Macaca ulna from new excavations at the Notarchirico Acheulean site (Middle Pleistocene, Venosa, southern Italy)

Keywords: Barbary Macaque, anatomy, Cercopithecoidea, Mediterranean area

Introduction

The taxonomy of European macaques

The first records of macaque in Eurasia are from the latest Miocene fossiliferous sites of Almenara-Casablanca-M (Spain) [START_REF] Köhler | Macaca (Primates, Cercopithecidae) from the Late Miocene of Spain[END_REF] and Moncucco Torinese (Italy) [START_REF] Alba | First joint record of Mesopithecus and cf. Macaca in the Miocene of Europe[END_REF], yielding remains assigned to cf. Macaca sp. (Table SD1). These data suggest that macaques probably dispersed out of Africa between 5.9 and 5.3 Ma, when the sea level dropped during the Messinian Salinity Crisis [START_REF] Agustí | Evidence for African-Iberian exchanges during the Messinian in the Spanish mammalian record[END_REF][START_REF] Alba | First joint record of Mesopithecus and cf. Macaca in the Miocene of Europe[END_REF][START_REF] Alba | First record of Mesopithecus (Cercopithecidae, Colobinae) from the Miocene of the Iberian Peninsula[END_REF][START_REF] Colombero | Late Messinian mollusks and vertebrates from Moncucco Torinese, north-western Italy. Paleoecological and paleoclimatological implications[END_REF]. Although there is consensus to consider that the European fossil macaques belong to the Macaca sylvanus lineage, uncertainty persists regarding their specific taxonomic status. European fossil remains of macaques are usually grouped in chronologically successive subspecies of Macaca sylvanus: the Pliocene M. sylvanus prisca [START_REF] Gervais | Zoologie et Paléontologie Françaises[END_REF]; the late Pliocene to Early Pleistocene M. sylvanus florentina [START_REF] Cocchi | Su di due Scimmie fossili italiane[END_REF]; and the Middle to Late Pleistocene M. sylvanus pliocena Owen, 1846 [START_REF] Szalay | History of the Primates[END_REF][START_REF] Delson | Fossil macaques, phyletic relationships and a scenario of deployment[END_REF][START_REF] Alba | Dentognathic remains of Macaca (Primates, Cercopithecidae) from the late early Pleistocene of Terrassa (Catalonia, Spain)[END_REF][START_REF] Alba | Fossil remains of Macaca sylvanus florentina (Cocchi, 1872) (Primates, Cercopithecidae) from the Early Pleistocene of Quibas (Murcia, Spain)[END_REF][START_REF] Alba | First record of macaques from the early Pleistocene of Incarcal (NE Iberian Peninsula)[END_REF]Marigó et al., 2014) (Table SD1). However, most of the taxonomic attributions are based on the chronology of the findings, with only subtle differences recognized between M. sylvanus subspecies [START_REF] Szalay | History of the Primates[END_REF][START_REF] Delson | Fossil macaques, phyletic relationships and a scenario of deployment[END_REF][START_REF] Alba | New macaque remains from the Middle Pleistocene of Gruta da Aroeira (Almonda karst system, Portugal)[END_REF]. Indeed, fossil remains of M. s. florentina are considered larger than those of M. s. prisca, but close to extant specimens of M. s. sylvanus [START_REF] Rook | Fossil Macaca remains from RDB quarry (Villafranca d'Asti, Italy): new data and overview[END_REF]; whereas specimens of M. s. pliocena are nearly identical to those of M. s. florentina,, except for slightly wider cheek teeth [START_REF] Delson | Fossil macaques, phyletic relationships and a scenario of deployment[END_REF].

Given the lack of adequate diagnostic characters, several attributions do not go beyond the species level [START_REF] Montoya | La fauna del Pleistoceno Inferior de la Sierra de Quibas (Abanilla, Murcia)[END_REF][START_REF] Zapfe | Zähne van Macaca aus dem Unterpleistozän von Untermassfeld[END_REF].

A second macaque lineage in Europe is represented by the extinct Macaca majori, an endemic primate from the Plio-Pleistocene of Sardinia (Italy) [START_REF] Rook | Fossil Macaca remains from RDB quarry (Villafranca d'Asti, Italy): new data and overview[END_REF][START_REF] Alba | First joint record of Mesopithecus and cf. Macaca in the Miocene of Europe[END_REF] (Table SD1). The rich sample collected from Capo Figari, Monte Tuttavista and Is Oreris [START_REF] Rook | A comparative study of adult facial morphology and its ontogeny in the fossil macaque Macaca majori from Capo Figari, Sardinia, Italy[END_REF][START_REF] Rook | The Italian fossil primate record: an update and perspectives for future research[END_REF][START_REF] Alba | First joint record of Mesopithecus and cf. Macaca in the Miocene of Europe[END_REF][START_REF] Zoboli | New remains of Macaca majori Azzaroli, 1946 (Primates, Cercopithecidae) from Is Oreris (Fluminimaggiore, southwestern Sardinia)[END_REF]) allows a clear diagnosis for this taxon (e.g., a short anteroposterior palatal length, reduced body size) (see [START_REF] Zoboli | New remains of Macaca majori Azzaroli, 1946 (Primates, Cercopithecidae) from Is Oreris (Fluminimaggiore, southwestern Sardinia)[END_REF].

Fossil macaques from the Italian record

The majority of the Italian fossil primates was described from 1870s to the beginning of 1900s (Gervais, 1872;[START_REF] Cocchi | Su di due Scimmie fossili italiane[END_REF]Forsyth Major, 1872a,b, 1875[START_REF] Forsyth Major | Observations sur la Faune des Mammifères quaternaires de Corse et Sardaigne[END_REF][START_REF] Ottolenghi | Nota sopra una scimmia fossile italiana[END_REF][START_REF] Seguenza | I vertebrati fossili della provincia di Messina. Parte II: mammiferi e geologia del piano Pontico[END_REF][START_REF] Seguenza | Nuovi resti di mammiferi fossili di Gravitelli presso Messina[END_REF][START_REF] Merciai | Sopra alcuni resti di vertebrati miocenici delle ligniti di Ribolla[END_REF][START_REF] Portis | Primi avanzi di Quadrumani del suolo di Roma[END_REF]. Nevertheless, this mammalian group was almost forgotten by Italian paleontologists during the last century, and as recently as in the last decades only a few studies were published [START_REF] Rook | A comparative study of adult facial morphology and its ontogeny in the fossil macaque Macaca majori from Capo Figari, Sardinia, Italy[END_REF][START_REF] Rook | The Italian fossil primate record: an update and perspectives for future research[END_REF][START_REF] Zoboli | New remains of Macaca majori Azzaroli, 1946 (Primates, Cercopithecidae) from Is Oreris (Fluminimaggiore, southwestern Sardinia)[END_REF]Bona et al., 2017).

However, one of the earliest occurrences of macaque in Europe is from Moncucco Torinese (dated approximately 5.41-5.33 Ma), where Mesopithecus pentelicus pentelicus is also documented (Alba et al., 2013;[START_REF] Colombero | Late Messinian mollusks and vertebrates from Moncucco Torinese, north-western Italy. Paleoecological and paleoclimatological implications[END_REF]. Except for M. majori, the Italian Plio-Pleistocene record of macaque was poorly investigated [START_REF] Gentili | The Italian fossil record: Recent finds and their geological context[END_REF][START_REF] Rook | The Italian fossil primate record: an update and perspectives for future research[END_REF]. The Pliocene macaque from Villafranca d'Asti is tentatively assigned to M. s. prisca [START_REF] Rook | Fossil Macaca remains from RDB quarry (Villafranca d'Asti, Italy): new data and overview[END_REF], whereas the fossil remains from the Early Pleistocene are generally ascribed to M. s. florentina (Barberino del Mugello, Monte Peglia, Pietrafitta, Valdarno Superiore) [START_REF] Rook | Macaca sylvanus florentina (Cocchi, 1872) from the Pleistocene Mugello basin (Northern Apennines, Italy)[END_REF][START_REF] Gentili | The Italian fossil record: Recent finds and their geological context[END_REF] (Table SD1). Many fossils recorded from the Middle Pleistocene sites of the Italian Peninsula are commonly referred to Macaca sylvanus ssp., but seldom described in detail (Table SD1). The latest occurrence of Barbary macaque from Italian Peninsula is currently attested by the early Late Pleistocene of Grotta degli Orsi Volanti [START_REF] Mazza | An unexpected Late Pleistocene macaque remain from Grotta degli Orsi Volanti (Rapino, Chieti, central Italy)[END_REF] (Table SD1).

Here the fossil macaque from recent fieldworks at Notarchirico (Venosa basin, southern Italy), which represents the southernmost and easternmost occurrence of Macaca sylvanus in the Italian fossil record, is described for the first time (Fig. 1a-c).

Geological and paleontological context

The Notarchirico site, discovered in 1979, has yielded a 7 m-thick sequence of fluvial sediments within eleven archaeological levels (Fig. 1b-c). The old excavations found a rich fossil sample, including a hominin femur fragment found in the upper part of the sequence (level α, [START_REF] Belli | Découverte d'un fémur acheuléen à Notarchirico (Venosa, Basilicate)[END_REF]. The faunal assemblage mainly collected from the upper part of the sedimentary succession was attributed to the Isernia faunal unit [START_REF] Sala | Notarchirico un sito del Pleistocene medio iniziale nel bacino di Venosa. Ediz. Osanna per Soprint. speciale al Museo Nazionale Preistorico Etnografico[END_REF]. 40 Ar/ 39 Ar ages and ESR dates have revised the chronology of the whole sedimentary sequence and constrained all archaeological levels between ca. 610 and 675 ka from the upper part of the sequence (level A to level F) excavated by M. Piperno in the 1980's [START_REF] Pereira | The earliest securely dated hominin fossil in Italy and evidence of Acheulian occupation during glacial MIS 16 at Notarchirico (Venosa, Basilicata, Italy)[END_REF]. Since 2016, new excavation campaigns started in the long archeological sequence of Notarchirico [START_REF] Moncel | A biface production older than 600 ka ago at Notarchirico (Southern Italy) contribution to understanding early Acheulean cognition and skills in Europe[END_REF]. New fieldwork focusing on the unknown base of the sequence have permitted the team to recognize four additional units (G, H, I and J), all older than 668 ± 6 ka [START_REF] Pereira | The earliest securely dated hominin fossil in Italy and evidence of Acheulian occupation during glacial MIS 16 at Notarchirico (Venosa, Basilicata, Italy)[END_REF][START_REF] Moncel | A biface production older than 600 ka ago at Notarchirico (Southern Italy) contribution to understanding early Acheulean cognition and skills in Europe[END_REF][START_REF] Moncel | The origin of early Acheulean expansion in Europe 700 ka ago: new findings at Notarchirico (Italy)[END_REF] (Fig. 1b-c). The 40 Ar/ 39 Ar ages calculated for levels I and J (695.2 ± 6.2 ka and 690.3 ± 5.8 ka) are undistinguishable, suggesting a rapid deposition of the mainly fluvial complex sediment during MIS 17 towards the MIS 16 transition [START_REF] Moncel | The origin of early Acheulean expansion in Europe 700 ka ago: new findings at Notarchirico (Italy)[END_REF]. Among large mammals, the main faunal association includes Palaeoloxodon antiquus, Hippopotamus antiquus, Dama cf. clactoniana and Bison schoetensacki, among others. Cervids are dominant among the large mammals, whereas equids, rhinocerotids and carnivorans are missing. The mammal association indicates the presence of humid zones in a varied landscape and a relatively warm climate. Biochronologically, it also permits to attribute the levels A and α, as well as the newly excavated levels G to I, to the Galerian period [START_REF] Moncel | The origin of early Acheulean expansion in Europe 700 ka ago: new findings at Notarchirico (Italy)[END_REF], consistent with the recent chronometric dating [START_REF] Moncel | The origin of early Acheulean expansion in Europe 700 ka ago: new findings at Notarchirico (Italy)[END_REF] and lithic artefacts [START_REF] Moncel | The origin of early Acheulean expansion in Europe 700 ka ago: new findings at Notarchirico (Italy)[END_REF]. Levels G, H and I yield evidence of hominin occupations with artefacts including bifaces in level G, pushing back the age of the biface production (and the Acheulean) in Southern Europe [START_REF] Moncel | The origin of early Acheulean expansion in Europe 700 ka ago: new findings at Notarchirico (Italy)[END_REF].

Materials and methods

The macaque specimen from Notarchirico is a proximal fragment of a right ulna. The B24GNC specimen was recovered from the level G and is stored at "Museo Nazionale Archeologico di Venosa" (Potenza, Basilicata) (Fig. 1b-c). To build our comparative dataset, we used the available literature on extant and Early -Late Pleistocene macaques, which include the fossil remains from Villafranca d'Asti (RDB quarry) and Pietrafitta, and extant specimens of M. s. sylvanus (Table 1) [START_REF] Gentili | The Italian fossil record: Recent finds and their geological context[END_REF][START_REF] Rook | Fossil Macaca remains from RDB quarry (Villafranca d'Asti, Italy): new data and overview[END_REF]. We also collected morphological data and measurements of Middle As comparative material, we included fossil specimen of Mesopithecus pentelicus penticulus from Moncucco Torinese (Alba et al., 2013) and Pikermi [START_REF] Delson | Fossil colobine monkeys of the Circum-Mediterranean region and the evolutionary history of the Cercopithecidae (Primates, Mammalia)[END_REF], and Dolichopithecus ruscinensis from Montpellier (=Perpignan) and Pestzentlorinc [START_REF] Delson | Fossil colobine monkeys of the Circum-Mediterranean region and the evolutionary history of the Cercopithecidae (Primates, Mammalia)[END_REF]. We also used morphological and biometric data on extant specimens of M. s. sylvanus housed at Civic Zoology Museum of Rome and Royal Belgian Institute of Natural Sciences, Brussels.

Standard measurements of greatest breadth of olecranon (OB), greatest breadth across the anconeal process (APB), greatest breadth across the coronoid process (CPB), greatest height of olecranon fossa (HOF), greatest depth across the anconeal process (APD), were taken to the nearest 0.1mm (Table 1) (Fig. SD1). Boxplot of CPB and bivariate plot of HOF vs. CPB were realized to visually assess postcranial size and proportions.

Description and comparison

The B24GNC is a proximal portion of the right ulna in a good state of preservation, which consists of about one-fourth of the shaft (Fig. 1d-f). Medially, the olecranon is relatively high, with the anterior border higher than the posterior one; its rough surface delineates the insertion area of the triceps brachii, brachialis and flexor carpis ulnaris muscles. In anterior view, the olecranon is mediolaterally compressed and its medial and lateral crests are similarly developed, and they are separated by a shallow groove. Laterally, the inner spongy bone of the olecranon is viewable due to the weathering, which obscures the attach area of the flexor carpis ulnaris muscles, in the proximal portion, and the epitrochleo-anconeus muscle, along the distal border. In medial view, at the level of the trochlear notch, the shaft surface is rough, which indicates the attach area of the flexor carpi ulnaris and extensor pollicis longus muscles. The anconeal process ends in a marked crest anteroposteriorly directed in medial view; its anterior border is disto-laterally inclined in anterior view; its process is robust and anteriorly projected in lateral view. The trochlear notch is quite reduced and its opening is semicircular in shape in medial and lateral views, whereas it is medio-laterally enlarged in anterior view, with a straight medial and a curved lateral outlines. The coronoid process is robust and anteriorly projected in medial and lateral views. The radial notch is well developed and its posterior border reaches the half of the shaft in medial view. Anteriorly, the radial notch is strongly concave, the border is inclined medially, and the facet is butterfly-shaped (according to [START_REF] Harrison | New postcranial remains of Victoriapithecus from the middle Miocene of Kenya[END_REF].

Despite the fragmentary nature of the specimen, its classification can be inferred since the morphology of the proximal epiphysis of the ulna is highly diagnostic. The overall morphology of the ulna from Notarchirico resembles the fossil remains of M. s. cf. prisca from Villanfranca d'Asti (V.J. 130) [START_REF] Rook | Fossil Macaca remains from RDB quarry (Villafranca d'Asti, Italy): new data and overview[END_REF]), M. s. florentina from Pietrafitta (IR-1703) [START_REF] Gentili | The Italian fossil record: Recent finds and their geological context[END_REF], M. sylvanus ssp. from Casal Selce (CS 520) and Fontana Ranuccio (FR 85), and extant specimens of M. sylvanus [START_REF] Harrison | New postcranial remains of Victoriapithecus from the middle Miocene of Kenya[END_REF] in several features: relatively long and straight olecranon with the anterior border higher than the posterior one in lateral and medial views; wide surface of articulation with the radius in lateral view; anteriorly-protruding anconeal process and coronoid process in lateral and medial views. The studied specimen differs from Me. pentelicus in the shape of the olecranon (e.g., a straight proximal outline in lateral and medial views), and in the profile of the trochlear and radial notches (e.g., less anteriorly directed anconeal and coronoid process in lateral and medial views and less developed articular surface with the radius in lateral view) [START_REF] Delson | Fossil colobine monkeys of the Circum-Mediterranean region and the evolutionary history of the Cercopithecidae (Primates, Mammalia)[END_REF]Alba et al, 2013).

Compared to Dolichopithecus ruscinensis from Montpellier (=Perpignan) and Pestzentlorinc [START_REF] Delson | Fossil colobine monkeys of the Circum-Mediterranean region and the evolutionary history of the Cercopithecidae (Primates, Mammalia)[END_REF], the studied ulna differs in the anteriorly-protruding coronoid process in lateral and medial views, the anterior border of the olecranon higher than the posterior one in lateral and medial views and a larger radial notch in lateral view. The ulnae of the extant colobines differ from that of Notarchirico since those possess an anterior concavity of the proximal shaft and a retroflected olecranon in lateral and medial views (Rose 1998;[START_REF] Egi | Distal humerus and ulna of Parapresbytis (Colobinae) from the Pliocene of Russia and Mongolia: phylogenetic and ecological implications based on elbow morphology[END_REF]. The ulnar morphology of M. sylvanus is strongly interrelated with that of the distal epiphysis of the humerus (with which forms the "humeroulnar joint"), and it is linked to its quadrupedal locomotion strategy, with the closepacking position of the elbow providing high stability during parasagittal movements [START_REF] Rose | Another look at the anthropoid elbow[END_REF].

The size of the macaque ulna from Notarchirico (Table 1) is compared with those of other extinct and extant macaques and Me. penticulus in Figure 2. The values of the CPB of the fossil specimens falls in the variation of the extant Barbary Macaque, with the exception of the studied fossil, which displays the largest size among the analyzed sample (Fig. 2a). In the figure 2b, the fossil specimens of M. sylvanus shows a relatively size variation, with the ulna from Notarchirico that occupies the extreme right of the plot. Furthermore, the fossil of Me. pentelicus pentelicus from Moncucco Torinese shows a different proportions of the ulna, with similar CPB value, but an higher trochlear notch compared with those of the fossil M. sylvanus (Fig. 2b). However, given the variation and the restricted sample available for extinct M. sylvanus and Me. pentelicus pentelicus , no taxonomic implication can be drawn from the ulna size.

Discussion and Conclusion

Taxonomy

In the last decades, new studies focused on the description of fossil macaques, involving both new finds and revisions of historical collections. Most of the available material of this primate consists of mandibular and maxillary remains, which are very similar to the extant North African subspecies.

Following the proposal by [START_REF] Szalay | History of the Primates[END_REF] and [START_REF] Delson | Fossil macaques, phyletic relationships and a scenario of deployment[END_REF], fossil European macaques are often grouped in three temporal-geographic subspecies of M. sylvanus, but the reliability of this division is debated (e.g., [START_REF] Rook | Fossil Macaca remains from RDB quarry (Villafranca d'Asti, Italy): new data and overview[END_REF][START_REF] Alba | Dentognathic remains of Macaca (Primates, Cercopithecidae) from the late early Pleistocene of Terrassa (Catalonia, Spain)[END_REF][START_REF] Alba | Macaque remains from the early Pliocene of the Iberian Peninsula[END_REF][START_REF] Alba | New macaque remains from the Middle Pleistocene of Gruta da Aroeira (Almonda karst system, Portugal)[END_REF][START_REF] Bona | Macaca sylvanus Linnaeus 1758 from the Middle Pleistocene of Quecchia quarry[END_REF]. Indeed, due to lacking clear-cut taxonomic criteria, classifications are often either chronologically-based (e.g., [START_REF] Alba | First record of macaques from the early Pleistocene of Incarcal (NE Iberian Peninsula)[END_REF][START_REF] Alba | Macaque remains from the early Pliocene of the Iberian Peninsula[END_REF][START_REF] Alba | New macaque remains from the Middle Pleistocene of Gruta da Aroeira (Almonda karst system, Portugal)[END_REF], or limited to M. sylvanus ssp. (e.g., [START_REF] Montoya | La fauna del Pleistoceno Inferior de la Sierra de Quibas (Abanilla, Murcia)[END_REF][START_REF] Zapfe | Zähne van Macaca aus dem Unterpleistozän von Untermassfeld[END_REF].

Moreover, this unresolved taxonomy is mainly established solely on craniodental remains. Indeed, postcranial ones are limited to a few findings across Europe, such as those from Untermassfeld (Zapfe, 1997), Deutsch-Altenburg [START_REF] Fladerer | Macaca (Cercopithecidae, Primates) im Altpleistozän von Deutsch-Altenburg, Niederösterreich[END_REF] and Sierra de Quibas [START_REF] Alba | Fossil remains of Macaca sylvanus florentina (Cocchi, 1872) (Primates, Cercopithecidae) from the Early Pleistocene of Quibas (Murcia, Spain)[END_REF]. Besides the Italian specimens, other ulnae of macaques were recorded from Swanscombe and Senèze [START_REF] Singer | Pleistocene Macaca from Hoxne, Suffolk, England[END_REF], but the material was not described or figured. The morphological analysis conducted in this work reveals a strong affinity between the studied material and the fossil and extant specimens of M. sylvanus [START_REF] Harrison | New postcranial remains of Victoriapithecus from the middle Miocene of Kenya[END_REF][START_REF] Gentili | The Italian fossil record: Recent finds and their geological context[END_REF][START_REF] Rook | Fossil Macaca remains from RDB quarry (Villafranca d'Asti, Italy): new data and overview[END_REF], and also stresses the differences with Me. pentelicus pentelicus [START_REF] Delson | Fossil colobine monkeys of the Circum-Mediterranean region and the evolutionary history of the Cercopithecidae (Primates, Mammalia)[END_REF]Alba et al, 2013), Dolichopithecus ruscinensis [START_REF] Delson | Fossil colobine monkeys of the Circum-Mediterranean region and the evolutionary history of the Cercopithecidae (Primates, Mammalia)[END_REF] and extant colobines [START_REF] Rose | Another look at the anthropoid elbow[END_REF]. Biometrically, the specimens from the Middle Pleistocene sites of the Italian Peninsula fall within the range of variation of the extant Barbary macaque subspecies. The ulna from Notarchirico is slightly larger than the other analyzed specimens, but considering the restricted sample, these differences may not to be significative.

Nevertheless, the only ulna known of Me. pentelicus pentelicus possesses different proportions, especially in the relatively higher trochlear notch if compared with that of extant and extinct Barbary Macaque. Thus, considering the lack of diagnostic features for Macaque subspecies, the ulna from Notarchirico is ascribed to M. sylvanus, avoiding the attribution to subspecies level based on chronological grounds.

According to recent literature (e.g., [START_REF] Bona | Macaca sylvanus Linnaeus 1758 from the Middle Pleistocene of Quecchia quarry[END_REF][START_REF] Alba | Macaque remains from the early Pliocene of the Iberian Peninsula[END_REF][START_REF] Alba | New macaque remains from the Middle Pleistocene of Gruta da Aroeira (Almonda karst system, Portugal)[END_REF], a more comprehensive analyses should clarify the alpha-taxonomy of extinct M. sylvanus subspecies.

Paleoenvironment

The Barbary macaque from Notarchirico was recovered from level G, associated with fossil remains of Palaeoloxodon antiquus, Hippopotamus antiquus and Bison schoetensacki [START_REF] Moncel | The origin of early Acheulean expansion in Europe 700 ka ago: new findings at Notarchirico (Italy)[END_REF].

Although only a few species were identified, these taxa indicate the presence of open-forest and bodies of water, and warm climatic condition. As such, the level G of Notarchirico was accumulated during an interglacial. Considering the revised chronology of the site, this very likely indicates the end of MIS 17 [START_REF] Moncel | A biface production older than 600 ka ago at Notarchirico (Southern Italy) contribution to understanding early Acheulean cognition and skills in Europe[END_REF][START_REF] Moncel | The origin of early Acheulean expansion in Europe 700 ka ago: new findings at Notarchirico (Italy)[END_REF].

The extant Barbary macaque inhabits a variety of environments from shrublands to evergreen forests, and even herbaceous-dominated rocky ridges (Fooden, 2007, and references therein). The presence of arboreal vegetation is nonetheless a limiting factor, since the species requires trees for sleeping,, feeding, and escaping from predators [START_REF] Delson | Cercopithecidae from the Pliocene of Saint-Vallier[END_REF][START_REF] Fooden | Systematic review of the Barbary macaque, Macaca Sylvanus (Linnaeus, 1758)[END_REF]. In addition, it needs to be near water [START_REF] Masseti | The primates of the western Palaearctic: a biogeographical, historical, and archaeozoological review[END_REF]. Thereby, considering the ecological flexibile of the Barbary macaque, paleoecologically its presence at Notarchirico could suggests the presence of wooded and wet areas. SD1. Question mark denotes uncertain age. Map produced with QGIS.

Paleobiogeographic and taphonomic implications

During the Pleistocene, European representatives of macaque subspecies mainly occurred in the Mediterranean area, being probably bound by both climatic and latitudinal factors [START_REF] Ardito | An overview of the geographic and chronologic distribution of west European cercopithecoids[END_REF]Elton and O'Regan, 2008) (Fig. 3). Indeed, a Mediterranean climate, with warm dry summers and relatively wet winters, also characterizes the current distribution range of the extant Barbary macaque populations, even though the species can tolerate winter snow at high elevations [START_REF] Delson | Cercopithecidae from the Pliocene of Saint-Vallier[END_REF][START_REF] Fooden | Systematic review of the Barbary macaque, Macaca Sylvanus (Linnaeus, 1758)[END_REF]. Considering the known ecological tolerance, a strong relation between the areal expansion and Plio-Pleistocene climatic fluctuations for the extinct M. sylvanus subspecies has been hypothesized [START_REF] Fooden | Systematic review of the Barbary macaque, Macaca Sylvanus (Linnaeus, 1758)[END_REF]Elton and O'Regan, 2008). According to this idea, the European fossil macaques extended their geographical ranges to much higher latitudes during the interglacials, whereas they retreated into Mediterranean glacial refugia during cold episodes [START_REF] Fooden | Systematic review of the Barbary macaque, Macaca Sylvanus (Linnaeus, 1758)[END_REF]Elton and O'Regan, 2008). Its past distribution indicates that this primate was widespread along the Mediterranean coasts of Europe, especially in the Italian and Iberian peninsulas (Fig. 3). On the contrary, the presence of the species in central Europe, as well as in Britain, was only documented from deposit chronologically referred to an interglacial stage (Fig. 3). An exception is the Late Pleistocene site of Hunas (Germany) (Rosendahl et al., 2011), where the macaque remains were found in levels K-H, chronologically referred to MIS 4, therefore representing the only occurrence in central Europe during a glacial episode. As shown in Figure 3, the presence of the Barbary macaque at Notarchirico is the southernmost and easternmost known in Italy.

At Notarchirico, the co-occurrence of macaques and hominins is documented [START_REF] Moncel | The origin of early Acheulean expansion in Europe 700 ka ago: new findings at Notarchirico (Italy)[END_REF], as in some other fossiliferous deposits (e.g., Gruta da Aroeira, Lezetxiki II and Grotta degli Orsi Volanti). Faunal remains coming from the levels F to I of the recent fieldworks are generally highly affected by abrasion (trampling and hydraulic transport) and bone surfaces are poorly preserved.

Therefore, even though neither cut marks nor other anthropic percussion marks have been observed, this could be a mere lack of documentation. Likewise, despite a relatively good state of legibility (moderate abrasion), the macaque ulna shows no cut-marks. Similarly, the ulna from Fontana Ranuccio was found associated with lithic artefacts [START_REF] Segre | Fontana Ranuccio: Italy's earlyest Middle Pleistocene hominid site[END_REF][START_REF] Strani | Ungulate dietary adaptations and palaeoecology of the Middle Pleistocene site of Fontana Ranuccio (Anagni, Central Italy)[END_REF], but there is no indication of human exploitation.

Despite the Barbary macaques being very widely distributed in Europe throughout the Pliocene and Pleistocene, the fossil record of this taxon is limited, and its presence in Europe was relatively rare.

The majority of the documented specimens in Europe is isolated teeth and there are relatively few postcranial elements. Plio-Pleistocene Macaca sylvanus could probably have inhabited both deciduous and coniferous forest, but it could also be adapted to transitional environments.

The European macaque became locally extinct in the early Late Pleistocene. [START_REF] Elton | Macaques at the margins: the biogeography and extinction of Macaca sylvanus in Europe[END_REF] pointed out that its extinction may be related to vegetation change, or increased predation from Homo, but also to stochastic factors occurring as a result of small population sizes. The early Late Pleistocene extinctions also included some megafauna (Palaeoloxodon antiquus, Hippopotamus amphibius, and

Stephanorhinus hemitoechus), and it can be related to the lowering of global and regional temperatures in the Last Glacial period that caused reduction in forest cover. Another plausible factor is the influence of hominins. Definitely, the potential Homo-Macaca interactions during the Pleistocene will deserve more emphasis from both paleoecological and archaeozoological viewpoints. During the Pleistocene in Europe Macaca sylvanus co-existed alongside earlier Homo species for over 1 million years, but the Barbary macaques became extinct before the earliest occurrence of Homo sapiens in Europe [START_REF] Elton | Macaques at the margins: the biogeography and extinction of Macaca sylvanus in Europe[END_REF]. Modern Macaca sylvanus distribution is negatively affected by human occupation. In many Middle and Late Pleistocene sites hominins and macaques were found together and, especially in resource-limited environments.

Although there is still no known evidence in paleontological literature, a possible scenario where Homo included primates in its diet cannot be ruled out [START_REF] Meloro | The evolutionary history and palaeo-ecology of primate predation: Macaca sylvanus from Plio-Pleistocene Europe as a case study[END_REF]. The environmental context given by this find of macaque provides informative data on the climatic conditions of the hominin occupations and the plant environments where hominins able to produce the earliest bifaces in southern Italy lived.

Fig. 2 -Proportions of the ulna of Macaca sylvanus from Notarchirico, compared to extinct and extant specimens of Barbary macaque and Mesopithecus pentelicus pentelicus. Specimens identified in more detail in Table 1. aboxplot of the CBP measurements (CBP = greatest breadth across the coronoid process); bplot of HOF (greatest height of olecranon fossa) vs. CBP (greatest breadth across the coronoid process). SD1. Question mark denotes uncertain age. Map produced with QGIS. HOF -greatest height of olecranon fossa; APD -greatest depth across the anconeal process.

Fig. 1 -

 1 Fig. 1 -Location map (1a), view and description of the stratigraphic succession of Notarchirico (southern Italy) (1b-1c). Monkey outline in 1a indicates the geographic position of the site. B24GNC, fragmented right ulna from Notarchirico in medial (1d), anterior (1e) and lateral (1f) views. Scale bar for Figs. d-f 3 cm.

  Pleistocene specimens of Barbary macaque from Fontana Ranuccio (FR; specimen 85; stored at Italian Institue of Human Paleontology) and Cava di Breccia di Casal Selceupper level (CS; specimen 520; stored at PaleoFactory laboratory, Department of Earth Sciences, Sapienza University of Rome (PF)).

  Institutional abbreviations CAM: Comparative Anatomy Museum, University of Modena; CZMR: Civic Zoology Museum of Rome; MGPT: Museum of Geology and Paleontology, Turin University; MHNG: Muséum d'Histoire Naturelle de Genève; MNAV: Museo Nazionale Archeologico di Venosa; NMB: Naturhistorisches Museum Basel; NHW: Naturhistorisches Museum Wien; PF: PaleoFactory laboratory, Department of Earth Sciences, Sapienza University of Rome; Italian Institute of Human Paleontology (IsIPU);

Fig. 2 -

 2 Fig. 2 -Proportions of the ulna of Macaca sylvanus from Notarchirico, compared to extinct and extant specimens of Barbary macaque and Mesopithecus pentelicus pentelicus. Specimens identified in more detail in Table 1. aboxplot of the CBP measurements (CBP = greatest breadth across the coronoid process); bplot of HOF (greatest height of olecranon fossa) vs. CBP (greatest breadth across the coronoid process).

Fig. 3 -

 3 Fig. 3 -Macaca remains in the late Miocene to Late Pleistocene of Europe, details are provided in TableSD1. Question mark denotes uncertain age. Map produced with QGIS.

Fig. 3 -

 3 Fig. 3 -Macaca remains in the late Miocene to Late Pleistocene of Europe, details are provided in TableSD1. Question mark denotes uncertain age. Map produced with QGIS.

Table 1 -

 1 Measurements of ulna of extinct and extant specimens of Barbary macaque and extinctMesopithecus pentelicus pentelicus. * estimated measurements from[START_REF] Rook | Fossil Macaca remains from RDB quarry (Villafranca d'Asti, Italy): new data and overview[END_REF].

	Species	Catalogue number	Site	Age	Reference	OB	APB	CPB	HOF	APD
	Macaca sylvanus cf. prisca	NMB VJ 130	RDB quarry	late Pliocene	Rook et al. (2001)			17.6	11.9*	8.3*
	Macaca sylvanus	PF -CS520	Cava di Breccia di Casal Selce -upper level	Middle Pleistocene	This work			15.7	12.9	7.8
	Macaca sylvanus	IsIPU -FR85	Fontana Ranuccio	Middle Pleistocene	This work	13.5	14.4	16.7	13.3	8.3
	Macaca sylvanus	MNAV -B24GNC	Notarchirico	Middle Pleistocene	This work	14.1	18.0	20.7	13.1	9.3
		CAM-1637			Rook et al. (2001)			15.0
		CAM-283/4			Rook et al. (2001)			18.0
		NHMW-34222			Rook et al. (2001)			14.0
	Extant Macaca sylvanus	NHMW-32990			Rook et al. (2001)			19.0
	sylvanus	MHNG-3383			Rook et al. (2001)			20.0
		MHNG-446•36			Rook et al. (2001)			18.0
		CZMR -MP 128 -Dx			This work			16.3
		CZMR -MP 128 -Sx			This work			15.5
	cf. Mesopithecus pentelicus pentelicus	MGPT-PU 130507	Moncucco Torinese	late Miocene	Alba et al. (2013) 14.1		16.0	14.4

Table Table 1

 Table -Measurements of ulna of extinct and extant specimens of Barbary macaque and extinct Mesopithecus pentelicus pentelicus. * estimated measurements from[START_REF] Rook | Fossil Macaca remains from RDB quarry (Villafranca d'Asti, Italy): new data and overview[END_REF]. Literature data on extinct of Barbary macaque from late Miocene to Late Pleistocene sites of Europe Fig.SD1-Standard measurements of ulna. Abbrevations: OB -greatest breadth of olecranon; APB -greatest breadth across the anconeal process; CPB -greatest breadth across the coronoid process;

	Supplementary
	Tab.SD1 -

Acknowledgments

This work was realized thanks to the financial and scientific support of the Leakey Foundation ("Early Evidence of Acheulean bifacial technology in Europe" grant, 2015Europe" grant, -2016) ) and the National Museum of Natural History, Paris, France (ATM Action Transversale du Muséum, 2016-2018). We are indebted to the Soprintendenza of Basilicata (Italy) for the agreement to study fossil collections of the site of Notarchirico, especially Dr. T.E. Cinquantaquattro, Dr. F. Canestrini, Dr. R. Pirraglia, and Dr. S. Mutino. We thank the Venosa Museum and Dr. A. Mantrisi for their assistance. We are grateful to R. Carlini (Civic Zoology Museum of Rome) and O. Pauwels (Royal Belgian Institute of Natural Sciences, Brussels) for their support in the study of the comparative material. The authors thank L. Bellucci J. Conti, D.A. Iurino and F. Strani for their useful suggestions and support. We are indebted to Eric Delson for the English revision of the submitted work. We thank Eric Delson and two anonymous reviewers, the Associate Editor and Editor Andrea B. Taylor for their helpful comments and suggestions, which were crucial to substantially improved our manuscript.

Tab. SD1