Vegetation reconstruction on the basis of pollen in Late Pleistocene hyena coprolites from San Teodoro Cave (Sicily, Italy)

Vegetation reconstruction based on pollen from coprolites of extinct spotted hyena (Crocuta crocuta spelaea, Goldfuss 1832) recovered from excavations carried out in 1998 at San Teodoro Cave (Sicily, Italy) supports previous indications of pre-Late Glacial conditions. Eight of the twelve coprolites analysed contained well-preserved pollen grains. There is a general similarity between the pollen contents from the coprolites but they show variability. They suggest a main vegetation type dominated by steppic taxa (Poaceae, Artemisia, Ephedra, Chenopodiaceae and Asteraceae) but also including arboreal taxa (Pinus and Cupressaceae). Low percentages of pollen of mesophilous woody taxa (Quercus, Betula, Abies, Alnus, Pistacia, among others) are noticeable, suggesting the existence of nearby refugia for temperate and Mediterranean vegetation. A reconstruction of the landscape, using the coprolite pollen record and other pollen records from Sicily and south Italy, shows the predominance, during the pre-Late Glacial, of a wooded steppe biome, with elements representing a variety of local environmental conditions.     

Which predators are responsible for faunal accumulations at the Late Pleistocene layers of El Harhoura 2 Cave (Témara,Morocco)?

El Harhoura 2 cave (Témara, Morocco) has yielded Aterian and Iberomaurusian lithic artifacts associated with faunal remains. Both humans and carnivores occupied this cave and non-human predator modifications occurred mainly at the end of the Late Pleistocene. Diverse faunal taxa have been identified, with a predominance of gazelles and various carnivores, particularly canids. The location of the cave and of the excavation area, at the bottom of a cliff of low elevation and in the entrance of the cave, does not correspond to a protected area for large raptor nests or a natural trap. Considering the consumed species, the type of carnivore remains, the skeletal representation of prey, taphonomic alterations such as tooth marks, semi-digested bones and destruction sequences, large canids would be the main cause for faunal modifications. However, North African fossil data attributed to hyena activities present similar results. This paper highlights the difficulty of discriminating between potential accumulators/consumers due to a lack of taphonomic reference data. This study thus demonstrates the necessity of compiling fossil records and neotaphonomic reference data for North African medium-large predators in order to better understand the taphonomic history of North African archaeological and paleontological sites.     

Late Pleistocene skeleton of Canis lupus l., 1758 from Grotta Mora Cavorso (Jenne,Latium, central Italy)

This paper describes the skeleton of Canis lupus found in Layer 7 of Grotta Mora Cavorso (Latium, central Italy), correlated with the MIS 3. Research on the deposition dynamics of this find is still in progress, but the action of human or other predators can be excluded. This specimen consists of a near complete articulated skeleton, lacking only the calvarium. The age at death was estimated at around six years old. Preliminary morphometric analyses and comparisons with samples of Italian Late Pleistocene wolf and extant Apennine wolf (C. lupus italicus) remains, show that the dimensions of the teeth and long bones are among the biggest known from the Italian Late Pleistocene and larger than the extant Apennine wolf.     

Coprolites as a source of information on the genome and diet of the cave hyena

We performed high-throughput sequencing of DNA from fossilized faeces to evaluate this material as a source of information on the genome and diet of Pleistocene carnivores. We analysed coprolites derived from the extinct cave hyena (Crocuta crocuta spelaea), and sequenced 90 million DNA fragments from two specimens. The DNA reads enabled a reconstruction of the cave hyena mitochondrial genome with up to a 158-fold coverage. This genome, and those sequenced from extant spotted (Crocuta crocuta) and striped (Hyaena hyaena) hyena specimens, allows for the establishment of a robust phylogeny that supports a close relationship between the cave and the spotted hyena. We also demonstrate that high-throughput sequencing yields data for cave hyena multi-copy and single-copy nuclear genes, and that about 50 per cent of the coprolite DNA can be ascribed to this species. Analysing the data for additional species to indicate the cave hyena diet, we retrieved abundant sequences for the red deer (Cervus elaphus), and characterized its mitochondrial genome with up to a 3.8-fold coverage. In conclusion, we have demonstrated the presence of abundant ancient DNA in the coprolites surveyed. Shotgun sequencing of this material yielded a wealth of DNA sequences for a Pleistocene carnivore and allowed unbiased identification of diet.     

Past distribution and ecology of the cork oak (Quercus suber) in the Iberian Peninsula: a pollen-analytical approach

Abstract.  This study presents pollen-analytical data from continental and offshore Iberian Peninsula sites that include pollen curves of Quercus suber , to provide information on the past distribution and ecology of the cork oak ( Q. suber ). Results centre on a new pollen record of Navarrés (Valencia, eastern Spain), which shows that the cork oak survived regionally during the Upper Pleistocene and was important during a mid-Holocene replacement of a local pine forest by Quercus -dominated communities. This phenomenon appears linked to the recurrence of fire and reinforces the value of the cork oak for reforestation programmes in fire-prone areas. In addition to Navarrés, other Late Quaternary pollen sequences (Sobrestany, Casablanca-Almenara, Padul, SU 8103, SU8113, 8057B) suggest last glacial survival of the cork oak in southern and coastal areas of the Peninsula and North Africa. Important developments also occur from the Late Glacial to the middle Holocene, not only in the west but also in the eastern Peninsula. It is suggested that, in the absence of human influence, Q. suber would develop in non-monospecific forests, sharing the arboreal stratum both with other sclerophyllous and deciduous Quercus and Pinus species.     

Animal resources and subsistence strategies

Several faunal assemblages excavated in deposits of different antiquity (from Lower Paleolithic to Bronze Age), located in Northern, Central and Southern Italy, were studied from the archeozoological and taphonomic point of view. Data obtained by different Authors allow reconstruction of subsistence strategies adopted by prehistoric humans in these areas and through time, in particular as far as the exploitation of animal resources is concerned. The following assemblages were considered: Isernia La Pineta (Molise; Lower Paleolithic), Grotta Breuil (Latium; Middle Paleolithic), Grotta della Ghiacciaia (Verona; Middle Paleolithic), Riparo di Fumane and Riparo Tagliente (Verona; Middle and Upper Paleolithic), Riparo Mochi (Liguria; Upper Paleolithic), Grotta della Continenza (L'Aquila; Upper Paleolithic and Mesolithic), Grotta dell'Edera (Trieste; Mesolithic and Neolithic), Grotta della Cala at Marina di Camerota (Salerno; Eneolithic), Contraguda (Sassari; Neolithic), Castellaro Lagusello (Mantova; Bronze Age). Exploitation of the vegetal resources has been analyzed in the Neolithic sites of Colle Santo Stefano (Fucino), Settefonti (L'Aquila) and Catignano (Pescara).     

Dento-alveolar lesions and nutritional habits of a Roman Imperial age population (1st–4th c. AD): Quadrella (Molise, Italy)

The study of teeth is very important in archaeoanthropology for reconstruction of the nutritional habits and living conditions of past populations. We have analysed dental lesions of pathological (caries, abscesses and ante mortem tooth loss) and non-pathological origin (calculus), linear enamel hypoplasia and tooth wear in 67 adults from the Roman Imperial age necropolis (1st-4th c. AD) of Quadrella (Molise, Italy). The high frequency of caries (likely giving rise to the abscesses and ante mortem tooth loss), the abundant calculus and the low frequency of heavy wear are probably due to a limited use of hard fibrous foods and a high consumption of carbohydrates. The high frequency of linear enamel hypoplasia suggests metabolic problems during growth. Comparison of these data with those for two coeval Italian necropoleis near Rome (Latium), Isola Sacra and Lucus Feroniae, indicates poorer living conditions in the Quadrella population.     

Hominid tool-making versus carnivore scavenging

The effects of hyena scavenging on the remains of 75 animals in the Awash National Park, Ethiopia, were examined to provide a modern analogy to the fossil assemblage from Makapan. Brain's recent work ('70, '76) suggests that the actions of carnivores may account for the characteristics of the Makapan assemblage, a theory which contrasts with Dart's ('49, '55, '57, '59) interpretation of the fossils as osteodontokeratic tools and weapons manufactured, collected, and used by australopithecines. The modern assemblage was like that from Makapan in terms of relative representation of skeletal elements and patterns of bone breakage. Observations of hyenas in the Awash also lend support to the hypothesis that the Makapan assemblage may represent the uneaten remains of carnivore kills. The representation of major vertebrate groups at Makapan was compared with their representation at Awash and with their representation in hyena diets in the Serengeti and Ngorongoro Crater (Kruuk, '72). This technique of faunal analysis was unable to detect differences between these groups, despite the fact that the three modern assemblages represent animals that died of different causes. Thus extreme caution must be used in deducing any specific information about the mode of death or taphonomic history of the animals represented in an assemblage on the basis of such faunal analyses.     

A Human Deciduous Tooth and New 40Ar/39Ar Dating Results from the Middle Pleistocene Archaeological Site of Isernia La Pineta,Southern Italy

Isernia La Pineta (south-central Italy, Molise) is one of the most important archaeological localities of the Middle Pleistocene in Western Europe. It is an extensive open-air site with abundant lithic industry and faunal remains distributed across four stratified archaeosurfaces that have been found in two sectors of the excavation (3c, 3a, 3s10 in sect. I; 3a in sect. II). The prehistoric attendance was close to a wet environment, with a series of small waterfalls and lakes associated to calcareous tufa deposits. An isolated human deciduous incisor (labelled IS42) was discovered in 2014 within the archaeological level 3 coll (overlying layer 3a) that, according to new ⁴⁰Ar/³⁹Ar measurements, is dated to about 583–561 ka, i.e. to the end of marine isotope stage (MIS) 15. Thus, the tooth is currently the oldest human fossil specimen in Italy; it is an important addition to the scanty European fossil record of the Middle Pleistocene, being associated with a lithic assemblage of local raw materials (flint and limestone) characterized by the absence of handaxes and reduction strategies primarily aimed at the production of small/medium-sized flakes. The faunal assemblage is dominated by ungulates often bearing cut marks. Combining chronology with the archaeological evidence, Isernia La Pineta exhibits a delay in the appearance of handaxes with respect to other European Palaeolithic sites of the Middle Pleistocene. Interestingly, this observation matches the persistence of archaic morphological features shown by the human calvarium from the Middle Pleistocene site of Ceprano, not far from Isernia (south-central Italy, Latium). In this perspective, our analysis is aimed to evaluate morphological features occurring in IS42.     

Ostracods from the marginal coastal Lower Cretaceous (Aptian) of the Central Tunisian Atlas (North Africa): Paleoenvironment,biostratigraphy and paleobiogeography

This work provides the first detailed taxonomic study of ostracod species from the Lower Cretaceous (Aptian) marginal coastal deposits of the Central Tunisian Atlas, the Kebar Formation, as well as biostratigraphic, paleoecological and paleobiogeographic implications. The ostracod fauna provides new insights into the depositional environment and biostratigraphic framework of the Kebar Formation, and is represented mainly by freshwater and brackish-water species among them, Perissocytheridea tunisiatlasica nov. sp., is newly described. The environmental setting of this formation comprises marginal-littoral conditions in its lower part, thus not exclusively non-marine/continental as assumed previously. The ostracod associations from the studied locality, Jebel Ksaïra, indicate a minimum age of Early Aptian as recently attributed to the lower member of the Kebar Formation based on charophytes (presence of Late Barremian to Early Aptian Globator maillardii var. biutricularis Vicente and Martín-Closas, 2012), whereas an Early Albian age had been previously assigned to the Jebel Kebar site. The relative sea-level fall documented in the lower member of the Kebar formation at Jebel Ksaïra might correspond to the 3rd order cycle major eustatic sea-level fall event starting at the base of the Aptian. Paleobiogeographically, the non-marine ostracod fauna newly discovered in the Kebar Formation shows some affinities to contemporaneous faunas of southern and western Europe, e.g. that of the uppermost Weald Clay Group of southern England (Cypridea fasciata subzone of Horne, 1995), West Africa, and possibly to eastern South America (Brazil). Hence, the studied ostracods further support the hypothesis of supraregional faunal exchange by passive ostracod dispersal during the Early Cretaceous – between Europe and Northern Africa on the one hand, and eastern South America/western Africa and North Africa on the other hand. This leads to the hypothesis that the Peri-Tethyan islands could have worked as effective bridges for non-marine ostracods to become widely dispersed passively by “island-hopping” of larger animals and thus, ultimately, facilitated intercontinental faunal exchanges between South America and Europe – potentially even Asia – via North Africa during the Early Cretaceous.     

Geopalaeontological setting,chronology and palaeoenvironmental evolution of the Baccinello-Cinigiano Basin continental successions (Late Miocene,Italy)

The Latest Miocene succession of the Baccinello-Cinigiano Basin in southern Tuscany (Italy) recorded a faunal turnover documenting the extinction of an older, insular, endemic faunal complex characterised by the extinct ape Oreopithecus bambolii and the setting of a new, continental, European faunal complex including the colobine monkey Mesopithecus. A similar turnover pattern (Late Miocene ape/Latest Miocene Cercopithecidae) is generally observed in Late Miocene continental successions of Eurasia, from Spain to central Europe, Southwest Europe, the near East, and Southwest Asia. Abundant literature reports that the Late Miocene Eurasian hominoid primate distribution closely tracks the climatic/environmental changes occurring during the 12–9 Ma interval, until their extinction in western Europe. In the primate record, the dispersion of Cercopithecidae and the contraction of hominids is interpreted as an event depicting a pattern of “continentalisation” in the Old World. The sedimentary succession of the Baccinello-Cinigiano basin, one of the longest continuous vertebrate-bearing continental successions in the Neogene Italian record, contributes to the debate on this hypothesis. This paper provides an overview of the main characteristics of the sedimentary succession, the chronological constraints (biochronology, radiometric datings, magnetostratigraphy), and the palaeoenvironmental evolution as derived from palaeobiological approaches and from the study of stable carbon and oxygen isotope contents along the entire sedimentary succession. The 2 myr geological history of the Baccinello Cinigiano Basin, which documents the evolutionary history of Oreopithecus and associated faunas, does not have a direct relation with the event of the Messinian Salinity Crisis. The evolutionary history of Baccinello-Cinigiano Basin and its palaeontological record have been mainly driven by the regional tectonism and palaeogeographic changes that affected the northern Tyrrhenian regions in Late Miocene (Latest Tortonian–Messinian) times.     

Palaeoloxodon and Human Interaction: Depositional Setting,Chronology and Archaeology at the Middle Pleistocene Ficoncella Site (Tarquinia,Italy)

The Ficoncella site in northern Latium (Italy) represents a unique opportunity to investigate the modalities of a short occupation in an alluvial setting during the Lower Palaeolithic. The small excavation area yielded a lithic assemblage, a carcass of Palaeoloxodon antiquus, and some other faunal remains. The main objectives of the study are to better characterize the depositional context where the Palaeoloxodon and the lithic assemblage occur, and to evaluate with greater precision the occupation dynamics. A 25 m-long well was drilled just above the top of the terrace of the Ficoncella site and faunal and lithic remains were analyzed with current and innovative techniques. The archaeological site contains floodplain deposits as it is located next to a small incised valley that feeds into a larger valley of the Mignone River. A tephra layer capping the site is ⁴⁰Ar/³⁹Ar dated to 441± 8 ka. Collectively, the geochronologic, tephrochronologic and geologic data, suggest the site was occupied during MIS 13. The new results should prompt further research at Ficoncella in order to improve our understanding of the dynamics of human settlement in Europe during the Early to Middle Pleistocene.     

New data on Late Pleistocene small vertebrates from northern France

Quaternary small vertebrate assemblages from northern France, and more generally from northwestern Europe, are poorly recognized, and studied less than those from more southern, Mediterranean areas. However, important sites of human occupation are often found in these northern areas, with significant issues related to the occupation of these sites by Neanderthals and previous humans, as well as faunal dynamics under climatic pressure. In this paper, we present preliminary results from the study of small vertebrate assemblages from two Late Pleistocene sites of northern France: Mutzig (Alsace) and Le Rozel (Normandy). Both are ancient rock-shelters that have been excavated recently and have yielded evidence of Neanderthal occupation, but in very different contexts: Le Rozel is located in a coastal area of northwestern France contemporaneous with a temperate period, while Mutzig is located at the foot of the Vosges mountains in northeastern France and was occupied during a cold period. Consequently, even if these two sites are chronologically close to each other, differences in faunal composition are observed and discussed in relation to the geographic, climatic and biochronological context, in comparing the results from small vertebrates with those from other disciplines (numerical dating, large mammals).     

On the paleopathologic findings exhibited by the late Homo erectus of Ceprano, Italy

In March 1994, during excavation work for the construction of a motorway, a fragmented calvaria was discovered in the “Campo Grande” area near Ceprano, a town in southern Latium (Central Italy) about 55 miles from Rome. After reconstruction, the remain was recognized as belonging to aHomo erectus; it has been estimated that it goes back to the lowest middle Pleistocene. The calvaria exhibits two pathologic findings. The first is a congenital malformation of the sphenoidal sinus consisting in a deep, wide recess penetrating into the left greater wing as far as the sphenotemporal suture. The lesion was examined with the help of a cast whose features were compared with those described in some early and recent cases reported in the literature. The second finding is a healed, depressed fracture of the right brow ridge. The cause and mechanism of this lesion are discussed, and a mechanical approach has been used to provide information pertinent to the specific reason for the injury. The possibility that the hominid was butted by a large animal appears to be the most likely cause of the fracture.     

The TD6.3 faunal assemblage of the Gran Dolina site (Atapuerca,Spain): a late Early Pleistocene hyena den

Ethological studies have shown that besides human groups, large-medium carnivores have bone-collecting habits. The research developed since the last half of the twentieth century has attempted to characterise the carnivore’s accumulations and to identify them in the archaeo-paleontological record. At present, we have diagnostic criteria that define the accumulations produced by hyenids (mainly, Crocuta crocuta), thereby allowing us to differentiate them from the other accumulating agents. The faunal assemblage recovered at the Early Pleistocene TD6.3 layer of the Gran Dolina site is characterised by the presence of typical elements described in hyena dens: presence of small carnivores remains, high bone breakage, low epiphysis survival and a high frequency (>30%) of specimens with carnivore induced modifications, including large amounts of digested bones. However, attritional mortality profiles, hyena’s cubs remains, mid-shaft bone cylinders or differential anatomical composition among different weight-sizes, have not been observed or are ambiguous. In addition, anthropic evidences are scarce and concentrated in the uppermost section of the layer. TD6.3 is the result of an accumulation produced by hyenas using the cave as a den, in alternation with sporadic occupations by hominin groups. TD6.3 shows that hyena fossil accumulations may present great anatomical and taxonomic variability.     

Role of man in the history of vegetation of the Ellero valley (maritime Alps, Italy)

The pollen analysis of a sediment core from a peat bog (Rifugio Mondovi) at the mountain belt (1760 m) in the Ellero Valley (Italian Maritime Alps) shows the postglacial vegetation history. The sequence starts at 12,000 BP during a peak of pine pollen; this first phase shows a low representation of birch and the presence of Tilia. Younger Dryas is characterised by increasing percentages of Artemisia, showing the presence of deciduous Quercus, fir and beech. Elm appears at the beginning of the Holocene during the second pine peak (9800 BP). A 3000-year hiatus is present. Sedimentation resumes at 6000 BP in a Rhododendron fir-wood. The present timberline at 1500 m, at the limit of the beech wood, is a result of the decline of the fir-wood at 2600 BP, which allowed an expansion of beech. During this period, there was a continual increase in Gramineae and deciduous oak and the first occurrences of evergreen oak are observed. The development of larch occurs at 1800 BP, together with walnut, chestnut, cereals and vine.     

The Miocene mammal record of the Vallès-Penedès Basin (Catalonia)

The land mammal record of the Vallès-Penedès Basin (Catalonia, NE Spain) ranges from the early Miocene (Ramblian) to the late Miocene (Turolian), that is from about 20 to 7 Ma. Here we present an updated review of the mammal succession focusing on biochronology as well as on environmental and faunal changes. Based on faunal similarities with central Europe, we interpret this basin as a transitional zone between the forested environments of northern regions and the more arid landscapes of the inner Iberian Peninsula. The quality of the Vallès-Penedès record and its chronostratigraphic control is clearly better for the late Aragonian and the Vallesian (between 12.6–9.0 Ma), especially for small mammals. Therefore, we analyze small mammal diversity dynamics during this interval. Contrary to previous analyses, which found an abrupt extinction event coinciding with the early/late Vallesian boundary (the Vallesian Crisis), our results show that this pattern is due to uneven sampling. Instead, taxonomic richness slowly decreased since the late Vallesian as a result of a series of extinctions that mostly affected forest-dwelling taxa.     

Small mammal diversity of Mt. Kenya based oncarnivore fecal and surface bone remains

Ecological dynamics and faunal diversity documentation is normally conducted by direct observation and trapping of live animals. However,surveys of carnivore scat prey and surface bone remains, which are relatively inexpensive, can provide complementary data that expand carnivore diet breadth and may improve accuracy regarding inferences of the ecological dynamics of a given ecosystem. We used this inexpensive method to document species diversity variation with elevation on the leeward(Sirimon) and windward(Chogoria)areas of Mt. Kenya. Bone and fecal specimens were opportunistically collected by walking 2 km in opposite directions from transect points selected at 200-m intervals along the elevational gradient of the study areas. We collected a total of 220 carnivore fecal and owl pellet specimens from both study sites, which were mainly deposited by the spotted hyena(Crocuta crocuta), leopard(Panthera pardus),serval(Leptailurus serval), genet(Genetta sp.), and Mackinder's Cape owl(Bubo capensis mackinderi).Serval scats were the most common, followed by those of the spotted hyena. Scats and bones were found at the lowest density at the lowest elevations,peaked at mid-higher elevations, and then declined at the highest elevations. Based on skeletal analysis only, there were more species in Sirimon(19) than in Chogoria(12). Small fauna(rodents to duiker size bovids) formed the bulk of the identified remains,representing 87.9% of the Sirimon fauna and 90.9% of the Chogoria fauna. The genus Otomys was the dominant prey of the owl and serval in both sites. Three giraffe teeth were found at 3 500 m a.s.l. in Chogoria on the edge of Lake El is, suggesting that it is an occasional visitor to such high elevations. This study underscores the value of fecal and bone surveys in understanding the diet and diversity of mammals in ecological ecosystems,but such surveys should be complemented with analysis of hairs found in scats to obtain a more complete list of carnivore prey at Mt. Kenya.     

Pollen from archaeological layers and cultural landscape reconstruction: Case studies from the Bradano valley (Basilicata,southern Italy)

Abstract

The article aims at presenting some aspects of environmental reconstruction through pollen analysis from archaeological contexts. The anthropogenic pollen transport into archaeological sites is regarded as an interesting tool to improve knowledge on flora and vegetation in the area of influence of sites. The zoophilous plants can be found more easily than in the regional airborne pollen rain where anemophilous pollen is generally overrepresented. Moreover, pollen from archaeological contexts is mainly a result of the cultural landscape shaped by human activities. Two case studies from the Bradano Valley (Basilicata, southern Italy), rich in archaeological sites dating altogether from the Middle Bronze Age to the Medieval age, are reported. Difesa San Biagio and its surroundings is one of the biggest settlements of the area, settled in early times by Enotrians. Altojanni is an extended area mainly frequented in Hellenistic, Roman late Imperial and Medieval times. A very open landscape, and clear signs of plant exploitation and cultivation, breeding and settlements were present in the two sites. Though samples are disturbed and preservation problems are sometimes observed, the main characters of pollen spectra are recurrent. High percentages of Poaceae and Cichorioideae, together with coprophilous fungal spores, strongly suggest a long tradition of pastoral activities. These case study examples suggest that human activities would have produced a fairly xeric environment.     

Palaeoenvironment and palaeoclimate in the western Liguria region (northwestern Italy) during the Last Glacial. The small mammal sequence of Riparo Mochi (Balzi Rossi,Ventimiglia)

Riparo Mochi is considered a key site for the Middle–Upper Palaeolithic chrono-cultural sequence in the Mediterranean area. Climatic and environmental conditions have been reconstructed after the study of the small mammal assemblage by means of linear regression method and the Habitat Weighting method. The climate proxies for the late Mousterian and Proto-Aurignacian (cultural units I to G) indicate an environment with a relative high percentage of forest component during two cold and one relative warm oscillations. Open environments with low temperatures are registered for the Gravettian (cultural Unit D) allowing a possible relation to the Heinrich 3 Event. The final Gravettian (cultural Unit C) shows a relative warm peak, tentatively related to Greenland Interstadial 2. The late Epigravettian (cultural Unit A) is characterized by a high percentage of forest-related environment suggesting that this horizon formed during the Bølling–Allerød Interstadial. Finally, the Riparo Mochi sequence contributes to reconstruct the processes of faunal changes along the northern Tyrrhenian coast in a period of climatic instability, especially during the end of Marine Isotope Stage 3 when foraging groups adopted different mobility strategies between southern France and the Italian peninsula.