LARGE MAMMALS FROM THE MIDDLE PLEISTOCENE (MIS 11) SITE OF FONTIGNANO 2 (ROME, CENTRAL ITALY), WITH AN OVERVIEW OF “SAN COSIMATO” ASSEMBLAGES. Alessio Iannucci 1, Beniamino Mecozzi 1, Raffaele Sardella 1 1 Dipartimento di Scienze della Terra, PaleoFactory, Sapienza Università di Roma, Roma, Italy Corresponding author: B. Mecozzi ABSTRACT: Here we describe fossil mammal remains recovered from the late Middle Pleistocene (MIS 11) of Fontignano 2 (Rome, central Italy). Two species are recognized: the aurochs Bos primigenius and the red deer Cervus elaphus. The presence of B. primigenius represents one of the earliest diagnostic evidence of the species. Conversely, remains of C. elaphus are not fully diagnostic at a subspecies level, despite being long considered among the reference occurrences of C. e. eostephanoceros in Italy. This reconsideration, concurrent with the revised chronology of several localities of the area of Rome, questions the validity of the chronosubspecific and evolutionary repartition of the red deer as often envisioned in the literature, i.e., C. e. acoronatus, C. e. eostephanoceros, C. e. rianensis, C. e. elaphus. Remains of Fontignano 2 are part of those recovered within the San Cosimato Formation, also including Via di Brava (MIS 13 or MIS 11), with Palaeoloxodon antiquus, and San Cosimato (MIS 11), with Steph- anorhinus sp., B. primigenius, Castor fiber, and Emys orbicularis. Keywords: Fossil vertebrate, Biochronology, Galerian, Quaternary, Italian Peninsula. Available online http://amq.aiqua.it ISSN (print): 2279-7327, ISSN (online): 2279-7335 Alpine and Mediterranean Quaternary, 34 (1), 2021, 155-164 1. INTRODUCTION The area of Ponte Galeria (Rome, Fig. 1), along the Tyrrhenian coast in central Italy, has yielded an ex- ceptional amount of fossil remains recovered from geo- chronologically calibrated deposits, providing a unique source of knowledge on Middle Pleistocene terrestrial ecosystems of Europe. Indeed, the progressive uplift of the area led to continental depositional conditions since about 0.8 Ma, in coincidence with the start of the activity of the Roman Magmatic Province (Conticelli & Pecceril- lo, 1992; De Rita et al., 1993; Karner et al., 2001). As a result, volcanic products are intercalated within the fluvi- al and lacustrine sedimentary successions deposited from this point onwards, providing radiometric age con- straints for the correlation of the several fossiliferous localities that are here represented (Ambrosetti, 1965; Ambrosetti & Bonadonna, 1967; Ambrosetti et al., 1972; Caloi & Palombo, 1978, 1979b, 1986; Kotsakis et al., 1992; Di Stefano et al., 1998; Petronio & Sardella, 1999; Milli et al., 2008; Palombo & Milli, 2011; Sardella & Petrucci, 2012; Marra et al., 2014; Sardella et al., 2015). The seminal work of Conato et al. (1980) recognized five informal lithostratigraphic units in the area, extend- ing from the late Early Pleistocene to the late Middle Pleistocene: Monte Mario, Ponte Galeria, S. Cosimato, Aurelia, and Vitinia Formations. Subsequent studies have improved this scheme and further developed its calibration with the Marine Isotopic Scale (MIS), in turn resulting in a refined chronological placement of several fossiliferous localities (Palombo & Milli, 2011; Sardella & Petrucci, 2012; Marra et al., 2014). The name “San Co- simato” has often been employed as a composite Local Fauna, including sporadic fossil findings recovered with- in the San Cosimato Formation (sensu Conato et al., 1980; e.g., Palombo, 2004). However, since deposits of the San Cosimato Formation are now known to repre- sent two distinct sedimentary cycles correlated to Valle Giulia (MIS 13) and San Paolo (MIS 11) Formations (Marra & Rosa, 1995; Funiciello & Giordano, 2008), there are some ambiguities in referring to these assem- blages (Fig. 1). Kotsakis et al. (1992) reported the presence of the two arvicolid rodents Prolagurus pannonicus and ?Predicrostonyx sp. from the “Blue-grey Helicella clays” (Ponte Galeria Formation) at Fontignano. These species are considered indicators of cold climate and are associated with an oligotypic assemblage of mol- luscs that also support this paleoclimatic inference. The Local Fauna constituted by Prolagurus pannonicus and ?Predicrostonyx sp. is the oldest in the area of Pon- te Galeria and we refer to it as Fontignano 1, to distin- guish it from remains recovered from the upper levels (previously considered as San Cosimato Formation) of the same site, which represent Fontignano 2. Here we provide description and taxonomic attribu- tion of the large mammal fossil remains of Fontignano 2, which, although mentioned in several works (e.g., Di https://doi.org/10.26382/AMQ.2021.07 156 Iannucci A. et al. Stefano & Petronio, 1993; Marra et al., 2014) have nev- er been described so far. Moreover, we discuss bio- chronological implications for the species recognized and we summarize the knowledge on San Cosimato fossil assemblages. 2. MATERIALS AND METHODS The fossil remains described in this work were recovered in the 1990s, during geological surveys car- ried out in the area of Ponte Galeria (Salami, 1992), and they are housed in the PaleoFactory Laboratory, De- partment of Earth Sciences, Sapienza University of Rome (PF). The morphological analysis is based on the comparison with other fossil remains as well as recent osteological material stored in the same institution and in the University Museum of Earth Science, Department of Earth Sciences, Sapienza, University of Rome (MUST). Measurements were taken with sliding calipers nearest to 0.1 mm, mainly following von den Driesch (1976). We also measured the maximum anteroposteri- or depth of the distal epiphysis of the humerus (DAPd). Uncertain values are preceded by “c.” throughout the text. We considered: Greatest length (GL); Greatest length from caput (GLC); Smallest breadth of diaphysis (SD); Anteroposterior distal diameter (DAPd). Other abbreviations employed in this work are as follows: Faunal Unit (FU); Formation (Fm); Local Fauna (LFA) 3. SYSTEMATIC PALAEONTOLOGY Order ARTIODACTYLA Owen, 1848 Family BOVIDAE Gray, 1821 Subfamily BOVINAE Gray, 1821 Genus BOS Linnaeus, 1758 Species Bos primigenius Bojanus, 1827 Material: right humerus (PF-Font2-1; Fig. 2a1-a2); left horn core (PF-Font2-2; Fig. 2b1-b2). Description and comparison: The left horn core (PF -Font2-2) also preserves part of the postero-lateral re- gion of the cranium. It is in a better condition than that of the other specimen, although its posterior portion is damaged near the base of the horn. The cornual pro- cess is separated from the frontal bone by a marked neck; from this point it extends first laterally, and then anteriorly and upwards. The cross section of the horn is oval in the proximal preserved part, more elongated antero-posteriorly than dorso-ventrally. The right humerus (PF-Font2-1) is almost complete but severely fractured, and its epiphyses are damaged at the level of the lateral tuberosity (proximally) and of the medial epicondyle (distally). The bone surface is partially encrusted by a grayish patina, with sporadic presence of small pebbles (1 - 5 cm in size). In the prox- imal epiphysis, the floor of the bicipital groove is swelling in anterior view. Laterally, the lateral tuberosity possess- es two shallow grooves. The diaphysis possesses a well -developed midshaft lateral imprint of the deltoid tuber- Fig. 1 - Location and geochronological setting of the studied area. The geological Formations formerly included in the San Cosimato For- mation are highlighted. Data from: Ambrosetti (1965); Ambrosetti & Bonadonna (1967); Ambrosetti et al. (1972); Caloi & Palombo (1978, 1979b, 1986); Kotsakis et al. (1978, 1992); Conato et al. (1980); Marra & Rosa (1995); Di Stefano et al. (1998); Petronio & Sardella (1999); Palombo (2004); Milli et al. (2008); Palombo & Milli (2011); Sardella & Petrucci (2012); Marra et al. (2014); Sardella et al. (2015); Pereira et al. (2018); Boschian et al. (2019); Mecozzi et al. (2021a). 157 Large mammals from Fontignano 2 osity in anterior view. In the distal epiphysis the lateral condyle is complete and appears laterally projected. Laterally, the ridge connecting the main shaft with the lateral condyle curves at the level of the junction with the main shaft. The margin of the lateral epicondyle is short and curve, and originates below the lateral epicon- dylar crest. There are objective difficulties in the attribution of postcranial remains of Bovidae to Bison or Bos (e.g., Masini et al., 2013). Nevertheless, PF-Font2-1 is large, measuring c. 460.0 mm in GL, 391.7 mm in GLC, and 78.8 mm in SD; and stout in appearance, especially at the level of the deltoid tuberosity. Morphologically, most of the traits observed in PF-Font2-1 fall in the variability of B. primigenius, whereas are different than those of Bison (Sala, 1986; Balkwill & Cumbaa, 1992). The mor- phological features, and the large size and robustness support an attribution to B. primigenius. PF-Font2-2 can be confidently assigned to the same species. Indeed, in Bison the horns are inserted in the cranium in a more anterior position, and they have a different development, protruding outward and back- ward; in Hemibos the horns are directed posteriorly, with a subtriangular cross section (Petronio & Sardella, 1998; Martínez-Navarro & Palombo, 2004; Masini et al., 2013). Family CERVIDAE Goldfuss, 1820 Subfamily CERVINAE Goldfuss, 1820 Genus CERVUS Linnaeus, 1758 Species Cervus elaphus Linnaeus, 1758 Material: right antler (PF-Font2-3; Fig. 2e); left dis- tal humerus (PF-Font2-4; Fig. 2c1-c4); right distal hu- merus (PF-Font2-5; Fig. 2d1-d2). Description and comparison: the right antler PF- Font2-3 preserves only the basal part of the beam, with the insertion of the brown tine and the bez tine, but these two tines are broken. The beam shows pearls and knots typical of the red deer. The beam has an elliptical outline, with the major axis anteroposteriorly oriented. At the level of the insertion, the brow and bez tines have an elliptical outline, flattened mediolaterally. The lack of the tines prevents to estimate the angle that these form with the beam. The bez tine is near to the brow tine, dis- Fig. 2 - Mammal remains from Fontignano 2: Bos primigenius right humerus PF-Font2-1 in caudal (a1) and cranial (a2) views, left horn core PF-Font2-2 in dorsal (b1) and frontal (b2) views; Cervus elaphus left humerus PF-Font2-4 in cranial (c1), caudal (c2), lateral (c3), and medial (c4) views, right distal humerus PF-Font2-5 in caudal (d1) and cranial (d2) views, right antler PF-Font2-3 in external view (e1). tanced 42.5 mm from the beam. Above the bez tine, the beam is long 236.9 mm, and there is no trace of the trez tine. Other measurements of the antler are: the antero- posterior breadth of the burr - 64.9 mm; the medio- lateral breadth of the burr - 52.7 mm; the antero- posterior breadth of the beam between the brow and the bez tines - 72.2 mm; the depth of the brow tine at the base - 52.1 mm; the breadth of the brow tine at the base - 30.4 mm; the depth of the bez tine at the base - 34.4 mm; the breadth of the bez tine at the base - 36.5 mm. The main differences between C. e. acoronatus and C. e. eostephanoceros is the terminal part of the antler, or crown (Fig. 3). The acoronate red deer lacks a real crown, possessing a terminal fork with only two points (Di Stefano & Petronio, 1992, 1993). On the con- trary, the eostephanoceros red deer shows a real crown formed by at least 5 tines (Di Stefano & Petronio, 1992, 1993). The most reliable features in the identification of red deer subspecies are missing in the antler from Fon- tignano, which suggests caution in the taxonomical at- tribution. The left humerus (PF-Font2-4) preserves part of the diaphysis and the distal epiphysis, but the medial condyle is fragmented. In anterior view, the radial pit is slightly larger than the coronoid pit; the epicondylar crest is damaged but appears robust; the capitulum is not preserved; the crest of the lateral condyle is more marked than that of medial condyle. Posteriorly, the medial condyle is more pronounced than the lateral condyle; the olecranon fossa is wide distally, whereas is narrower, but deeper, proximally. PF-Font-5 is a frag- mented humerus diaphysis, lacking epiphyses. The morphology and size of the shaft resemble those of PF- Font2-4. The morphology of the humerus in fallow and red deer is quite similar, especially in the early Middle Pleis- tocene taxa. Nevertheless, the relative development of the radial and coronoid pit is one of the diagnostic char- acters to differentiate the modern fallow deer and red deer (Di Stefano, 1995; Lister, 1996). Following this, PF- Font2-4 can be attributed to C. elaphus. Considering the biometry, Di Stefano (1995) noticed that the angle de- scribed by the orientation of the trochlea is usually more acute in Dama (<18°) than in Cervus (>20°). In PF- Font2-4 this angle spans 20° and the DAPd measures 53.8 mm, values that support an attribution to C. ela- phus. 4. DISCUSSION 4.1. Biochronological implications Bos primigenius According to Masini et al. (2013) B. primigenius is first documented in Isernia FU, even though not from the type locality, but as a component of other LFAs of the Venosa Basin in Basilicata, namely Notarchirico (Cassoli et al., 1999) and Loreto (Caloi & Palombo, 1979a). How- ever, remains from these localities are scanty and main- ly limited to postcranial material, elements not fully diag- nostic in Bovidae (Moncel et al., 2020). Indeed, Masini et al. (2013, p. 54), discussing the morphological differ- ences between B. priscus and B. primigenius, remarked that: “Although a number of authors try to tell the differ- ence between the postcranial elements, the objective difficulty of firmly identify either no diagnostic or imper- fectly preserved remains of the large bovids, makes it difficult to ascertain the actual extent in time, space and number of findings of the steppe bison fossil record in Italy”. Moreover, the possibility that some remains of the enigmatic Hemibos galerianus (Petronio & Sardella, 1998) may have been misidentified cannot be ruled out, since up to now the dental and postcranial morphology of this Middle Pleistocene bovid are still unknown. For instance, Caloi & Palombo (1979a) underlined the slen- derness of postcranial remains of B. primigenius from Loreto, but for reconciliate this evidence with the species attribution they argued that the sample likely consists mainly of female individuals. It is worth highlighting that the presence of H. galerianus in Italy is documented only by two fragmentary horn cores collected from the lower level of Cava di Breccia di Casal Selce (Petronio & Sardella, 1998; Martínez-Navarro & Palombo, 2004) and Ponte Molle (= Ponte Milvio) (Martínez-Navarro & Pal- ombo, 2007). Martínez-Navarro et al. (2010) stressed the importance of the coincidence between the dispersal of Bos and the Acheulean culture in Eurasia. However, at Notarchirico remains attributed to B. primigenius are only reported from the upper levels of the stratigraphic succession (Cassoli et al., 1999), whereas the occur- rence of the species cannot be confirmed in lower levels that document the earliest evidence of an Acheulean settlement in Italy (Moncel et al., 2020; Mecozzi et al. 2021b). A fragment of a frontal bone and a fragmentary proximal femur referred to B. primigenius were also found at Monte Oliveto (Berzi, 1972), but a revision of 158 Iannucci A. et al. Tab. 1 - Middle Pleistocene Cervus taxa from Europe considered in the text as established by authors. part there is a bifurcate fork (Fig. 3a-b) (Di Stefano & Petronio, 1992). The term acoronatus was chosen to indicate an antler where the terminal crown lacks, as opposite to the crown that instead characterizes the modern C. elaphus. The second taxon is C. e. eostepha- noceros, recognizable for a first crown-like antler, even if different from that of the modern red deer (Fig. 3c-e) (Di Stefano & Petronio, 1993). As exemplified by the de- scription above, the identification of red deer subspecies rests exclusively on antler morphology, which compli- cate the reconstruction of the evolutionary history of C. elaphus. Indeed, antler remains are not ubiquitously represented in the fossil record, and underestimation of ontogenetically related morphological changes (e.g., presence/absence and number of tines, development of the crown) may led to misidentifications (Heintz, 1970; Lister, 1987). In Italy, antler remains were collected from Fontana Ranuccio (Di Stefano & Petronio, 1993), Cava Nera Molinario (type locality of C. e. eostephanoceros) (Di Stefano & Petronio, 1993), Ponte Molle (Mecozzi et al., 2021a), Via Flaminia 8.2 Km (Kotsakis et al., 1978), Riano (Leonardi & Petronio, 1974), Torre del Pagliaccet- to (Caloi et al., 1978), and Castel di Guido (Sala & Barbi, 1996). In Europe, C. e. acoronatus was identified from the Middle Pleistocene sites of Süssenborn and Mosbach (type locality) (Kahlke, 1960, 1969; Lister, 1990; Di Stefano & Petronio, 1992). Two other putative red deer species were established on the sample of Mosbach: “Cervus” elaphoides Kahlke, 1960 and this material is needed to con- firm its classification. B. primi- genius was also reported by Valdemino (Tozzi, 1969; Noc- chi & Sala, 1997), but the material was never described or figured. Finally, a nearly complete hemimandible from Cava Rinaldi was referred to B. primigenius by Ambrosetti (1965), but later reassigned to Bison sp. (Sala, 1986; Petronio & Sardella, 1998). Unequivocal fossil re- mains of B. primigenius are only known since Fontana Ranuccio FU, as exemplified by remains from Fontana Ra- nuccio (Cassoli & Segre Naldini, 1993; Strani et al., 2018), Ponte Molle (Mecozzi et al., 2021a), and the horn core from Fontignano 2 de- scribed in this work. In this scenario, an early dispersal of B. primigenius (older than Fontana Ranuccio FU) in the Italian Peninsula cannot be confirmed. In younger depos- its, B. primigenius became soon a common element, as testified for example by the late Middle Pleistocene sites of Malagrotta (Caloi & Pal- ombo, 1979b) and Castel di Guido (Sala & Barbi, 1996). Cervus elaphus The red deer, C. elaphus is a typical Middle Pleis- tocene species but is first documented from a few late Early Pleistocene sites (Atapuerca-TD4-6, van der Made et al., 2017; Bermúdez de Castro et al., 1999; Dorn Dürkheim, Franzen et al., 2000; Happisburgh HSB3, Parfitt et al., 2010). Many chronological and/or geographical subspe- cies of C. elaphus were named (Tab. 1), with a general agreement in recognizing the simpler antler morphology of earlier forms, but several alternative opinions on the validity and relationships of subsequent taxa (e.g., Heintz, 1970; Di Stefano & Petronio, 2002; van der Made et al., 2014; Croitor, 2018). In Italy, largely relying on remains recovered from the area of Rome, the red deer lineage is often considered to be represented by four different chronosubspecies: C. e. acoronatus (Beninde, 1937), C. e. eostephanoceros Di Stefano & Petronio, 1993, C. e. rianensis Leonardi & Petronio, 1974, C. e. elaphus (Leonardi & Petronio, 1974; Di Stefano & Petronio, 1992, 1993, 2002), notwithstanding the occurrence of endemic island forms and of C. e. aretinus Azzaroli, 1961 from Tuscany (Azzaroli, 1948, 1961; Di Stefano & Petronio, 2002; Croitor, 2018). The antler of the acoronatus red deer possesses five points, with two basal tines and a third tines in the high position on the beam, all directed anteriorly; while in the upper 159 Large mammals from Fontignano 2 Fig. 3 - Antler morphology of Cervus elaphus from localities of the area of Rome: a - Ponte Molle, b - Via Flaminia Km 8.2, c - Cava Nera Molinario, d - Ponte Molle, e - Fontana Ranuccio, f - Fontignano 2, g - Riano, h - Torre del Pagliaccetto, i-l - Castel di Guido; a-b - C. e. acoronatus, c-e - C. e. eoste- phanoceros, f - C. elaphus ssp., g-l C. e. elaphus. Figures not in scale. “Cervus” reichenaui Kahlke, 1995. However, Lister (1990) recognized that specimens of “C.” elaphoides represented young individuals of C. e. acoronatus; and Pfeiffer (1997) reassigned to Dama (Pseudodama) reichenaui the other taxon. Other evidences of C. e. acoronatus were reported by Di Stefano & Petronio (1992), but these were then reconsidered as Dama (e.g., Caune de l’Arago, Saint Prest; Lister, 1990) or the lack of antlers limits the attrib- ution at the species level (e.g., Atapuerca, Azanza & Sanchez, 1990). From the Middle Pleistocene locality of Mauer was instead reported the subspecies C. e. priscus (Kaup, 1839), a form considered different from C. e. acoronatus in having an incipient crown, with six points (instead of the maximum of 5 points in C. e. acor- onatus). C. e. eostephanoceros was only reported from Hundsheim (Di Stefano & Petronio, 1993). Subsequent recovery and description of new material confirm the occurrence of C. e. acoronatus at Caune de l’Arago, and also of C. e. priscus (or C. e. eostephanoceros), providing evidence of a turnover of two forms between MIS 13 and MIS 12 (Magniez et al., 2013). The authors, however, referred the younger form to C. e. priscus (or C. e. eostephanoceros), reflecting the close morphologi- cal affinity and uncertain relationship between these two taxa. Of considerable interest for red deer evolution is the sample recovered at Steinheim, which was referred to C. e. angulatus Beninde, 1937. Eleven complete or nearly complete antlers were illustrated by Beninde (1937), showing a huge morphological variability. Anoth- er taxon was proposed, based on the sample from the Middle Pleistocene site of Riano, C. e. rianensis (Leonardi & Petronio, 1974). As reported by the authors, the four antlers recovered from this site show a large variability in number of points (five or six) and develop- ment of the crown. Although there are slight differences between the antler from Riano and those of the modern red deer, no subspecies separation would seem sup- ported (Fig. 3g-l). In this respect, the antler found in the Via Flaminia Km 8.2 deposit, initially referred to C. e. rianensis (Kotsakis et al., 1978) is particularly relevant (Fig. 3b). In fact, the chronostratigraphic revision of this deposit was carried out by Marra et al. (2018), who re- ferred the fossil level to MIS 13. The authors, in addition, noticed that the antler is closer to C. e. acoronatus ra- ther than to C. e. rianensis. Considering that the type material of C. e. eostephanoceros from Cava Nera Moli- 160 Iannucci A. et al. Fig. 4 - Mammal remains from Via di Brava: Palaeloxodon antiquus molar PF-Brava-1 in occlusal view (a); and San Cosimato: Bos primi- genius right upper first molar MPUR sn019/LJ in occlusal view (b), Stephanorhinus sp. left lower third or fourth premolar MPUR sn023/LJ in buccal view (c), Castor fiber left lower fourth premolar MPUR sn021/LJ in occlusal view (d). nario is also correlated to MIS 13 but placed in a lower stratigraphic position (Marra et al., 2014, 2018), this suggests a chronological overlap between the two forms. 4.2. San Cosimato assemblages Apart from B. primigenius and C. elaphus from Fontignano 2, there are several other remains recov- ered from the San Cosimato Fm of Conato et al. (1980) (Fig. 4). Palombo (2004) provided a faunal list, even though without further details on the provenance or de- scription of the material: Stephanorhinus sp., ?Dama sp., Cervus cf. C. e. eostephanoceros, B. primigenius, and Castor fiber. However, the fossils reported by Cona- to et al. (1980) (Emys orbicularis, beavers, and rhinocer- os) and other associated remains are housed in MUST and catalogued as from “Via della Pisana - San Cosima- to”, suggesting that they were recovered from the out- crops of the San Cosimato Fm at Via della Pisana, now referred to the MIS 11 sedimentary cycle (San Paolo Fm; Marra et al., 2014). Salami (1992) also mentioned the recovery of a fragmentary molar of Palaeoloxodon antiquus from an outcrop of the San Cosimato Fm at Via di Brava (Fig. 4a). This specimen is housed in PF, but it is not accompanied by more precise stratigraphic infor- mation that would allow a correlation with updated Fms, and therefore it could be either MIS 13 or MIS 11 in age (Valle Giulia or San Paolo Fms). Finally, Palombo & Ferretti (2005) reported undescribed remains of Mam- muthus trogontherii from Fontignano (Rome) housed in the Geological Museum “Giovanni Capellini” of Bologna. However, this is a mistake resulting from a case of ho- monymy. In fact, according to Verri (1886) the material stored in Bologna was recovered from Fontignano in Umbria (near Perugia). 5. CONCLUSION Three distinctive LFAs can be recognized under the name San Cosimato: Fontignano 2 (MIS 11), with B. primigenius and C. elaphus; Via di Brava (MIS 13 or MIS 11), with P. antiquus; and San Cosimato (MIS 11), with Stephanorhinus sp., B. primigenius, C. fiber, and E. orbicularis. According to Palombo (2004), to this list can be added ?Dama sp. We argue that the name San Cosimato should be restricted to the LFA. Indeed, for bichronological correla- tions and paleoecological inferences it would be more desirable to refer to updated Fms (e.g., Funiciello & Giordano, 2008), or to geochronologically constrained localities withing them (Milli et al., 2008; Marra et al., 2014; Sardella et al., 2015). The study of the fossils collected from Fontignano 2 reveals the presence of B. primigenius and C. ela- phus. An overview of the bovid sample of the Middle Pleistocene sites of the Italian Peninsula suggests a first dispersal of B. prigeminius in Fontana Ranuccio FU, with no clear evidence of an earlier diffusion (Isernia FU). The geological revision of many fossiliferous sites of the city of Rome and its surroundings allowed to up- date their chronology. In this new scenario, the two red deer subspecies, C. e. acoronatus and C. e. eostepha- noceros, seem to overlap in time during MIS 13, as tes- tified by the lower stratigraphic position of the type mate- rial of C. e. eostephanoceros from Cava Nera Molinario than that of C. e. acoranatus from Via Flaminia Km 8.2. The coexistence of two red deer subspecies within a so restricted geographical area is hardly conceivable and may reflects the shortcomings of a dichotomic separa- tion in the absence of a clear understanding of popula- tion variability. In this respect it is worth to remark that the replacement between acoronated and coronated forms occurring at Caune de l’Arago during the MIS 13 - MIS 12 transition is not abrupt (Magniez et al., 2013), and the presence of more advanced morphologies with- in the samples of Mosbach and Mauer was also noted (Lister, 1987; Di Stefano & Petronio, 1993). More in general, complete antler remains of C. elaphus from the Pleistocene of Europe are scanty, and other morphologi- cal and biometric analyses are needed to complement our knowledge on the evolution of the species (Lister et al., 2010; Magniez et al., 2013; van der Made et al., 2014; Iannucci et al., 2020). The description of the fossil remains from Fon- tignano 2 highlights the importance of adding faunal element with a clear chronological context for the recon- struction of the Quaternary terrestrial ecosystem of the Euro-Mediterranean area. ACKNOWLEDGEMENTS We thank the Soprintendenza Speciale di Roma, Archeologia, Belle Arti e Paesaggio for the agreement of the field activities on the area of Rome. We are grateful to Linda Riti and Michele Macri for the help and support during the study of the fossil collection of MUST. We wish to thank Ilaria Mazzini, Luca Bellucci, Dawid Adam Iurino, Flavia Strani and Jacopo Conti for their useful suggestions and support. We also thank Gbola Salami for having carried out the geological and stratigraphical reconstruction of selected sedimentary successions of the Ponte Galeria area and for collecting fossil remains from these deposits. We are thankful for helpful com- ments and suggestions provided by two anonymous reviewers. 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