. The surface overlying a Haustator vermicularis shell topoli (Lindsay et al., 1980). Highstand marine sandstones are char- bed (Danise, 2010; Tinelli, 2013) forms the MFS separating the lower acterised by the recovery of two mysticete skeletons, at Montopoli (EB, shoreface sandstone from an open marine mudstone with scattered see Capellini, 1905) and Sinalunga (CB, Fig. 7). Fully continental en- shells (facies F5), marking a sudden and prolonged deepening of the vironments were established throughout the Gelasian in all basins here basin (biozone MPL2: Sorbi et al., 2012; Tinelli, 2013). The succession under study (e.g., Benvenuti and Del Conte, 2013; Benvenuti et al., is topped by deposits from shallower depths, expression of the FSST, 2014; Bianchi et al., 2015). below the upper SB (Fig. 4). Similar small-scale depositional sequences, expressed through fining-upward cycles ≥40 m-thick, are also present 4.2. Sedimentary facies and facies associations in synthem S3 at Case al Poggio, near Siena (biozones MPL3-MPL4a: Bianucci et al., 2001) and at Castelfiorentino (biozone MPL4b: Seven groups of siliciclastic facies and one group of carbonate fa- Benvenuti et al., 2014). cies, with very different fossil content, have been recognised (Table 1). – Siliciclastic facies form a paleoenvironmental gradient from terrestrial 4.3.2. Piacenzian EDS (synthems S4 S6) to marine and, in the case of marine facies, from shallowest to deepest As depth of deposition shallows during the Piacenzian, and ampli- (Fig. 8a). Facies types are fluvial conglomerate and sandstone, and al- tude of cycles of sea level variation increases, the cyclic stacking of EDS luvial mudstone (F1); intertidal to very shallow subtidal mudstone and becomes the typical depositional theme (Benvenuti et al., 2007, 2014; muddy sandstone (F2); shallow subtidal coarse- and medium-grained Dominici et al., 2008). In FB, the northwesternmost basin, depths re- sandstone with sparse conglomerate (F3); deep subtidal muddy fine- main considerable and facies change is more subtle. Pliocene at Orciano “ ” grained sandstone (F4); open shelf sandy mudstone (F5); outer shelf to Pisano traditionally belongs to the Blue Clays formation (Bossio et al., upper bathyal mudstone (F6); outer shelf and bathyal turbidite sand- 1997), but two distinct bodies where evidenced since the late nine- stone and conglomerate (F7). An eighth group is formed by facies de- teenth century (D'Ancona, 1871). The lower one is formed by grey fi posited subtidally in limited mixed carbonatic-siliciclastic ramps (F8), claystone with very rare shells, the upper one by muddy, very- ne irrespective of depth (Nalin et al., 2016). Facies F6–F7 are restricted to grained grey sandstone richly fossiliferous. The lower part of the latter Zanclean deposits (synthems S1–S3); facies F5 and F8 characterise the interval outcrops at the foothill of the small town of Orciano Pisano, upper Zanclean-middle Piacenzian interval (synthems S3–S4); facies around the locality Case Nuove (Bianucci and Landini, 2005; Berta F1–F4 characterise the upper Piacenzian (synthems S5–S6). The OOB et al., 2015). Here a laterally persistent shell bed is dominated by the succession, characterised by facies F1–F4, is an exception within the turritellid Archimediella spirata, overlain by a 25 m-thick monotonous Zanclean. sandstone interval with intercalated shell beds or sparse shells, be- coming muddier upward. The surface overlying the Archimediella shell bed is interpreted as the TS of an EDS, coinciding with the lower SB of a 4.3. Elementary depositional sequences (EDS) lower Piacenzian EDS. The overlying muddy fine-grained sandstone is the TST and HST (biozone MPL5a: Dominici et al., 2009; Fig. 6). At Both fluvial (Benvenuti and Del Conte, 2013) and marine facies shallower settings, in all basins to the East and South of FB, EDS of groups (siliciclastics: Benvenuti et al., 2007, 2014; carbonates: Nalin synthem S4 take the form of an alternation of mudstone and carbonate et al., 2016) are stacked to form facies associations which record cyclic (facies F5 and F8: Nalin et al., 2016), or mudstone and sandstone (facies variations of depositional and environmental conditions in response to F5 and F3–F4: Benvenuti et al., 2014; biozone MPL5a). Middle and a change in accommodation space. Physical surfaces and the inter- upper Piacenzian EDSs form an alternation of coastal mudstone and vening deposits allowed subdividing depositional sequences in systems sandstone (facies F1-F2-F3: Benvenuti and Dominici, 1992; Dominici, tracts (Benvenuti et al., 2007; Dominici et al., 2008). At the simplest 1994), with MF interval and HST marked by a shell bed topped by a scale, these hybrid facies associations form elementary depositional lower shoreface sandstone, or a shelf mudstone (facies F4-F5: Benvenuti sequences, up to 10–20 m thick, in their turn stacked to form composite et al., 2007; Dominici et al., 2008). sequences (original concepts from Mutti et al., 1994). This hierarchy is particularly evident in synthems S5–S6, formed at a time of pronounced 5. Distribution of large marine vertebrates glacio-eustatic oscillations and expressed around coastal settings, where maximum facies contrast allows for the expression of subtle cycle of sea 5.1. Geographic distribution level variation (e.g., Benvenuti and Dominici, 1992; Benvenuti et al., 2007; Dominici et al., 2008). Analogue sharp facies contrast within The MM geographic distribution is listed in Table 2. All MM that is Zanclean EDS in OBB (Tinelli, 2013), but otherwise absent in deeper geographically located, irrespective of stratigraphy, is plotted in Fig. 3. sediments (facies F5 and F7). EDSs have different expressions de- The largest number is recovered in FB in northwestern Tuscany, with a pending on the time interval and the sedimentary basin. peak at a few sites around the small town of Orciano Pisano, in the Pisa

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Fig. 6. Taphonomy of a 10 m-long, undetermined balae- nopterid at Orciano Pisano (see Fig. 4 for the sequence stratigraphic and sedimentary context), synthem 4. A: Pla- nimetry of the quasi-articulated and nearly complete ske- leton. B: Detail of the central part of the skeleton in the field. The cortex layer of vertebrae and flipper bones is badly consumed, whereas some of the costae are still pris- tine. C: Lateral view of a turritellid shell bed, below, and the surface where the whale skeleton lied (dashed line), about 15 cm above the shell bed. The sediment is a very fine-grained silty sand, completely bioturbated (large ver- tical burrows are visible). D: Top view detail of the turri- tellid shell bed. At the center a valve of Yoldia nitida, sur- rounded by a few specimens of the turritellid Archimediella spirata.

province. This coincides with the highest number of known species, 5.2.2. Synthem S3 including mammals, elasmobranchs, turtles, and large bony fishes. MM The upper part of S3 yielded an articulated and well-preserved ba- is also abundant in the province of Siena, around Volterra (VEB; lenopterid skeleton found in a 30 m-thick mudstone succession at Bianucci and Landini, 2005), and around San Quirico, particularly rich Castelfiorentino, in the Elsa basin (EB, WEl3-1). These strata are richly in elasmobranchs according to the available data (SRB; Marsili, 2006). fossiliferous, with several shell beds with epifaunal cemented taxa, such A fourth basin with a consistent number of findings is OOB, where as vermetid gastropods, oysters and corals (Facies F5), in a normal- cetaceans, sharks, sirenians and large teleost fishes have been un- polarity magnetostratigraphic interval (Gauss chron: Andrea Albianelli, earthed. personal communication, 1999). Based on physical stratigraphic cor- relation, this can be assigned to a lower Piacenzian HST. A skull of Hemisyntrachelus sp. (Alderighi et al., 2011) and partly articulated 5.2. MM stratigraphic distribution vertebrae and costae of a dolphin skeleton (Arbeid et al., 2015) were recently excavated a few hundred meters apart one from the other, 5.2.1. Synthems S1–S2 along a monotonous mudstone S3 succession intercalated with several In the vicinity of Saline di Volterra, one of the sites with remains of Ostrea and Serpulorbis shell beds, near Certaldo (facies F5, DEI3-1-2 in Pliophoca etrusca (Berta et al., 2015) and sperm whale (Table 2), S1 is Table 2). Overall, specimens of marine mammals attributed to synthem represented by a bathyal mudstone (facies F6, biozone MPL1). Lower- S3 are eleven (some are uncertain and may come from the upper part of middle Zanclean MM is otherwise absent, with the exception of S2 in S2: Table 2). OOB, where large marine vertebrates are concentrated at the maximum flooding interval, outcropping at Poggio alle Mura (Danise, 2010), Camigliano (Sorbi et al., 2012) and Arcille (Tinelli et al., 2012; Tinelli, 5.2.3. Synthem S4 2013), in biozone MPL2. At Poggio alle Mura a slightly disarticulated Synthem S4 yielded the highest abundance and species-richness of balaenid whale skeleton (WOm2-1 in Table 2) has been excavated in the Tuscan MM. In the Fine Basin, the locality of Orciano Pisano is contact with the laterally-persistent Haustator shell bed at the maximum represented in Table 2 by 18 records of whales, dolphins (a partial flooding interval (Figs. 4, 5; Danise, 2010; Tinelli, 2013). At Camigliano skeleton: Bianucci, 1996; Bianucci et al., 2009b), seals (Berta et al., and Arcille, tens of km from Poggio alle Mura, the Haustator shell bed is 2015), tens of other unidentified cetacean elements, hundreds of shark associated with other articulated skeletons and isolated MM remains, teeth and vertebrae (Fig. 3), sea turtles, large bony fishes, and sea birds including several specimens of the sirenian Metaxytherium sub- hosted in museum collections (Bianucci and Landini, 2005; Marsili, apenninum (MOm2-1-5), large bony fishes, rays, and sharks Carcharias 2007; Cioppi and Dominici, 2011). In locality Case Nuove, a single taurus, Carcharhinus sp., Galeocerdo cuvieri, and Squatina sp. (Sorbi middle Piacenzian transgressive surface has yielded a whole, articu- et al., 2012; Tinelli et al., 2012; Tinelli, 2013). A partial skull of the lated whale skeleton (Figs. 4, 6), teeth of blue and white sharks and delphinid Etruridelphis giulii, with right and left dentaries fractured, but bones of sea birds (Dominici et al., 2009). From the same site comes a nearly complete and with most teeth still in their alveoli (DSi2-1 in skeleton of the monk seal Pliophoca etrusca (Berta et al., 2015), and Table 2), was recovered southeast of Siena, near Chianciano Terme possibly many other museum specimens labelled “Orciano Pisano”, (SRB) at the top of a mudstone (facies F6) intercalated with turbiditic suggesting that this interval forms the most prolific bonebed of the sandstone beds (Facies F7), topped by a monotonous mudstone interval region (Table 2). A few cm above the Archimediella shell bed, glauco- (upper Zanclean, uppermost part of biozone MPL3: Bianucci et al., nitic and deeply bioeroded whale bones (Danise, 2010), associated with 2009b). In the same basin, isolated bones of two undetermined beaked ichnological evidence of the activity of Osedax bone-eating worms (see whales and bones of four different specimens attributed to Metax- Higgs et al., 2012), lie in a bioturbated muddy, fine-grained sandstone ytherium subapenninum were found in deltaic sandstones, in the middle with a complex boxwork of Ophiomorpha and Thalassinoides trace fossils of a fining-upward succession, at the boundary between biozones MPL3 (6Fig.), associated with a diverse paleocommunity of molluscs and and MPL4a (Bianucci et al., 2001; WSi2-1-2 and MSi2-1-4 in Table 2), other benthic invertebrates with complex trophic connections suggesting that findings at the two SRB localities belong to the same MF (Dominici et al., 2009; Danise et al., 2010). At the boundary between interval, in the upper part of S2. Three specimens of marine mammals FB and VEB, an incomplete skull and skeleton of the dolphin Etrur- were identified in synthem S1, seven in synthem S2. idelphis giulii was recovered near Lorenzana (Lawley, 1876; Bianucci,

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Fig. 7. Taphonomy of an incomplete, undetermined mys- tecete (WEl4-1 in Table 2) at Castel San Gimignano, syn- them 4, comprising articulated torso elements. A: Plani- metry of the articulated elements. B: Detail of one of the limbs in the field (trowel for scale = 22 cm): humerus, ra- dius and ulna are in anatomical relationship; the cortex layer is well preserved, suggesting quick burial of the car- cass. C: Bones of the chest region; on the background the massive sandstone associated with the fossil whale. Ar- ticulated shoreface pectinid bivalves (Pecten flabelliformis) were interspersed in the sandstone.

1996; Bianucci et al., 2009b), at a locality associated with a muddy a shelfal deposit: Martini et al., 2016). Judging from historical accounts sandstone interval (facies F4) in synthem S4. An incomplete and ar- (Capellini, 1883), the killer whale Orcinus citonensis (DCh4-1) was re- ticulated mysticete was recovered in a sandstone at San Gimignano, covered in a sandy unit lying on top of a thick mudstone interval (S3 or associated with pectinid bivalves (Fig. 7; facies F3; Elsa Basin, EB: S4) and is tentatively assigned to S5. Specimens of marine mammals Danise and Dominici, 2014), here tentatively assigned to the uppermost attributed to synthem S5 are 10. part of the synthem (WEl4-1 in Table 2). Overall, specimens of marine mammals attributed to synthem S4 are 28. 5.2.5. Synthem S6 A fairly complete juvenile skeleton of the large balaenid Idiocetus 5.2.4. Synthem S5 guicciardinii was recovered in the second half of the 19th Century in the The next MM-rich stratigraphic interval is Piacenzian deposits of EB near Montopoli Valdarno (Capellini, 1905), in open shelf strata at- fl synthem S5. An incomplete and disarticulated balaenid skeleton was tributable to the interval of maximum ooding of S6 (WEl6-1 in found in intertidal deposits of the lower part of the synthem, at Table 2). The MFS of S6, of uppermost Piacenzian age, is marked in EB Casenuove (facies F2, EB; Bianucci, 1995; Collareta et al., 2016; WEl5-1 by a laterally continuous Pteria phalenacea shell bed, with a high-di- in Table 2). A large balaenid was recovered higher up section, a few versity association of macroinvertebrates, including a rich decapod meters above a laterally-continuous very thick and complex Haustator paleocommunity (Garassino et al., 2012). In the vicinity, near Palaia, a vermicularis shell bed, up to 2 m-thick (Benvenuti et al., 1995a), traced right whale (Eubalena sp., WEl6-2 in Table 2) was recovered in 1974 in laterally for 2 km to the east of San Miniato (“Turritella strata”, De sandy mudstones, in association with mollusc shells and teeth of the Stefani, 1874), and forming a surface of transgression within the TST of great white shark (Carcharodon carcharias: Bisconti, 2002; Sorbini et al., S5 (Benvenuti et al., 2007, 2014; Dominici et al., 2008). The MF in- 2014). A tightly articulated balaenopterid skeleton was found at Sina- terval is formed by a Glycymeris insubricus shell bed, separating around lunga (WCh4-1 in Table 2), in deltaic sandstones and conglomerates San Miniato shoreface sandstone (facies F3 or F4) from offshore mud- (Fig. 8). Marine mammal specimens attributed to synthem S6 are thus stone (F5; Benvenuti et al., 2007). The balaenid skeleton is almost ar- three. ticulated and bioeroded, closely associated with teeth of the great white shark and other scavengers (Borselli and Cozzini, 1992; Bianucci et al., 5.3. MM facies type distribution 2002; Danise and Dominici, 2014), in the early HST of synthem S5 (Benvenuti et al., 2007; Dominici et al., 2008; WEl5-2 in Table 2). The In an ideal deepening-up gradient, multi-element findings of marine Glycymeris shell bed can be traced laterally for several km. In the vi- mammals are very rare in intertidal and very shallow subtidal pa- cinity of Fiano, it includes bioeroded and encrusted gravels inherited leoenvironments (facies type F2, 1,5%: Fig. 9A), moderately re- from underlying successions, interpreted as ravinement deposits. The presented in delta or shoreface sandstones (facies type F3, 4,6%), most interval of maximum flooding is marked by the stacking of at least three abundant in sandy mudstone of open shelf settings (facies type F5, distinct shell beds, all including a high-diversity association with 70%), rare in outer shelf and bathyal sediments (facies type F6–F7, bioeroded and encrusted shells. This situation suggests that balaenid 1,5%). The most pristine and complete skeletons are associated with WEl5-2 lies in correspondence to an interval of low rates of sedi- gravelly well-sorted sands from event sedimentation, suggesting a ne- mentation. In SRB, near Castelnuovo Berardenga, shelfal mudstones gative relationship between taphonomic loss and sedimentary processes (facies type F5) have yielded MM remains at a few localities. Delphinid at delta fronts. In the tightly articulated and pristine Sinalunga balae- remains were found at the “I Sodi” quarry and at Troiola (DSi5-1-2, in nopterid (WCh4-1 in S4-CB), the cortical surface of the tightly-con- Table 2). Bones of a beaked whale and undetermined mysticetes are nected vertebrae is practically intact, and carpal, metacarpal and pha- reported from Guistrigona (Manganelli and Benocci, 2014) and a langes of the flipper are in perfect anatomical connection, as if a sudden fragmentary specimen of the monk seal Pliophoca etrusca from Cas- depositional event buried a fresh carcass (Fig. 8; similar pristine ske- telnuovo Berardenga (Berta et al., 2015). A very rich shark fauna, in- leton are found in deep water turbiditic successions: Stinnesbeck et al., cluding sawsharks, thresher, frilled, bluntnose sixgill, bramble, gulper, 2014). Another pristine and tightly articulated skeleton, belonging to a kitten, sand tiger, shorten mako, basking and requiem sharks, an as- killer whale (WCh3-1 in CB), was collected at Cetona in the second half sociation suggesting an upper slope paleoenvironment for the sur- of the 19th century, in a locality associated with sandstones, also pos- roundings of Castelnuovo Berardenga Scalo (Cigala Fulgosi et al., 2009; sibly of deltaic origin. Large vertebrates embedded in fine-grained, the same mudstone interval at the same locality has been interpreted as muddy matrix (shelf deposit formed below storm wave base) and those

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Fig. 8. Taphonomy of a 5 m-long, undetermined balae- nopterid at Sinalunga (WCh6-1 in Table 2; see Fig. 4 for a tentative sequence stratigraphic interpretation), synthem 6. A: Oblique view of the fully articulated skeleton lying in a gravelly sandstone, stratified in the lower part, massive in the upper. Vertebrae are tightly connected as if in life. B: Plan view of the flipper, with carpals, metacarpals and phalanges in perfect anatomical connection. C: The gravelly sandstone lies above a bioturbated muddy sandstone, with vertical burrows (Ophiomorpha). D: The lower part of the unit with the whale skeleton is formed by three fining-up- ward beds. Each bed grades from gravel to medium-grained sand. Articulated and empty shoreface bivalves (e.g., Cal- lista chione), not in life position, are interspersed with the coarse gravel.

associated with laterally-persistent shell beds (condensed deposits) are maximum near Orciano Pisano (S = 27), with a second peak in S5, slightly disarticulated and fairly complete, showing signs of long per- around Castelnuovo Berardenga (SRB, S = 16). Differently from the manence on a low energy, well‑oxygenated seafloor before the final marine mammals, showing a complete turnover from the Pliocene to burial. Bioerosion of bones caused by phototrophic cyanobacteria and the recent (Table 3), 34% of Pliocene shark species are still extant in the algae, heterotrophic fungi and bacteria (Orciano balaenopterid WFi4-1 NWMS (Tables 5–6). Marine mammal frequency and diversity gradually in S4-FB), and eventually by whalebone-eating siboglinid worm of decreases in S5–S6. genus Osedax (on a ziphiid humerus, WFi4-14: Higgs et al., 2012), oc- curred in condensed intervals, in association with glauconite (Danise, 6. Paleoenvironment-fauna relations 2010). In one instance, a condensed shelly interval is traced for a few kilometers, connecting bioturbated, lower shoreface sandstones The paleoecology of Pliocene MM of Tuscany can be reconstructed yielding slightly disarticulated sirenian skeletons (MOm2-1, MOm2-2) by comparison with the ecology of their modern relatives. This ap- and other MM (Tinelli, 2013) with open shelf deposits yielding slightly proach can be applied at the family or genus level for marine mammals disarticulated whale remains (WOm2-1 in S2-OOB). (Table 3), at the genus or species level for sharks (Table 5). With the Regarding a sequence stratigraphic interpretation, pristine skeletons only exception of the sirenian Metaxytherium subapenninum, feeding on from delta front sediments can be part of the FSST (Fig. 4; in an al- seagrasses (Domning, 2001), and small demersal sharks (e.g., catsharks, ternative interpretation they may belong to the early TST, when incised frilled sharks), all MM studied here are pelagic forms that had no direct valleys are filled with coastal deposits). TST deposits account for 6,2% connection to conditions at the seafloor (Tables 4 and 6). The paleoe- of cases (Fig. 9B). Much more frequently, articulated or slightly dis- cology of benthic habitats informs however on the situation of the articulated skeletons are associated with the HST of the corresponding overlying water column in terms of factors that matter on the dis- depositional sequence, lying above the MFS (56,9% of cases), or within tribution of pelagic organisms, such as water depth, salinity and nu- the maximum flooding interval, above a laterally-persistent shell bed trient levels. In the second place, since all MM remains after death ul- (10,8%). Bone bioerosion is less pronounced in the late HST, when timately sink to the seafloor, benthic paleoecology is also a means to sedimentation rate increases (Castelfiorentino whale WEl3-2 in S3-EB). understand taphonomic controls on MM distribution. Cluster analysis based on the distribution of mollusc species in 72 samples resulted in 5.4. MM abundance and species-richness the identification of four main groups of samples, roughly corre- sponding to the four main facies types recognised based on lithology Some taxa need revision, but a conservative estimate of the different and sedimentary structures (F2–F5 in Table 1). Clusters are formed by morphologies suggest that at least 17 marine mammal species lived in samples from outer shelf and upper slope (three samples), open shelf the NWMS during the Pliocene (possibly > 20, an estimate for the (34 samples), shoreface (23 samples), and transitional settings, such as whole epoch, i.e. 5.332-2.588 Ma), against nine presently living in the brackish-water coastal lagoons and tidal flats (12 samples: see Sup- same area (and two occasional visitors). Among the cetaceans, six fa- plement material, Fig. S1). ANOSIM confirms that sedimentary facies milies were present, against only five presently living in the Ligurian type can broadly predict what benthic assemblage it will yield (Table 7; Sea (Table 3). The most abundant Pliocene species of Tuscany are the general R = 0,632). The difficulty to discriminate between upper and sirenian Metaxytherium subapenninum (N = 10), the dolphin Etrur- lower shoreface facies, and between shoreface and open shelf facies is idelphis giulii (N = 7) and the delphinid Hemisyntrachelus cortesii confirmed by overlaps in sample distribution in the NMDS ordination (N = 5: Table 4). Overall abundance and species richness are not ran- diagram (Fig. 10A). Samples AG1, MON1 and MON2 allow re-inter- domly distributed, but are maximum in Piacenzian strata of synthem preting the associated sandstones, originally included in upper shore- S4, dated at 3.2–3.0 Ma (Table 2, Fig. 2, 9C–D), particularly in the FB face facies type, as offshore deposits. The presence of gravels and and VEB (Table 4). A species list of marine mammals summing up cobbles intensely bioeroded by bivalves and polychaete (ichnofossils fossils found around Orciano Pisano and at La Rocca, near Volterra, include Gastrochaenolithes, Meandripolydora and Caulostrepsis) and en- yields a species richness (S) of 13. This Piacenzian peak in marine crusted by balanids, oysters, serpulids, and bryozoans, suggests they are mammal diversity is matched by the fossil record of sharks, also part of condensed beds resulting from transgressive pulses (hiatal

18 S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx

6.1. Coastal lagoon, tidal flat and embayment

Intertidal faunas are always associated with facies type F2 and are characterised by low-diversity associations, sometimes with < 10 taxa and dominated by one or two species, including species today living in brackish waters of the Mediterranean, at intertidal or very shallow subtidal depths (e.g. Cerastoderma edule, Nassarius reticulatus, Scrobicularia plana: Pérés and Picard, 1964). Facies type F2 is also as- sociated with samples having a species richness higher than the pre- ceding and including species typical of seagrass bottoms and known to withstand moderate changes of salinity. In only one instance a large vertebrate was associated with intertidal deposits (a balaenid, lying on top of large wood fragments: Bianucci, 1995; Collareta et al., 2016).

6.2. Upper shoreface

Facies type F3 is associated with a high-diversity assemblage re- presenting a paleocommunity dominated by suspension feeders adapted to shifting sandy bottoms, with bivalves typical of modern shoreface sandy bottoms (e.g., families Glycymeridae, Tellinidae, Donacidae and Veneridae). Among extinct species of this recurring assemblage, some are large-sized or have very thick shells. Some species of this group indicate the presence of vegetated bottoms. Small pyramidellid gas- tropods are parasitic on echinoderms, also typical of the upper shore- face.

6.3. Lower shoreface

Species richness further increases in collections associated with fa- cies type F4 (lower shoreface). Species typical of this recurring as- semblage include both suspension-feeding and detritus-feeding bivalves and gastropods. The following gastropod families are usually re- presented by several species: Trochidae, Rissoidae, Cerithiidae (from vegetated bottoms), Naticidae, Muricidae, Turridae, Conidae, Terebridae, Bullidae, Cylichnidae (carnivores), Pyramidellidae (echi- noderm parasites). Many bivalve species occur in both facies types F2 and F3. At three different sites and at different stratigraphic units large marine vertebrates, including mysticetes, sirenians and sharks, were recovered in association with shell beds dominated by the gregarious turritellid gastropod Haustator vermicularis (Fig. 5C–D).

6.4. Offshore and upper slope

Another important set of species recurred in facies type F5 (mud- stone deposited in offshore bottoms at shelf depths). Among char- acterising gastropods are the suspension feeders (Turritella tricarinata, Archimediella spirata and Petalochoncus intortus), deposit feeders (Aporrhais uttingeriana) and carnivores or scavengers (Epitonium fron- diculoides, Nassarius semistriatus, Mitrella nassoides). Also the bivalves occupy many different ecological niches (e.g., infaunal detritus feeders, epifaunal suspension feeders, either free-living, byssate, or cemented). Outer shelf and uppermost bathyal sediments from F6 mudstones, Fig. 9. Quantitative analysis of the facies type and sequence stratigraphic distribution of studied at only one location, are characterised by a separate set of large marine vertebrates, Pliocene of Tuscany (N = 39, see Table 2). A: The vast majority of cases (77%) are associated with fine-grained muddy sediments of the shelf, a few are carnivorous gastropods and by a few small bivalve species. Ubiquitous found in deltaic coarse-grained strata. B: Most MM (69%) is found in highstand deposits, a molluscs include species found from intertidal to outer shelf depths few in proximity of the maximum flooding interval, or in transgressive deposits. (e.g., Corbula gibba), and those preferential of open marine waters, from lower shoreface to outer shelf. Most multi-element findings of marine concentrations). The relationship between facies type and mollusc as- mammals are associated with sandy mudstones from open shelf set- sociation is broadly summarised in the following paragraphs (see online tings, below storm wave base, also in association with an Archimediella – Supplement information for a list of characterising species). spirata shell bed (Fig. 6C D). No molluscs were recovered in facies type F7, with the exception of bathyal mudstone in the lowermost Pliocene of FB, associated with sparse specimens of the gryphaeid epifaunal bi- valve Neopycnodonte navicularis (not sampled).

19 .Dmnc tal. et Dominici S. Table 3 Paleoecology of Pliocene large mammals and ecology of modern NWMS analogues.

Suborder Family Species Pliocene Modern Foraging Frequency Feeding Preferred prey Trophic level Occurrence in areas of References for living Reference for Tuscany NWMed depth ecology enhanced primary species extinct species productivity

Odontoceti Delphinidae Globicephala 1 Raptorial Cephalopods, fish 4,4 Hocking et al., etruriae feeder 2017 Globicephala 1 600–800 Common Raptorial Cephalopods, fish 4,4 Provençal basin, Ligurian Sea Pauly et al., 1998; Praca melas feeder and Gannier, 2008 Etruridelphis giulii 1 Raptorial Cephalopods, fish 4,3 Hocking et al., feeder 2017 Delphinus delphis 1 Oceanic Rare Raptorial Cephalopods, fish 4,2 Pauly et al., 1998 feeder Hemisyntrachelus 1 Raptorial Cephalopods, fish 4,3 Hocking et al., cortesii feeder 2017 Stenella sp. 1 Raptorial Cephalopods, fish 4,2 Hocking et al., feeder 2017 Stenella 1 Oceanic Common Raptorial Cephalopods, fish 4,2 Pauly et al., 1998 coeruleoalba feeder Tursiops osennae 1 Raptorial Cephalopods, fish 4,2 Hocking et al., feeder 2017 Tursiops truncatus 1 Neritic Common Raptorial Cephalopods, fish 4,2 Pauly et al., 1998 feeder Orcinus citonensis 1 Raptorial Fish/ 4,5 Young et al., 2012; feeder cephalopods/ Hocking et al., marine mammals 2017 Orcinus orca 1 Very rare Raptorial Fish/ 4,5 Balearic Sea, Provençal basin, Pauly et al., 1998;

20 feeder cephalopods/ Ligurian Sea Notarbartolo di Sciara, marine mammals 1987 Grampus griseus 1 300–1500 Common Raptorial Fish, cephalopods 4,3 Provençal basin, Ligurian Sea Pauly et al., 1998; Praca feeder and Gannier, 2008; Azzellino et al., 2016 Ziphiidae Tusciziphius 1 Suction Cephalopods 4,3 Hocking et al., crispus feeding 2017 Ziphiidae indet. 1 Suction Cephalopods 4,3 Hocking et al., feeding 2017 Ziphius cavirostris 1 1000–2000 Rare Suction Cephalopods, fish 4,3 Ligurian Sea Pauly et al., 1998; feeding Moulins et al., 2007 Physeterid- Physeter 1 1000–2000 Common Suction Cephalopods, fish 4,4 Balearic Sea, Provençal basin, Pauly et al., 1998; Praca ae macrocephalus feeding Ligurian Sea and Gannier, 2008; Pirotta et al., 2011 Physiteridae 1 Suction Cephalopods, fish 4,4 Hocking et al., indet. feeding 2017 Kogiidae Kogia pusilla 1 Suction Fish, cephalopods 4,4 Hocking et al.,

feeding 2017 Earth-Science Reviews176(2018)xxx–xxx Mysticeti Balaenidae Balaena 1 Ram filter Krill, fish 3,2 Berta et al., 2016; forsythmajori feeding/ Hocking et al., continuous 2017 Balaena 1 Ram filter Krill, fish 3,2 Berta et al., 2016; montalionis feeding/ Hocking et al., continuous 2017 Balaenotus insignis 1 Ram filter Krill, fish 3,2 Berta et al., 2016; feeding/ Hocking et al., continuous 2017 Balaenotus 1 Ram filter Krill, fish 3,2 Berta et al., 2016; orcianensis feeding/ Hocking et al., continuous 2017 (continued on next page) .Dmnc tal. et Dominici S. Table 3 (continued)

Suborder Family Species Pliocene Modern Foraging Frequency Feeding Preferred prey Trophic level Occurrence in areas of References for living Reference for Tuscany NWMed depth ecology enhanced primary species extinct species productivity

Balaenula 1 Ram filter Krill, fish 3,2 Berta et al., 2016; balenopsis feeding/ Hocking et al., continuous 2017 Eubalaena sp. 1 Ram filter Krill, fish 3,2 Berta et al., 2016; feeding/ Hocking et al., continuous 2017 Idiocetus 1 Ram filter Krill, fish 3,2 Berta et al., 2016; guicciardinii feeding/ Hocking et al., continuous 2017 Balaenopte- Cetotheriophanes 1 Ram filter Krill, fish 3,4 Berta et al., 2016; ridae cf. C. capellinii feeding/ Hocking et al., continuous 2017 Balaenopteridae 1 Ram filter Krill, fish 3,4 Berta et al., 2016; indet. feeding/ Hocking et al., continuous 2017 Balaenoptera 1 400–2500 Common Ram filter Krill, fish 3,4 Ligurian Sea, Strait of Sicily, Pauly et al., 1998; physalus feeding/ Celtic Sea Greenpeace continuous International, 2009; Panigada and Notarbartolo di Sciara, 2012; Notarbartolo di Sciara et al., 2016a, 2016b; Volkenandt et al.,

21 2015 Balaenoptera 1 Very rare Ram filter Krill, fish 3,4 acutorostrata feeding/ continuous Pinnipedia Phocidae Pliophoca etrusca 1 Semi- Fish, cephalopods 4,0 Hocking et al., aquatic/ 2017 raptorial/ pierce feeding Monachus 1a 1–350 Rare Semi- Fish, cephalopods 4,0 Black Sea (Danube delta) Sergeant et al., 1978; monachus aquatic/ Adam and Berta, 2002; raptorial/ Karamanlidis et al., pierce feeding 2015; Kienle and Berta, 2016; see also Parrish, 2009,onMonachus schauinslandia Sirenia Dugongidae Metaxytherium 11–10 Herbivore Small and mid- 2,0 Domning, 2001 subapenninum sized seagrasses

Total species 21 11 Earth-Science Reviews176(2018)xxx–xxx richness: Pliocene-Modern shared species: 0

a Data relative to pre-industrial times. S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx

6.5. Carbonate platform MM gradually rises in S3, in the late Zanclean-early Piacenzian, whereas it suddenly peaks in synthem S4, where species richness of A recurring benthic assemblage associated with the highly fossili- both marine mammals and sharks is highest. S5 reflects a lower di- ferous facies type F8, not included in the quantitative analysis, consists versity of marine mammals, but still a high diversity of sharks, while of the large pectinid bivalves Gigantopecten latissima and Hinnites crispus values of both groups drop to the lowest abundance and species rich- and by a mixture of photozoans (large benthic foraminifera), hard- ness in S6, during the late Piacenzian-early Gelasian (Fig. 9D). substrata dwellers (brachiopods, echinoderms), encrusters (red algae, The S4 diversity peak coincides with the middle part bryozoa) and bioeroders (clionaid sponges). All fossil-rich carbonates (3.264–3.025 Ma) of the Piacenzian, a time interval in which the Earth are associated with synthem S4, in the mid-Piacenzian (Fig. 2, see also experienced global average temperature 1.84–3.60 °C warmer than the Nalin et al., 2016). pre-industrial period (Dowsett et al., 2013). Climatic impact is testified by the widespread occurrence of carbonate deposits in S4 (Fig. 2), with 7. Paleodepths sedimentary facies indicative of warm-temperate to subtropical condi- tions, with summer sea-surface temperature considerably warmer than Multivariate techniques are usefully applied to stratigraphic and 20 °C and winter temperatures colder than 20 °C (Nalin et al., 2016). paleobiologic analysis (Scarponi and Kowalewski, 2004). We used the This suggests a causative link between global climate and biodiversity, results of the ordination analysis to estimate depths of the final resting S4 diversity peak recording a global phenomenon, possibly an increase place of some large vertebrates listed in Table 2. Samples in the NMDS of speciation rate connected with global warming. Similarly, we pro- ordination plot following a water depth gradient, with shallower sam- pose that the lower diversities recorded at S5-S6 are the regional ex- ples to the left (low values of NMDS axis 1) and deeper samples to the pression of an increase in extinction rate related to climatic cooling and right (high values of axis 1). Therefore, NMDS values of axis 1 can be global sea-level drop, ultimately leading to the global MM extinction used as a proxy for relative water depth. We calculated palaeodepths by event recorded on a coarser scale at the Pliocene-Pleistocene boundary fitting a logarithmic regression curve between paleodepth of 23 modern (Pimiento et al., 2017): the finer stratigraphic resolution adopted here species (data from MARBEF database), common in our dataset, and the suggests a stepwise extinction event. NWMS data also point to a se- values on NMDS axis 1 (Supplement material: Tables S1–S4). The re- lective effect, extinction being recorded by marine mammals, with a gression analysis, with R2 = 0.813, indicates that scores along the 100% regional turnover between Pliocene and Recent (Table 3), but not NMDS main axis are a good predictor of the preferred depth for the 23 as much by the shark fauna, with 34% of the species still living in the modern species (Fig. 10B, inset), thus supporting the bathymetric in- Mediterranean, while still others are today extant, although restricted terpretation. This allowed to estimate the depth of the 72 samples, to tropical and subtropical latitudes. Also the Piacenzian teleost fish which ranged from 0.4 m to 365 m, i.e., from intertidal to upper slope fauna (Cigala Fulgosi et al., 2009) and benthic molluscs (Raffi et al., depths (in accordance with a previous estimate of paleodepths in the 1985) show a high percentage of holdovers, suggesting that marine upper part of the Pliocene of EB, based on counts of foraminifera: mammals have been particularly subject to climatic change (see also Dominici et al., 2007). Facies type F2 is deposited at 0–5 m depth, F3 at Steeman et al., 2009). 3–30 m, F4 at 10–100 m, and F5 at 40–300 m, F5 at around 350 m Estimates of body size in Pliocene mysticetes of Tuscany, with depth (only one sample: Fig. 10B). The position of fossil cetaceans, several specimens reaching 10 m (Danise and Dominici, 2014: seven sirenians, pinnipeds and large sharks (respectively W, M, P and S, specimens in Table 1), are comparable to global values derived from the N = 13) was plotted near the corresponding shell bed in the NMDS literature for this time interval (Lambert et al., 2010), confirming that ordination. The resulting pattern shows that all MM considered, asso- NWMS baleen whales were larger than their Miocene analogues, and ciated with open shelf settings and with facies types F4–F5, cluster smaller than modern forms. Within odontocetes, the most common around −100 m, spanning −30–300 m. Pliocene delphinid, Etruridelphis, was larger than the modern analogue Stenella (Bianucci et al., 2009b). The same is true for Hemisyntrachelus 8. Factors in the Pliocene NWMS marine megafauna fossil record cortesii, larger than modern Tursiops (Bianucci, 1997a). On the other hand, the largest extant delphinid Orcinus orca, reaching 9 m, is about The detailed sequence-stratigraphic framework and the abundant twice as long as Orcinus citonensis (Heyning and Dahlheim, 1988). The shell beds, offering an independent check on sedimentary facies dis- high percent of holdover points to a more conservative figure for larger tribution by benthic paleoecology, allows also to explore factors in MM sharks (Table 5), but the presence in the Pliocene of the gigantic geographic and stratigraphic distribution, and to sort out evolutionary, Carcharocles megalodon and some large thermophilic species today re- ecological, and taphonomic drivers of this particular fossil record. stricted to lower latitudes suggests that impoverishment of the fauna is NWMS Pliocene distribution can be compared with the results of ana- coupled with an average decrease in size (Marsili, 2008). Comparing logous studies carried out in different settings and at different times, to sizes suggests an overall restructuring of NWMS MM during the last draw conclusions on the lessons to be learned about MM evolution, 3 million years (see also Bisconti, 2009). ecology, and taphonomy. 8.2. Ecological control 8.1. Evolutionary control The composition of the Pliocene NWMS MM is affected in the first Available data allow a meaningful comparison of NWMS MM di- place by the availability of food. At the lowest trophic level, inferring versity across the Pliocene, particularly detailed for marine mammals from the ecology of the modern MM (Tables 3, 5), we find herbivore (Fig. 9D). Although the study suggests a facies control, with MM re- sirenians feeding on seagrasses (trophic level, TL = 2,0), all others mains being generally associated with late TST-HST muddy sandstones being carnivores, thus having TL > 3,0 (Fig. 11). The lowermost levels and mudstones from lower shoreface and offshore shelf paleosettings, among the carnivores were occupied by baleen whales and whale the temporal pattern of biodiversity recorded on a regional basis likely sharks (TL = 3,2–3,4), filter-feeding on pelagic crustaceans (krill). reflects also a paleobiologic phanomenon, since lower shoreface and Roughsharks, catsharks, and houndsharks, with many species living in offshore shelf sediments are represented in all synthems. Marine the modern Mediterranean, have a relatively species-poor Pliocene re- mammals are unrecorded in the thickest part of synthem S1-2 (Zan- cord, probably due to a preservation bias related to their small size. At clean, mostly bathyal mudstone and turbiditic sandstone), but they are TL = 3,7–4,0 were one species of catshark and a monk seal, both present in S2, in OOB (Zanclean lower shoreface and open shelf de- feeding near the seafloor on crustaceans, teleost fishes and cephalo- posits) and SRB (Zanclean, both in deltaic and in upper slope deposits). pods. At TL = 4,1 were sandbar, tiger and blue sharks, feeding on

22 S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx teleost fishes, cephalopods and on marine mammals. At the same level, in horizontal (during seasonal migration) and vertical direction (during in slope environments, kitefin sharks mainly fed on other sharks. The daily feeding migration: Roman and McCarthy, 2010), a process par- majority of MM species were found at TL = 4,2, with smaller odonto- ticularly important in oligotrophic seas (Allgeier et al., 2017), like large cetes (three species) and 11 species of sharks, including several requiem sectors of the Mediterranean. Finally (literally, after death), MM be- sharks, a houndshark and a hammerhead. Larger delphinids, sperm comes a detrital sources of energy and habitat for deep sea whale-fall whales, beaked whales, together with mackerel, sand and sand tiger communities (Roman et al., 2014; Smith et al., 2015), with paleonto- sharks, occupied high trophic levels (TL = 4,3), followed at the top of logical evidence available for the NWMS (Dominici et al., 2009; Higgs the global NWMS food web by white shark, megalodon shark, one et al., 2012; Baldanza et al., 2013). species of sevengill shark (genus Notorynchus), and killer whales, all Much evidence suggests that a bottom-up control on the structure of feeding on marine mammals and smaller sharks (TL = 4,4–4,7). With NWMS MM community was exerted by wind-driven upwelling currents, no exception, all highest levels encountered in modern NWMS offshore through enhanced oceanic productivity and concentration of preys. pelagic and nearshore communities were occupied during the Pliocene Over geologic time and on a global scale, these factors may have ex- by an analogous MM, often by the same species (large sharks), or by erted a selection favouring large size, triggering the Plio-Pleistocene congeneric or confamilial species (marine mammals: Table 3, 5). The emergence of whale gigantism in several lineages (Slater et al., 2017). Pliocene pelagic ecosystem, typified by the mid-Piacenzian S4 asso- The largest among marine top predators can travel long distances and ciation, must have been however trophically more diversified (Fig. 11), cross oceans, but tend to congregate in shallow waters with abundant including aquatic megaherbivores, several balaenid filter feeders, larger prey. Baleen whales, abundant in boreal eutrophic waters (Woodley and more diverse dolphins, and shark species today extinct (e.g., and Gaskin, 1996), with population size under the control of food Carcharocles megalodon) or restricted to tropical seas (e.g., Galeocerdo availability (Croll et al., 2005), always require high prey density for cuvier). With the exception of TL = 2, all Pliocene NWMS MM were efficient bulk filter feeding (Goldbogen et al., 2011). Six different spe- either apex predators of their community, or mesopredators, occupying cies of mysticetes currently foraging in the Southern Ocean, among trophic positions below apex predators. The definitions of apex pre- which the largest animals that have ever lived in world oceans, exploit dators (or top predators) and mesopredators are relative, and to an the high biomass of Antarctic krill, their main food resource. Great extent context-dependent (species that in a context are apex predators, white sharks are abundant in the offshore of California (Jorgensen in another are mesopredators, e.g., Estes et al., 1998). Since predation is et al., 2010), Australia-New Zealand and South Africa (Bonfi l et al., a trophic interaction in which one animal (predator) consumes another 2005), clustering in proximity of seal colonies, including monk seals. (prey) as a source of energy (food), irrespective of the means by which The distribution in the modern Mediterranean is no exception to gen- this is accomplished (Lourenço et al., 2013), filter-feeding baleen eral rule, only on a smaller scale, with large sharks and marine mam- whales can be considered apex-predators of their community (e.g., mals congregating in productive areas, such as the Strait of Sicily and Lewison et al., 2004; Notarbartolo di Sciara et al., 2016b). Among the Balearic Archipelago (Tables 3, 5), with the second largest animal sharks, 68% of living Mediterranean elasmobranches are ranked as top on Earth, the fin whale, showing a movement pattern that parallels predators, with a trophic level of 4 or more (Goffredo and Dubinsky, seasonal variability in available feeding habitat (Notarbartolo di Sciara 2014, including superorder Batoidea, against 22% of teleost fishes), an et al., 2016b), contributing to the horizontal transfer of nutrients. estimate that can be extended to Pliocene NWMS MM. Relationships Tuscany faces the Ligurian Sea, where a deep-water upwelling current between apex predators and mesopredators are complex and hard to coming from Southwest convects nutrients to the water surface, leading define in ecology, involving predation on other predators (intraguild to high levels of primary productivity in its western sectors, extending predation, combining competition and predation: Polis et al., 1989), westward to the Provençal and Balearic Seas, with spring algal blooms. where consumption and competition need to be proved (Lourenço These waters host different trophic regimes in an otherwise oligotrophic et al., 2013). In the Mediterranean Pliocene, paleontological evidences Mediterranean Sea (Lazzari et al., 2012; Melanotte-Rizzoli et al., 2014; of carnivores serving as food to MM include inferred killing of prey Stambler, 2014). Supporting a conspicuous biomass of zooplankton (Bianucci et al., 2010) and scavenging (Cigala Fulgosi, 1990; Bianucci (Cuzin-Roudy, 2011), the Ligurian Sea sustains large populations of fin et al., 2002; Dominici et al., 2009). The occurrence of intraguild pre- whales (Balaenoptera physalis) and striped dolphins (Stenella coer- dation must have been far more extended than what taphonomy can uleoalba: Notarbartolo di Sciara et al., 2008). Mediterranean-resident prove, however, given a higher diversity of raptorial feeders, this guild fin whales have adapted to exploit localised mesoscale hotspots of including during the Pliocene the killer whale Orcinus citonensis, the productivity that are highly variable in space and time (Notarbartolo di large delphinid Hemisyntrachelus cortesii (phylogenetically related to the Sciara et al., 2016b), feeding behaviours possibly mediated by the modern killer whale: Murakami et al., 2014), the largest shark of all depth of prey and species-specific behaviours, allowing to minimise times, Carcharocles magalodon (Marsili, 2008), and a diverse association competition with other large filter feeders (see Friedlaender et al., of smaller carnivores, including monk seal, delphinids and sharks with 2014). The sperm whale Physeter macrocephalus uses habitat across a 4,0 < TL < 4,2 (Table 3, 5), candidate prey for larger raptorial fee- range of depths and a specialised diet (Rendell and Frantzis, 2016), ders. This interaction likely exerted in its turn a control on community gathering along NWMS steeper slopes, where water currents allows for structure at lower trophic levels through processes like “mesopredator higher trophic level biomass (Pirotta et al., 2011). Beaked whales are release” and trophic cascades (Roemer et al., 2009), eventually linking particularly abundant in the Ligurian Sea and central Tyrrhenian Sea, pelagic and nearshore communities, including benthic animals and preferring submarine canyons at slope depths (Podestà et al., 2016). plants, like in many modern ecosystems (Estes et al., 2011, 2016), down The largest among common Mediterranean delphinids, Grampus griseus, to slope depths (e.g., Parrish, 2009). Large raptorial feeders could exert is frequent in the Ligurian and Thyrrenian seas where it forages on a control on the diversity of the filter-feeding MM, like it has been cephalopods at depths 300–1500, where upwelling currents are most suggested on a global scale since the Miocene (Lambert et al., 2010), effective (Azzellino et al., 2016). and for the Pliocene by Bisconti (2003), when baleen whales were more The NWMS existed as a Liguro-Provençal back-arc oceanic basin diversified, both in terms of species richness, size range, and feeding since the late Miocene, when Sardinia rotated to its present position strategies, including both skim- and lunge-filter feeding (respectively (Gattacceca et al., 2007), and in coincidence with the formation of the balaenid and balaenopterid whales: Berta et al., 2016; Hocking et al., hinterland basins here under study (Muttoni et al., 2001), justifying the 2017), minimising competition for food and diversifying spatial niches assumption that the modern NWMS trophic regime is a feature that (see also Marx et al., 2017, for the upper Miocene). A further important dates back at least to the upper Miocene. Consistenly, diatomites de- top-down control on community structure is suggested by modern posited in Tuscany during the Messinian, before the salinity crisis studies on the role of baleen and sperm whales as nutrient vectors, both (Bossio et al., 1997; Roveri et al., 2014), indicate that high productivity

23 S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx

Table 4 Geographic distribution of Pliocene large mammals in Tuscany.

Suborder Family Basin OOB VEB FB SRB EB EB CB Per species Frequency % abundance Locality Arcillea Volterrab Orciano Sienae,d San Miniatof Certaldog,d Sinalungah Pisanoc,d

Odontoceti Delphinidae Globicephala etruriae 1 1 1.6 Etruridelphis giulii 1 5 1 7 10.9 Hemisyntrachelus cortesii 1 3 1 5 7.8 Orcinus citonensis 1 1 1.6 Stenella sp. 1 1 1.6 Tursiops osennae 1 1 1.6 Delphinidae indet. 2 1 3 4.7 Ziphiidae Tusciziphius crispus 1 1 1.6 Ziphiidae indet. 1 3 4 6.3 Physeteridae Physeteridae sp. 2 2 3.1 Kogiidae Kogia pusilla 1 2 3 4.7 Mysticeti Balaenidae Balaena forsythmajori 1 1 1.6 Balaena montalionis 1 1 1.6 Balaenotus insignis 1 1 1.6 Balaenotus orcianensis 2 2 3.1 Balaenula balenopsis 1 1 1.6 Balaenula sp. 2 2 3.1 Eubalaena sp. 1 1 1.6 Idiocetus guicciardinii 1 1 1.6 Balaenidae indet. 1 3 2 6 9.4 Balaenopteridae Cetotheriophanes cf. C. 1 1 1.6 capellinii Balaenopteridae indet. 1 2 1 4 6.3 Pinnipedia Phocidae Pliophoca etrusca 2 1 1 4 6.3 Sirenia Dugongidae Metaxytherium 5 1 4 10 15.6 subapenninum Per area abundance: 6 9 21 17 5 4 2 64

a Includes Camigliano, Poggio alle Mura. b Includes La Rocca, Saline di Volterra, Poggiarone. c Includes Santa Luce, Pieve Santa Luce, Greppoli. d One or more specimens tentatively attributed to the species. e Includes Sarteano, Pienza, Castelnuovo Berardenga, Val di Pugna. f Includes Montaione, Ricavo, Poggio Tagliato, Casenuove. g Includes Castelfiorentino, Castel San Gimignano. h Includes Cetona. was a primitive feature of the Ligurian Sea. Paleontological evidences with two exceptions, one associated with delta-front, coarse-grained include the high diversity and abundance of Pliocene MM in FB, the beds (WCh6-1, Fig. 8), another from outer shelf or upper slope sedi- closest to modern upwelling areas (Fig. 3), and the association of sev- ments (DSi2-1). These data point to a strong environmental control on eral MM with “Turritella beds”, turritelline gastropods showing gre- the quality of the MM fossil record, at least concerning articulation and garious habit and high abundance in areas of high primary productivity completeness of skeletons, very shallow and very deep (bathyal) depths (Allmon, 1988). Both a comparison with the modern and paleoecologic being generally unfavourable to the preservation of MM bones data strongly suggest that a wedge of NWMS nutrient-rich waters in- (Fig. 9A). To explain these results, hypotheses are based on the avail- tersecting the Tuscan shelf exerted a strong bottom-up control on able data on carcasses of MM in modern marine environments. community structure through mixing and upwelling of nutrients, sti- The biostratinomy of lung-breathing marine mammals depends on mulation of phytoplankton blooms, followed by zooplankton increase, water depth (Allison et al., 1991; Smith, 2006). The vast majority of while nekton and vertebrates tracked plankton concentrations, as is mammals are negatively buoyant and sink after death, but in shallow typical of modern upwelling systems throughout the world (Polis et al., waters the low hydrostatic pressure allows putrefaction gases to de- 1997). High surface primary productivity in the study area during the velop and carcasses to resurface. Skeletons are scavenged (Dicken, Pliocene would have caused a concentration of detritus-falls, sup- 2008) and disintegrate while floating, leading to the preferential de- porting a diverse community of deep-sea scavengers. position of isolated bones. At deeper settings, where high hydrostatic pressure allows the carcass to lay relatively undisturbed on the seafloor, the skeleton becomes only slightly disarticulated after soft tissue re- 8.3. Taphonomic control moval (Reisdorf et al., 2012). The subsequent fate of deep-water, dis- articulated skeletons depends on the nature of the scavenging fauna and Taphonomic data on articulation and completeness of MM speci- other elements of the whale-fall community, and on the time of ex- mens are available for large mammals, the shark record being formed posure on the seafloor (Boessenecker et al., 2014). At depths deeper mainly by isolated teeth collected through surface picking during the than the slope breaks, where sedimentation rate is very low, carcasses years (Cigala Fulgosi et al., 2009), with only a few contextualised stu- are exposed for a long time and the skeleton is rapidly disintegrated. dies (e.g., Bianucci et al., 2002; Dominici et al., 2009). Our record in- Time-series analyses carried out on modern whale-fall communities at cludes several marine mammals with a high degree of articulation slope depths (range 382–2893 m: Lundsen et al., 2010) suggests that (14%) and completeness (27%), or with at least cranial and post-cranial carcasses up to 17 m are rapidly degraded, with the deepest whale elements of the same individual (38%). All articulated specimens and carcasses disappearing after only seven years of exposure on the sea- the most complete skeletons are associated with shelf mudstones, floor. Larger skeletons may persist on deeper settings for decades, but if usually forming the HST of the relative depositional sequence (Table 2),

24 .Dmnc tal. et Dominici S. Table 5 Paleoecology of Pliocene sharks and ecology of modern NWMS analogues.

Order Family Species Pliocene Modern Preferred prey Trophic level Occurrence in areas of enhanced References NWMed NWMed primary productivity

Hexanchiformes Chlamydoselachidae Chlamydoselachus lawleyi 1 Teleost and chondrichthyan 4,2 Suruga Bay Cigala Fulgosi et al., 2009 (based on extant fishes, cephalopods congeneric) Echinorhinidae Echinorhinus brucus 1 1 Teleost and chondrichthyan 4,4 Cortés, 1999 fishes Hexanchidae Hexanchus griseus 1 1 Cephalopods, teleost fishes, 4,3 Balearic Sea Cortés, 1999; Barría et al., 2015 marine mammals Heptranchias perlo 1 Teleost fishes, cephalopods, 4,2 Cortés, 1999 crustaceans Notorynchus lawley 1 Chondrichthyan and teleost 4,7 Cortés, 1999 (on genus Notorynchus) fishes, marine mammals Centrophoridae Centrophorus granulosus 1 1 Teleost and chondrichthyan 4,1 fishes, cephalopods Dalatiidae Dalatias licha 1 1 Teleost and chondrichthyan 4,1 Balearic Sea Cortés, 1999; Navarro et al., 2014 fishes, cephalopods Pristiophoriformes Pristiophoridae Pristiophorus sp. 1 4,2 Squatiniformes Squatinidae Squatina sp. 1 4,1 Squatina aculeata 1 Teleost fishes, crustaceans, 4,2 Cortés, 1999 cephalopods Squatina oculata 1 Teleost fishes, crustaceans, 4,0 Cortés, 1999 cephalopods Squatina squatina 1 Teleost fishes, crustaceans, 4,0 Cortés, 1999 cephalopods Lamniformes Odontaspidae Carcharias taurus 1 1 Teleost and chondrichthyan 4,4 Cortés, 1999

25 fishes Odontaspis ferox 1 1 4,4 Odontaspis acutissima 1 4,4 Alopiidae Alopias superciliosus 1 1 Cephalopods, teleost fishes, 4,2 Cortés, 1999; Boldrocchi et al., 2017 marine mammals Alopias vulpinus 1 Cephalopods, teleost fishes 4,2 Strait of Sicily Cortés, 1999; Greenpeace International, 2009; Barría et al., 2015 Cetorinidae Cetorhinus maximus 1 1 Krill 3,2 Strait of Sicily Cortés, 1999; Greenpeace International, 2009 Lamnidae Carcharodon carcharias 1 1 Teleost and chondrichthyan 4.5 Ligurian Sea, Thyrrenian Sea, Cockcroft et al., 1989; Compagno, 2002; fishes, marine mammals Balearic Sea, Strait of Sicily, Boldrocchi et al., 2017 Adriatic Sea Carcharocles megalodon 1 4,5 Lamna nasus 1 Teleost fishes, cephalopods 4.2 Strait of Sicily Greenpeace International, 2009 Isurus oxyrinchus 1 1 Teleost and chondrichthyan 4.3 Strait of Sicily Greenpeace International, 2009 fishes, cephalopods Isurus xiphodon 1 4,3

Paradotus benedeni 1 4.2 Earth-Science Reviews176(2018)xxx–xxx Carcharhiniformes Carcharinidae Carcharhinus altimus 1 Teleost and chondrichthyan 4.3 fishes, cephalopods Carcharhinus brachyurus 1 1 4.2 Carcharhinus brevipinna 1 Teleost fishes, cephalopods 4.2 Cortés, 1999 Carcharhinus falciformis 1 Teleost fish, cephalopods 4.2 Cortés, 1999 Carcharhinus leucas 1 Teleost and chondrichthyan 4.3 Cockcroft et al., 1989, Cortés, 1999 fishes, marine mammals Carcharhinus limbatus 1 Teleost and chondrichthyan 4.2 Cortés, 1999 fishes, cephalopods Carcharhinus longimanus 1 Teleost fishes, cephalopods, 4.2 Cortés, 1999 marine mammals Carcharhinus obscurus 1 1 4.2 Cockcroft et al., 1989; Cortés, 1999 (continued on next page) .Dmnc tal. et Dominici S. Table 5 (continued)

Order Family Species Pliocene Modern Preferred prey Trophic level Occurrence in areas of enhanced References NWMed NWMed primary productivity

Teleost and chondrichthyan fishes, cephalopods, marine mammals Carcharhinus perezi 1 4.2 Carcharhinus plumbeus 1 1 Teleost fishes, crustaceans, 4.1 Cortés, 1999; Ellis and Musick, 2007 cephalopods Galeocerdo cuvier 1 Teleost fishes, marine mammals, 4.1 Cockcroft et al., 1989; Cortés, 1999; crustaceans Boldrocchi et al., 2017 Prionace glauca 1 1 Teleost fishes, cephalopods, 4.1 British Columbia, English Cortés, 1999; Greenpeace International, crustaceans, marine mammals Channel, western Atlantic coast, 2009; Williams et al., 2010; Queiroz et al., Strait of Sicily, Balearic Sea 2012; Barría et al., 2015 Sphyrna zygaena 1 1 Cephalopods, teleost fishes 4.2 Scyliorhinidae Pachyscyllium dachiardii 1 3.7 Scyliorhinus canicula 1 Crustaceans, benthic 3.6 Balearic Sea Cortés, 1999; Barría et al., 2017 invertebrates Scyliorhinus stellaris 1 Cephalopods, teleost fishes, 4 Cortés, 1999 crustaceans Galeus melastomus 1 Crustaceans, benthic 3.7 Cortés, 1999 invertebrates, teleost fishes Triakidae Galeorhinus galeus 1 1 Teleost fishes, cephalopods 4.2 Balearic Sea Cortés, 1999; Barría et al., 2015 Mustelus asterias 1 Crustaceans, teleost fishes, 3.7 Cortés, 1999 cephalopods Mustelus mustelus 1 Crustaceans, cephalopods, teleost 3.8 Cortés, 1999 fishes

26 Mustelus punctulatus 1 Crustaceans, teleost fishes, 3.8 Cortés, 1999 cephalopods Squaliformes Oxynotidae Oxynotus centrina 1 Crustaceans 3.9 Balearic Sea Barría et al., 2015 Squalus acanthias 1 Teleost fishes, crustaceans, 3.9 Balearic Sea Barría et al., 2015 cephalopods Species richness: 30 33 Pliocene-Modern shared species: 16 (34% holdover) Earth-Science Reviews176(2018)xxx–xxx S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx

Table 6 Geographic distribution of Pliocene sharks in Tuscany.

Order Family Basin VEB FB FB SRB SRB SRB EB EB Number of occurrences Species/locality Volterra Orciano Parlasciob San Quirico Castelnuovo Sienad San Miniatoe Certaldo Pisanoa Berardengac

Hexanchiformes Chlamydoselachidae Chlamydoselachus 11 2 lawleyi Echinorhinidae Echinorhinus 11 2 richiardii Hexanchidae Hexanchus griseus 11 1 1 3 Notorynchus lawley 11 2 Centrophoridae Centrophorus 11 2 granulosus Dalatiidae Dalatias licha 111 3 Pristiophoriformes Pristiophoridae Pristiophorus sp. 1 1 1 2 Squatiniformes Squatinidae Squatina sp. 1 1 1 2 Lamniformes Odontaspidae Carcharias taurus 11 Odontaspis ferox 111 3 Odontaspis 11 1 1 11 5 acutissima Alopiidae Alopias superciliosus 11 Cetorinidae Cetorhinus maximus 11 2 Lamnidae Carcharodon 11 1 1 1 11 6 carcharias Carcharocles 112 megalodon Isurus oxyrinchus 11 1 1 1 1 5 Isurus xiphodon 11 1 1 3 Paradotus benedeni 11 1 1 3 Carcharhiniformes Carcharinidae Carcharhinus 1 1 brachyurus Carcharhinus 1 1 falciformis Carcharhinus leucas 112 Carcharhinus 112 longimanus Carcharhinus 11 2 plumbeus Carcharhinus 1 1 obscurus Carcharhinus perezi 11 2 Galeocerdo cuvier 11 1 3 Prionace glauca 11 1 1 1 4 Sphyrna zygaena 1 1 Scyliorhinidae Pachyscyllium 1 1 dachiardii Triakidae Galeorhinus galeus 1 1 Species richness 9 27 4 11 16 9 2 1 Total per area/locality: S=30

a Includes Santa Luce, Pieve Santa Luce. b Includes San Frediano, Lari. c Includes Sodi, Guistrigona. d Includes Troiola, Pienza, Coroncina, Trequanda. e Includes Poggio Tagliato, San Maiano. not buried, they also ultimately undergo complete destruction. Al- (Alfaro-Lucas et al., 2017). The general paucity of novel taxa on though environmental forcing triggered by higher temperatures, active shallow-water whale falls suggests that species-rich, specialised whale- currents and sediment transport, clearly plays a role, a specialised fall communities develop only in the food-poor deep sea. Accordingly, whale-fall fauna rapidly consumes both soft and mineralised tissues. among new animal species described in the recent literature from whale This fauna is characterised by low diversity and high abundance of falls, only about 10% have been found on whale remains at depths microorganisms, most likely as a result of both specialisation to nutrient of < 260 m (12 out of 129 new species: Smith et al., 2015). Of all MM enrichment and high growth rates. Whale-fall habitats likely undergo a investigated so far, only whales are known to host a whale-fall com- temporal microbial succession from primarily heterotrophic to more munity, but also carcasses of large elasmobranchs undergo rapid de- heterotrophic/chemosynthetic metabolisms until the whale biomass is struction at bathyal depths, teeth being all that eventually remains completely exploited (Smith et al., 2015). Of all the specialised taxa, (Higgs et al., 2014a; teeth are also lost by sharks during feeding: bone-eating polychaetes of genus Osedax, with their soft root-like tis- Pokines and Symes, 2013). As a consequence, over geological time no sues that erode the bones to access nutrients (Tresguerres et al., 2013; large marine vertebrate is expected to be recovered at depths greater Miyamoto et al., 2017), are the primary cause of bone disintegration, than the shelf break. On the opposite side, the lack of a biota specialised particularly of denser bones (Higgs et al., 2011). Found also at shelf in exploiting large organic falls, coupled with higher rates of deposition depths, but invariably in low abundance (Huusgaard et al., 2012; Higgs in proximity of sediment sources, make it more probable that the most et al., 2014b), bone-eating worms occur in high numbers in the deep articulated and complete whale skeletons become part of the fossil re- sea (Smith et al., 2015) where they act as biodiversity regulators cord in shelf settings below storm wave base.

27 S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx

Fig. 10. NMDS ordination of bulk samples (N = 72, see Fig. 2 for their stratigraphic position), based on the dis- tribution of standardised abundances of 329 mollusc spe- cies (further explanation on multivariate techniques in the main text). A: Samples are subdivided based on the asso- ciated sedimentary facies types (F2–F5 in Table 1). The main axis ordinates samples along a paleodepth gradient, from shallowest to deepest, moving from the left to the right side of the bivariate plot. B: Same ordination, with an estimate of absolute paleodepth of each sample based on score along the main axis and calibrated through the average modern depth distribution of 23 extant species characterising the Pliocene dataset (abundance > 0.15%, see text; regression logarithmic curve in the inset). Verte- brates recovered in proximity of some of the samples (N = 13) are plotted on the diagram, confirming that, on average, the MM fossil record is concentrated on the open shelf at an estimated depth of 30–300 m (M = sirenians; S = sharks; P = pinnipeds; W = whales).

9. Comparison with other studies deposited in the North American seaway during an interval of max- imum flooding (Schmeisser McKean and Gillette, 2015). Upper Cre- The study of sedimentary facies uncovers some environmental fac- taceous mosasaur remains are particularly concentrated in fine-grained tors that directly control the taphonomy of large marine vertebrates. shelf deposits in Europe (Jagt and Jagt-Yazykova, 2016). In the Pria- Such abiotic drivers include water pressure, wave energy and sedi- bonian late TST of Egypt, complete, partially articulated whale skele- mentation rate — three factors summarised by water depth, and up- tons of archaeocetes, together with bones and teeth of sirenians and welling, bringing to the surface deep sea nutrients and concentrating sharks, are abundant at transgressive surfaces, whereas well-articulated preys. The taphonomic pathway of large marine carcasses is also driven whales are associated with rapidly accumulating shoreface sediments of by biotic factors that change in geological time in response to coevo- the FSST, comprising “Turritella shell beds” (Peters et al., 2009). During lution between bacteria, scavengers and their substrates. We now ex- the Oligocene, eomysticete whale bones were deposited at shelf depth plore the multifaceted nature of MM taphonomy by reviewing Mesozoic below storm wave base. The occurrence of sparse traces attributed to and Cenozoic studies where sufficient data for stratigraphic paleo- Osedax and the association with a glauconitic limestone testify to the biology are available. exposure of bones on the seafloor without undergoing complete de- struction (Boessenecker and Fordyce, 2014), in a manner similar to MM at Orciano (see Danise, 2010; Higgs et al., 2012). Burdigalian/Langhian 9.1. Abiotic factors MM recovered in the Antwerpen Sands, in Belgium (including isolated remains of a baleen whale, several odontocetes and a pinniped) are According to one of the few previous studies on the sequence stra- fragmented, worn and associated with clayey sandstone rich in glau- tigraphic distribution of MM, Jurassic ichthyosaurs, plesiosaurs, and conite, suggesting long exposure on the seafloor (Louwye et al., 2010). pliosaurs of the Sundance Seaway, in North America, display facies Bones are concentrated at the base of a coarsening-upward succession, control and are found primarily in offshore mudstone and at condensed on top of shallow marine, coarse-grained sandstone, suggesting this is a intervals at the maximum flooding surface, rather than shoreface and surface of maximum flooding. An association of MM taxonomically estuarine sandstone (McMullen et al., 2014). Taphonomic data on comparable to that here under study is encountered in the Mio-Pliocene Upper Cretaceous marine reptiles and large fishes suggest that partially of the Purisima Formation, in Central California. Taphofacies differ in articulated and disarticulated skeletons are associated with little bio- some aspects. The Pliocene of California yields laterally persistent logical activity and relatively rapid burial by muddy sediments,

28 S. Dominici et al. Earth-Science Reviews 176 (2018) xxx–xxx bonebeds with polished and phosphatised bones, and abundant phos- phate nodules that are absent in Tuscany, indicating times of higher sediment starvation during transgressive pulses, in an area of much stronger nutrient content (the California Current system is a northern- hemisphere analogue of the Peruvian upwelling system, associated with the economically most important fish stocks in the world: Mann and Lazier, 2006). Shoreface deposits indicate stronger wave energy, and the preferential absence of molluscs in bonebeds indicates chemical destruction of carbonate shells (Boessenecker et al., 2014, in a quan- titative MM taphonomic study). Episodic sedimentation, however, causes the preferential preservation of articulated remains in the Cali- fornian offshore as in the Pliocene of NWMS. The late Miocene Pisco Formation in Perù offers another, more extreme example of MM taph- onomy in a eutrophic setting. Here hundreds cetaceans, pinnipeds, and sharks were described in an exceptional state of preservation (Bianucci et al., 2016a, 2016b), within a monotonous succession of finely lami- nated white diatomites (Di Celma et al., 2015), suggesting very high primary productivity in an area of intense upwelling and volcanic ac- tivity. Algal blooms sustained high biomass of apex predators (see Marx and Uhen, 2010), triggering at the same time anoxic conditions at the Fig. 11. Occupancy of trophic levels by Pliocene marine mammals and sharks in the seafloor where MM carcasses remained intact (Brand et al., 2004; see north-western Mediterranean, expressed by number of species per trophic level (see Tables 3, 5 for explanation and references). This figure, summing up data for the whole also Gioncada et al., 2016)inoffshore paleosettings (Marx et al., 2017). epoch, spanning circa 2,8 My, closely matches the association found in one single syn- Finally, the stratigraphic distribution of Pliocene cetaceans in western them S4, of much shorter duration (lower-mid-Piacenzian, 400–500 ky). Emilia (Italy) shows an uneven distribution of findings (N = 24, dol- phins and baleen whales) and a strong positive correlation with off- occurrence of large marine reptiles and teleost fishes in the oceans, long shore mudstones (no findings in shoreface sandstones, rare occurrences before whale evolution, and although their distribution is not limited to in epibathyal mudstones: Freschi and Cau, 2016), paralleling the dis- large carcasses (Pyenson and Haasl, 2007), the radiation of ocean-going tribution of Tuscan Pliocene MM. mysticetes at the Oligocene onset of the Antarctic Circumpolar Current (Fordyce, 2003) clearly increased available substrates worldwide. The 9.2. Biotic factors steady increase of cetacean size during the Neogene, with a dramatic pulse in the last 5 million years, when Neoceti surpassed 10 m length Many reviews of Triassic (Camp, 1980; Hogler, 1992; Motani et al., and reached 30 m in the Pleistocene (Lambert et al., 2010; Slater et al., 2008; Hu et al., 2011; Liu et al., 2014) and Early Jurassic marine rep- 2017), would have thus triggered a second and more massive radiation tiles (Benton and Taylor, 1984) report good preservation and a high of bone-eating worms (Kiel and Goedert, 2006). Consistent with this degree of completeness and articulation of skeletal material. This re- hypothesis, the preferential distribution of modern Osedax in high la- cord might be partially controlled by prevailing anoxic or dysoxic titude settings worldwide (Taboada et al., 2015) suggests that biodi- conditions in the bottom waters of many Mesozoic fossiliferous deposits versity hotspots coincide with the feeding grounds of larger cetaceans. (e.g., Middle Triassic Besano and Guangling Formations, Lower Jurassic Other bone-eaters of modern deep water whale-fall ecosystem belong to Blue Lias and Posidonia Shale Formations), which precluded organism the group of abyssochrysoid snails, with fossils found on Late Cretac- activity within the sediment, and prevented predation or scavenging of eous plesiosaur (Kaim et al., 2008) and sea turtle bones (within a the carcasses on the sea bottom (Beardmore and Furrer, 2016). Evi- chemosynthesis-based association: Jenkins et al., 2017). Modern abys- dence of advanced levels of disarticulation or bone degradation sochrysoid whalebone-eaters of genus Rubyspira, hosting a specific and (Martill, 1985; sauropterygians, crocodilians, ichthyosaurs and fishes exclusive microbiome (Aronson et al., 2017), split during the late Eo- from the Middle Jurassic Oxford Clay Formation) is mostly attributed to cene/early Oligocene (Johnson et al., 2010). Species of Rubyspira physical factors (e.g., weathering on the seafloor). Up to the Early-Late benefited too from the radiation of ocean-going whales. Although Cretaceous, biological activity is testified by circumstantial evidence of scanty, available evidence on the geological history of bone-eaters thus scavenging (Hybodus teeth associated with marine reptile skeletons, makes the ephemeral nature of large carcasses in modern deep seas — Martill et al., 1994), and by the more common occurrence of microbial and their absence in bathyal deposits of the Pliocene of NWMS — an mats, grazers and encrusters (Martill, 1987; Meyer, 2011; Danise et al., unsuitable model for the Mesozoic and the early Paleogene. 2014; Reolid et al., 2015), but lack traces of bone-eating worms and sulphophilic fauna typical of modern whale falls. The siboglinid Osedax 10. Conclusions is an evolutionary novelty in possessing a root system that hosts het- erotrophic mutualists and secretes bone-dissolving acids (Tresguerres Sedimentary facies in the Pliocene of Tuscany are vertically stacked et al., 2013; Miyamoto et al., 2017), and an ecosystem engineer (Alfaro- • to form small-scale depositional sequences particularly in the upper Lucas et al., 2017). Osedax is today associated with whale falls world- half, Piacenzian part of the succession, with laterally-continuous wide (Taboada et al., 2015), but its impact on MM has changed in time. shell beds marking transgressive surfaces and intervals of maximum The oldest trace fossils attributable to a bone-eating fauna are found on flooding. Small-scale sedimentary sequences are stacked to form six Early-Late Cretaceous plesiosaur and sea turtle bones (about 100 Ma: major, unconformity-bounded stratigraphic units (synthems) of re- Danise and Higgs, 2015). Time estimates suggest that Osedax diverged gional extension, forming a high-resolution framework to study the from other siboglinids in the Middle Cretaceous (around 108 Ma: chronostratigraphic distribution of marine megafauna (MM). Taboada et al., 2015). However, if the bone-eating worm lives also on Benthic biotopes, identified through a quantitative study of a large the bones of birds and terrestrial mammals (Rouse et al., 2011), its • mollusc dataset, can be arranged to form an ideal onshore-offshore, global nature and high species diversity in modern oceans suggest that bathymetric gradient, connecting terrestrial environments with whale falls, as complex and species-rich habitats, have been the most deep sea epibathyal bottoms, consistent with the distribution of important biodiversity generators (Higgs et al., 2014b; Smith et al., sedimentary facies. MM remains and shell beds are present in all 2015). Although the first appearance of Osedax is concomitant with the

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