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The long sedimentary sequence of Notarchirico has yielded evidence of one of the earliest Acheulean manifestations in Europe and of recurrent hominin occupation, spanning from the end of the interglacial MIS 17 to the glacial MIS 16 (~695–610 ka). Here, we report the new discovery of a lion, Panthera spelaea, from the site, based on a metatarsal from layer A. This part of the sequence dates to ~660–612 ka (MIS 16, 40Ar/39Ar age). Therefore, Notarchirico's lion represents the earliest confirmed occurrence of the species in southwestern Europe, although older findings are known from adjacent areas. Lions and several other large mammal species dispersed into Europe during the Early–Middle Pleistocene Transition, which also witnessed the spread of the Acheulean. Ecological and behavioural adaptability was probably key, for hominins and other species, to cope with the intense and recurrent environmental fluctuations that occurred during this period. [ABSTRACT FROM AUTHOR]

Our work presents an updated overview of the Italian Middle Pleistocene records of hippopotamuses, including the two species Hippopotamus antiquus and Hippopotamus amphibius. In addition to reviewing several well-known fossils in the literature, a large number of samples are described herein for the first time. Following the recent results published in the literature, where the skull from the Middle Pleistocene of Cava Montanari (ca. MIS 13) was confirmed to belong to H. amphibius, one of the aims of this work was to investigate the H. antiquus–H. amphibius transition. A morphological analysis applied to a large sample confirmed the validity of the arrangement of the enamel ridges of the external surfaces of the lower canines as a diagnostic character for specific identifications. Finally, biometric analyses allowed us to test the size variability during the Middle Pleistocene, which confirmed that H. antiquus was generally larger than H. amphibius. Nevertheless, the remains of H. antiquus dated to ca. 600 ka show a reduced size when compared to older fossils of the same taxon, probably as a response to severe glacial conditions that occurred during MIS 16. [ABSTRACT FROM AUTHOR]

A skull of Hippopotamus recovered from the area of Tor di Quinto, within the urban area of Rome (central Italy) is here redescribed. Despite being one of the most complete specimens of hippopotamuses of the European Pleistocene, the Tor di Quinto skull did not attract much research interest, due to long-standing uncertainties on its provenance. This work begun in 2021, when the skull was restored, within a large renovation project on the vertebrate exposed at the Earth Science University Museum of Sapienza University of Rome. Original sediments were found inside the cranial and mandible cavities during the restoration work, which were sampled for petrographic analyses. By combining a review of the old paleontological, archeological and geological literature published during the 19th and 20th century on the Rome basin and the correlation of these new sedimentological and petrographic information with the lithostratigraphic and synthemic units of the national geological cartography, we clarify that the Hippopotamus skull was most likely to have been collected from a quarry called Cava Montanari, from a formation dated between 560 and 460 ka. Morphological and biometric analyses clearly support an attribution of the Cava Montanari specimen to the extant species Hippopotamus amphibius. The reassessment of the stratigraphic and geological data on Cava Montanari implies that the studied specimen is the earliest confirmed occurrence of Hippopotamus amphibius in the European fossil record. [ABSTRACT FROM AUTHOR]

Here, we describe a partial cranium of a large canid dated at 406.5 ± 2.4 ka from the Middle Pleistocene of Ponte Galeria (Rome, Italy). The sample represents one of the few Middle Pleistocene remains of a wolf-like canid falling within the timeframe when the Canis mosbachensis–Canis lupus transition occurred, a key moment to understand the spread of the extant wolf (Canis lupus) in Europe. CT-based methods allow studying the outer and inner cranial anatomy (brain and frontal sinuses) of a selected sample of fossil and extant canids. Morphological and biometric results allowed to: (I) ascribe the cranium from Ponte Galeria to an adult Canis lupus, representing the first reliable occurrence of this taxon in Europe; (II) provide the content for a biochronological revision of the Middle Pleistocene record of European wolves. [ABSTRACT FROM AUTHOR]

During the last century, Grotta Romanelli (Southern Italy) has been a reference site for the European Late Pleistocene stratigraphy, due to its geomorphological setting and archaeological and palaeontological content. The beginning of the sedimentation inside the cave was attributed to the Last Interglacial (MISs 5e) and the oldest unearthed evidence of human occupation, including remains of hearths, was therefore referred to the Middle Palaeolithic. Recent surveys and excavations produced new U/Th dates, palaeoenvironmental interpretation and a litho-, morpho- and chrono-stratigraphical reassessment, placing the oldest human frequentation of the cave between MIS 9 and MIS 7, therefore embracing Glacial and Interglacial cycles. These new data provide evidence that the sea reached the cave during the Middle Pleistocene and human occupation occurred long before MISs 5e and persisted beyond the Pleistocene- Holocene boundary. [ABSTRACT FROM AUTHOR]

The Ponte Galeria area within the city of Rome has yielded numerous fossiliferous localities that represent a reference point for the study of the European Middle Pleistocene ecosystems. Within Ponte Galeria a rich collection of fossil mammals has been unearthed from Cava di Breccia – Casal Selce 2 (MIS 15) thus the site represents an optimal laboratory to investigate the palaeoenvironments of a defined territory during the Middle Pleistocene. We investigate the feeding behaviours of the ungulate community of Cava di Breccia – Casal Selce 2 to reconstruct the MIS 15 habitats and also compare the data with those of the nearby site of Fontana Ranuccio (MIS 11) which shares similar faunal composition with Cava di Breccia – Casal Selce 2 to test if ungulates occupied the same niches during two different interglacials. Open habitats with scattered woodlands characterised the Ponte Galeria area during MIS 15, whereas woodlands were more widespread during MIS 11 at Fontana Ranuccio. Ungulates display similar diets in both localities, suggesting that cervids, large bovids and equids adopted the same niche partitioning strategies during both interglacials. [ABSTRACT FROM AUTHOR]

The site of Coste San Giacomo, in the Anagni Basin, is renowned for its mammal fauna of Early Pleistocene (Gelasian) age, including 32 taxa of small and large mammals. Adding to these, a small amount of remains of amphibians and reptiles was also recovered during recent excavations focused on collecting small vertebrates as well. The analysis of these remains resulted in the identification of a total of at least six different taxa: three anurans (Bufonidae indet., Pelophylax sp., and Rana sp.) and three reptiles (Lacertidae indet., Pseudopus sp., and Natrix sp.). This assemblage is comprised of taxa that are common in the Italian Quaternary, but also presents the first occurrence of the large anguid Pseudopus from the Apennine Peninsula in this period. On the whole, the herpetofauna indicates the presence of humid conditions in CSG at the moment of deposition of the bone bed that yielded the vertebrate remains. [ABSTRACT FROM AUTHOR]

The phyletic relationship between Canis lupus and the Early-Middle Pleistocene Canis mosbachensis is widely accepted among scholars, although the taxonomy of several European fossil specimens is still debated. In the last decades, many studies focused on the evolution of Pleistocene wolves have been proposed considering new materials as well as specimens belonging to historical collections. The canid remains recovered during the last century and housed in museums, undoubtedly represent a valuable source of biometric and morphological data which require, in some cases, an updated review of the fossil material. Indeed, cases of stratigraphic and taxonomic misinterpretations represent a potential loss of information for understanding the palaeobiology of these iconic predators. Here we provide the first updated description of the specimen belonging to the collection of the Italian geologist and palaeontologist Alessandro Portis dating back to the early last century. Previously reported as Canis lupus, the cranium from the historical excavation of Ostiense palaeontological site (Rome, Italy) is ascribed to the Early-Middle Pleistocene wolf Canis mosbachensis. Moreover, the reassessment of the stratigraphic and geological data of the Ostiense site led to consider the studied specimen as the last occurrence of the Mosbach wolf in the Italian fossil record. [ABSTRACT FROM AUTHOR]

The karst fissures known as "ventarole", located in the Salentine Peninsula (southernmost part of Apulia, Italy), were first studied by Mirigliano in 1941. These fissures are generally filled with reddish sediments or "terre rosse" in the lower part, and with brownish sediments or "terre brune" in the upper one. Both deposits are particularly rich in vertebrate remains. The mammal assemblages collected within the "terre rosse" from Melpignano and San Sidero are chronologically referred to the early Late Pleistocene (MIS 5), whereas those from the "terre brune" are referred to the Late Pleistocene - Early Holocene (MIS 2/1) transition. These ages were estimated from correlation with the similarly reddish and brownish sediments cropping out in Grotta Romanelli and with other Apulian local faunal assemblages. However, no detailed textural or mineralogical characterization has been carried out on the "ventarole" deposits. Moreover, the presence of several species led to hypothesize a persistence of Middle Pleistocene taxa during the Late Pleistocene in Apulia. In addition, the fauna of Melpignano was proposed as a Faunal Unit of the late Aurelian Land Mammal Age. In the last two decades, a team from the Department of Earth Sciences of Sapienza University of Rome have carried out fieldwork in this fossiliferous area. The rich fossil sample recovered is here presented, coupled with a reassessment of the remains collected since the 1900s. The updated faunal lists of Melpignano and San Sidero include several new species here identified for the first time, in particular Dama clactoniana, Equus mosbachensis and Lynx pardinus. New biochronological and paleoenvironmental data for southern Italy are presented, deriving from the analyses of the mammal remains from Melpignano and San Sidero and their comparison with those from other Middle-Late Pleistocene Apulian sites. Finally, textural and mineralogical analyses performed on several "ventarole" samples allow for a better description of the characteristics of the "terre rosse" and "terre brune". [ABSTRACT FROM AUTHOR]

Grotta Romanelli, located on the Adriatic coast of southern Apulia (Italy), is considered a key site for the Mediterranean Pleistocene for its archaeological and palaeontological contents. The site, discovered in 1874, was re-evaluated only in 1900, when P. E. Stasi realised that it contained the first evidence of the Palaeolithic in Italy. Starting in 1914, G.A. Blanc led a pioneering excavation campaign, for the first-time using scientific methods applied to systematic palaeontological and stratigraphical studies. Blanc proposed a stratigraphic framework for the cave. Different dating methods (C14 and U/Th) were used to temporally constrain the deposits. The extensive studies of the cave and its contents were mostly published in journals with limited distribution and access, until the end of the 1970s, when the site became forgotten. In 2015, with the permission of the authorities, a new excavation campaign began, led by a team from Sapienza University of Rome in collaboration with IGAG CNR and other research institutions. The research team had to deal with the consequences of more than 40 years of inactivity in the field and the combined effect of erosion and legal, as well as illegal, excavations. In this paper, we provide a database of all the information published during the first 70 years of excavations and highlight the outstanding problems and contradictions between the chronological and geomorphological evidence, the features of the faunal assemblages and the limestone artefacts. [ABSTRACT FROM AUTHOR]

Fossil mammal assemblages found in various localities of the Italian Peninsula provide significant information to create a detailed biochronological framework for the middle–late Villafranchian and Epivillafranchian of Europe and to reconstruct the evolution of early Pleistocene terrestrial ecosystems, when the earliest dispersal of Homo in Europe occurred. Here, we provide an updated critical overview on three Italian sites that in the last few years have provided the most interesting information on this crucial time interval: Coste San Giacomo (Latium; about 2.1 million years, Ma), Pantalla (Umbria; about 1.9–1.7 Ma), and Pirro Nord (Apulia; about 1.6–1.3 Ma). [ABSTRACT FROM AUTHOR]

Various species and ecomorphotypes of the genus Equus are recorded in several southern European Early Pleistocene local faunal assemblages (LFAs), though their taxonomy, phylogenetic relationships, and chronological distribution are still a matter of debate. This article aims to increase knowledge on the European pre-Olduvai stenonoid horses by describing and discussing the equid sample from the middle Villafranchian (Gelasian) Italian site of Coste San Giacomo (CSG; Anagni). Although horse remains from CSG are scanty, the morphological traits, dimensions, and proportions of teeth and some limb bones suggest some affinities with middle-sized European stenonoid horses, in particular with Equus senezensis, as supported by statistical analysis. This opens a new window on the possible phylogenetic relationships of the middle-sized, slender middle Villafranchian horses, although some questions about their actual taxonomic rank cannot be firmly answered based on available data. The complex relationship among environment, structure of the accompanying mammalian fauna, and the presence in the Early Pleistocene LFAs of only one equid, large or middle-sized, or even of more horse species with different size is briefly discussed. [ABSTRACT FROM AUTHOR]

European large mammal assemblages in the 1.2–0.9 Ma timespan included Villafranchian taxa together with newcomers, mostly from Asia, persisting in the Middle Pleistocene. A number of biochronological schemes have been suggested to define these “transitional” faunas. The term Epivillafranchian, originally proposed by Bourdier in 1961 and reconsidered as a biochron by Kahlke in the 1990s, is at present widely introduced in the literature. This contribution, after selecting the most representative European large mammal assemblages within this chronological interval, provides a new definition proposal for the Epivillafranchian as a biochron included within the Praemegaceros verticornis FO/ Bison menneri FO, and Crocuta crocuta FO. [ABSTRACT FROM AUTHOR]

Recent fieldwork in the Coste San Giacomo site (Early Pleistocene – Gelasian; central Italy) led us to discover new fossils of the Antilopini bovids Gazella borbonica and Gazellospira torticornis that are presented and discussed in this paper. These taxa have important palaeoenvironmental and biochronological significance during the Early Pleistocene. They were the last Antilopinae that inhabited the Italian peninsula, characterizing the middle Villafranchian large mammal assemblages living in arid and open environments. Gazella is one of the most diverse and widespread genus among Antilopinae subfamily, living today in Africa and in Asia. G . borbonica was the last species that inhabited Europe. In the Italian peninsula, it has been found in three sites: Montopoli, Dianella and Coste San Giacomo. The spiral horn-cored G . torticornis , larger in size than G . borbonica , has been found in the Italian sites of Montecarlo and Coste San Giacomo. Here, we present the study of the Italian record, based in particular on the horn cores. Finally, this material is compared with the remains coming from the coeval European localities. [ABSTRACT FROM AUTHOR]

Abstract: Canis lupus dispersed into Europe in the late Middle Pleistocene. The phylogenetic origin of C. lupus from Canis mosbachensis is widely accepted in the literature although their relationships and taxonomy are debated. In this paper, canid remains coming from the so-called “terre rosse”, Level G at Grotta Romanelli (Apulia, Southern Italy) are described. The Level G was dated between 69,000 and 40,000 ± 3250 years using the 230Th/238U method. Despite the great archeological importance of the site, some of the vertebrate fossil remains have been never described in detail. Among these are cranial remains of a canid that different authors referred to Canis aureus, C. lupus, C. mosbachensis, Canis aff. mosbachensis, or Canis sp.. The skull remains from Grotta Romanelli Level G were analyzed using Computed Tomography and 3D virtual modeling to provide a detailed reconstruction of the specimens and investigate inner structures. In addition, the Grotta Romanelli material was compared with (1) fossil wolves from other Middle-Late Pleistocene sites of Apulia (Melpignano/San Sidero, Ingarano and Grotta Paglicci) and France, (2) C. mosbachensis remains from various Early-Middle Pleistocene European sites, (3) a large sample of the extant Italian subspecies C. lupus italicus. The Late Pleistocene wolves from Apulia can be included in a single group, morphologically and morphometrically homogeneous. Although the wolf from Grotta Romanelli shares some similarities with the Early-Middle Pleistocene C. mosbachensis, its overall morphology and proportions fall into the wide variability of the extant C. lupus. Moreover, this determination is much more parsimonious than the taxonomic attribution of the “Apulian wolves” (including the Romanelli specimens) to C. mosbachensis, a typical late Early-Middle Pleistocene species, whose survival in Apulia during the Late Pleistocene is not adequately supported. [Copyright &y& Elsevier]

An outstanding sample of Canis etruscus has been found within the faunal assemblage from the early Pleistocene site of Pantalla (Italy), which is referred to the early late Villafranchian. Canis etruscus appeared in Europe about 2 Ma ago. It is regarded as an important taxon for biochronology, as its first occurrence (the 'wolf event') has been used to define one of the Villafranchian faunal turnovers. The discovery of four crania from Pantalla prompted a revision of C. etruscus, in order to better describe its cranial morphology. Since early studies, the distinction between C. etruscus and the coeval C. arnensis has been based mainly on mandibular traits. For this reason, our study is aimed at highlighting differences in craniodental characters between the two species. Canis arnensis has been conventionally considered a jackal-like dog, while C. etruscus is regarded as a wolf-like dog. Consequently, we decided to use jackals for comparison, in addition to C. lupus. Although the jackal group has been traditionally considered as quite homogenous (different species are partially sympatric and similar in both size and ecology), recent genetic studies demonstrate that jackals are not monophyletic. Considering the model offered by extant species, our goal is to delineate the degree of intra- and interspecific variability among the basal forms of the genus Canis. [ABSTRACT FROM AUTHOR]

Abstract: Hyaenas are relevant members of the Plio-Pleistocene carnivore guilds in Africa and Eurasia. The spotted hyena Crocuta crocuta is an important element of the dispersal events from Africa and Asia to Europe occurred at the Early–Middle Pleistocene transition (about 0.8Ma). The presence of Crocuta crocuta remains from the Ponte Galeria formation at Casal Selce (Roma, Italy) from sands of the Ponte Galeria Formation referable to the earliest Middle Pleistocene (MIS 18), close to the Brunhes/Matuyama boundary, is one of the earliest occurrence of the genus in Europe. [Copyright &y& Elsevier]

Abstract: The revision of the Early Pleistocene collection from the site of Monte Argentario (southern Tuscany, Italy), housed at the Istituto Italiano di Paleontologia Umana (Roma), revealed the occurrence of a mostly complete left horn core, attributable to a small-middle size bovid species corresponding to the genus of Asian origin Soergelia (Soergelia sp. cf. Soergelia minor). This is the first finding of the genus in the Italian Peninsula. The Monte Argentario fauna associated with Soergelia includes the African origin saber-toothed tiger Megantereon whitei and other Late Villafranchian species. [Copyright &y& Elsevier]

The pardel lynx Lynx pardinus is today restricted to small populations living in southern Iberian Peninsula. However, this endangered species was widely spread throughout Iberia until historical times and is currently the subject of intense conservation programs. Paleontological data suggest that its past geographical range was much wider, including also southern France and northern Italy. Here, we report exceptionally preserved fossil remains of L. pardinus from the Late Pleistocene (about 40′000 years) of Ingarano (Italy), which represent the largest sample of fossil lynx currently known in Europe. This new evidence allows (1) to revise the taxonomy of European fossil lynxes, (2) to extend far southeast the paleobiogeographical distribution of L. pardinus , and (3) to offer new insights on the evolutionary history (e.g., relationships with other extinct and extant lynx species) and paleobiology (e.g., intraspecific variation, body mass) of this iconic European felid. • We describe and analyze the Lynx sample from the Ingarano site (southern Italy). • It is the largest and best preserved European sample of Late Pleistocene lynxes. • The sample is referred to Lynx pardinus Temminck (1827). • The new data allow to revise the taxonomy of European fossil lynxes. • Compared to the extant form, the fossil L. pardinus was larger with a wider paleobiogeographical distribution. [ABSTRACT FROM AUTHOR]

Resource competition and niche partitioning among the exceptionally high number of sympatric ungulates of the Early Pleistocene site of Coste San Giacomo (Central Italy) is here examined through the study of their dietary proclivities and body size. The main aim of this study is to investigate the niche differentiation mechanisms that let the fossil ungulates coexist in the same region. We also provide information about the complementarity of two different methodologies that observe diet variation at a different time scales (inner and outer mesowear) in the study of dental wear patterns of fossil ungulates. Results from analyses of dental wear degree and body masses predictions show that a wide range of feeding behaviours were adopted by the taxonomical groups (i.e., cervids, bovids and equids) in order to avoid competition. Among larger ungulates diet ranges from strict browsing ( Eucladoceros sp., Gazellospira torticornis ), to mixed feeding ( Gallogoral meneghinii , Leptobos sp.) to pure grazing ( Equus stenonis ), whereas smaller taxa are more selective feeders ( Axis cf. lyra , Croizetoceros cf. ramosus ) with only one exception ( Gazella borbonica ). When taxa with the same feeding behaviour occurred in the same habitat, competition was minimised by differences in body size. [ABSTRACT FROM AUTHOR]

Cave hyenas (genus Crocuta) were among the most representative large carnivorans of Eurasian ecosystems during the Middle-Late Pleistocene. They are close relatives of the extant African spotted hyenas, the only extant member of the genus Crocuta. However, the taxonomy of these forms is still debated. Using palaeogenetic and palaeogenomic evidence, previous studies suggested a complex evolutionary history between spotted and Eurasian cave hyenas. Up to now, no molecular data have been retrieved from their Mediterranean counterparts, leaving a gap in the understanding of population dynamics of the genus Crocuta. To investigate the relationships of the Sicilian hyena to other cave and extant spotted hyenas we obtained palaeogenomic data from a Late Pleistocene coprolite chronologically dated between 32 and 21 Ka. Surprisingly, we found that the Sicilian genome forms a basal lineage of cave hyena. Moreover, results show that the Sicilian cave hyena is less admixed with African spotted hyena than the German and Russian specimens are, thus rejecting any large scale gene flow from Africa to Sicily. Our findings show that the genetic structure of European cave hyenas was more complex than previously documented, and offers new insights into the phylogeny of the genus Crocuta. • First palaeogenomic data from a Late Pleistocene Crocuta coprolite from Mediterranean area. • Sicilian specimen forms a basal lineage of Eurasian cave hyena. • Sicilian cave hyena is less admixed with African spotted hyena than mainland European specimens are. • Palaeogenomic data support the role of Mediterranean islands as source of diversity of Pleistocene mammals. [ABSTRACT FROM AUTHOR]

Quaternary glacial/interglacial alternations, influenced by orbital obliquity cycles with a 41-ka long periodicity, started in the northern hemisphere around 2.6 Ma ago. Such alternations affected the terrestrial ecosystems, especially those of the Mediterranean region, with changes in the floristic communities and the dispersal and radiation of a number of large mammal open dwellers. Analyses of tooth wear patterns of ungulates from the Early Pleistocene site of Coste San Giacomo allow for a more objective reconstruction on the paleoenvironments and the climate in the Italian Peninsula during this epoch. Our results show that this area was composed by a mosaic of biomes, in particular by steppe and woodlands/wetlands. Evidence of such heterogeneity is provided by the wide spectrum of feeding behaviours found among the numerous ungulate herbivores here recorded, with cervids ( Axis cf. lyra , Croizetoceros cf. ramosus and Eucladoceros sp.) exhibiting browser diets, most of the bovids ( Gazella borbonica and Leptobos sp. and Gallogoral meneghinii ) being intermediate feeders and the equid Equus stenonis showing a strict grazer behaviour. These results provide new insights for a timing of changing ecosystems in Southern Europe and reveal the environmental legacy of this global climatic shift, which is essential for understanding the early occupation of Homo in Europe. Thus, our data provide new evidence that such an environmental heterogeneity and a wide spectrum of available food resources could have been the main factors favouring the settlement of early species of Homo in this area. [ABSTRACT FROM AUTHOR]

Among fossil vertebrates, oral pathologies are of particular interest, because of their considerable effect on teeth and maxillary/dentary bones and, as a consequence, on mastication and feeding behaviour. This study focused on a pathological left hemimandible referred to the mustelid Meles meles unearthed from a Late Pleistocene karst filling deposit at San Sidero (Apulia, South Italy). This fossil shows unusual marked abnormalities related to a rare case of nonodontogenic chronic suppurative osteomyelitis. Clinical diagnosis of the disease and the timing of its development have been defined on the basis of a veterinary approach and X-ray analyses. Such a pathological condition can be explained as a consequence of a wound due to a porcupine quill. The analysis of the injury also provides information about the biomechanics of the bite and on the feeding behaviour. The study case confirms how palaeopathological analyses can be considered valuable tools to reconstruct the physiology of animals that lived in the past and to depict in detail the interactions among Late Pleistocene mammals, thus allowing a more accurate reconstruction of the ecology in fossil mammals. [ABSTRACT FROM AUTHOR]

Abstract: The site of Pantalla (central Italy) yielded a rich late Villafranchian (Early Pleistocene) faunal assemblage, which includes some well-preserved large mammal skulls. We describe here two nearly complete crania and a left hemimandible of Acinonyx pardinensis from this locality, representing the most complete cranial material of this species in Europe. These finds allowed us to define more clearly the craniodental morphology of A. pardinensis. Similarly to the forms from North Africa and China, the giant cheetah from Pantalla has a more generalized skull than the living Acinonyx jubatus, showing some primitive, pantherine-like features such as the less domed dorsal outline of the cranium, the more developed sagittal and nuchal crests and the less bowed zygomatic arches. High-resolution CT scans of the specimens were used to obtain the first 3D model of a cranium with articulated mandible of A. pardinensis. Starting from the insertion areas on this model we reconstructed the jaw muscles of the Pantalla felid, confirming its affinities with pantherine felines. In the light of the musculoskeletal skull anatomy and the average body mass (about 80 kg), it is likely that A. pardinensis could kill large prey through a hunting strategy more similar to pantherine cats than to the living cheetah. [Copyright &y& Elsevier]

Medium-sized deer are frequently found in Early Pleistocene fossiliferous deposits of Southern Europe, and hence they represent a reference group for the Villafranchian fauna. Their evolutionary history is still highly debated, as multiple classifications and systematic revisions have been proposed with specimens being ascribed to several genera such as Pseudodama , Metacervoceros , Euraxis , Praeelaphus , Axis , Dama , or Cervus. Whereas most studies focus on the taxonomy of the group, a few have addressed how these cervids evolved in relation to the major Early Pleistocene climatic events such as the onset of the Quaternary glaciations. A remarkably rich collection of Dama -like deer has been unearthed from the Early Pleistocene site of Pirro Nord (Apricena, south-eastern Italy). Here we analyse palaeoecological and palaeoneurological data of the Pirro Nord sample to investigate both habitat occupation and the evolutionary history of this group during the Early Pleistocene. To do so, we integrate dietary proxies (dental wear patterns and stable isotope signal) with morphological data from a virtual endocast of a well-preserved male specimen, a largely unexplored research path in cervid palaeobiology studies. Moreover, palaeoneurological data may also provide clues to solve the systematic issue of this group. Dental mesowear results point to a long-term mixed diet for the Pirro Nord deer, while a leaning towards a grazing behaviour is suggested by dental microwear patterns. The range of the stable isotope δ13C ratios suggests that it foraged on abrasive water-stressed C3 vegetation in warm woodland and semi-open habitats. Palaeoneurological data seems to confirm a closer affinity in the endocranial morphology of this taxon to the extant fallow deer Dama compared to other Early Pleistocene forms. Our research represents a novel approach to the study of Early Pleistocene fossil deer palaeoecology that can be extended also to other groups to investigate their evolutionary history in relation to climate changes. • Pirro Nord Dama -like deer display adaptations for grass-rich diet in open habitats. • Stable isotope δ13C ratios indicate a mixed diet based on water stressed C3 plants. • Dental microwear patterns are compatible with consumption of grit-infested plants. • In terms of brain morphology a close affinity to modern fallow deer is observed. • Brain " Dama -like" features were gradually developed in this group. [ABSTRACT FROM AUTHOR]

• One of the most complete Pleistocene cetacean skeletons ever found is reported. • Taphonomic data detail the story of the carcass of this whale before its final burial. • This fossil helps to reconstruct the Mediterranean Quaternary baleen whale fauna. • Diversity and disparity of this fauna is strictly linked to palaeoceanographic changes. An almost complete and partially articulated skeleton of an Early Pleistocene baleen whale is here described. The fossil, measuring 11 m in length, was discovered in the Calcarenite di Gravina Formation at Lama Lamasinata site (Bari, southern Italy) in 1968. The bifurcated first rib combined with other characters supports the identification of the fossil whale as a possible undescribed species of Balaenoptera (Mysticeti, Balaenopteridae), close to or nested in the B. borealis – B. edeni – B. ricei clade. However, the limited number of preserved diagnostic characters suggests a prudent assignment of the Bari whale to Balaenoptera sp. The associated molluscs suggest a mid-shelf setting deposition near to the boundary between infralittoral and circalittoral environments, probably 40–60 m deep. An associated Carcharodon carcharias tooth (the first case of a possible trophic interaction between white shark and cetaceans in the Pleistocene) and shark bite marks on a rib support the hypothesis that an early scavenger action prevented the rising of the whale carcass because of the removal of abdominal tissues and the consequent reduction of the decomposition gas accumulation. The occurrence of chemosymbiotic bivalves near the skeleton could testify the development of the sulphophilic stage during decay. Overall, the Bari whale skeleton and its associated fossil fauna represent the first well-documented case of Pleistocene whale fall community. The Bari specimen sheds new light on the diversity and disparity of the mysticete fauna in the Mediterranean Pleistocene also related to the geodynamic, palaeoclimatic and palaeoceanographic conditions that favoured upwelling events and nutrients supply into the southern Adriatic basin. [ABSTRACT FROM AUTHOR]

Abstract: The puma-like cat Puma pardoides (Owen, 1846) is reported from several Eurasian localities dated to the Late Pliocene–Early Pleistocene, but its Italian fossil record is really poor; in fact, the only possible occurrence comprises some fragmentary postcranial material from the Late Villafranchian locality of Pirro Nord (1.6–1.3Ma). In the present paper, we describe an isolated left upper carnassial of a medium-sized felid belonging to the collection of fossil mammals from Montopoli (Tuscany, Italy), the type locality of the Middle Villafranchian Faunal Unit (∼2.6 Ma). All the morphological and morphometric characters of the tooth are consistent with an attribution to Puma pardoides (Owen, 1846), which comes out as the second felid species from Montopoli together with Acinonyx pardinensis (Croizet et Jobert, 1828) and is, possibly with the specimens from Perrier-Étouaries, the earliest occurrence in western Europe. [Copyright &y& Elsevier]

The giant, short-faced hyena Pachycrocuta brevirostris was the largest Hyaenidae ever existed and the one that perfectly embodied the distinctive bone-cracking adaptations of this mammal family. Its dispersal into Europe is regarded as a biochronological marker of the Late Villafranchian at ∼2.0 Ma, and its potential ecological interactions with other carnivorans and early Homo populations diffusing Out of Africa have given rise to extensive discussions. Nevertheless, our comprehension of the extinction of P. brevirostris remains vague. Here, we first critically evaluate the European fossil record of the species and then we review the whole Epivillafranchian and Galerian Hyaenidae record, including P. brevirostris , Crocuta crocuta and " Hyaena " prisca. Biometric comparisons with other extinct and extant bone-cracking hyenas are carried out. In contrast to a common view, we recognize that there is neither evidence of a persistence of P. brevirostris beyond the Early-Middle Pleistocene boundary, nor of a coexistence between the giant hyena and C. crocuta. The replacement between the two species, which was also accompanied by the arrival of " H." prisca , occurred at ∼0.8 Ma and can serve as a marker of the Epivillafranchian–Galerian turnover, part of the faunal renewal that reflects the response of mammal communities to the Early–Middle Pleistocene Transition. Moreover, we clarified that Pliocrocuta perrieri and " H. " prisca were different species, and that the latter was relatively more widespread than often assumed, being recorded from localities spanning in age almost the whole Middle Pleistocene and even the early Late Pleistocene. • The Epivillafranchian and Galerian European record of hyenas is reviewed. • Pachycrocuta brevirostris did not persist beyond the Early-Middle Pleistocene boundary, nor it coexisted with Crocuta crocuta • The Pachycrocuta - Crocuta replacement occurred ∼0.8 Ma, accompanied by the arrival of " Hyaena " prisca. • Pliocrocuta perrieri and " Hyaena " prisca were different species. • " Hyaena " prisca was documented throughout the Middle to early Late Pleistocene. [ABSTRACT FROM AUTHOR]

Sus strozzii is the only species of Suinae occurring in the first part of Early Pleistocene in Europe and western Asia. It is characterized by a large body size and several morphological similarities with extinct and extant pigs from Island South East Asia (ISEA), such as the Javan Warty Pig S. verrucosus . In this study, a new mandible of S. strozzii from the Early Pleistocene of Pantalla (central Italy) is described and the diagnosis of the species is emended. Based on 52 craniomandibular and dental morphological characters, the first phylogenetic analysis of both living and fossil Eurasian and African Suinae is carried out to reconstruct their relationships. Our results are in agreement with the literature based on molecular and/or morphological analyses. The late Miocene Microstonyx is the first taxon to branch off the monophyletic Suinae clade. The separation between the African ( Kolpochoerus / Hylochoerus and Metridiochoerus / Phacochoerus ) and the Eurasian ( Sus ) clades occurred early, probably even in the late Miocene. The inclusion of Potamochoerus in the latter group is questionable, being probably due to the retaining of similar plesiomorphic characters in this African pig and in Sus . The Wild Boar S. scrofa and the Early–Middle Pleistocene S. lydekkeri are the species of Sus that display the most numerous plesiomorphic character states. They are followed by the monophyletic group of suines that show verrucosic lower canines, which includes the Pliocene S. arvernensis , S. strozzii , and the extinct and extant verrucosic pigs from ISEA. [ABSTRACT FROM AUTHOR]

Climatic oscillations at the Gelasian/Calabrian transition modified terrestrial palaeoenvironmental settings in the European region. A gradual drop in global temperatures beginning about 2.7 Ma led to drier conditions and to a reduction in, and subsequent disappearance of, sub-tropical vegetation in the central Mediterranean area by ca 1.2 Ma. Large ungulates are sensitive to vegetation changes and faced with harsher environmental settings may shift their feeding strategies to exploit available food resources in different ecosystems. In fossil assemblages such dietary adaptations are reflected by tooth morphology (a phylogenetic signal) and tooth wear degree (a direct signal of the species' diet). In this paper, we investigate how large herbivores responded to palaeoenvironmental changes that occurred at the passage between the Gelasian and Calabrian ages in the Italian Peninsula, analysing the dental wear patterns and hypsodonty of the early Pleistocene fossil ungulates assemblage of Olivola (Aulla, Central Italy). We found that while ungulate feeding behaviours during the Gelasian spanned from browsers to grazers, in the locality of Olivola this group of herbivores display a narrower range of diet types with many taxa adopting a mixed feeding behaviour. Cervids in particular, whose fossils are often associated only with wooded environments, as a response to the reduction of covered sub-tropical vegetation, shifted from a strict browsing diet in mostly closed habitats to a more abrasive one taking advantage of the spread of open landscapes. We also provide new data on the feeding behaviour of the rare fossil caprines, Procamptoceras and Gallogoral meneghinii , suggesting a grass-rich mixed diet for the former and a certain degree of dietary plasticity for the latter. Our research thus contributes to the better understanding of how ungulates adapted in the past to exploit different resource types during pivotal climatic changes and how environments changed in Central Italy on the onset of colder and more arid conditions. [ABSTRACT FROM AUTHOR]

The Middle Pleistocene site of Fontana Ranuccio (Anagni Basin, Central Italy) has yielded nearly 25,000 remains belonging to large and small vertebrates, including four isolated teeth of Homo sp., and a rich lithic assemblage containing bone tools. Here we provide new dental wear analyses of the ungulate community to improve the palaeoenvironmental reconstruction for the site. Analyses indicate a mostly browsing diet for the cervids and a grazing diet for the perissodactyls Stephanorhinus sp. and Equus cf. E. mosbachensis , suggesting a range of habitats that spanned from forests to open grasslands. A discrepancy between mesowear and microwear results is observed in the case of the bovid, Bos primigenius ; this can be tentatively explained as resulting from a temporal switch towards a sub-optimal diet, possibly reflecting the effects of marked seasonality. Findings shed new light on the structure of the 400 kyr ecosystems that existed during the early occupation of Homo in Europe. [ABSTRACT FROM AUTHOR]

The late Villafranchian is one of the pivotal timespans in the succession of Pleistocene European faunal assemblages, setting the bases for the major faunal renewal that characterized the continent during the Epivillafranchian. Bison is one of the most important and successful large mammals to spread in Europe at the latest stages of the Early Pleistocene. Here we describe the remains of a large bovid from the late Villafranchian site of Pietrafitta (ca. 1.5 Ma), previously attributed to Leptobos. Our analyses allow to refer this sample to the genus Bison. The primitive characters featured by this sample suggest the attribution to the subgenus Eobison , a long-debated taxon which includes all the earliest forms of Bison. At Pietrafitta, thus, one of the largest and most complete record of primitive Bison is recorded. The vague diagnosis and confused taxonomic history of Eobison called for a reappraisal of its status. We present a re-definition of the diagnostic characters and a review of the Eurasian record of Eobison with a focus on the late Villafranchian samples from the Mediterranean area. We recognize at least three valid species of Eobison remarking, however, the extreme morphological variability of this group. The comparative analysis of Eobison and its closest relatives (i.e., Leptobos and Bison s.s.), confirms that both the leptobovine and bisontine clades underwent to an increase of stoutness of the appendicular skeleton in response to the shrinking of forested habitats and the onset of colder, arid climate during the Pleistocene. • We describe the remains of a primitive form of Bison from Pietrafitta (Italy, 1.5 Ma). • We reappraise the status and fossil record of the debated subgenus B. (Eobison). • At least three valid species of Eobison are here recognized. • The diagnostic characters of Leptobos , B. (Eobison) and B. (Bison) are re-defined. • Body size and proportions of these taxa are correlated with environmental changes. [ABSTRACT FROM AUTHOR]

The putative competition between large predators and humans during the Pleistocene, especially in cave environments, has been a hotly debated topic during the last decades. Here, with the aid of digital paleoneurological techniques, we study the brain morphology and proportions of two fossil hyenas from the Eurasian Pleistocene, Crocuta spelaea and Crocuta ultima . Our results show that the brain morphology of these extinct species closely resembles that of the extant Crocuta crocuta and differs from those of the other extant genera ( Hyaena and Parahyaena ). Our analysis of brain proportions, however, shows a lesser development of the anterior brain in the extinct taxa compared with C . crocuta , thus more closely resembling the condition of both Hyaena and Parahyaena . The latter fact indicates that the greater anterior brain development of C . crocuta is a derived and recently-acquired trait, and suggests that extinct species of Crocuta displayed less-developed social abilities and/or a more restricted adaptability to new environments compared to the former. These behavioral differences between extinct and extant spotted hyenas might contribute to a better understanding of the taphonomical evidence obtained from Eurasian Pleistocene sites in relation to the ecological role played by humans and hyenas in the taphocenosis. [ABSTRACT FROM AUTHOR]

The Fontana Ranuccio hominin teeth (FR, Latium, Italy) are dated to the Middle Pleistocene. In previous studies these teeth were classified as two lower (left and right) second molars, one lower left central incisor and a badly worn incisor crown, the exact position of which could not be determined. In 2012 these remains were acquired by the Anthropological Service of S.B.A.L. (Italian Ministry of Culture) and for this reason re-analysed. In a thorough revision we have reassessed them both morphologically and dimensionally as two lower (left and right) first molars, one lower left lateral incisor and a possible upper left canine. The comparison with penecontemporaneous and diachronic samples shows that the Fontana Ranuccio teeth are morphologically similar to Atapuerca-Sima de los Huesos, Arago XIII and Neanderthal samples. [ABSTRACT FROM AUTHOR]

Volcanic sediments are often unsuitable for the fossilization of both hard and soft organic tissues, however, in some circumstances, they can provide unusual conditions for the preservation of remains. Here we report an exceptional case of soft tissue fossilization of a Late Pleistocene Eurasian griffon vulture (Gyps fulvus) in the pyroclastic sequence of the Alban Hills volcanic region (SE Rome, Italy). CT analyses have revealed an exceptional natural cast of the complete head and neck that preserve extraordinary detail including the fossilized everted tongue, beak, feather insertions and the first record of the nictitating membrane of the eye. This fossilization (superior in detail even to the victims of the AD 79 Plinian eruption of Vesuvius) reveals no evidence of burning and requires re-evaluation of the thermal constraints in operation for the preservation of organic materials within pyroclastic sediments. The analysis of the external morphological features has provided key information regarding the taphonomic processes in operation, the emplacement temperatures of distal pyroclastic flow deposits and the relationships between organic materials and low temperature phreatomagmatic flows. This sheds light not only on the extremely rare situation of fossilization in volcanic contexts but also provides a new perspective on taphonomic studies of highly detailed casts of fossil vertebrates. [ABSTRACT FROM AUTHOR]

Pollen analyses have been carried out on the infilling deposits of Grotta Romanelli (Apulia, Italy), a reference site for the Middle and Upper Palaeolithic of Italy. The analysis focused on Terre rosse , a fine unit till now ascribed to an interstadial phase following the Würm acme, and on the uppermost unit (Terre brune) , recently dated to the latest Late Pleistocene-Early Holocene. Despite the diffuse barrenness and low pollen concentration of many levels, pollen data from Grotta Romanelli gives insights into the palaeoenvironmental setting of the deposits and their chronological attribution. The presence of Olea in all levels of Terre rosse strongly suggests their attribution to the Last Interglacial (Eemian), during which this plant was diffused in the Mediterranean area. The Terre brune deposition occurred when the environment was open, with rare trees and shrubs and prevailing steppe elements. This association reflects the climatic conditions of the Lateglacial, with evidence of both the warm interstadial Bølling/Allerød and the cold stadial Younger Dryas. Mediterranean, mesophilous and riparian arboreal elements are present, especially in the Early Holocene levels. Comparison with modern pollen material allowed some fossil grains, found in high amounts and in clusters, to be tentatively ascribed to the species Crithmum maritimum (Apiaceae), Muscari comosum and to Asparagus maritimus / Ornithogalum (Asparagaceae). The significant occurrence of such entomophilous plants reveals differential transportation inside the cave and, since most of them are edible and/or have medicinal properties, suggests an intentional introduction and possible use during time, by both Neanderthals and modern humans. • Pollen was preserved in Late Pleistocene/Early Holocene deposits of Grotta Romanelli. • Olea pollen constraints the deposition of Terre rosse to the Eemian (MIsS 5e). • Steppe herbs and rare arboreal pollen in Terre brune mark the Lateglacial. • Clusters/high amounts of entomophilous pollen may reveal the use of plants by humans. [ABSTRACT FROM AUTHOR]

Abstract: The Pirro Nord palaeontological and archaeological locality has been the object of recent field research to investigate the fossil content of its Early Pleistocene vertebrate palaeocommunities, and to understand the genesis of the karst network and related deposits that delivered the lithic industry representing the earliest human occupation of Europe known so far, spanning 1.7–1.3Ma. The succession of the Pirro 10 site has been described in terms of facies and fossil vertebrate content of the 37 distinguished Sedimentary Units (SU). On the whole, the succession is made of chemical and clastic sediments that include debris fall and alluvial deposits. SU are distinguished according to lithology, colour, discontinuities, lamination, fossil concentration, and taphonomical parameters. Palaeomagnetic data and biochronological characteristics refer the Pirro 10 site to the latest Late Villafranchian Mammal Age of the Italian biochronological mammal scale. The fossil content is quite homogenous. Common taxa are Axis eurygonos, Hystrix refossa, Ursus etruscus, Canis mosbachensis together with small mammal remains mainly represented by Chiroptera and the arvicolid Microtus (Allophaiomys) ex gr. ruffoi. A significant change in the taxonomic record occurs in SU 22 and 108, where a clear increase of Equus altidens and Bison (Eobison) degiulii remains indicates shifting towards drier and more open environments and possibly deteriorating local climatic conditions. In general, for all phases, taphonomic evidence suggests that (1) bone accumulation results from carcass transport into the cave system by accident, (2) skeletons were decomposed and partially disarticulated near the cave entrance, (3) the cave environment caused corrosion, desquamation and pitting, and (4) bones were moved within the cave system by water flow, usually for short distances. Pirro 10, Pirro 13 and all karst structures of the Pirro Nord locality during Early Pleistocene were part of a complex interconnected karst system. Dissolution was more effective along the highly fractured core-zones of the Pliocene fault that bounded the “Apricena horst” to the south, where Pirro 10 gallery and Pirro 13 shaft are located. At that time, Pirro 10 was developed within the vadose zone whereas Pirro 13 represented a vertical structure connecting the surface to an underground karst floor. In general, the filling of Pirro 10 indicates fluctuations of the base level: water flow circulation in phreatic and epiphreatic conditions led to the deposition of clastic sediments, whereas flowstones reflect base level lowerings. [Copyright &y& Elsevier]

Abstract: Facies and ostracod analyses have been performed on a continental core drilled at Coste San Giacomo (Anagni, central Italy), where a middle Villafranchian faunal assemblage was first discovered in 1978. The core intercepted the palaeontological level and penetrated the underlying sedimentary succession, previously largely unknown, for a total sediment recovery of 40 m. In addition, new field surveys led to the discovery of Hippopotamus sp. remains at the faunal site, thus pre-dating the dispersal of this large ungulate into Europe to the middle Villafranchian. The new sedimentological and palaeoecological data allowed to define a palaeoenvironment mainly ascribable to an alluvial plain, characterized by a marsh evolving into a floodplain with overbank deposits and a sand-bed fluvial-channel. This scenario is supported by the reassessed Coste San Giacomo faunal assemblage, consistent with running and/or clear waters as well as prairies and grasslands, under mild climatic conditions. [Copyright &y& Elsevier]

Abstract: The western L’Aquila Basin recorded in Quaternary times two well distinguished phases of intramontane basin fill, concomitant with uplift of Central Apennines and characterized by diverse Large Mammal assemblages. The first phase was dominated by alluvial fan and lacustrine-palustrine sedimentation, respectively Gelasian and Calabrian in age, that took place within an internally-drained, fault controlled, hydrographic system. A late Villafranchian-earliest Galerian Large Mammal assemblage, approximately 1.3–1.1 Ma, pertains to this phase and includes: Mammuthus meridionalis, Stephanorhinus etruscus and the large cervid Arvernoceros aff. A. giulii. The second phase of filling, Middle Pleistocene to Holocene in age, is dominated by fluvial sedimentation, which is related to a through-going fluvial system developed after the emptying of the former lake. A middle Galerian Large Mammal assemblage, approximately 600ka, comes from the oldest deposits of this phase. Biochronological data help date steps of geological and morphological evolution in an intramontane setting, and permit to temporally constrain the major change in local hydrography and sedimentation at the Early-Middle Pleistocene transition. This change was likely forced by an increment of uplift rates, which is well recorded along the axial range and peripheral belts of the Apennines. [Copyright &y& Elsevier]

Abstract: The Pirro 13 karst fissure (Cava dell’Erba, Apricena, Foggia, Italy) shows a significant vertebrate assemblage (Pirro Nord Faunal Unit) associated with lithic artifacts attributable to mode 1. The Faunal Unit, corresponding to the latest Villafranchian, is dated on biochronological basis between 1.3 and 1.6 Ma and accordingly the artifacts represent among the earliest evidence of human presence in Europe. The vertebrate assemblage includes many species of amphibians, reptiles, birds and mammals, and attests the earliest occurrence of many taxa in Italy such as Bison degiulii, Capreolus sp., Equus altidens, and Meles meles. The lithic assemblage is characterized by a high degree of opportunism and by the exclusive presence of debitage reduction sequences. [Copyright &y& Elsevier]