Paleoecological Insight Into the Straight-Tusked Elephant Population from the Late Middle Pleistocene Site of Poggetti Vecchi

Home , Dwarf elephant, Mammuthus meridionalis, Palaeoloxodon, Primelephas, Sinomastodon

D016 - Scienze della Terra e del Mare DiSTeM - Dipartimento di Scienze della Terra e del Mare Settore Scientifico Disciplinare – GEO/01

Paleoecological insight into the straight-tusked elephant population from the late Middle Pleistocene site of Poggetti Vecchi

IL DOTTORE IL COORDINATORE Chiara Capalbo Alessandro Aiuppa

IL TUTOR CO TUTOR Federico Masini Paul Mazza

CICLO XXX 2014-2017 ANNO CONSEGUIMENTO TITOLO 2018

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Paleoenvironmental context of the early Neanderthals of Poggetti Vecchi for the late middle Pleistocene of Central Italy

Marco Benvenutia*, Jean-Jacques Bahainb, Chiara Capalboa,c, Chiara Caprettid, Francesco Cianie, Carmine D’Amicof, Daniela Esug, GIanna Giachih, Claudia Giulianie, Elsa Gliozzii, Simona Lazzerif, Nicola Macchionif, Marta Mariotti Lippie, Federico Masinic, Paul Peter A. Mazzaa, Pasquino Pallecchii, Anna Revedink, Andrea Savorelli1, Marco Spadij, Lorena Sozzif, Amina Viettib,k, Mario Voltaggiol, Biancamaria Arangurenh aDipartimento di Scienze della Terra, Università di Firenze, 50121 Florence, Italy bDépartement de Préhistoire du Muséum National d’Histoire Naturelle, UMR 7194 du CNRS, 75013 Paris, France cDipartimento di Scienze della Terra e del Mare (DISTEM), Università di Palermo, 90123 Palermo, Italy dCNR - IVALSA, 50019 Florence, Italy eDipartimento di Biologia, Università degli Studi di Firenze, 50019 Florence, Italy fDipartimento di Bioscienze e Territorio, Università degli Studi del Molise, 86090 Pesche (Isernia), Italy gDipartimento di Scienze della Terra, Università Sapienza, 00185 Rome, Italy hSoprintendenza Archeologia della Toscana, 50121 Florence, Italy iDipartimento di Scienze, Università Roma TRE, 00146 Rome, Italy jIstituto Italiano di Preistoria e Protostoria, 50121 Florence, Italy kDipartimento di Chimica, Università degli Studi di Torino, 10125 Turin, Italy lCNR –IGAG, 00015 Rome, Italy

(RECEIVED December 1, 2016; ACCEPTED June 8, 2017)

Abstract Work on thermal pools at Poggetti Vecchi in Grosseto, Italy, exposed an up to 3-meter-thick succession of seven sedimentary units. Unit 2 in the lower portion of the succession contained vertebrate bones, mostly of the straight-tusked elephant, Palaeoloxodon antiquus, commingled with stone, bone, and wooden tools. Thermal carbonates overlying Unit 2 are radiometrically dated to the latter part of the middle Pleistocene. This time span indicates that early Neanderthals produced the human artifacts from Poggetti Vecchi. The elephant bones belong to seven individuals of different ages. Sedimentary facies analysis and paleoecological evidence suggest a narrow lacustrine-palustrine embayment affected by water-level ﬂuctuations and, at times, by hydrothermal water. Cyclic lake-level variations were predominantly forced by the rapid climatic ﬂuctuations that occurred at Marine Isotope Stage (MIS) 6–7 transition and throughout the MIS 6. Possibly an abrupt, intense, and protracted cold episode during the onset of MIS 6 led to the sudden death of the elephants, which formed an unexpected food resource for the humans of the area. The Poggetti Vecchi site adds new information on the behavioral plasticity and food procurement strategies that early Neanderthals were able to develop in Italy during the middle to the late Pleistocene transition. Keywords: Paleoenvironment; Paleoecology; Early Neanderthals; Late middle Pleistocene; Central Italy

INTRODUCTION ecosystems of interacting geological, climatic, and biological factors. A Paleolithic site discovered in Grosseto in Italy during Increased collaboration between natural sciences, archeology, the spring of 2012 illustrates the value of detailed geoarchaeo- and humanities has been bridging conceptual and methodo- logical studies. The site is located at Poggetti Vecchi that is a few logical gaps over the last four decades; this has led to the creation kilometers north of Grosseto, and is an open-air, stratiﬁed, of geoarcheology (Rapp and Hill, 1998; Ghilardi and Desruelles, Paleolithic site that was unexpectedly brought to light at a depth 2009). Geoarchaeology is the most effective means for revealing 2 of around 2.5 m while digging for a new 160 m thermal swim- how past human populations dealt with the impact on ming pool located northward of other preexisting Paleolithic sites. Emergency excavations in a 60 m2 portion of the pool led to the unearthing of human artifacts and vertebrate bones of *Corresponding author at: Dipartimento di Scienze della Terra, Uni- versità di Firenze, 50121 Florence, Italy. E-mail address: ma.benvenuti@ various mammal species, mostlyfromthestraight-tusked uniﬁ.it. (M. Benvenuti). elephant Palaeloxodon antiquus.Researchersfromthe

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Ministry of Cultural Heritage, Activities, and Tourism conducted widespread in southern Tuscany and generate a high geother- the stratigraphic excavation, carefully recording the position of mal gradient (Baldi et al., 1995). all the finds in order to produce GIS maps of each level. The Poggetti Vecchi assemblage of faunal remains and artifacts is radiometrically dated to the latter part of the Data description middle Pleistocene, which indicates that the tools found were This section reports the results of the stratigraphic, paleon- made by early Neanderthals. At transition from the middle to tological, and dating analyses. For the methods applied, see the late Pleistocene, the only evidence of human presence in Supplement Item 1. Italy is the human skeleton from Altamura, which is dated to the Marine Isotope Stage (MIS) 6–7 boundary (Lari et al., Stratigraphy of the site 2015). In fact, most of the Italian sites with evidence of human activity on bones of Paleoloxodon antiquus are The east–west trending, 20-m-long swimming pool excavation assigned to the Lower Paleolithic, e.g., Notarchirico (Piperno (Fig. 1C and 2) exposed vertebrate bones and stone artifacts and Tagliacozzo, 2001), Castel di Guido (Boschian and (Fig. 3A) embedded in a few meters of largely muddy sedi- Sacca, 2014), Atella (Abruzzese et al., 2015), Ficoncella ments of the plain. Several hydrothermal springs were found at (Aureli et al., 2015), and La Polledrara (Santucci et al., 2015). the bottom of the excavation. The abundant discharge of the The Poggetti Vecchi site is particularly interesting from the springs had to be drained with water pumps to keep the sections geoarcheological perspective, because it provides snapshots clear and to recover the specimens. Seven lithostratigraphic of the local environment at a time when early Neanderthals units were encountered in the excavated section; they will be occupied the area. The site is especially valuable because described here, starting from the lowest (Figs. 2 and 3). it demonstrates the considerable skills of Neanderthals to opportunistically reconfigure and vary their activities Unit 1 according to their circumstances and resource availability. Unit 1 consists of about 0.5 m of massive, dark grayish The present study is a part of a wider, multidisciplinary mudstone (Fig. 3B). The fossils present in this unit comprise research project, which aims to reconstruct the behavior and abundant mollusks, ostracods, micro-vertebrates (mammals, ways of life of the early Neanderthal populations in the amphibians, fishes), and plant remains, the latter including region. In particular, this paper focuses on the environmental either unaltered (seeds, carophites, pollen grains) or charred conditions under which middle Pleistocene humans lived in elements. The fossils were found uniformly dispersed in the the area around Poggetti Vecchi. sediment matrix.

Unit 2 GEOLOGICAL BACKGROUND AND The lower boundary of this bone bed is marked by an ARCHEOLOGY OF THE SITE erosional contact. The unit is from 20 to 40 cm thick and mostly yielded remains of Palaeoloxodon antiquus mixed with bone Geomorphic setting and stone artifacts, scattered lithic production waste, but also The Poggetti Vecchi site is located on the northeastern side wooden rods, which are presumably tools (Fig. 3A). Unit 2 is of Grosseto’s plain in southern Tuscany, Italy (Fig. 1A). This starved of sediment from the marginal source areas, and the wide, coastal lowland is surrounded by low elevations com- skeletal remains accumulated to form a condensed layer of posed of Triassic–Neogene clastic and carbonate deposits. The fauna. The top of the bone bed is an irregular surface formed by lowland was a lake, known as Lacus Prilis, through Etruscan the assemblage of large bones and wooden remains. and Roman times, before it became a wetland reclaimed by the nineteenth century (Stea and Tenerini, 1996). Its long-lasting Unit 3 lacustrine-palustrine system resulted in the sandy-muddy sur- Unit 3 is a lithologically composite unit, up to 20 cm thick, face sediments that extend for a few hundred meters under- including two distinct divisions (Fig. 2): a lower one, indi- ground, forming the thick Quaternary coastal basin inﬁll cated as 3a, made up of dark grayish mudstone with gastro- (Bravetti and Pranzini, 1987; Censini and Costantini, 2002). pod shells dispersed at the bottom; and an upper portion, Poggetti Vecchi stands at the foot of a hill some 11 m above the called 3b, which consists of a few centimeters of laterally plain that has developed on Mesozoic carbonate bedrock, discontinuous, whitish, carbonate mudstone. which is extensively exposed on the slopes leading to the town of Roselle to the east (Fig. 1B). The lowland is affected Unit 4 by diffuse karstiﬁcation that results in catastrophic subsoil collapses; one of these sinkholes formed in January 1999 at Unit 4 lays on an erosive surface and consists of millimeter- to Bottegone, about 2 km north of Poggetti Vecchi (Censini and centimeter-sized, clast-supported, coated, carbonate particles Costantini, 2002). The area has also many thermal springs known as pisoliths, with scattered gastropod shells (Fig. 3C). (Bencini et al., 1977), which suggests a local geothermal Pisoliths are typical particles that grow by accretion of bio- anomaly. Small magmatic Neogene-Quaternary intrusions are chemical precipitates in stagnant thermal water; they still form

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Figure 1. (color online) (A) Location of the study area and schematic geological map of the Grosseto plain and surrounding elevations. (B) Oblique aerial view of the Poggetti Vecchi site from Bing Map (https://www.bing.com/maps/?cc=it). (C) Detailed oblique view of the excavation area, with annotated location of cores S1 and S2 and logs A and B reported in Figure 2.

in the artiﬁcial pools that were built at Poggetti Vecchi. yielded microfossils dominated by ostracod shells. These Remains of Palaeoloxodon antiquus associated with stone sediments form a lenticular body up to 30 cm thick, which tools were found at the base of this unit. Sediment samples also tapers towards the northern end of the excavation.

Log B S1 Anthropogenic debris a) b) Unit 7: Carbonate mudstone and Log A sandstone Unit 6: Dark massive mudstone

Unit 5: Pebble-sized conglomerate

Unit 4: Thermal pisolithes Unit 3: a) Dark massive mudstone; a) b) PV9 b) Withish carbonate mudstone PV8 Unit 2: Framework of vertebrate bones, lithic artefacts and woods PV7 Unit 1: Dark massive mudstone PV2 PV1 Paleobiological sample PV5-6 PV4 PV3 ESR/U-series dating sample U/Th dating sample Lithic artefacts ? ? Reworked lithic artefacts ? Vertebrate bones Pisolithes

mudstone, sandstone and breccia predating the site 1 meter

2 meters

Figure 2. (color online) Correlation of stratigraphic data along the cross section indicated in Figure 1C. Samples analysed for mollusk and ostracod contents and for radiometric dating are shown.

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B C

anthropogenic debris anthropogenic unit 7b debris

unit 7a

unit 6 unit 6

unit 4 unit 5

unit 1

unit 3b

Figure 3. (A) Plan view of Unit 2 during the archaeological excavation showingtheframeworkofbones,stones(most of which artifacts), and wooden rods (indicated by red arrows). (B) Part of the section excavated along the northern side of the site where log A was measured. (C) The section excavated on the southern side (log B); note two molars and bones of Palaeloxodon antiquus on top of Unit 4. Ten centimeters divisions for scale in all photographs. (For interpretation of the references to color in this ﬁgure legend, the reader is referred to the web version of this article.)

Unit 5 abundant gastropod shells and plant remains, together with occasional stone implements at the base, associated with a An erosional surface separates this unit from the underlying Unit small amount of wooden items, possibly tools. 4. Unit 5 includes pebble-sized, clast-supported conglomerates with an intervening sandy-muddy matrix, arranged in a lenticular bed up to 25 cm thick (Fig. 3B). Clasts are subangular to sub- Unit 7 rounded and mostly composed of micritic grayish limestone with Unit 7, up to 2 m thick, starts with an erosive base. This unit subordinate pisoliths and gastropod shells. Sparse vertebrate consists of two layers (Fig. 2): the lower one, called 7a, is remains and stone tools were also recovered from this unit. 0.4m of massive, whitish, calcareous mudstone (Fig. 3C), and contains the skull of a juvenile Palaeoloxodon antiquus; Unit 6 the upper one, 7b, consisting of a set of beds made Unit 6 is up to 0.9 m thick and is a massive, dark, grayish up of massive/graded calcareous sandstone and mudstone, mudstone similar to Unit 1 (Fig. 3B and C). The unit contains separated from one another by erosive surfaces (Fig. 3B).

An anthropogenic deposit up to 2 m thick, containing remains Archeological aspects of the site and their from the Roman Era, covers Unit 7 (Fig. 2, 3B–C). consequence The stratigraphic framework was constructed by correlat- fi ing the stratigraphic logs recorded in the southern and The open-air, strati ed Paleolithic site of Poggetti Vecchi includes several anthropic layers: the oldest one is Unit 2, northern portions of the excavation, and two shallow cores, which yielded the richest collection of archaeological items. S1 and S2, that were drilled just outside the excavated The excavated portion of this unit has an area of approximately area (Fig. 2). This revealed that the site used to be in a small 60 m2. As explained above, Unit 2 developed upon the valley extending from the Poggetti Vecchi hill down erosional surface that formed on top of Unit 1, when the lake to the plain. The seven lithological units are enclosed level dropped. Animals (mostly elephants) and groups of within a section, between cores S1 and S2, which widens a hunter-gatherers visited the valley bottom during the short few meters north–south from the excavated area. Cores S1 and S2 were retrieved at depths up to 10 m. The sediments period that this unit represents. The presence of humans is demonstrated by the occurrence of numerous lithic tools and in the upper parts of the cores correspond to the deposits lithic manufacturing waste, obtained from siliceous fluvial of Units 7 and 6, whereas the clastic succession beneath pebbles or cobbles deriving from the erosion of various is dominated, from the top downwards, by massive Apennine mountain geological formations. The most common mudstone with subordinate sandstone, which abruptly passes raw materials used at the site are different types of chert, into reddish mudstone. The latter overlies a calcareous radiolarite, and quartzite. The stone tools are only broadly breccia present at the bottom of the cores. Both successions – indicative of an early middle Paleolithic age. About 200 lithic accumulated before the formation of Units 1 7. Summing up, fl our site consists of deposits piled up in a small valley carved objects (cores, akes, and unworked raw material) and several waste materials (debris and fragments) have been recovered. at the side of a wide lacustrine area, whose water table Most of the lithic implements, mostly scrapers but also notches repeatedly fluctuated, and which was ultimately affected by and choppers, had been retouched. Dozens of tiny flakes the geothermal heat flux. have also been found at the site: these are considered to be by-products of retouching that took place on site. Radiometric dating In virtually all the European middle Pleistocene sites where remains of Palaeoloxodon antiquus are associated Unit 4 contains pisoliths and vertebrate bones suitable with human artifacts, hunting or scavenging is normally for radiometric dating (see Supplement Item 1: Numerical suggested when just one individual elephant is present (e.g., dating). Pisoliths yield a U-series age of 171 ± 3 ka, and a left Weber 2000; Villa et al., 2005; Wenban-Smith et al., 2006; lower molar of Bos primigenius provided an ESR/U-series Yravedra et al., 2010; Aureli et al., 2015; Santucci et al., age of 170 ± 13 ka (Table 1). Based on these ages, the 2015). In contrast, when many elephants are involved, they Poggetti Vecchi succession accumulated during a period are generally assumed to have died due to natural causes spanning the latest MIS 7 interglacial and the whole MIS 6 (Gaudzinski, 1996; Villa et al., 2005; Boschian and Sacca, glacial. 2014; Santucci et al., 2015).

Table 1. (A) Activity ratios and age of the pisolite carbonatic fraction from Unit 4. All errors reported as 1σ. Age calculated using the Isoplot/ EX 3.0 program proposed in Ludwig (2003). (B) ESR/U dating of Auroch’s tooth. A k-value (Da α Internal (mGy/a)) of 0.13 ± 0.02 was used following the procedure outlined in Grün and Katzenberger-Apel (1994). The enamel thickness (Da β (μGy/a)) removed during preparation process was used to calculate the beta contribution. Cosmic dose (Da α (γ +cosm) (μGy/a)) obtained using the formulae of Prescott and Hutton (1994). Uncertainties on the ESR/U-series datings (ESR/U-series ages (US or AU) (ka)) were calculated using Monte-Carlo approach following Shao et al. (2014).

234U/238U 230Th/234U 230Th/232Th (activity (activity (activity 238U 232Th Age ratio) ratio) ratio) (ppm) (ppm) (ka) 1.414 ± 0.014 0.841 ± 0.008 77 ± 5 1.48 ± 0.01 0.049 ± 0.003 171 ± 3

Paleodose U uptake Da α Internal Da β Da(γ +cosm) Da total ESR/U-series Sample Tissue U (ppm) DE (Gy) p-parameter (μGy/a) (μGy/a) (μGy/a) (μGy/a) ages (US) (ka)

PGV1502 dentine 1.84 ± 0.12 91.41 ± 3.03 −0.84 ± 0.06 36 ± 37 69 ± 16 433 ± 20 538 ± 45 170 ± 13 (c n°5485) enamel 0.11 ± 0.01 −0.53 ± 0.08

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Table 2. List of mollusk species and abundance matrix of samples collected from the different layers of the Poggetti Vecchi archaeological site. Ecological classes (EC): AF, Aquatic freshwater; TH, terrestrial hygrophilous; TW, terrestrial woodland; TO, terrestrial openland; TC, terrestrial catholic; TU, terrestrial unassigned.

Samples

Mollusc species EC PV1 PV2 PV3 PV4 PV5 PV6 PV7 PV8 PV9 Viviparus sp. AF – 1 ––––––– Melanopsis etrusca AF 43 33 14 13 9 1 96 17 35 Bithynia leachii AF 15 35 4 58 21 55 30 11 52 Bithynia tentaculata AF 180 191 41 6 55 129 3 – 3 Bithynia sp. AF 106 70 26 – 82 105 3 7 13 Belgrandia thermalis AF 8 7 263 – 166 1 9 14 13 Pseudamnicola moussonii AF 2021 1931 642 2485 458 327 1925 1200 1896 Islamia sp. AF –––––––1 – Valvata piscinalis AF 3 12 4 1 –– – – – Physa acuta AF –––1 –– – – – Galba truncatula AF 2 –––––––1 Radix peregra AF 3 2 6 4 – 2 ––7 Stagnicola palustris AF 2 – 621– 15– Planorbis moquini AF 23 3 – 163 1 – 53 77 243 Planorbis planorbis AF 7 14 15 48 10 19 15 – 5 Gyraulus crista AF – 13– 1 –––– Acroloxus lacustris AF 13 4 13 – 59 58 ––1 Pisidium sp. AF 1 ––––1 ––– Oxyloma elegans TH 1 –––––––1 Vertigo antivertigo TH 2 4 – 6 –– 451 Carychium tridentatum TH ––9 – 2 – 478 Vitrea subrimata TW – 13––– 2 –– Daudebardia rufa TW – 1 ––––––– Vertigo pygmaea TO – 1 ––––––– Pupilla muscorum TO ––––––216 Vallonia pulchella TO 3 2 3 – 1 – 132 Punctum pygmaeum TC –––––––12 Euconulus fulvus TC –––––––1 – Discus sp. TU ––––––––1 Monacha sp. TU ––––––––2 Hygromiidae indet. TU 3 4 – 10 ––15 7 24 Abundance 2436 2317 1052 2797 866 698 2163 1357 2316 Species Richness (S) 18 19 15 12 13 10 15 15 20

Unit 2 at Poggetti Vecchi is the only European site that Poggetti Vecchi were found in a poor state of preservation: is dated to the latter part of the middle Pleistocene (MIS their restoration and analysis are still under way. 6–7 transition) that has yielded the remains of several In Unit 4 of the same site, on the other hand, in the small individuals of Palaeoloxodon antiquus of different onto- area (25 m2) that was excavated, stone implements were genetic ages that were found together with stone, bone, found associated with the tusk and other remains of Palaeo- and wooden artifacts, mostly concentrated in a relatively loxodon antiquus. This Unit is radiometrically dated to about small area at the western end of the excavation. Some of 171 ka, the terminus ante quem for Unit 2. Based on this the implements found were horizontally oriented and dating, the human evidence from the site can be confidently commingled with the elephant remains (Fig. 3A), whereas attributed to the early Neanderthal period. others were found under the bones. The wooden artifacts Unit 5 also revealed stone implements and faunal remains. present unequivocal evidence of human modification, as Yet, only part of the lithic industry is fresh; the remainder in, for example, the removal of small lateral branches and is polished and could, therefore, possibly derive from a the removal of bark, and are therefore of particular inter- secondary context. est, as such finds are very rarely encountered in prehistoric In the case of Unit 6, the lithic industry found at the base of deposits due to the low durability of this material the unit was found scattered with faunal and wood remains. (Schniewind, 1990; Blanchette, 2003; Schoch et al., 2015, This suggests that the shores of the lake were visited at a time and references therein). All the wooden remains of when the water levels were starting to rise. The recurrent

PV9 PV9

PV8 PV8

PV7 PV7

PV6 PV6

PV5 PV5

PV4 PV4

PV3 PV3

PV2 PV2

PV1 PV1

Figure 4. Malacological diagram (percentages of species) from the Poggetti Vecchi archeological site.

association of vertebrate remains and human implements in Paleontological analyses different layers (Units 2-4-5-6) of the Poggetti Vecchi stratigraphic succession indicates that humans visited the area on Mollusks a regular basis, and were hunting and exploiting large mam- Overall, the samples analyzed for this study come from Units mals, especially Palaeloxodon antiquus. 1, 3, and 6. They have yielded rich and well-diversiﬁed The microenvironment generated by the hot thermal assemblages of non-marine mollusks in a good state of springs at Poggetti Vecchi at a time when the climate was preservation. A total of 29 freshwater and terrestrial gastro- cooling on a global scale probably attracted both the local pod species (17 aquatic prosobranchs and pulmonates plus 12 fauna and their human pursuers, and no doubt these land pulmonates) and one bivalve have been identiﬁed. circumstances helped create Poggetti Vecchi’s unusual and Moreover, a few undetermined species of land Hygromiidae interesting co-occurrence of faunal remains and human were also recorded. In many cases, the species found at implements. The site is undoubtedly important because it Poggetti Vecchi are still present today in Europe, Italy dates to the middle-late Pleistocene transition, which is very included. The complete list of species, together with the poorly studied in Europe owing to the lack of evidence abundance matrix, is reported in Table 2. The aquatic found thus far. In Italy, following the recently revised dating gastropods dominate both in number of species (varying from of the remains from Saccopastore to around 250 ka (Marra 8 to 13) and specimens (up to 2781 in sample PV4), whereas et al., 2015), the only prehistoric evidence from the time of the land gastropods are generally represented by lower spe- the MIS 6–7 transition is the human skeleton from Altamura cies diversity (varying from 0 to 9) and especially by a lower (Lari et al., 2015). Sites from this period are also rare in other number of specimens (up to 47 in sample PV9; Table 2). parts of Europe, probably because of the concurrent climate There is, therefore, a much higher number of aquatic than of cooling (Sanchez Yustos and Diez Martin, 2015). In this terrestrial species in the excavated interval. The details of the context, therefore, the Poggetti Vecchi site offers a note- molluscan assemblage from Units 1, 3, and 6 are given worthy chance of gaining new insights into early Neanderthal below. Starting from Unit 1 (PV1-2), the total number of behavioral plasticity and food procurement strategies (cf., shells ranges from 2317 to 2436, and species richness from Binford, 1987; Weber, 2000; Surovell and Waguespack, 18 to 19. Pseudamnicola moussonii dominates among the 2008; Rabinovich et al., 2012; Yravedra et al., 2014; Scott aquatic species (ca. 83%), followed by Bithynia tentaculata et al., 2014). (7–8%), and Bithynia sp. (3–4.5%; Fig. 4), whereas Melanopsis

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Figure 5. Selected examples of mollusc species from the sediment samples collected at Poggetti Vecchi archaeological site. Aquatic species: (A) Melanopsis etrusca Brot; (B) Pseudamnicola moussonii (Calcara); (C) Heleobia stagnorum (Gmelin); (D) Planorbis planorbis (Linnaeus). Land species: (E) Pupilla muscorum (Linnaeus). Scale bar = 1 mm.

etrusca (1.5–2%), Bithynia leachii (1–1.5%), Belgrandia occurrence of B. thermalis and A. lacustris. Land gastropods thermalis (<1%), Valvata piscinalis (<1%), Galba truncatula are very poorly represented occurring with only occasional (<1%), Radix peregra (<1%), Stagnicola palustris (<1%), specimens of V. antivertigo and Hygromiidae indet. Planorbis moquini (<1%), P. planorbis (<1%), and Acroloxus In Unit 6 (PV7-9 samples), the total number of shells ranges lacustris (<1%) are poorly represented. Freshwater bivalves from 1357 to 2316, with species richness ranging from 15 to 20. include a single shell of Pisidium sp., and terrestrial gastropods P. moussonii continues to dominate, with very high percentages consist of occasional hygrophilous taxa (Oxyloma elegans (82–89%), followed by P. moquini,which,inturn,showsan and Vertigo antivertigo), and very rare woodland species increasing upward trend within the unit (2.5–10.5%). The other (Vitrea subrimata and Daudebardia rufa), and openland aquatic taxa are found in much lower frequencies. The spring species (Vertigo pygmaea and Vallonia pulchella). water species Islamia sp. was recorded (with only one specimen) The total number of shells and species richness (1052 for the ﬁrst time in the succession in this unit. Terrestrial species and 15, respectively) plummeted in Subunit 3a (PV3). show a higher shell abundance (25–47) and species richness P. moussonii still dominates, but at a lower percentage (61%) (6–9) than in the other units. In particular, the open-landscape than in the previous samples. B. thermalis becomes pro- Pupilla muscorum,thecatholictaxaPunctum pygmaeum and portionally more abundant (25%) than other aquatic species. Euconulus fulvus,andtheecologicallyunassignedspecies, Land gastropods include only occasional specimens of Discus sp. and Monacha sp., ﬁrst appear in the succession here. Carychium tridentatum, V. subrimata, and V. pulchella. Summing up, the molluscan fauna recovered from the In Subunit 3b, the number of shells ranges from up to 2797 deposits exposed by the excavation, but also from cores S1-2, (PV5 sample) to 698–866 (PV7-9 samples), whereas species is characterized by the dominance of freshwater taxa (Fig. 5). richness varies from 10 to 13. P. moussonii strongly dominates The most common and abundant freshwater species, P. mous- with a high percentage in PV4 (89%), just as in the PV1-2 sonii (from 30% to nearly 90% of abundance), colonizes not only samples. In contrast, its frequency declines abruptly in samples spring water, or stream and river waters not far from the PV5–6 (47%–53%). Among the other aquatic taxa, Planorbi- spring, but also rocky and sandy substrata (Giusti and dae are the best represented (7.5%), with the subordinate Pezzoli, 1980; Giusti et al., 1995; Cianfanelli, 2009). Members

A B C

D E F G

H I J

K L M

Figure 6. Scanning electron micrographs of fossil ostracod valves recovered in the sediments of Poggetti Vecchi archaeological site. (A) Candona (Candona) improvisa Ostermeyer. (B) Candona (Neglecandona) neglecta Sars. (C) Cyprideis torosa (Jones). (D) Darwinula stevensoni (Brady and Robertson). (E) Cypria ophtalmica (Jurine). (F) Cyclocypris ovum (Jurine). (G) Vestalenula cylindrica (Straub). (H) Ilyocypris getica Masi. (I) Ilyocypris monstriﬁca (Norman). (J) Pseudocandona marchica (Hartwig). (K) Scottia pseudobrowniana Kempf. (L) Paralimnocythere messanai Martens. (M) Heterocypris salina (Brady). White bar = 0.1 mm.

of Bithyniidae (B. leachii and B. tentaculata, and Bithynia sp.), 2000; Cianfanelli, 2010). In addition, a single individual of the Lymnaeidae (R. peregra and S. palustris), Planorbidae spring water genus Islamia (Islamia sp.) was recorded in (P. moquini and P. planorbis), and Acroloxidae (A. lacustris), PV7-9. The generally rare occurrence of land hygrophilous which are almost constantly present in the samples (Fig. 4), taxa, such as C. tridentatum, V. antivertigo, and the presence of require slow moving or still, clear, shallow waters found in one species of Succineidae (O. elegans), indicates restricted well-vegetated lakes and ponds (Ložek, 1964; Girod et al., marshland surroundings. Instead, P. muscorum (Fig. 5) and 1980). Furthermore, two species typical of thermal spring V. pulchella in the PV7-9 samples denote an open landscapes waters and slow flowing waters, in this case, M. etrusca (Fig. 5) such as dry grassland and exposed areas (Ložek, 1964; Kerney, and B. thermalis, are frequent, with variable percentages in 1999) in the later stages of the site. Units 1–4 and 6, as well as in core S2. These indicate the presence of thermal springs either nearby (Units 1 and 6) or at Ostracods the site itself (Units 3 and 4). M. etrusca requires running water fed by thermal springs with a high mineral content, All the ostracod valves are well preserved and, generally, each temperatures between 18° and 35°C, and an average pH species includes both adults and juveniles. A population age of 6.8–7.2 (Bartolini et al., 2010). Population densities decrease structure of this type indicates an autochthonous thanatocoenosis moving away from the spring and with faster flowing water typical of an environment with moderate energy (Boomer et al., (Cianfanelli et al., 1991; Manganelli et al., 2000; Bartolini et al., 2003). Thirteen species of ostracods from eleven genera have 2010). M. etrusca, which is often associated with T. fluviatilis, been identified in the nine analyzed samples (Fig. 6, Table 3). B. tentaculata, P. moussonii, B. thermalis, S. palustris, These include: Darwinula stevensoni (Brady and Robertson), P. planorbis, and A. lacustris (Giusti and Pezzoli, 1980; Vestalenula cylindrica (Straub), Candona (Candona) Cianfanelli et al., 1991, 2010; Manganelli et al., 2000; Bartolini improvisa Ostermeyer, Candona (Neglecandona) neglecta et al., 2010), is endemic to Tuscany. Abundant populations of Sars, Pseudocandona marchica (Hartwig), Cypria ophtalmica the species are still present today in 7 sites with thermal waters (Jurine), Cyclocypris ovum (Jurine), Ilyocypris monstrifica between Livorno and Grosseto. One of them is the Poggetti (Norman), Ilyocypris getica Masi, Heterocypris salina (Brady), Vecchi spring. M. etrusca is currently an endangered species Scottia pseudobrowniana Kempf, Paralimnocythere messanai due to the overexploitation of thermal waters (Manganelli et al., Martens, and Cyprideis torosa (Jones).

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Table 3. Percentage frequencies of the ostracod species recovered in the different units of the Poggetti Vecchi archaeological site.

Samples H. salina C. torosa P. marchica C. neglecta C. improvisa C. ovum C. ophtalmica PV9 43 8 40 0 0 4 4 PV8 46 7 17 22 0 2 2 PV7 69 20 0 0 0 0 3 PV6 79 5 12 0 0 0 3 PV5 83 0 7 4 0 0 0 PV4 72 0 9 4 0 2 0 PV3 4 6 8 0 77 2 0 PV2 22 19 7 22 0 4 4 PV1 23 15 5 28 0 15 3

Samples I. monstriﬁca I. getica D. stevensoni V. cylindrica S. pseudobrowniana P. messanai

PV9 4 1 0 0 0 0 PV8 2 2 0 0 0 0 PV7 3 3 0 0 3 0 PV6 3 1 0 0 0 0 PV5 0 0 0 0 0 0 PV4 0 1 2 0 0 0 PV3 0 0 0 2 0 0 PV2 4 4 4 4 0 4 PV1 3 2 3 3 0 0

Almost all samples (except PV3) are dominated by Europe Database (NODE; Horne et al., 1998) and WorldClim H. salina, which is accompanied by P. marchica (particularly (version 1.3; Hijmans et al., 2005) datasets. The inferred abundant in the upper part of the succession), with sub- paleotemperatures were then compared with today’s mean July ordinated amounts of C. torosa and C.(N.) neglecta. All the and January temperatures at Poggetti Vecchi. The results of the other species show very low frequencies and are found scat- MOTR analysis are reported in Figure 8, and they show that tered throughout the succession, except for C.(C.) improvisa, July and January paleotemperatures of 15–22°C for July and which is the dominant species (77%) in the low diversity and −1–4°C for January were both signiﬁcantly below the assemblage recovered in sample PV3. present day mean values, which are 25° C and 7.6°C, Cluster analysis was used to produce the dendrogram as illustrated in Figure 7, with a correlation coefﬁcient of 0.98. Two clusters, A and B, emerge at the very low similarity PV1 PV2 PV7 PV4 PV5 PV6 PV8 PV9 PV3 value of about 0.07. Cluster A includes all samples from the 1.0 archeological excavation site, except one from Unit 3a; these samples are mainly dominated by H. salina and include 0.9 species typical of permanent shallow waters of ponds and lakes (Meisch, 2000; Furhmann, 2012). The presence of 0.8

C. torosa with both noded and smooth shells, combined with 0.7 the predominance of H. salina, indicate slightly brackish waters, with salinity values below the osmoregulation 0.6 threshold of C. torosa (8 psu; Keyser, 2005; Frenzel et al., 0.5 2012). The co-occurrence of P. marchica restricts the likely Similarity salinity range to a maximum of 4 psu (Grifﬁths and Holmes, 0.4 2000). Such a low salinity is testiﬁed also by the abundance of Chara gyrogonites. Cluster B, on the other hand, includes 0.3 only sample PV3: it is dominated by C.(C.) improvisa, 0.2 a species typical of seasonal pools (Meisch, 2000). A B The Mutual Ostracod Temperature Range (MOTR) method 0.1 was applied to the assemblages of cluster A in Figure 1 to estimate the paleotemperature ranges at the time of the 0.0 deposition of the sediment. Horne (2007) and Horne et al. Figure 7. Dendrogram resulting from cluster analysis in Q-mode (2012) created this method for determining paleotemperature using the UPGMA method and the Morisita–Horn distance ranges by combining the Nonmarine Ostracod Distribution in applied to the samples from the Poggetti Vecchi archeological site.

Mean July air temperature ranges Mean January air temperature ranges

H. salina

P. marchica

C. neglecta

C. ovum

C. ophtalmica

I. monstrifica

D. stevensoni

S. pseudobrowniana

5° 10° 15° 20° 25° 30° 35° -30° -25° -20° -15° -10° -5° 0° 5° 10° 15°

25° 7.6° Figure 8. (color online) Estimated average air palaeotemperatures during the deposition of the sediments in the Poggetti Vecchi archaeological site based on the recovered fossil ostracods. Black lines indicate the tolerance temperature ranges for each species. The pink and light blue ﬁelds indicate the estimated July and January average temperature ranges in which the recoverd ostracod species could cohexist. The red arrows indicate the present July and January average temperatures for the Poggetti Vecchi area. (For interpretation of the references to color in this ﬁgure legend, the reader is referred to the web version of this article.)

respectively (these being the average values for the period (Bos primigenius), predominate in the excavation. Based on 2005–2014; data retrieved from the hydrological station NISP 67% and 32.6% were found for the two species, respec- Grosseto -Servizio Idrologico Regionale Toscana). tively. Browsing herbivores, such as the red deer (Cervus elaphus), roe deer (Capreolus capreolus), and fallow deer Vertebrates (Dama dama)aremuchrarer.Cervus elaphus,forexample,is the only middle-sized species present in the Unit 2 assemblage; Unit 2 was particularly rich in vertebrate remains, whereas whereas the remains of roe deer and fallow deer total 1.4% and several isolated bones were found in Units 4 and 6. A total of 1.2% of the bones of the small-sized species, respectively. 634 specimens were retrieved from Unit 2: 149 (23.5%) of AveryadvancedUrsus deningeri (transitional to U. spelaeus; them are undetermined. Table 4 and Figure 9 report the list of Auguste, 1992; García et al., 1997; Pacher and Stuart, 2008) is the species, with their relative frequencies and abundances attested to by a right upper canine and a left m2. The mammal expressed in Number of Identified Specimens (NISP) counts. remains were accompanied by some scanty remains of the Eur- The remains of very large-sized grazers, i.e., the straight- opean pond turtle (Emys orbicularis), together with a vertebra of tusked elephant (Palaeloxodon antiquus)andtheaurochs the European whip snake (Hierophis viridiflavus), and two fragmental bones of birds. Table 4. Vertebrate taxa from Poggetti Vecchi, with relative Only three specimens, in this case the lateral portion of frequencies expressed in Number of Identified Specimens (NISP) diaphysis of a left humerus of Bos primigenius and the basal counts. portion of a shed right antler and a fragmental diaphysis of a

Taxa NISP NISP % Palaeoloxodon antiquus 283 39.70 45.00 40.00 Bos primigenius 138 19.35 35.00 Capreolus capreolus 12 1.70 30.00 Dama dama 1 0.14 25.00 20.00 Cervus elaphus 38 5.80 NISP % 15.00 Ursus deningeri-spelaeus 2 0.28 10.00 Arvicola terrestris 62 8.70 5.00 0.00 Terricola sp. 2 0.28 Microtus cf. arvalis 1 0.14 Aves

Dama dama Terricola sp. Aves 2 0.28 Indetermined Bos primigenius Cervus elaphus Emys orbicularis Emys orbicularis 4 0.56 Arvicola terresMicrotus tris cf. arvalis Doubtful attribution C apreolus capreolus Hierophis viridiflavus Palaeoloxodon antiquus Hierophis viridiﬂavus 1 0.14 Ursus deningeri-spelaeus Doubtful attribution 74 10.37 Undetermined 93 13.04 Figure 9. (color online) Relative frequencies of the vertebrate taxa Total 713 of Poggetti Vecchi, expressed in Number of Identiﬁed Specimens (NISP) counts (data reported in Table 4).

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Figure 10. Pollen diagram of samples collected on the wall excavation of log A: selected taxa. Exaggeration 15 ×.

tibia of Cervus elaphus, show evidence of human modiﬁcation. Particularly striking is the presence of at least nine left and three No such evidence has been observed on the elephant remains. right shed antlers of this species. Also, remains of a single adult The terms of micromammals in Unit 2 (Table 4), these are individual of Capreolus capreolus were found, alongside those dominated in NISP counts by the watervole Arvicola of a fetus, a juvenile, and an adult of Bos primigenius.Allthe amphibius. This species, which is characterized by indivi- other taxa include one individual each. duals with undifferentiated dental enamel (sensu Heinrich, 1978), is typical of waterside habitats. This water vole totals Pollen approximately 95% of the small mammal counts. The genus Microtus, on the other hand, is represented in Unit 2 by only All the samples contained a fairly rich amount of well- three specimens: a fragmental left m1, attributed to Microtus preserved pollen grains, yielding a total of more than 100 taxa cf. arvalis, a left m1 of Microtus (Terricola) ex gr. savii, and (Fig. 10). In all of the samples, herbaceous plants (NAP) a right M1 ascribed to Microtus sp. The ﬁrst lower molars dominate, whereas arboreal, shrubby and climbing plants found show a relatively short and wide anteroconid as well as (AP) never exceed 22%, indicating the persistence of an a wide anterior cap, which are features suggesting a Microtus open environment throughout the whole time period under (Terricola) species of the savii group. examination. A cross examination of Units 4 and Unit 6 shows that they Noteworthy, among the arboreal taxa is the presence of yielded fossil assemblages very similar to that found in Tertiary elements in the lowermost samples of Unit 1, such as Unit 2. Indeed, the relative frequencies in Unit 4 are virtually Tsuga, cf. Keteleeria, and Carya. Two different morphotypes the same as those observed in Unit 2. In Unit 6, however, the (i.e., with different exine thickness) of Ulmaceae suggest the frequency of Arvicola amphibius drops to 88%. Microtus possible presence of Zelkova in addition to Ulmus. These spp. also grow progressively less abundant from Unit 6 to morphotypes occur throughout the sequence but always in Unit 2. Although the cheek teeth (M3 and m1) diagnostic of low amounts. Buxus is always present in low concentrations Microtus cf. arvalis are missing in the arvicolid assemblages throughout the succession. Its presence is unusual because from Units 6 and 4, we cannot exclude its undetected Buxus grains are easily degraded. presence in either unit. Worth noting is the occurrence of a Poaceae, on the other hand, are dominant and represented fragmental mandible of Crocidura cf. suaveolens and a by different morphotypes, and this indicates the presence fragmental maxillary bone of Oryctolagus sp. in Unit 6. of extensive grasslands on the plain. The most frequent In terms of large mammals, based on the evidence of the morphotype (A) is quite small, ranging from 22 to 28 µm, skull, mandible, molar, and tusk remains from Unit 2, Palaeo- with a thick annulus measuring about 5 µm in diameter. loxodon antiquus is represented by four juveniles in the range of A noteworthy variety of pollen grains of hygrophytes 1–8yearsofage,togetherwitha14–15 year-old subadult, and and hydrophytes were recorded, indicating habitats with two 25–30 year-old adults. Cervus elaphus is also represented high biodiversity. They include Poaceae t. Glyceria type, by at least four individuals, ranging from 3 to 12/13 years of age. Cyperaceae, Juncaceae, Alismataceae, Lythrum, cf Butomus,

Figure 10. Continued.

Callitriche, Potamogeton, cf. Sparganium t., Sagittaria, The Boxwood remains above all consist of straight Typha spp., cf Lemna, Nymphaea, and Nuphar. Their branches, most of whose surfaces of most of these branches are amounts change throughout the stratigraphic sequence. partially or entirely blackened. Buxus is a small, slow-growing evergreen tree or shrub. It usually lives in forests of larger trees, associated with Juniperus.Itcanalsobefoundnextto Wood remains xero-termophilus taxa, such as Quercus,inshrubsorthicketson An isolated fragment of Ulmus sp. (elm) was recovered from well-drained calcareous substrates on plains to lower slopes. Unit 1, at the base of the stratigraphic succession. Unit 2 The high concentration of straight boxwood branches found in contained several horizontally oriented wood fragments the Units 2 and 6 may indicate that they were intentionally interspersed with the elephantremains.Mostofthewooditems selected by humans to make wooden implements. The study of were concentrated in a relatively small area of the excavation. these purported wooden tools is presently under way. AfewwoodfragmentswerealsofoundatthebaseofUnit6. Buxus sempervirens L. (boxwood) is dominant throughout DISCUSSION the succession, whereas decidous Quercus sp. (deciduous oak), Fraxinus sp. (ash), Juniperus sp. (juniper), and Populus/Salix Paleoenvironmental reconstruction at the (poplar/willow) are barely represented. The relative number of archeological site of Poggetti Vecchi fragments for each of these trees is summarized in Table 5. The integration of the multiproxy paleobiological data with the stratigraphy at the site has allowed for a detailed reconstruction Table 5. Taxa identification of the wood fragments found in the of the landscape and an understanding of its evolution during succession of Poggetti Vecchi. the deposition of Units 1–6. The stratigraphic data together Lithological Unit Taxa No. of fragments with the information provided by the fossil mollusk and ostracod assemblages indicate four different flooding events sepa- 6 Buxus sempervirens 9 rated by regressions. During the flooding episodes, organic Decidouos Quercus 1 muds were deposited in a protected embayment of a shallow Populus/Salix 2 lake with rather homogeneous physicochemical characteristics. 2-3 Buxus sempervirens 36 In contrast, concentrations of vertebrate remains and human Decidouos Quercus 2 artifacts accumulated on the erosive surfaces that formed dur- Fraxinus sp. 2 Juniperus sp. 1 ing the regressions. The succession of the events is as follows. 1 Ulmus sp. 1 A narrow valley was carved in the oldest deposits found in cores S1–S2, along the edge of a lake. A very shallow

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bay formed in this valley during the first episode of lake- red deer (Cervus elaphus) and the roe deer (Capreolus level rise, and subsequently lacustrine sediments started to capreolus) browsed in sparse groves. The abundant fre- accumulate. The deposits of Unit 1 record the infrequent quency of freshwater fens, together with a high variety of sedimentation of lacustrine mud on a poorly oxygenated wetland plants, denotes the presence of seeps and water bottom that favored the preservation of organic matter. bodies. Given the flux of wetland vegetation throughout the Abundant and diversified mollusk and ostracod assemblages, sedimentary succession, the bodies of water must have dominated by P. moussonii and H. salina, respectively, repeatedly expanded and contracted. Trees likely grew on the inhabited the bottom and this indicates fresh or slightly saline hills but also in well-drained areas of the plain; the pre- waters (<4 psu). When the bay dried up an erosive surface dominance of Buxus, in particular, indicates well-drained, formed, separating Unit 1 from Unit 2. As the lake receded, calcareous soils. It is often associated with travertines. elephant carcasses and various human artifacts accumulated on its now-exposed shores. Chronological and paleoclimatic implications Unit 3 records a second lacustrine phase. A new rise in the water table initially created a shallow, mud-dominated pool The paleontological information reported above, but also the in the small embayment (Unit 3a); the predominance of the radiometric dating, indicates that the Poggetti Vecchi site ostracod C.(C.) improvisa in this particular horizon may developed during the late Quaternary, a period characterized reflect seasonal fluctuations of the water level. This event was by frequent climate fluctuations. Although infrequent, the then followed by the deposition of carbonate mud (3b), which presence of Tertiary pollen grains in the lowermost samples of is the first significant evidence of thermal activity at the site. Unit 1 is striking because these plants were thought to have The pisoliths of Unit 4 document the formation, during the disappeared from central Italy during the early-middle Pleis- late middle Pleistocene (MIS 6), of a thermal pool similar tocene. Specifically, Tsuga and Carya had a marked decline in to the man-made ones that were built in modern times. This northern and Central Italy during the early middle Pleistocene, MIS 6 thermal pool was visited by both humans and large disappearing between 0.75 and 0.45 Ma, although they may mammals, which included Palaeoloxodon. Although the have survived longer in Southern Italy (Corrado and Magri, invertebrate assemblages were still dominated by P. moussonii 2011; Orain et al., 2013). Zelkova rapidly reduced in cold-arid and H. salina at this point, B. thermalis and M. etrusca reach periods and expanded again in warm-moist periods and finally their highest frequency in Unit 3, confirming the thermal became extinct in peninsular Italy before the latest glacial influence on the waterbody. maximum (Follieri et al., 1986). Based on these data, the lower The second lacustrine phase ended with a new lake-level fall, part of Unit 1 could not be younger than 0.4 Ma. We cannot and the older deposits were dissected by fluvial downcutting exclude the possibility that the Tertiary pollen grains are (Unit 5). In fact, a short creek flowing down from the Poggetti actually allochthonous elements that have been reworked from Vecchi hill drained the valley. In this stage, runoff flow over the older stratigraphic levels and that they may have migrated into valley floor was of sufficient energy to transport the calcareous higher beds via the capillary rise of mud. pebbles that formed the channel lag of Unit 5. The numerical ages of 171 ± 3 ka and 170 ± 13 ka A new flooding (third phase) once again restored the lake obtained, respectively, from the pisoliths and Auroch’s tooth embayment, as attested to by the deposits of Unit 6. In this found in Unit 4 locate the studied succession within an case, the depositional conditions were virtually identical to interval bracketing the transition between the latest MIS 7 those when Unit 1 was formed. and the whole of MIS 6. The physical stratigraphy of the Yet another drop in lake-level fall led to a new, deep incision transgressive and regressive deposits that formed during the of the valley floor. The erosive surface separating Unit 7 from time interval indicated by the age determination of Unit 4 older units remained exposed for a while, as indicated by the suggests the following tentative calibration (Fig. 11). occurrence of the isolated skull of Palaeoloxodon,aswellasthe Unit 1 transgression: lake highstand during the warm abundant presence of terrestrial gastropods. substage MIS 7a or during the following interstadial at the Finally in the fourth phase, the valley floor was once again transition to MIS 6 (between 200–191 ka). Under either of carved and infilled with calcareous mud (Subunit 7a). This these hypotheses, the Tertiary flora recorded in the lower part recreated the lake bay, but this time most likely under thermal of Unit 1 would have survived in this area more than 0.4 Ma. influence. The valley was infilled with calcareous sand and Unit 2 regression: lake lowstand and subaerial erosion at subordinate mud (Subunit 7b), these last reworked from the onset of MIS 6. thermal carbonates by the water flowing toward the lake. The Units 3 and 4: lake transgression and activation of the thermal carbonate composition of Unit 7 indirectly indicates that the system in the first interstadial of MIS 6 (around 170 ka). thermal activity had finally stabilized. Unit 5 regression: lake lowstand at the successive stadial. Pollen, terrestrial mollusks, and vertebrate remains provide Unit 6 transgression: lake highstand during the second further information about the overall environmental condi- interstadial. tions surrounding of the site. At the time when the fossili- Unit 7 regression: lake lowstand at the acme of MIS6 ferous site was formed, the Poggetti Vecchi area was (around 130 ka). Deposition of Unit 7 may have occurred dominated by extensive open grassland; Palaeoloxodon during the climatic rise in temperatures culminating with the antiquus and Bos primigenius grazed the prairie, while the peak interglacial interval MIS5e.

Figure 11. (color online) Tentative correlation of the Poggetti Vecchi Stratigraphy, calibrated for Unit 4 with radiometric datings to the Late Quaternary Marine Isotope Stages.

The paleobiological inferences based upon the Poggetti Vec- as P. muscorum and V. pulchella from Unit 6, indicates that chi small mammal remains providefurtherinformationsup- this Unit formed under cool climatic conditions (Rousseau, porting the tentative synchronization of the succession with the 1989; D’Amico et al., 2014). These conditions are compa- main climatic events of the associated time period. The dominant tible with the global cooling trend that culminated in the Arvicola amphibius is a typical inhabitant of moist environ- glacial maximum of MIS 6. ments. The presence of Microtus cf. arvalis in Unit 2, the The ostracod fauna that was collected from Units 1–6 and decreasing abundances of M. (Terricola)exgr.savii and the analyzed with the MOTR method suggest there were mean absence of Crocidura and Oryctolagus in the following units, annual temperatures about 6°C lower than today, which is in might indicate cooler conditions moving upwards in the suc- line with the temperatures at the onset of MIS 6 based on cession. Modern Italian populations of Microtus cf. arvalis are estimates from the Vostok Ice core in Antarctica, which confined to the northern regions of the peninsula and are absent indicates 7.5°C lower than today (Petit et al., 1999). In light from the central Apennine further south. Therefore, its presence of Ayalon et al.’s (2002) study, the humid period dated at in southern Tuscany at very low altitudes is indicative of colder 171 ka, which is also the calculated age of Unit 4, was likely conditions at the time of Unit 2 than those existing today. the first interstadial of the glacial interval MIS 6. In contrast the occurrence of Crocidura cf. suaveolens, Oryctolagus sp., and Microtus (Terricola) savii in Unit 6 is CONCLUSIONS indicative of warmer habitats. The large mammal assemblage and the herpetofauna from The radiometric dating of about 171 ka obtained from the Unit 2 including thermophilic taxa, such as the roe deer pisoliths and third lower molar of Bos primigenius of Unit 4 Capreolus capreolus,theEuropeanpondturtleEmys orbicularis, places the Poggetti Vecchi archaeo-paleontological site in and the European whip snake Hierophis viridiflavus,showawide a very crucial time span. This time period tells us that distribution ranging from the warmperi-Mediterraneanarea,all early Neanderthal populations produced the implements. the way into the much cooler northern European contexts Archaeo–paleontological sites attesting to the middle to late (Lelièvre et al., 2010a, 2010b). The European pond turtles are Pleistocene transition are scarcely documented in Europe. usually only active above 5°C and so are another indicator Poggetti Vecchi therefore offers the chance of obtaining an of temperatures across the region (Novotnýet al., 2004). unprecedented amount of information on the behavior of Palaeoloxodon antiquus is typical of temperate contexts but early Neanderthals under the circumstances that have been also occurred in cooler conditions (Mazza et al., 2006). documented through from this study. On the whole, the fauna of Unit 2 comprises of species with Our analysis shows that at the time when the bone accumu- varied climatic distributions that seem to be well-tuned with the lation was formed the area consisted of widespread, open, wet fluctuations that occurred within the relatively short period grassland, scattered with water bodies, woods, and, perhaps, between the onset of MIS 6 (lake regression and erosion at the top riverine belts of trees. The hot thermal springs in one of these of Unit1), and the next interstadial (lake transgression and water bodies located in a small embayment, probably created an deposition of Units 3 and 4). Intense seasonal oscillations can attractive recess for the resident fauna, especially in a time period reasonably be expected to have occurred during the time span when the climate was globally deteriorating. It would appear recorded by the Poggetti Vecchi succession. The occurrence of from the artifacts found that the area could have been routinely Microtus cf. arvalis is not at odds with the hypothesis that perhaps patrolled by early Neanderthal hunter-gatherers who were mon- acoldsnapoccurredduringthetimeofthedepositionofUnit2, itoring the movements of the fauna that tended habitually to which might have caused the sudden death of the elephant clan. congregate in the warmer valley of Poggetti Vecchi. We can The molluscan fauna discloses relevant information that speculate that during a cold snap, large amounts of game ani- may assist in defining the climate regime during the deposi- mals would gather in the recess. Many of them were possibly tion. The recorded presence of open-landscape species, such debilitated by the harsh, inhospitable circumstances, or most

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probably already lay dead, as perhaps happened in the case of Piombanti, P.P, Santinelli, M. (Eds.), Codice Armonico, Terzo the elephants. Their concentrated presence represented a valu- Congresso di Scienze Naturali, Ambiente toscano, Castiglioncello able resource that could have attracted the local human pur- (LI)1, sezione Scientifica.EdizioniETS,Pisa,Italy,pp.63–70. suers. If this reconstruction is correct, the collected evidence Bencini, A., Duchi, V., Martini, M., 1977. Geochemistry of thermal indicates that these early Neanderthals were particularly skilled springs of Tuscany (Italy). Chemical Geology 19, 229–252. at modifying their survival strategies to take advantage of the Binford, L.R., 1987. Data, relativism and archaeological science. Man 22, 391–404. particular environmental circumstances and opportunities pre- Bischoff, J., Rosenbauer, R., Tavoso, A., de Lumley, H., 1988. A sented to them in a changing environment. test of uranium-series dating of fossil tooth enamel: results from Tournal Cave, France. Applied Geochemistry 3, 145–151. Blanchette, R.A., 2003. Deterioration in Historic and Archaeological ACKNOWLEDGMENTS Woods from Terrestrial Sites. In: Koestler, R.J., Koestler, V.R., Charola, A.E., Nieto-Fernandez, F.E. (Eds.), Art, Biology and We are particularly indebted to the owner of the site A. Ceccarelli, Conservation: Biodeterioration of Works of Art.TheMetropolitan who funded the excavation. We wish to thank A. Pessina and Museum of Art, New York, pp. 328–347. G. Poggesi for their support during the completion of this study, Boomer, I., Horne, D.J., Slipper, I.J., 2003. The use of ostracods in S. Pozzi, S. Caloni and S. Caramiello for the restoration of the fossil palaeoenvironmental studies, or what can you do with an ostracod bones. We are very grateful to M.-H. Moncel, D. Bridgland, and shell? In: Park, L.E., Smith, A.J. (Eds.), Bridging the Gap: Trends C. Marean for significantly improving the manuscript. We thank in the Ostracode Biological and Geological sciences. The K. Eadie and J. A. Thonn for the language editing. This study was Paleontological Society Papers 9, Boulder, pp. 153–180. financially supported by the Italian Ministry of Cultural Heritage, Boschian, G., Sacca, D., 2014. In the elephant, everything is good: the “Fondazione Banca Federico del Vecchio” and the “Ente Cassa carcass use and reuse at Castel di Guido (Italy). Quaternary di Risparmio di Firenze”. Additional funding for the research International 361, 288–296. was provided by Italian MIUR (the Italian Ministry of Education, Bravetti, L., Pranzini, G., 1987. L’evoluzione quaternaria della pianura University and Research) ex60% grants to the Department of Earth di Grosseto (Toscana): prima interpretazione dei dati del sottosuolo. 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'RZQORDGHGIURPKWWSVZZZFDPEULGJHRUJFRUH7HDFKHUV&ROOHJH/LEUDU\&ROXPELD8QLYHUVLW\RQ$XJDWVXEMHFWWRWKH&DPEULGJH&RUHWHUPVRIXVHDYDLODEOHDW KWWSVZZZFDPEULGJHRUJFRUHWHUPVKWWSVGRLRUJTXD 344 M. Benvenuti et al.

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