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... A long trench of more than 30 m has been opened, exposing a series of levels, called F, G, H, I1, I2 and J, with faunal and lithic remains. The results of the numerous multidisciplinary studies have been published in numerous papers [16,20,21,[31][32][33][34][35]. ...
... Drawing from the paleontological material obtained from the new excavations, Mecozzi et al. [34] described a partial ulna of Macaca sp., providing the first evidence of this species at Notarchirico. ...
... This group is represented by two remains recovered from level G ( Fig 5). The first, B24GNC, is a proximal portion of a well-preserved right ulna of Macaca sp., described by [34]. During the 2022 fieldwork, an isolated tooth was found, identified as a right deciduous upper canine. ...
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Here we revise all the paleontological sample of Notarchirico, including historical collections and new findings collected during 2016–2023 excavations. Notarchirico is one of the most significant sites for the study of human evolution and terrestrial ecosystem dynamics during the Early-Middle Pleistocene Transition, preserving nearly 100.000 years of environmental and climatic changes constrained between 695 ± 6 ka and 614 ± 12 ka. The deposit yielded the oldest human fossil of the Italian Peninsula, and one of the oldest European evidence of Homo heidelbergensis, as well as one of the earliest evidence of bifacial tools in western Europe, commonly associated with the Acheulean techno-complex. Our paleontological results revealed the presence of three different mammal complexes, documenting faunal dynamics in response of climatic driven-changes recognized during the early Middle Pleistocene. The lower complex (levels I2-G) indicates the dominance of wooded spaces, sparse steppes, and the existence of water bodies (lakes or ponds), indicating a deterioration of the fully interglacial conditions recorded during the end of MIS 17; the middle complex (levels G-C) with a low number of mammal remains can be attributed to the glacial conditions of MIS 16; the upper complex (levels B-above α) indicates an improvement in climate, transitioning towards the full interglacial conditions of the of MIS 15. The faunal sample of Notarchirico, based on its firm chronological setting, offers important data for the Biochronological Scheme of European Land Mammals, including one of the oldest records of Palaeoloxodon antiquus and Cervus elaphus in Europe, Panthera spelaea in southwestern Europe, Dama cf. roberti in Italian Peninsula, and one of the latest occurrences of Bison schoetensacki in Europe.
... Macaques (genus Macaca) have been widely distributed throughout Europe from the Late Miocene to the Late Pleistocene (Mecozzi et al., 2021). Specifically, the first European record of this taxon is found in the Iberian Peninsula at the Almenara-Casablanca M site (MN13, 5.9e5.3 ...
... Despite the scarce anatomical evidence available, the specimen analyzed here from Barranc de la Boella is included in this chronological and geographical range. This record, together with the findings from southern France, Italy, and Greece (Mecozzi et al., 2021), could contribute to the consideration of M. sylvanus populations as representatives of the Mediterranean fauna during the Early Pleistocene (Elton and O'Regan, 2014). Nevertheless, the synchronous records from the TE9 level of the Sima del Elefante (Atapuerca; Rosas et al., 2006) demonstrate a wider Iberian distribution during the Early Pleistocene that would reach latitudes farther away from the meso-Mediterranean influence. ...
... In particular, European populations of this taxon are considered to be part of the so-called Pleistocene warm-adapted faunas, with range expansions northward during interglacial periods and contraction toward southern refugia during glacial periods (Fooden, 2007;Alba et al., 2011;Elton and O'Regan, 2014). The apparently continuous record of macaques on the Iberian Peninsula during the Pleistocene (Mecozzi et al., 2021), as well as that of other components of faunas associated with warm climates (e.g., hippopotamuses; Fidalgo et al., 2023), point to the potential of this area as a climatic refuge in the cold and arid phases that worsen from the Early-Middle Pleistocene transition onwards (Head and Gibbard, 2015;Maslin and Brierley, 2015;Strani et al., 2019;Berends et al., 2021). ...
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The BB specimen is an upper right canine belonging to a male of the primate M. sylvanus. The agreement among the qualitative characteristics of the tooth and the chronology of the site suggest its identification as M. sylvanus cf. florentina, although the lack of consensus on the validity of the proposed subspecies and the large biometric overlap do not allow certainty of this subspecific assignment. Macaca cf. sylvanus fossils are known across Iberia and throughout Europe in the Pliocene and Pleistocene, mainly in warmer intervals. Data suggest that the species is limited to areas that are more southern in colder phases, whereas in warm phases, this taxon could extend its distribution further north. Predation by carnivore species is likely, while interaction with early humans is less definite.
... Since the Quaternary, the area of Rome was shaped by the action of the Tiber and Aniene Rivers, combined with the volcanic activity of the Alban Hills and the Sabatini Mounts. This could be schematized as follows: the rivers hollowed out the fluvial valleys and coastal plain incisions during a glacial stage, filled with deposits, which often included pyroclastic products, during the following phases (e.g., [124][125][126]). The presence of volcanic materials (pumice, tephra, as primary deposition or reworked) in alluvial-deltaic sediments allows a reliable age to be obtained for many of these localities (obtained by radiometric dating or through correlations with previously dated deposits). ...
... The names of most of these deposits commonly correspond to those of the quarries (Cava means quarry in Italian), for example, Cava Nera Molinario or Cava Montanari. In the Tor di Quinto area, the fossil remains were collected by gravel and sand deposits quarried for the extraction of building materials [40,125]. The most important toponyms are Cava Montanari (described in Section 2), Cava Nera Molinario, Monte Antenne, Ponte Molle, and Tor di Quinto; all of these yielded hippopotamus remains. ...
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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.
... For 2016-2018, the majority of remains comes from levels F and I1, and amounts to 4081 remains (NRT: number of total remains), including 289 anatomically identified elements (NISPa: number of anatomically identified specimen) that allow an inference of 24 minimum number of individuals (MNI) (Tables 2 and 3). Preliminary mammal faunal list was reported by Moncel et al. (2020), and the only remain of macaque was studied by Mecozzi et al. (2021). The analysis has been enriched by the material of 2019-2021 where 138 remains are anatomically identified (total of 1068 remains) These fossils allow us to expand our knowledge on the mammal faunal assemblage of the lower levels of Notarchirico and to identify the presence of new taxa. ...
... This faunal list is similar to that of the youngest levels α and A of Notarchirico ; Table 2), even if mammal fossils recovered during ongoing excavations, unfortunately, are generally poorly preserved and scarcely diagnostic. Two species have been documented during the new excavations, not found in the upper levels (α and A): Hippopotamus antiquus in levels G and I1, and Macaca sp. in level G Mecozzi et al. 2021). ...
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New excavations in the lower part of the sequence dated between 670 and 695 ka by ⁴⁰Ar/³⁹Ar and ESR-U-Th at Notarchirico revealed layers with lithic and bone remains attesting several phases of human occupations. Some of these occupations are located at the top of residual pebble/cobble lags along former water channels, while others are more disturbed. All the layers yield faunal and lithic remains. Here, we aim to discuss the interpretative limits of traces of hominin occupations in such Early Palaeolithic sites through a multidisciplinary approach focusing on depositional and post-depositional processes in sedimentary units applied on the micro/macro-mammal remains, artefacts (surfaces, micro-wear traces), and spatial distribution of the archaeological material. These data are then compared with those from M. Piperno’s previous excavations in the upper part of the sequence (610–670 ka). As is often the case in open-air deposits and wetland environments, the majority of the bone surface modifications are related to natural abrasion caused by trampling and water flow. By contrast, the lithic material provides more relevant results both on taphonomic processes before and after the hominin occupations and function of the site. Despite the strong impact of post-depositional processes on archaeological material, evidence of hominin activities can nonetheless be inferred, shedding light on early hominin occupations of western Europe at the MIS 17/MIS 16 transition. The discussion takes penecontemporaneous open-air sites into consideration.
... European primates and carnivoran records are quite scarce, and the majority of the medium-and largesized taxa became extinct during the Middle Pleistocene, like the large felid Panthera gombaszoegensis (Sardella, 2007). The Barbary Macaque, Macaca sylvanus, is recorded in a few Italian Middle Pleistocene localities (Mecozzi et al., 2021c). Its presence offers important data for the reconstruction of the Mediterranean palaeoenvironments, and the findings associated with lithic artifacts require particular attention to the potential Homo-Macaca interactions during the Pleistocene (Strani et al., 2018;Mecozzi et al., 2021c). ...
... The Barbary Macaque, Macaca sylvanus, is recorded in a few Italian Middle Pleistocene localities (Mecozzi et al., 2021c). Its presence offers important data for the reconstruction of the Mediterranean palaeoenvironments, and the findings associated with lithic artifacts require particular attention to the potential Homo-Macaca interactions during the Pleistocene (Strani et al., 2018;Mecozzi et al., 2021c). In addition to iconic predators like Crocuta crocuta and Panthera sensu latu, other carnivorans characterized the Middle 8 Strani F. et al. ter, 2013;Bellucci et al., 2021;Stefanelli et al., 2021), which were replaced later by Praemegaceros solilhacus and Dama clactoniana (e.g., Sardella et al., 2006;Breda & Lister, 2013;Breda et al., 2015). ...
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The Middle Pleistocene was a crucial stage for the evolution of European mammals, a time when the majority of the modern taxa appeared in the continent for the first time. It is also in this interval that periodicity and intensity of glacial-interglacial cycles changed, an event that strongly impacted on Mediterranean marine and terrestrial ecosystems, and on vertebrate communities. This area can thus be considered an important laboratory to investigate how major climatic events influenced mammals' communities (among which also hominin populations) and the habitats they occupied. The state of art of the Middle Pleistocene palaeontological, archaeological and palaeoanthropological record of north Mediterranean region, and of the Italian Peninsula in particular, is here discussed.
... Furthermore, in the Italian Peninsula, the presence of M. sylvanus suggests temperate and wet climatic conditions (e.g. Mazza et al. 2005;Bona et al. 2016;Mecozzi et al. 2021). At Visogliano, Macaca sp. ...
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The Italian Peninsula is commonly interpreted as a Quaternary refuge, with a particularly important role for taxa requiring humid environments. This is testified by a relatively high diversity of amphibians on the peninsula nowadays, and also by the regional fossil record. Here we provide a complete description of the palaeoherpetofauna from Fada Nana Cave, in Verona province. It includes at least four amphibians (Salamandra salamandra, Bufo gr. bufo, Bufotes gr. viridis, Rana temporaria) and six reptiles (Lacertidae indet., Anguis gr. fragilis, Zamenis gr. longissimus, Natrix sp., Vipera gr. aspis, and an indeterminate small Colubroides) from Middle Pleistocene deposits. Fossils from Recent levels within the cave also reveal the presence of brown frogs and a green lizard, Lacerta gr. viridis. Rather interesting is the absence of the large anguid Pseudo-pus, found in other Early to Late Pleistocene-age sites in Veneto, and whose lack may be explained by the higher elevation of Fada Nana Cave. Based on the preliminary analysis of the small mammal assemblage, pre-Recent deposits in Fada Nana Cave are Middle Pleistocene in age and are tentatively assigned to the MIS 13 or 15 Interglacial, within the early Toringian Small Mammal Age. The Middle Pleistocene fauna recovered in the cave indicates a forested and moist environment, with water bodies and open meadows, for the nearby area.
... Fossil remains of Macaca sp. (Alba et al., 2014;Mecozzi et al., 2021) and the first appearance of the genus Sciurus (Colombero & Carnevale, 2016) in Europe are taxa worth to be mentioned. ...
Article
Geo-palaeontological sites are essential for scientific inquiry, education, conservation, cultural preservation, and even economic development. This study revisits 30 geo-palaeontological sites in the Piemonte region, originally assessed in 2004 thanks to the collaboration of the Department of Earth Sciences at the University of Turin, the Museo Paleontologico Territoriale dell’Astigiano and the Museo Regionale di Scienze Naturali of Turin. The updated site descriptions here provided incorporate new discoveries. The sites were re-examined using an innovative analytical framework recently developed by the palaeontological team of the University of Florence. This re-evaluation not only provides a deeper understanding of each site condition but also offers practical guidelines for the use of this method, making the results accessible to non-specialists involved in geosite management. The findings indicate that while few sites are at immediate risk of degradation, several warrant further conservation efforts. The study aims to inform and support the development of new regional policies for enhancing the preservation and promotion of both historical and newly identified geosites in the Piemonte region.
... On the other hand, the finding from Layer A of Notarchirico described herein is the earliest in southwestern Europe with a robust chronological constraint (Fig. 3), and only dates to~660-612 ka. At Notarchirico, some large mammal species, like hippopotamuses and macaques, are currently documented only in the lowest portion of the sequence, which correlates with the end of the interglacial MIS 17 (Moncel et al., 2020;Mecozzi et al., 2021), while hominins regularly occupied Notarchirico, probably taking advantage of the presence of raw materials, water, plants, meat and wood (Moncel et al., 2023). Large carnivorans have long been known only from an unstratified finding of a canid humerus (Piperno, 1999), but their presence is now documented throughout the sequence, including the lion remains described herein (Layer A), recently recovered canid material (Layer I), and indirectly evidenced by tooth marks on bones (Layers G, H and I). ...
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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.
... The archaeological record includes a human femoral fragment, and a rich faunal and lithic sample (e.g. Mallegni et al., 1991;Cassoli et al., 1999;Piperno, 1999;Santagata, 2016;Pereira et al., 2015;Moncel et al., 2019Moncel et al., , 2020Moncel et al., , 2023Santagata et al., 2020;Mecozzi et al., 2021;Carpentieri et al., 2023). Notarchirico represents one of the ancient sites of southern Europe with bifacial tools manufacturing, being a key site for gaining greater insight into the behaviors and evolution of biface-producing hominins during the Middle Pleistocene. ...
... Two new species were reported: Hippopotamus antiquus (layers G and I1) and Macaca sylvanus spp. (layer G; Mecozzi et al., 2021). Overall, the faunal assemblage of these layers matches the attribution to the Isernia faunal unit made for the upper part of the sequence. ...
Article
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The earliest evidence of bifaces in western Europe is dated to the initial phase of the Middle Pleistocene (la Noira, Notarchirico, Moulin Quignon, 700–670 ka), with the findings of Barranc de la Boella (1.0–0.9 Ma) considered to be an earlier local evolution. No transition assemblages are recorded during this time frame, and the “abrupt” appearance of bifaces during this time frame is associated with significant cognitive shifts in human technological behaviours (Acheulean techno-complex). The new investigations conducted at the site of Notarchirico unearthed 30 ka of repeated human occupation (695–670 ka, layers F-I2) during MIS 17, with evidence of bifacial tools in layer G (680 ka) and F along with other heavy-duty implements (LCTs, pebble tools, etc.). Massive production of debitage products realised on local raw materials collected in situ through simple and efficient core technologies characterises a large part of the lithic assemblage with a high ratio of diversified light-duty tools, including modified chert nodules. Despite core and flake assemblages being a recurrent trait of Lower Pleistocene contexts, the increase in retouched implements recorded at the onset of the Middle Pleistocene has been considered a significant technological shift. The technological analysis of the debitage products presented in this work highlights recurrent and systematic technological behaviours of the hominins of Notarchirico—who proved to efficiently overcome the raw materials dimensional constraints—even in the layers without bifaces. This may shed light on the meaning of cultural and behavioural innovation that the Acheulean techno-complex is thought to bring over Europe. It is plausible that given the substantial homogeneity of the lithic strategies within the sequence of Notarchirico, which only the “introduction” of the bifaces in the upper layers seems to interrupt, a supposed behavioural or cultural change in the site might have already occurred in the lowermost portion of the sequence. In this work, we evaluate the degree of change—if any—from a technological perspective by analysing the debitage reduction sequences.
... Of course, species that are genuinely rare in nature might be disproportionally affected by all the aforementioned factors. On the other hand, "rarity" in the fossil record does not necessarily imply a low demographic density, rather it can result from a taphonomic bias, which in turn can relate to several factors, for instance body size or environmental preferences [58,[61][62][63][64][65][66]. These biases can affect the representation of a taxon within a specific geographic area due to its biological and ecological characteristics, but also between areas that hosted different ecosystems A striking example is the paucity of the fossil record concerning the evolution of African great apes (chimpanzees and gorillas) in comparison to that of East African hominins [67]. ...
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The dispersal of primitive elephantines and monodactyl equids in Eurasia has long been regarded as representative of a substantial turnover in mammal faunas, denoting the spread of open environments linked to the onset of cold and dry conditions in the Northern Hemisphere. During the 1980s, this event was named the “Elephant-Equus event” and it was correlated with the Gauss-Matuyama reversal, today corresponding to the Pliocene-Pleistocene transition and the beginning of the Quaternary, dated at ~2.6 Ma. Therefore, the Elephant-Equus event became a concept of prominent biochronological and paleoecological significance, especially in western Europe. Yet, uncertainties surrounding the taxonomy and chronology of early “elephant” and “Equus”, as well as conceptual differences in adopting (or understanding) the Elephant-Equus event as an intercontinental dispersal event or as a stratigraphic datum, engendered ambiguity and debate. Here, we provide a succinct review of the Elephant-Equus event, considering separately the available evidence on the “elephant” and the “Equus”. Elephantines dispersed out of Africa during the Pliocene (Piacenzian). Their earliest calibrated occurrences from eastern Europe date at ~3.2 Ma and they are usually referred to Mammuthus rumanus, although the allocation of several samples to this species is tentative. Available dating constraints for other localities do not resolve whether the dispersal of Mammuthus was synchronous across Eurasia, but this possibility cannot be ruled out. The spread of Mammuthus was part of an intercontinental faunal exchange between Africa and Eurasia that occurred during the Piacenzian, but in this scenario, Mammuthus is quite unique in being the only genus of African origin dispersing to western Eurasia. The arrival of monodactyl equids from North America coincides with the Pliocene-Pleistocene transition, with several occurrences dated or calibrated at ~2.6 Ma and no compelling evidence prior to this age. In Europe, early monodactyl equids are often aligned to Equus livenzovensis, but the material from the type locality of this species is chronologically time-averaged and taxonomically heterogeneous, and western European samples are seldom abundant or informative. Regardless, this does not diminish the biochronological significance of the “Equus event”. Indeed, while the term “Elephant-Equus event” should no longer be used, as the appearance of elephantines in the European fossil record markedly precedes that of monodactyl equids, we endorse the use of the “Equus event” as a valid alternative to refer to the intercontinental dispersal event that characterizes the middle Villafranchian faunal turnover, epitomized by but not limited to monodactyl equids.
... perrieri e which provides indirect further support for rejecting a synonymy between the two forms e nor as C. crocuta e which probably reflects its different ecology. None of these considerations argue against the potential presence of "H." prisca in continental settings and/or at northern latitudes in the event of favorable conditions, as occurred for other faunal elements and hominin populations (Parfitt et al., 2005;Mecozzi et al., 2021d). ...
Article
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.
... Throughout its extant geographic range, the Barbary macaque is associated with a Mediterranean climate (dry summers and wet winters), and the species also require presence of arboreal cover, with preference for evergreen forest of cedar and oak (Fooden, 2007;Masseti and Bruner 2009). There seems to be a marked relation between climate and range fluctuations in the species, with most findings concentrated in the Mediterranean area, some records at northern latitudes favoured by warmer conditions, and a few occurrences during glacial phases (Elton and O'Regan 2014;Mecozzi et al. 2021 (Moncel et al. 2020). ...
Article
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.
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A nondestructive technological, petrographic and granulometric analysis of 75 lithic artifacts excavated by M. Piperno in the 1980s in the Early Paleolithic site of Notarchirico (Venosa, Southern Italy) was carried out to check their compatibility with the lag deposit of level B (MIS 15) and the hypothesis of lithological selection for knapping. A total of 289 clasts selected via a virtual grid superimposed to orthophotographs of three areas documenting the palaeosurface of level B (MIS15) were petrographically analyzed in situ to identify the petrofacies of the lag deposit from which the artifacts were collected. The results obtained proved the lithic artifacts are compatible with clasts of level B from a granulometric, petrographic and morphological point of view and that only some of the available lithotypes in level B were used to knap artifacts. Although limestones, cherts, marls and sandstones were available in level B, the latter are not present among lithic artifacts as lithotype. Moreover, the size and rock texture resulted to be stronger selection criteria than lithology. In fact, since the maximum length of stone tools are higher than those of level B’s clasts, this points to a supply area of lithic raw material larger than the investigated paleosurface.
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The urban area of Rome and its surroundings (the Roman Campaign, "Campagna Romana") yielded an exceptional amount of fossil remains of terrestrial large mammals, which are of prime relevance for biochronological correlations and for reconstructing the paleoenvironmental conditions in the Italian Peninsula of the Middle Pleistocene. Apart from the mere quantity, the scientific importance of the Roman fossil record rests on the geochronological constraints that allow to date or correlate several findings, as well as on the dense chronological cover of continental deposits, for the last ~0.8 Ma. On the other hand, the case of Rome is emblematic of the challenges posed by investigating historical fossil collections, for instance, the necessity of a thoughtful integration of historical documentation (e.g., geological maps, museum labels) and indirect geological information (e.g., borehole lithology, the correlation between obliterated fossil sites and extant outcrops), owing to the intense urbanization occurred especially since the 1800s. Fossil-rich deposits of Rome have supplied collections housed in the city's major museums, geosites, and universities, creating a valuable and unique paleontological heritage. Synthesis and perspective on these entwined aspects are provided herein, offering a geological and historical background alongside an overview of Middle Pleistocene mammal faunas of the area of Rome, with special emphasis on recent results that offer examples of-and how to deal with-different kinds of recoveries (from sporadic finds to systematic excavations), and ongoing work on the collection of the University Museum of Earth Sciences of Sapienza University of Rome (MUST). Reviewing the history of the MUST collection underlines the profound link between the history of the research on large mammal faunas of Rome and the history of the collection itself. The management of the paleontological heritage of Rome, consisting of thousands of remains spanning from isolated teeth to complete skeletons, is a crucial task for providing new data and support for research and dissemination, both of which are carried out at MUST accompanying traditional and yet fundamental efforts, such as cataloging and restoration, with the digital enhancement of the collection.
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In this article, we describe an almost complete macaque mandible from the Middle Pleistocene locality Marathousa 1 in the Megalopolis Basin of southern Greece. The mandible belonged to a male individual of advanced ontogenetic age and of estimated body mass~13 kg. Comparative metric analysis of its teeth permits its attribution to the Barbary macaque Macaca sylvanus, a species that was geographically widely distributed in Western Eurasia during the Plio-Pleistocene. The dental dimensions of the Marathousa 1 macaque fit better within the variation of the Early Pleistocene M. s. florentina and the Middle to Late Pleistocene M. s. pliocena rather than with the extant representative M. s. sylvanus. Moreover, principal component analysis reveals a better match with M. s. pliocena. However, because no clear-cut diagnostic criteria have been defined to differentiate these European fossil subspecies, we attribute the Marathousa 1 specimen to M. s. cf. pliocena, in agreement with the chronology of the locality. Previously known only from the Early Pleistocene of Greece by some isolated teeth, this is the first record of Macaca in the Middle Pleistocene of the country and one of very few in the eastern sector of the peri-Mediterranean region. We discuss the presence of macaques in the paleolake environment of Marathousa 1, as well as their predation risks from both carnivores and hominins present at the locality.
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Systematic excavation and multidisciplinary research undertaken over three decades have deepened our understanding of the early Palaeolithic archaeology at Cueva Negra del Estrecho del Río Quípar (Caravaca de la Cruz, Murcia, Spain). New results from biochronology and combined ESR and U-series dating corroborate previous magnetostratigraphy, placing the entire excavated sequence between the Jaramillo sub-chron and the Matuyama-Brunhes boundary (i.e. ca. 990-772 thousand years ago (ka)); palaeontological and palynological findings reflect temperate environmental conditions. A bifacially flaked limestone hand axe was excavated 1 m below the top of the Pleistocene sequence. The Equus cf. altidens tooth that provided the ESR estimate was excavated 1 m below the hand axe. Throughout its 5-m-deep sedimentary sequence, small nodules, fragments and struck flakes make up the bulk of the Palaeolithic assemblage. Stratigraphical analysis points to undisturbed continuous sedimentary deposition above a layer of ashy sediment, encountered 4.5 m below the top of the Pleistocene sequence, which contained thermally altered bone and heat-shattered chert cores and flakes. Cueva Negra is among the earliest European sites with firm evidence of combustion.
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Notarchirico (Southern Italy) has yielded the earliest evidence of Acheulean settlement in Italy and four older occupation levels have recently been unearthed, including one with bifaces, extending the roots of the Acheulean in Italy even further back in time. New 40Ar/39Ar on tephras and ESR dates on bleached quartz securely and accurately place these occupations between 695 and 670 ka (MIS 17), penecontemporaneous with the Moulin-Quignon and la Noira sites (France). These new data demonstrate a very rapid expansion of shared traditions over Western Europe during a period of highly variable climatic conditions, including interglacial and glacial episodes, between 670 and 650 (i.e., MIS17/MIS16 transition). The diversity of tools and activities observed in these three sites shows that Western Europe was populated by adaptable hominins during this time. These conclusions question the existence of refuge areas during intense glacial stages and raise questions concerning understudied migration pathways, such as the Sicilian route.
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In this article we briefly review primate interactions with predators throughout their evolutionary history. Like today, predators of past primates were taxonomically diverse, including crocodilians, aquatic mammals, hyaenids, raptors and other primates. There is strong evidence for felid predation of extinct primates, with most work undertaken on the African Plio-Pleistocene fossil record. Felid predation of Plio-Pleistocene primates from other areas, including Europe, is much less well understood, so we explored co-occurrence and potential interaction between carnivorans (with particular reference to felids) and Macaca sylvanus, which was widespread and present in Europe from the late Miocene to the late Pleistocene. Over its tenure in the fossil record, M. sylvanus co-occurred with a diverse array of carnivorans, including canids and hyaenids, but medium-sized felids probably posed the most significant predation risk. It is likely, however, that human predation was a major factor contributing to macaque extinction in Europe.
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The Barbary macaque, Macaca sylvanus (Linnaeus, 1758), is systematically reviewed, based on a study of 103 specimens and survey of relevant literature. This review includes analyses of external characters, cranial characters, DNA variation, and karyology. Information also is presented concerning natural history, reproduction, fossils, and taxonomic history. Current knowledge of geographic variation in M. sylvanus is insufficient to warrant recognition of subspecies. Although M. sylvanus is now restricted to northwestern Africa, fossil evidence suggests that this species is a relict of the ancestral macaque stock that originated in Africa ca. 7-6 million years ago (Ma) and dispersed to Eurasia ca. 6–5 Ma. Macaques formerly were widely distributed in Europe, but became extinct there ca. 0.100–0.020 Ma; macaques have survived in Asia and are now represented there by ca. 20 species. An annotated gazetteer lists 166 localities where M. sylvanus has been collected, observed, or reported.
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Functional features of the articular surfaces of the joints of the elbow region are described for a number of extant and fossil anthropoids. At the humeroradial and proximal radioulnar joints of extant hominoids a complex of features occurs that guarantees stability throughout extensive pronation-supination movements. By contrast, in extant non-hominoids the morphology of these joints is such as to permit more limited movement and to provide a close-packed position in full pronation. Ramapithecines and Oreopithecus resemble extant hominoids in these features, while Proconsul exhibits some features of both groups of extant anthropoids. The other Oligocene and Miocene anthropoids investigated all resemble extant non-hominoids. Major differences in the humeroulnar joint relate mostly to different means of stabilizing the joint. In extant hominoids, Oreopithecus, and ramapithecines the joint is truly trochleiform and is stabilized against forces acting in any direction. In other Oligo-Miocene anthropoids except Proconsul, and in extant non-hominoids, the main joint surface is more cylindrical, and is stabilized by an anterodistally projecting flange medially and a posteriorly projecting flange laterally. These features are best developed in cercopithecoids. It is suggested that in cebids the humeroulnar joint is translatory, but that it is non-translatory in all the extant and fossil catarrhines considered.
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Recent collections at the middle Miocene site of Maboko Island in Kenya have yielded a relatively large sample of isolated postcranial bones that can be assigned with some degree of confidence to the early cercopithecid. Victoriapithecus. This paper provides a detailed description of the postcranial morphology of Victoriapithecus, and offers some general conclusions concerning its inferred locomotor behavior and phylogenetic status. The range of metrical and morphological variation exhibited by the material does not exceed that seen in modern species of Old World monkeys. This findings is consistent with recent re-interpretations of the cranio-dental material that indicate that only a single species of Victoriapithecus may be represented at Maboko Island. Victoriapithecus appears to be a relatively small (with an estimated average body weight of 3·5–4·0 kg), agile and active quadrupedal monkey, adapted to moving effectively in both arboreal and terrestrial settings. In its overall morphology, and probably also in its inferred locomotor behavior. Victoriapithecus is most closely similar to modern-day semi-terrestrial Old World monkeys.
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The macaque material from the Early Pleistocene site of Quibas (Albanilla, Murcia, Spain), including dentognathic remains, isolated teeth and some postcranial bone fragments, is described. Both metrically and morphologically, this sample must be attributed to Macaca sylvanus (the Barbary macaque). This species is currently distributed through North Africa and Gibraltar, but was much more widely distributed during the Plio-Pleistocene, being represented by several European fossil subspecies. Metrical comparisons of dental size and proportions between extant M. s. sylvanus and fossil Macaca sylvanus florentina from the type locality and other Italian sites are undertaken, in order to classify the remains from Quibas at the subspecies level. The results show that the Quibas sample not only fits the range of variation of M. s. florentina from the type locality, but also differs from the extant Barbary macaque condition in several regards. This permits us to formally attribute the material from Quibas to M. s. florentina. The material described in this paper therefore significantly improves the knowledge of this fossil taxon, particularly regarding the upper dentition, and further confirms the taxonomic distinctiveness of this extinct taxon at the subspecies rank. Taken as a whole, M. s. florentina largely overlaps in dental dimensions with M. s. sylvanus, but differs from the latter by displaying (on average): (1) absolutely longer upper molars (especially M(1) and M(3)); (2) relatively wider upper molars (especially M(1) and M(2)); (3) longer M(3) as compared with the M(2); (4) absolutely longer M(1) and M(3); and (5) relatively narrower M(3).
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The thick continental record in the Fortuna Basin enables the to establishment of an accurate chronology for the main mammalian events in the Western Mediterranean between 7.6 and 4.5my, thus covering the whole Messinian interval. A first mammalian event at about 7.2my records a significant turnover in the muroid rodent associations. A second mammalian event is characterized by the dispersalofthemuridParaethomysandanumberoflargemammalswhichindicatesafirstAfrican-Iberianexchangeat about6.2my. Finally, a third mammalian event dated between 5.9 and 5.3my involved the dispersal from northern Africa of the gerbil rodents. This event is clearly associated with the Mediterranean sea level drop following the Messinian Salinity Crisis.
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Pleistocene macaque remains are uncommon in Great Britain. In addition to three specimens previously known from three sites, four teeth from Hoxne, Suffolk are recorded here. Closely resembling teeth of modern Macaca sylvanus (Barbary ape), they were recovered from a Middle Pleistocene stratum containing stone artifacts (including handaxes) and fauna including cold-climate lemmings. Pleistocene macaque remains are uncommon in Great Britain. In addition to three specimens previously known from three sites, four teeth from Hoxne, Suffolk are recorded here. Closely resembling teeth of modern Macaca sylvanus (Barbary ape), they were recovered from a Middle Pleistocene stratum containing stone artifacts (including handaxes) and fauna including cold-climate lemmings.