NTOLOG EO I L CA E TA ' PA Bollettino della Società Paleontologica Italiana, 61 (2), 2022, 223-247. Modena I TA L I A CI O N S A S. P. I. Vigna Nuova: the first Middle Villafranchian mammal assemblage from the Valdichiana Basin, Perugia (Italy) Beatrice Azzarà*, Marzia Breda, Omar Cirilli, Joan Madurell-Malapeira, Federica Ruzza, Leonardo Sorbelli, Domenico Tancredi & Marco Cherin B. Azzarà, Dipartimento di Fisica e Geologia, Università degli Studi di Perugia, Via Pascoli, I-06123 Perugia, Italy; beatrice.azzara@studenti.unipg.it *corresponding author M. Breda, Centro di Ateneo per i Musei (CAM), Università di Padova, Via Giotto 1, I-35121 Padova, Italy; marzia.breda@unipd.it O. Cirilli, Laboratory of Evolutionary Biology, Department of Anatomy, College of Medicine, Howard University, 20059 Washington DC, USA; Dipartimento di Scienze della Terra, Paleo[Fab]Lab, Università degli Studi di Firenze, Via La Pira 4, I-50121 Firenze, Italy; omar.cirilli@howard.edu J. Madurell-Malapeira, Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, E-08193 Cerdanyola del Vallès, Spain; joan.madurell@uab.cat F. Ruzza, Dipartimento di Fisica e Geologia, Università degli Studi di Perugia, Via Pascoli, I-06123 Perugia, Italy; federica.ruzza@studenti.unipg.it L. Sorbelli, Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, E-08193 Cerdanyola del Vallès, Spain; leonardo. sorbelli@icp.cat D. Tancredi, Dipartimento di Chimica, Biologia e Biotecnologie, Università degli Studi di Perugia, Via dell’Elce di Sotto 8, I-06123 Perugia, Italy; domenico.tancredi@studenti.unipg.it M. Cherin, Dipartimento di Fisica e Geologia, Università degli Studi di Perugia, Via Pascoli, I-06123 Perugia, Italy; marco.cherin@unipg.it KEY WORDS - Canis, Croizetoceros ramosus, Early Pleistocene, Equus senezensis, fossil mammals, Umbria. ABSTRACT - The region of Umbria (Central Italy) represents one of the most interesting areas for the study of Mediterranean Plio- Pleistocene mammal faunas due to the occurrence of numerous paleontological deposits. Most of the fossils have been discovered within the Tiberino Basin, which extends across the centre of the territory for ca. 1800 km2, and the small Tavernelle-Pietrafitta Basin, south of Lake Trasimeno. Herein, we provide an additional report on an assemblage within continental deposits from the locality of Vigna Nuova (Piegaro, Perugia) in the Valdichiana Basin, which are observed to crop out in only a few areas of western Umbria. Taphonomic and geological evidence, coupled with rare excavation notes, have allowed the sample to be divided into two assemblages. The younger was collected from the conglomerates (here called “upper layer”) which are still visible on the outcrop and is composed only of remains of cf. Leptobos. The data obtained from this fragmentary sample together with some sedimentological-stratigraphic considerations suggest that it can be correlated with other well-known Valdichiana assemblages (e.g., Selvella, Farneta), dated at ca. 1.5 Ma. The older sub-sample comes from a putative paleosol (here called “lower layer”) no longer visible in the field. It includes cf. Megantereon, Canis sp., Mustelidae indet., Proboscidea indet., Equus cf. senezensis, cf. Leptobos, Croizetoceros ramosus, Pseudodama sp., Cervidae indet. (large size), and Sus cf. strozzii. This assemblage can be referred to the Coste San Giacomo Faunal Unit (late Middle Villafranchian; ca. 2.2-2.1 Ma). It represents the first well- preserved sample of this age in Umbria and one of the few recorded in Italy, opening new interesting research perspectives on the distribution of mammals within the Peninsula and also on the stratigraphy and evolution of the Valdichiana Basin. “Aggiungeva che per scavare le poche ossa che mi avrebbe subito inviate aveva dovuto faticare enormemente e lottare per due giorni contro ogni sorta di difficoltà. Ricevuto il graditissimo invio mi affrettai a consolidare e restaurare con ogni cura quelle primizie ed oggi sono lieto di poter annunziare, in così solenne adunanza, la interessante scoperta…” (“He added that in order to excavate the few bones that he would immediately send me he had had to work enormously and struggle for two days against all sorts of difficulties. Having received the very welcome shipment, I hastened to consolidate and restore those fruits of the soil with every care and today I am pleased to be able to announce, in such a solemn meeting, the interesting discovery…”). Giovanni Capellini, 1890 INTRODUCTION NW-SE trending grabens related to the eastward migration of the Apennine extensional deformation (Martini & Sagri, In spite of its small geographic extension (ca. 1993; Pascucci et al., 1999; Collettini et al., 2006). These 8500 km2), Umbria represents a treasure trove of very depressions were filled by marine sediments on the west interesting discoveries for Plio-Pleistocene vertebrate and by continental sediments on the east, with the transition paleontology. Mentioned since the first half of the 19th between the two realms (i.e., the Tyrrenian paleocoast) century (Cuvier, 1821), in the following decades these crossing western Umbria from Città della Pieve south-east discoveries began to draw the attention of the Italian to Orvieto (Ambrosetti et al., 1987; Bizzarri & Baldanza, scientific community (e.g., Verri, 1879; Pantanelli, 1886; 2020). The main intermontane basin in the central-eastern Tuccimei, 1891, 1895; Clerici, 1894, 1895; Meli, 1895), part of the region is the Tiberino Basin (Fig. 1a), which including eminent scholars such as Giovanni Capellini, extends for ca. 1800 km2 from Sansepolcro to Terni who published at least three contributions on Umbrian (south-west) and Spoleto (south-east) (Basilici, 1997). This mammals (Capellini, 1888, 1889, 1890). extensional basin, once erroneously interpreted as a single From the paleogeographic point of view, during the huge lake (“Tiberino Lake”; Lotti, 1917), has actually Plio-Pleistocene, Central Italy was intersected by a set of been occupied at least over the last 3 Ma by a complex ISSN 0375-7633 doi:10.4435/BSPI.2022.12 224 Bollettino della Società Paleontologica Italiana, 61 (2), 2022 Pantalla, Torre Picchio, Podere San Lorenzo, Villa San Faustino, among others) and for the Tavernelle-Pietrafitta Basin (see the aforementioned site of Pietrafitta) and, following the steps traced by the famous 19th century researchers, they have been the subject of many modern studies and revisions (e.g., Sardella et al., 1995, 2018; Girotti et al., 2003; Cherin et al., 2016, 2019a). On the contrary, in the western part of Umbria (i.e., in the Valdichiana Basin), the occurrence of marine vertebrates is known above all (fossil cetaceans from the Allerona area; Baldanza et al., 2018). This knowledge gap is filled by the Vigna Nuova local faunal assemblage (LFA), which represents the only record of continental mammals from this portion of the regional territory. The collection was briefly mentioned by Gentili (1993). A faunal list was published by Petronio et al. (2002) and copied-and-pasted by Argenti (2004), who also stated that the Vigna Nuova LFA should be referred to the Tasso faunal unit (FU) “for its stratigraphic position with respect to Pietrafitta and its paleontological assemblage (mainly for the presence of Megantereon cultridens)” (Argenti, 2004, p. 70). Here we provide the first systematic study of the Vigna Nuova LFA, ascertaining that, in fact, it is made up of two distinct assemblages, one of which offers some surprising biochronological information. GEOLOGICAL SETTING AND TAPHONOMY The Vigna Nuova LFA was unearthed in 1989 during excavation works on the front of a hill (named “Greppo Lidolfo”) for the planting of a vineyard (hence the name, which means “new vineyard”). The site is located near Piegaro (Perugia; 42°58’04.2”N, 12°04’04.7”E) at the easternmost border of the Valdichiana Basin (Fig. 1a). Some inconsistencies exist as regards the stratigraphy of the deposits cropping out in the area. In the description of the Geological Map of Italy in 1:50,000 scale (Foglio 310, Passignano sul Trasimeno), Barchi & Marroni (2010) Fig. 1 - Location of the study area. a) The extensional basins cited in mention the Vigna Nuova assemblage as recovered from the text are highlighted. Red star: Vigna Nuova; black star: Selvella. the Fosso Cigne Unit (Tiberino Basin, Nestore Synthem). b) Detail of the Vigna Nuova outcrop. Following Argenti (2004), the assemblage, hence the unit, is referred to the Tasso FU (ca. 1.8 Ma). Conversely, the most recent Umbrian geological mapping projects and dynamic system of lacustrine, marshy and riverine (Regione Umbria, 2013) include the Vigna Nuova locality environments interconnected to each other (Basilici, 1997). in the Meletto Unit (Valdichiana Basin, Sanfatucchio Among the smaller basins, it is worth mentioning the Synthem), also referred to the Tasso FU. The Meletto Tavernelle-Pietrafitta Basin (Fig. 1a), located south of the Unit is very similar to and probably partially heteropic Trasimeno Lake, characterized alternatively by lacustrine/ with the nearby Gioiella Unit, which yielded important palustrine and alluvial plain conditions during the Plio- samples of terrestrial mammals in the surroundings of the Pleistocene. This basin hosts the most important Umbrian homonymous town, in particular at the locality of Selvella fossiliferous site, and one of the most relevant in Europe, (De Giuli, 1987), which is located only 16 km north-west that is, Pietrafitta (Martinetto et al., 2014; Sorbelli et al., of Vigna Nuova (Fig. 1a). The Selvella LFA, originally 2021, and references therein). The Tavernelle-Pietrafitta dated at ca. 1 Ma (De Giuli, 1987), is today referred to the Basin is interposed between the Tiberino Basin to the east Farneta FU (ca. 1.5 Ma; Alberdi & Palombo, 2013; Cirilli and the Valdichiana Basin to the west (Fig. 1a). The latter et al., 2020). In fact, the lithological and sedimentological develops from north-west to south-east along the Tuscany- features of the Fosso Cigne and Meletto Units are quite Umbria-Latium boundary and has been filled by shallow similar. Both are mainly composed by conglomerates marine and continental sediments since the Early Pliocene with a reddish medium-coarse sand matrix. Clasts are (Bizzarri & Baldanza, 2020). poorly sorted and predominantly made of sandstone Reports on fossil continental mammals are abundant in the first unit and carbonate in the second. The main in the literature for the Tiberino Basin (e.g., localities of difference is that the Fosso Cigne conglomerates have B. Azzarà et alii - Mammals from Vigna Nuova, Perugia (Italy) 225 channeled geometry, while those of Meletto show no clear structures. Unfortunately, the Vigna Nuova outcrop (about six m in height) today is almost completely covered by vegetation, so it is not possible to describe the local stratigraphic succession. The few exposed portions (Fig. 1b) are made of conglomerates with rounded, polygenic clasts, sometimes with imbricate bedding, alternating with thin layers of medium-coarse sand. The current evidence therefore does not allow to resolve the doubts about the stratigraphic attribution of the outcrop, for which further geological studies will be necessary. Nonetheless, the taphonomic features of the Vigna Nuova mammal collection, coupled with excavation notes and labels, offer some hints on the stratigraphic context. Based on the lithology of the embedding sediment (when available), color of fossils, and their mineralization, it is possible to clearly separate the collection into two groups. The first includes fossils embedded by or maintaining traces of a whitish-yellowish sand with carbonate matrix, rich in large-sized quartz crystals and spots of black organic matter. Calcite crystals can occur as coating and infills within larger bone cavities (e.g., medullary cavity in long bones). Bones are both mineralized and impregnated (i.e., minerals and secondarily organic matter fill the smaller cavities such as trabeculae; Fig. 2a), thus being very heavy. Some show root traces. Few specimens (SABAP_UMB 129847, 129849, 129852, 129853, 129869, 129881) are encrusted by a yellow-brown carbonate coating with reticular concretions. This assemblage is characterized by considerable variability in taphonomic attributes, ranging from poorly preserved isolated specimens (very abraded and broken bones) to well-preserved ones, sometimes still in anatomical connection. The weathering stages (Behrensmeyer, 1978) are heterogeneous, ranging from 0 to 4 (Fig. 2b). Several taxa are recognized in this first assemblage (Supplementary Online Material, SOM), with about half of the identifiable specimens referred to Cervidae (see Systematic Paleontology). On the contrary, only a bovid taxon (probably a single individual) is recognized in the second group of bones. These latter show traces of a dark-yellow to reddish sand. Mineralization is much less advanced and the inner cavities of the bones, regardless of their size, are empty. Most of the remains show evidence of abrasion and breakage typical of fluvial transport, whereas weathering is low (Fig. 2b) suggesting a little time spent in the sub-aerial environment prior to burial. The above observations support the hypothesis that the two assemblages come from different stratigraphic layers. Fig. 2 - Taphonomy of the Vigna Nuova assemblages. a) Detail of the inner part of a proboscidean bone (SABAP_UMB 22.3.264) The remains of the second group certainly belong to the showing heavy mineralization and filling of trabeculae by minerals conglomerates cropping out at Vigna Nuova. A putative and organic matter. b) Distribution of weathering stages (following fluvial depositional environment for these sediments is Behrensmeyer, 1978) in the fossils from the two fossiliferous layers compatible with the taphonomic features of bovid bones. of Vigna Nuova. On the other side, no traces of the sediments associated to the fossils of the first group are visible today in the field. Based on original notes and labels, we know that the MATERIALS AND METHODS paleontological material was recovered near the bottom of the hill, therefore from a portion of the succession The studied material is curated by Soprintendenza that is no longer exposed today. On the basis of the few Archeologia Belle Arti e Paesaggio dell’Umbria, Italy lithological and taphonomic elements available, we can (SABAP_UMB) and was prepared/restored and studied hypothesize that this lower layer was a paleosol, but only at the Dipartimento di Fisica e Geologia of the Università future studies in the area could confirm this hypothesis. degli Studi di Perugia (Italy). For preparation details see The fossils of the two assemblages are treated as separate Cherin et al. (2019a). The identification codes of the (lower layer and upper layer) hereinafter. SABAP_UMB are of two types: those assigned in the past 226 Bollettino della Società Paleontologica Italiana, 61 (2), 2022 have six digits (e.g., SABAP_UMB 129842), while those Description - SABAP_UMB 129842 (Fig. 3a) recently assigned are composed of an alphanumeric code preserves the root and a small part of the crown without such as SABAP_UMB 22.S241-3.263 (the complete list appreciable crenulations on the edges. The root is is provided in the SOM). In the manuscript the identifier moderately curved in mesiodistal direction. In mesial “S241-” is omitted, thus obtaining a code like SABAP_ view, the tooth appears overall stouter and only slightly UMB 22.3.263. compressed labiolingually. Identification of paleontological specimens is based on anatomical descriptions and morphological Comparisons and discussion - Sabertoothed felids comparative analysis. Biometric/statistical comparisons (subfamily Machairodontinae) represent a widespread are performed is some cases. In particular, for equids, group of top predators in the Plio-Pleistocene carnivoran bivariate plots and PCA were built in R Environment guilds of Europe and several other parts of the world v. 1.4.1103 (R Core Team, 2013), using the packages (Antón, 2013). The fragmentary nature of the specimen prcomp() v. 3.6.2 (Venables & Ripley, 2008) and ggplot2() from Vigna Nuova does not permit detailed comparisons. v. 3.3.3 (Wickham, 2016). PCA were performed on The absence/presence of crenulations along the mesial and third metatarsals, using the parameter scale = T to have distal edges of the crown is the more reliable criterium a unit variance before starting the analyses in R. The to differentiate Smilodontini and Homotherini, with the Log10 Ratio diagrams on third metatarsals are based latter normally showing clear crenulations. However, the on the log-transformed mean values of the Höwenegg absence of crenulations in the small preserved portion Hippotherium primigenium von Meyer, 1829 (Bernor of the crown of SABAP_UMB 129842 could be due et al., 1997). The Höwenegg H. primigenium sample to preservation issues. The morphology of the crown is represents a well-studied, homogeneous sample of a single another diagnostic character, being normally stouter and species of Equinae, for which the full range of descriptive less compressed in Homotherium and slenderer and more statistics (mean, standard deviation, confidence limits, and labiolingually compressed in Megantereon. Unfortunately, minimum, maximum, and median values) are available this character cannot be assessed too. The curvature of the for each bone, including the cranium, mandible, and teeth. root in SABAP_UMB 129842 is intermediate between All measurements are taken with digital calipers and that of Megantereon cultridens Cuvier, 1824 from are given with a precision of 0.1 mm. Measurements are Villarroya and that of Homotherium crenatidens Fabrini, provided as SOM. Sources of data used for comparisons 1890 from the Incarcal complex as can be seen in Fig. and measurement and anatomical abbreviations are 4a. The morphology of the tip of the root is not rounded indicated in the relative figure/table captions. as in Smilodontini, but more pointed as in Homotherium Anatomical and/or measurement nomenclature follows (Fig. 4a). However, the taxonomic value of this character Eisenmann et al. (1988) and Bernor et al. (1997) for is not discussed in the literature and we do not know its equids; Sala (1986), Masini (1989), and Duvernois (1990) variability other than in the samples we have selected for for bovids; Heintz (1970) for cervids; Fujita et al. (2010) comparison. and Cherin et al. (2018) for suids. To solve these issues, we built a violin plot of the ratio between labiolingual width and mesiodistal length Institutional abbreviations of the upper canine of Old World Homotherium and AUT: Aristotle University of Thessaloniki, Greece; Megantereon (Fig. 4b). Our results show that the studied IGF: Museo di Storia Naturale, Sezione di Geologia e specimen fully falls into the range of Megantereon species, Paleontologia, Università degli Studi di Firenze, Italy; IPS: although it is also close to the earliest representatives of Institut Català de Paleontologia Miquel Crusafont, Spain; the Homotherium lineage (i.e., Early Pliocene of Odessa LAMC: Natural History Museum of Los Angeles County, Catacombs), for which however, biometric variation is USA; MCL: Musée des Confluences de Lyon, France; virtually unknown due to poor record. Accordingly, we NHMB: Natural History Museum Basel, Switzerland; confidently refer the studied specimen to cf. Megantereon. SABAP_UMB: Soprintendenza Archeologia Belle Arti e Paesaggio dell’Umbria, Italy. Family Canidae Fischer von Waldheim, 1817 SYSTEMATIC PALEONTOLOGY Genus Canis Linnaeus, 1758 (LOWER LAYER) Class Mammalia Linnaeus, 1758 Canis sp. Order Carnivora Bowdich, 1821 (Fig. 3b-f) Family Felidae Fischer von Waldheim, 1817 Referred material - Articulated cervical vertebrae 3-7 Genus Megantereon Croizet & Jobert, 1828 (SABAP_UMB 129850); rib (SABAP_UMB 129875); thoracic vertebra 1 and rib fragments (SABAP_UMB 129877); articulated thoracic vertebrae 2-3 (SABAP_ cf. Megantereon UMB 129879). (Fig. 3a) Description - The canid material from Vigna Nuova Referred material - Fragmented right upper canine is represented only by elements of the axial skeleton, (SABAP_UMB 129842). all belonging to a single individual. The articulated B. Azzarà et alii - Mammals from Vigna Nuova, Perugia (Italy) 227 Fig 3 - Carnivorans from Vigna Nuova (Italy), lower layer. a) cf. Megantereon: right upper canine (SABAP_UMB 129842) in labial (a1) and distal (a2) views. b-d) Canis sp.: cervical vertebrae 3-7 (SABAP_UMB 129850) in dorsal view (b); left rib (SABAP_UMB 129875) in lateral view (c); thoracic vertebrae 2-3 (SABAP_UMB 129875) in lateral view (d). Scale bar: 2 cm. e-f) Comparison between the vertebrae C3-C4 from Vigna Nuova (SABAP_UMB 129850) (e) and those of Canis etruscus from Olivola (IGF 4523) (f) in left lateral view. Vertebrae are shown in anatomical connection, as they were discovered; the relative position of the vertebrae in the two specimens is slightly different (C4 is rotated clockwise with respect to C3 in SABAP_UMB 129850, while the two vertebrae are almost aligned in IGF 4523). Red asterisks indicate broken portions. The morphology of the posterior edge of C4 in SABAP_UMB 129850 is estimated (dotted line) because it is partially covered by C5. Scale bar: 2 cm. g) Mustelidae indet., left upper canine (SABAP_UMB 22.3.263) in mesial view. Scale bar: 1 cm. 228 Bollettino della Società Paleontologica Italiana, 61 (2), 2022 Fig. 4 - Comparisons between Machairodontinae upper canines. a) From left to right: Smilodon fatalis Leidy, 1868 from La Brea Tar Pits (LAMC Hc1142); Megantereon cultridens from Villarroya (IPS36785); M. cultridens from Villarroya (IPS36784); cf. Megantereon from Vigna Nuova (SABAP_UMB 129842; reversed); Homotherium crenatidens from Incarcal-I (IPS IN-I1542); H. crenatidens from Incarcal-I (IPS IN-I-421). Scale bar: 2 cm. b) Violin plot for the width/length ratio of the upper canine (measured at the cervix) in Pliocene to Middle Pleistocene Homotherium, M. cultridens, Megantereon adroveri Pons-Moyà, 1987, and the Vigna Nuova specimen. Epi: Epivillafranchian; EVil: Early Villafranchian; Gal: Galerian; MVil: Middle Villafranchian; LVil: Late Villafranchian;. Sources of data: Ballesio (1963); de Bonis (1976); Ficcarelli (1979); Hemmer (2001); Sardella & Iurino (2012). cervical vertebrae 3-7 (C3-7; SABAP_UMB 129850) is still visible, but half broken (Fig. 3d). The two most have retained their original shape, even if they miss some complete ribs (SABAP_UMB 129875 and SABAP_UMB portions (Fig. 3b). The spinous process is still present only 129877 partim; Fig. 3c) probably correspond to the first in C3, while the right transverse process is preserved only pair, as they perfectly articulate into the articular facets in C4-6. The vertebrae have a long and stout body, which on T1-T2. gradually became shorter and narrower towards C7 (see measurements in the SOM). The first thoracic vertebra Comparisons and discussion - The first occurrence (T1; SABAP_UMB 129877 partim) preserves only the of Canis in Europe has long been regarded as one of body and few fragments of the neural arch. T2 and T3 are the main biochronological markers of the onset of the still articulated (SABAP_UMB 129879) and in the former, Late Villafranchian, i.e., the so-called “wolf event” a slender, long, and posteriorly directed spinous process (Azzaroli, 1983; Azzaroli et al., 1988; Torre et al., 1992, B. Azzarà et alii - Mammals from Vigna Nuova, Perugia (Italy) 229 2001). Nevertheless, further discoveries such as those of absent curvature of the crown) of the left side (due to the Canis sp. in the Early Villafranchian of Vialette (France; position of furrows and keels), the field can be restricted Lacombat et al., 2008) and in the Middle Villafranchian to small-sized carnivorans (the preserved portion of the of Coste San Giacomo (Italy; Bellucci et al., 2012, crown is about 1 cm in height). We can exclude felids 2014), point to an earlier arrival of Canis in western and canids, characterized by upper canines with more Europe and support a model of diachronous dispersal marked curvature, very pointed apexes, and labiolingual across the continent (Sotnikova & Rook, 2010). Aside flattening (i.e., elliptical section). The greatest similarities from the aforementioned records of Canis sensu stricto are found with the mustelids, which instead have straight in the Early-Middle Villafranchian, several species are upper canines, with an approximately circular section and recognized in later times, including Canis etruscus Forsyth a complex of furrows corresponding, in relative position Major, 1877, C. arnensis Del Campana, 1913, and C. and development, to that of SABAP_UMB 22.3.263. The mosbachensis Soergel, 1928, among others (Cherin et al., overall size of the latter is approximately between that 2014; Bartolini-Lucenti & Rook, 2016; Bartolini-Lucenti of a marten and that of a badger, i.e., it approaches that et al., 2017, 2020 and references therein). of extinct forms such as Pannonictis (see Colombero et The C3 and C4 from Vigna Nuova show striking al., 2012). morphological resemblance with the same vertebrae of C. etruscus from Olivola (Italy; IGF 4523) figured by Torre (1967, pl. VIII, fig. 8). In turn, the same author stated that Order Proboscidea (Illiger, 1811) the postcranial morphology of C. etruscus does not differ significantly from that of the extant wolf Canis lupus Linnaeus, 1758, except in relatively smaller dimensions Proboscidea indet. (Torre, 1967, p. 131). The vertebrae from Vigna Nuova are extremely similar to those of Olivola, especially in the Referred material - Rib fragment (SABAP_UMB relative development of the pre- and postzygapophyses 22.3.264). with respect to the neural arch and body, in the inclination (ca. 45° anterodorsally to posteroventrally) of the anterior Description - The only available proboscidean material and posterior articular facets in lateral view, and in the is a small fragment of a rib. ventral projection of the posterior portion of the body relative to the anterior (i.e., the ventral edge of the body Comparisons and discussion - At least four is inclined posteroventrally in lateral view) (Fig. 3e-f). In proboscideans occur in the European Villafranchian addition to the morphological similarities, the vertebrae mammal assemblages: Mammut borsoni (Hays, are also very similar in size (e.g., the length of the body 1834), Anancus arvernensis (Croizet & Jobert, 1828), is 35.6 versus 34.7 mm in C3 and 31.4 versus 32.1 mm in Mammuthus rumanus (Stefanescu, 1924), and M. C4 in SABAP_UMB 129850 and IGF 4523, respectively). meridionalis (Nesti, 1825) (approximately in stratigraphic However, despite these similarities with C. etruscus, order, from older to younger). Due to the fragmentation the sample from Vigna Nuova does not offer sufficient of the available specimen, it is not possible to refer it to diagnostic characters for identification at the species rank a specific taxon. and is therefore referred to Canis sp. Order Perissodactyla Owen, 1848 Family Mustelidae Fischer von Waldheim, 1817 Family Equidae Gray, 1821 Genus Equus Linnaeus, 1758 Mustelidae indet. (Fig. 3g) Equus cf. senezensis Prat, 1964 Referred material - Crown of left upper canine (Fig. 5) (SABAP_UMB 22.3.263). Referred material - Articulated left metacarpals II-III- Description - An almost complete canine crown from IV, proximal part (SABAP_UMB 129840); left metacarpal the Vigna Nuova collection can be attributed to a small- III, proximal epiphysis (SABAP_UMB 129857); cervical sized carnivoran (Fig. 3g). The crown is straight and has vertebrae (SABAP_UMB 129867, SABAP_UMB a roughly circular section along all its height. The lingual 129874); left astragalus (SABAP_UMB 129869); wall is smooth, whereas the labial one shows a shallow mandible with left and right c1-i3 and left p2, divided into vertical furrow in distolabial position and another groove, two pieces (SABAP_UMB 129835 + 129837); partial left markedly deeper, in mesiolabial position. Neither of them hindlimb in anatomical connection with distal tibia, tarsus, reaches the apex of the crown. The latter is relatively metatarsals II-III-IV (SABAP_UMB 129856 + 129861 + rounded. A well-developed keel is present along the mesial 129862 + 129868). margin of the tooth and a weaker one along the distal one. Description - The mandible SABAP_UMB 129835 + Comparisons and discussion - The incompleteness of 129837 includes a well-preserved left p2 with medium- the specimen hinders any attempt at taxonomic attribution. high stage of wear. It shows a short and blunt anterostylid, Assuming that it is an upper canine (due to the virtually the metaconid is small and rounded, and the metastylid is 230 Bollettino della Società Paleontologica Italiana, 61 (2), 2022 B. Azzarà et alii - Mammals from Vigna Nuova, Perugia (Italy) 231 pointed distally. The linguaflexid is V-shaped, a feature et al., 1997, 1998; Azzaroli, 2000; Bernor et al., 2018, diagnostic of stenonian horses. The ectoflexid is very 2019; Rook et al., 2019; Cirilli et al., 2021a, b, c). This shallow. The preflexid is short whereas the postflexid is biochronological event represents the dispersal of the longer with simple ornamented morphology. This last monodactyl horses from North America into the Old feature indicates an advanced wearing stage. A small World, with the species Equus simplicidens Cope, 1892 pli caballinid is present on the mesial border of the as the most likely original pool for the Eurasian species. hypoconid. The anterior portion of the mandible exhibits a During the Gelasian, monodactyl equids diversified in slender symphysis with the left mandibular body partially different species such as Equus stenonis Cocchi, 1867, E. preserved. The lower incisors show a high degree of wear, major Depéret in Delafond & Depéret, 1893 ex Boule, E. with the infundibulum not always present (e.g., left i1). senezensis, and E. stehlini Azzaroli, 1964. Equus stenonis The lower canines are visible as small blunt enamel knobs represents the most widespread species during this time that emerge from the mandible about 1 cm behind the i3s. frame, with its first occurrence in the Early Pleistocene SABAP_UMB 129840 includes a left third metacarpal site of Saint Vallier (France, 2.45 Ma; dating in Nomade (MC3) preserving about half diaphysis and the proximal et al., 2014). This species dispersed in western Eurasia, epiphysis. It has a narrow appearance in anterior view, from Caucasus to the Iberian Peninsula, and diversified although not slender. The proximal epiphysis has a flat into different populations/ecomorphotypes (Palombo & articular surface for the magnum and large articular facets Alberdi, 2017; Cherin et al., 2021; Cirilli et al., 2021a). for the second and fourth metacarpals (MC2, MC4). The The distribution of E. major is still not well defined, but at latter are almost complete and preserved in anatomical the present time it seems mostly present in Central, North, connection. and Eastern Europe (Palombo & Alberdi, 2017). Cirilli The most complete specimen is the articulated left (2022) reported some new insights on the evolution and hindlimb SABAP_UMB 129856 + 129861 + 129862 + dispersals of the Gelasian medium-small sized horses. 129868. The tibia preserves the diaphysis and the distal Recent research on the Middle Villafranchian localities epiphysis, lacking the proximal one. Its general appearance of Senèze (France) and Coste San Giacomo (Italy) is rather bulky. Much information about the morphology have confirmed the presence of a medium-sized horse, of the distal epiphysis cannot be observed due to the intermediate between E. stenonis and E. stehlini (Palombo presence of the astragalus and calcaneum in anatomical et al., 2017; Cirilli et al., 2021a; Cirilli, 2022), identified connection, with some sediment in between. The latter as E. senezensis. The species was originally described as a bones are complete. The calcaneum has a short and robust subspecies of E. stenonis, namely E. stenonis senezensis, corpus, with a well-preserved massive tuber calcanei. The by Prat (1964, 1980) from the locality of Senèze (France, morphology of the other ankle elements (central, first + 2.2-2.0 Ma; dating in Nomade et al., 2014). Subsequently, second, third, and fourth tarsal bones) cannot be described Alberdi et al. (1998) questioned its subspecific rank, due to anatomical connection. The second, third, and suggesting a better attribution to the species level. This fourth metatarsals (MT2-4) are articulated and complete. interpretation was supported by the revision of the The MT3 appears robust, due to the reduced maximal European E. stenonis and E. stehlini samples undertaken length, the relatively large proximal and distal epiphyses, by Cirilli et al. (2021a) and Cirilli (2022), suggesting that and the wide circular diaphysis. The distal epiphysis has the medium-sized horse from Senèze truly represents a small medial and lateral protuberances, which are not different species during the Early Pleistocene. Beside larger than the medial and lateral trochleae. The latter are Senèze, E. senezensis has been reported also in Italy, round, with a well-developed crista sagittalis in between. from the Early Pleistocene localities of Coste San In posterior view, the long lines for the attachment of the Giacomo and Montecarlo (Upper Valdarno Basin). interosseus ligaments connecting the MT2 and MT4 are These sites have been dated at ca. 2.2-2.1 Ma (Ghinassi partially preserved. et al., 2005; Bellucci et al., 2012, 2014; Fidolini et al., SABAP_UMB 129869 is a left astragalus, with 2013; Florindo et al., 2021), hence similar in age with incomplete and weathered articular surface for the Senèze. The European medium-sized horses of the late calcaneum. The astragalus has a short and massive aspect, Early Pleistocene underwent a turnover at the Gelasian- with small medial and lateral trochleae and a large medial Calabrian boundary. Indeed, the first occurrence of Equus tuberculum tali for the attachment of the ligamentum altidens von Reichenau, 1915 has been recognized in the collaterale. The trochlear groove is large. paleoanthropological site of Dmanisi (Georgia; 1.85-1.76 Ma), predating the first Italian occurrences of Farneta Comparisons and discussion - In Europe, the Equus and Pirro Nord (Bernor et al., 2021). Equus altidens Datum is marked by the appearance of the large-sized would appear to be the most geographically widespread Equus livenzovensis Bajgusheva, 1978, occurring at species in the late Early Pleistocene, sharing its range the beginning of the Pleistocene (ca. 2.6 Ma) in the with the last representatives of E. stenonis and the earliest localities of Liventsovka (Russia), Montopoli (Italy), representatives of Equus suessenbornensis Wüst, 1900, Roca-Neyra (France), and El-Rincón 1 (Spain) (Alberdi and being part of an extensive faunal turnover occurring at Fig. 5 - Equus cf. senezensis from Vigna Nuova (Italy), lower layer. a) Left hindlimb with distal tibia, tarsus, metatarsals II-III-IV (SABAP_ UMB 129856 + 129861 + 129862 + 129868): specimen in anatomical connection, lateral view (a1); metatarsals in anterior (a2) and posterior (a2) views. b) Left astragalus (SABAP_UMB 129869) in anterior (b1) and medial (b2) views. c) Left metacarpals II-III-IV (SABAP_UMB 129840) in anterior (c1) and posterior (c2) views. d) Mandible (SABAP_UMB 129835 + 129837): incisor arcade in occlusal (d1) and ventral (d2) views; left corpus fragment with p2 in labial (d3) and occlusal (d4) views. Scale bar: 5 cm. 232 Bollettino della Società Paleontologica Italiana, 61 (2), 2022 ca. 1.8 Ma in Eastern Europe and slightly later in Central Gerakarou-1 (Mygdonia Basin, Greece) overlaps the and western Europe (Alberdi & Palombo, 2013; Palombo morphometric range of variation of E. stehlini and E. & Alberdi, 2017; Cirilli et al., 2020, 2021a; Bernor et al., altidens. 2021; Bartolini-Lucenti et al., 2022). More interesting results are depicted in the multivariate Morphometric comparisons on the Vigna Nuova equid analyses on the MT3s. Figure 6b shows the results for the remains are here provided based on the most relevant MT3 PCA, with PC1 and PC2 accounting for 85.4% of the elements, namely the p2 and the complete MT3. Figure 6a total variance (PC1 = 80.4%; PC2 = 5.0%). The loading plots the maximum width (M6) versus maximum length distribution is reported as an inset in Fig. 6b and in the (M1) for p2, comparing the Vigna Nuova specimen with SOM within the variance components. PC1 separates the European Early Pleistocene species. SABAP_UMB species by maximal length from negative to positive 129835 is included in the range of variation of E. values (less to more elongate), whereas PC2 denoted a senezensis from Senèze, and plots close to the largest slenderer morphology from positive to negative values. E. stehlini (Upper Valdarno Basin) and the smallest E. The Vigna Nuova specimen is included in the range of altidens (Dmanisi and Venta Micena). It is well separated variation of E. senezensis, distinct from the E. stehlini from the European E. stenonis, thus far highlighting sample from Upper Valdarno and close to the smallest its attribution to a medium-small sized species. Equus E. altidens individuals from Dmanisi. Equus senezensis suessenbornensis represents the largest species considered occupies an intermediate position between E. stenonis in the plot, whereas E. stenonis mygdoniensis from and E. stehlini, confirming the previous results obtained Fig. 6 - Biometric comparisons between the horse from Vigna Nuova and several Equus samples from the Plio-Pleistocene of Eurasia. a) Bivariate plot comparing the maximum width (M6) versus maximum length (M1) of the p2. b) PCA performed on ten measurements of the third metatarsal (M1, maximum length; M3, midshaft width; M4, depth of the diaphysis at level of M3; M5, proximal articular width; M6, proximal articular depth; M10, distal maximal maximum supra-articular width; M11, distal maximum articular width; M12, distal maximum keel depth; M13, distal maximum depth of the lateral condyle; M14, distal maximum depth of the medial condyle). c-d) Log10 ratio diagrams of third metatarsal measurements in European Gelasian (c) and Calabrian (d) equids. In addition to the measurements used in the PCA, this analysis includes M7 (maximum diameter of the articular facet for the third tarsal) and M8 (diameter for the anterior facet for the fourth tarsal). B. Azzarà et alii - Mammals from Vigna Nuova, Perugia (Italy) 233 by Cirilli et al. (2021b) and Cirilli (2022). Equus stehlini Referred material - Horn core fragment (SABAP_ is well separated from the rest of the comparative sample UMB 129851). and shows a narrow and massive MT3 morphology, with the largest E. stehlini specimens overlapping the smallest Description - The only bovid fossil from the E. senezensis. On the other hand, the medium-sized E. Vigna Nuova lower layer is a fragment of horn core. altidens shows a slenderer morphology, with a more Unfortunately, it is so heavily weathered and abraded that elongated maximal length and a reduced diaphysis. The the surface is no longer visible. Faint parallel furrows are Dmanisi and Pirro Nord populations overlap their range visible along one of the edges. The horn core is virtually of variation, whereas the sample from Venta Micena straight and does not show clear spiralization. (Spain) is somewhat larger. The Gerakarou-1 medium- sized E. stenonis mygdoniensis is included in the range Comparisons and discussion - Based on the absence shown by the European E. altidens samples. The European of curvature and spiralization, specimen SABAP_UMB E. stenonis samples (Saint Vallier, Olivola, Matassino, 129851 is here referred to cf. Leptobos (see detailed and Upper Valdarno Basin) overlap in their range of discussion in the section on the bovid remains from the variation, characterized by larger dimensions and more upper layer of Vigna Nuova). robust morphology than the medium-sized E. senezensis and E. altidens. Equus eisenmannae Qiu et al., 2004 from Longdan (China) and E. livenzovensis exhibit again Family Cervidae Goldfuss, 1820 longer MT3s when compared with E. simplicidens and E. stenonis, whereas E. major and E. suessenbornensis Genus Croizetoceros Heintz, 1970 remain the largest equid species from the European Early Pleistocene. These results are also confirmed in the Log10 ratio Croizetoceros ramosus (Croizet & Jobert, 1828) diagrams on MT3. Fig. 6c plots the mean values of (Fig. 7b) the European Gelasian Equus species, whereas Fig. 6d shows the Calabrian ones. Both diagrams include Referred material - Left maxilla with P2-M3 (SABAP_ the complete MT3 from Vigna Nuova (SABAP_UMB UMB 129863). 129856 + 129861 + 129862 + 129868 partim). As shown in Fig. 6c, the Vigna Nuova MT3 has a pattern close to Description - Specimen SABAP_UMB 129863 that of E. senezensis from Senèze, characterized by a belongs to an adult individual with teeth in moderate reduced maximum length (M1) and robust diaphysis, stage of wear. The upper premolars are relatively large proximal, and distal epiphyses (M3-M14). It is overall and similar in size to each other. In occlusal view, the P3 larger than that of E. stehlini (Upper Valdarno), whereas and especially the P2 stand out for the pointed lingual the incomplete MT3 from Montecarlo (Upper Valdarno margin. The P4 has a more squared lingual margin, with Basin) plots in between. The Vigna Nuova MT3 is a slight hint of molarization (i.e., a faint vertical groove smaller than that of E. stenonis, although showing a crosses the lingual wall of the tooth). On the labial side, similar pattern. Fig. 6d compares the Vigna Nuova MT3 the three premolars show pointed parastyle and metastyle with the E. altidens samples and E. suessenbornensis. and, between them, a strong pillar of the paracone, which Although SABAP_UMB 129856 + 129861 + 129862 + is mesially shifted and points forward. The upper molars 129868 partim shows similar dimensions in the proximal are similar to each other in overall morphology and exhibit and distal epiphyses than E. altidens, its M1-M3 pattern lingual cones with trapezoidal shape. The protoconal fold is much different form the E. altidens samples, which (pli protoconal) is noticeable in the M3, present but not show a slender morphology with a more elongated M1 marked in the M2, and not visible in the M1 due to the and reduced M3. SABAP_UMB 129856 + 129861 + more advanced wear. A sharp entostyle is developed in all 129862 + 129868 partim exhibits the same M3 values molars, while no lingual cingula are present. of E. altidens from Venta Micena, the largest among the E. altidens samples. As reported in the PCA, E. stenonis Comparisons and discussion - Croizetoceros is an mygdoniensis from Gerakarou-1 is included in the extinct cervid with uncertain phylogenetic relationships European E. altidens samples. and body size smaller than the Dama-like deer (see section Considering this evidence, we ascribe the medium- on Pseudodama below). The genus is reported in Europe sized equid from Vigna Nuova to E. cf. senezensis, since the Late Miocene with scanty remains (Azanza, recognizing their close similarities, especially in the 2000) and becomes a relatively common element of the postcranial elements. Early-Middle Villafranchian assemblages with the species C. ramosus, ranging from Spain to Greece, with different samples sometimes referred to different subspecies (Valli, Order Artiodactyla Owen, 1848 2004 and references therein). Croizetoceros sp. is reported Family Bovidae Gray, 1821 based on a single tooth from the Pliocene of Burdur (Turkey; Mayda et al., 2017). Genus Leptobos Rütimeyer, 1877-1878 The main dental features of C. ramosus are well described by Heintz (1970) on the basis of several samples from France and Spain, and resumed by Valli cf. Leptobos (2004) with reference to the richest European collection, (Fig. 7a) namely that of Saint Vallier (France). Most of the 234 Bollettino della Società Paleontologica Italiana, 61 (2), 2022 B. Azzarà et alii - Mammals from Vigna Nuova, Perugia (Italy) 235 Fig 7 - Artiodactyls from Vigna Nuova (Italy). a) cf. Leptobos: horn core fragment (SABAP_UMB 129851). b) Croizetoceros ramosus: maxilla with P2-M3 (SABAP_UMB 129863) in occlusal (b1) and labial (b2) views. c-g) Pseudodama sp.: right hemimandible with p2-m3 (SABAP_UMB 129848) in occlusal (c1) and labial (c2) views; left hemimandible with dp4 (fragmented), m1-m2 (SABAP_UMB 129865 + 129860) in occlusal (d1) and labial (d2) views; left frontal fragment with antler basis (SABAP_UMB 129845) in medial (e1) and anterior (e2) views; right antler fragment (SABAP_UMB 129880) in lateral (f1) and anterior (f2) views; right M1 (SABAP_UMB 129870) in occlusal view (g). h) Cervidae indet. (large size): left maxilla with dP2-M2 (SABAP_UMB 129866) in occlusal (h1) and labial (h2) views. i) Sus cf. strozzii: maxilla with P4-M2 and M3 fragment (SABAP_UMB 129847) in occlusal view. Scale bar: 2 cm. recognized features can be found also in the Vigna Nuova same applies to the ratio between the premolar and molar specimen, including the asymmetrical development of length (LP/LM), which is 0.70 in SABAP_UMB 129863 the strong pillar of the paracone in the premolars, the and between 0.69 and 0.79 in C. ramosus from several presence of the protoconal fold in the molars, the relative localities (Heintz, 1970; Kostopoulos, 1996; Kostopoulos development of the styles on the labial wall of the molars. & Athanassiou, 2005). In particular, the maxilla from Vigna Nuova shows The overall size of SABAP_UMB 129863 is also striking morphological similarities with some known similar to the average values of small-sized cervids of specimens of C. ramosus such as NHMB Prr.346 from the roe deer lineage (i.e., Procapreolus, Capreolus). the type locality of Les Etouaires (France), MCL QSV- However, these latter taxa are well recognizable in the 766 from Saint Vallier, AUT GER-188 from Gerakarou more articulated occlusal pattern of the upper teeth (e.g., (Greece), IPS28245/740b from La Puebla de Valverde numerous enamel folds that develop into the fossae), (Spain) (see Fig. 8 for comparisons). Dimensionally, the unlike the relatively simple morphology observed in Vigna Nuova specimen falls into the lower part of the SABAP_UMB 129863. Moreover, roe deer have more range of C. ramosus, approaching in particular to the elongated premolar series with respect to molars. For material from Gerakarou (measurements in the SOM). instance, the average LP/LM is 0.92 in Procapreolus However, it must be noted that the apparently larger cusanus from Les Etouaires (Heintz, 1970), 0.89 in size of the specimens from the French localities of Saint Capreolus cusanoides from Untermassfeld (Germany) Vallier, Pardines, and Les Etouaires could be due to a and 0.84 in Capreolus suessenbornensis from Süssenborn slightly different measuring technique (see SOM). The (Germany) (personal data). The above lines of evidence allow us to confidently refer the Vigna Nuova maxilla to C. ramosus, for which it represents one of the most complete and better-preserved specimens in Italy. Genus Pseudodama Azzaroli, 1992 Pseudodama sp. (Fig. 7c-g) Referred material - Left i2 (SABAP_UMB 129841); right frontal bone fragment with basal antler (SABAP_ UMB 129843); left frontal bone fragment with basal antler (SABAP_UMB 129844); left frontal bone fragment with basal antler (SABAP_UMB 129845); right frontal bone fragment with basal antler (SABAP_UMB 129846); right hemimandible with p3-m3 (SABAP_UMB 129848); antler fragments (SABAP_UMB 129849, SABAP_UMB 129853, SABAP_UMB 129881); left hemimandible fragment with dp4 (broken), m1, and m2 (SABAP_UMB 129860 + 129865), right M1 (SABAP_UMB 129870); broken left m3 (SABAP_UMB 129871); left i3 (SABAP_ UMB 129872); left i1 (SABAP_UMB 129873); right antler fragment (SABAP_UMB 129880). Description - Fossils of a medium-sized deer are relatively common in the Vigna Nuova collection. The Fig. 8 - Comparisons between Croizetoceros ramosus upper antler remains SABAP_UMB 129843, SABAP_UMB tooth rows. a) SABAP_UMB 129863 from Vigna Nuova. b) 129844, SABAP_UMB 129845, and SABAP_UMB NHMB Prr.346 from Les Etouaires (France; reversed). c) MCL QSV-766 from Saint Vallier (France). d) AUT GER-188 from 129846 are quite similar to each other both in shape and Gerakarou (Greece). e) IPS28245/740b from La Puebla de Valverde dimensions (measurements in the SOM). All show a basal (Spain; reversed). The fossils are drawn not to scale to highlight portion of the beam with an almost circular section and the morphological similarities. insertion of the first tine (unfortunately never preserved) 236 Bollettino della Società Paleontologica Italiana, 61 (2), 2022 at a certain distance above the burr. The pedicles are also attribution at the species rank, for which more complete roughly circular in section (measurements in the SOM). craniodental remains (especially antlers) would have been The most complete specimen (SABAP_UMB 129880) needed. The antlers from Vigna Nuova differ from those is a long fragment of right antler with basis of the basal of derived species of Pseudodama such as P. farnetensis tine, but missing the burr. The basal tine branches at and P. vallonnetensis, in which the basal tine is normally about 3 cm from the bottom edge of the antler. Above the inserted very close to or even leaning against the burr, as basal tine, the beam runs roughly vertical, with a slight in the extant Dama. Similarly, the sub-vertical morphology lateral curvature in the first part and a medial curvature of the best-preserved antler (SABAP_UMB 129880) also in the second. recalls the earlier forms of Pseudodama rather than the The only upper tooth is the isolated right M1 later ones, in which the antlers tend to diverge greatly SABAP_UMB 129870. It is virtually unworn and shows posteriorly and laterally. However, these features must mesiodistally compressed lingual cones, between which be considered with thoughtfulness because they can also a small entostyle develops. Mesiolingual, lingual, and be subject to ontogenetic variation. distolingual cingula are present but weak. The labial Other primitive characters of the Dama-like deer from styles are instead very prominent, as are the pillars of Vigna Nuova lie in the teeth and include the development the labial cones. of labial cones and styles and lingual cingulum in the upper The right hemimandible SABAP_UMB 129848 bears molar (SABAP_UMB 129870), enlarged i1 (SABAP_ the complete cheek tooth row, but in a very advanced UMB 129873), and un-molarized p4 (SABAP_UMB stage of wear (the m1 is almost completely worn 129848) (Azzaroli, 1947, 1992; Croitor, 2006; Pfeiffer, out). This prevents a detailed description of the dental 2016). Another primitive condition is the high ratio morphology. In occlusal view, the lingual wall of the between the lower premolar and molar lengths measured paraconid is open, that is, the paraconid and metaconid in SABAP_UMB 129848 (Lp/Lm = 0.69), which exceeds are not fused (i.e., “unmolarized” condition). Lingual the average values available for P. lyra from Montopoli cingula and a faint ectostylid are still visible in the m2 (Italy; Lp/Lm = 0.64, n = 2), P. rhenana from Saint Vallier and m3, but not in the m1 due to wear. The lower molar (Lp/Lm = 0.68, n = 18) and Senèze (Lp/Lm = 0.65, n = morphology is better recognizable in the hemimandible 11), P. nestii from Pantalla (Italy; Lp/Lm = 0.68, n = 3), fragment SABAP_UMB 129865 + 129860, bearing a Olivola (Italy; Lp/Lm = 0.62, n = 5) and Upper Valdarno fragmented dp4 and complete m1 and m2. Both molars (Italy; Lp/Lm = 0.63, n = 16). On the contrary, later forms show a mesial cingulum and a prominent ectostylid in of Pseudodama show relatively shorter premolar rows, as labial view. can be seen in P. farnetensis from Pietrafitta (Italy; Lp/Lm The lower incisors SABAP_UMB 129841, SABAP_ = 0.63; n = 5) and Pirro Nord (Italy; Lp/Lm = 0.58; n = UMB 129872, and SABAP_UMB 129873 may belong to 2) and P. vallonnetensis from Untermassfeld (Germany; the same individual based on preservation, relative size, Lp/Lm = 0.59; n = 21) and Vallparadís Estació (Spain; and absent wear. Lp/Lm = 0.57; n = 9) (Breda, 2015; Cherin et al., 2022; this paper). Comparisons and discussion - Pliocene and In summary, the material of Pseudodama of Vigna Pleistocene cervids with overall morphology and size Nuova is suggested to belong to a basal species of the similar to the extant fallow deer are known as Dama-like genus (e.g., P. lyra, P. rhenana), but the incompleteness deer. Villafranchian and Epivillafranchian forms from of the specimens requires a cautious attribution to Europe are characterized by un-palmated antlers with Pseudodama sp. three or four points. These forms have been allocated into a variety of genera (e.g., Cervus, Dama, Axis, Rusa, Metacervocerus, Praeelaphus, etc.) and species, whose Cervidae indet. relationships with each other and with present-day species (Fig. 7h) are not clear (see a summary in Cherin et al., 2019a). Azzaroli (1992) proposed to refer all these species to the Referred material - Left maxilla with dP2-M2 genus Pseudodama, including the “Italian” lineage - with (SABAP_UMB 129866). the species Pseudodama lyra Azzaroli, 1992, P. nestii (Azzaroli, 1947), and P. farnetensis Azzaroli, 1992 - and Description - Specimen SABAP_UMB 129866 is the “French” lineage - with the species P. pardinensis particularly interesting. It is a left maxillary fragment of (Croizet & Jobert, 1828), P. rhenana (Dubois, 1904), a large-sized deer with deciduous premolars dP2-dP4, and P. perolensis (Bout & Azzaroli, 1952). Although slightly worn M1, and freshly erupted M2. The dP2 we believe that some of the aforementioned species is heavily worn and shows a distal lobe which is only could probably be synonymized, we follow Azzaroli’s slightly wider than the mesial one. Labially, the parastyle, concept, recognizing significant similarities between paracone, and metastyle are pointed, while the metacone them all (Breda & Lister, 2013; Breda, 2015), but also does not form a pillar on the labial wall. In the dP3, the including the Epivillafranchian P. vallonnetensis (de distal lobe is significantly wider and shorter than the Lumley et al., 1988) (Breda et al., 2015, 2020; Cherin et mesial one. On the labial wall, the pillar of the paracone al., 2019a, 2022). and the mesostyle are particularly prominent and are The morphology and size of the identifiable material oriented mesially. A similar morphology characterizes of middle-sized deer from Vigna Nuova are clearly the labial wall of the dP4, in which, however, the two akin to the genus Pseudodama. On the other hand, the lingual cones are almost equally developed and constricted available fossils do not retain characters that allow for an mesiodistally. In the M1, a low and rounded entostyle B. Azzarà et alii - Mammals from Vigna Nuova, Perugia (Italy) 237 develops between the protocone and hypocone. The protocone is pointed, tiny and slightly visible. The labial parastyle and mesostyle are very strong and stretch cingulum of the mesial lobe is broken but it is possible in labial direction. The M2 looks like a slightly larger to recognize that a hypoectoconid was present. The version of the M1, although the entostyle is replaced by transverse valley between the paracone and metacone is a lingual cingulum. In all teeth, the enamel on the lingual wide lingually and ends exactly halfway through the tooth; walls is rough. no tetrapreconule is visible. The tetracone is central and rectangular. In all the teeth no styles are visible in lingual Comparisons and discussion - The most common large- and labial view. sized deer occurring in the Villafranchian is Eucladoceros, No descriptions can be provided for SABAP_UMB characterized by spectacular comb-like antlers (Azzaroli 129852 as the specimen preserves only a distal splinter & Mazza, 1992; Croitor, 2018). Nevertheless, other of the M3, which however is dimensionally comparable groups of cervids of comparable size can be found in the to the corresponding tooth of SABAP_UMB 129847 same time interval, although their record is poorer. These and shows a similarly high enamel thickness. Based on include the enigmatic Arvernoceros, whose systematic preservation features, it is likely that the two specimens status and chronological occurrence are debated (Heintz, belonged to the same individual. 1970; Petronio & Pandolfi, 2011; Croitor, 2018). The genus Praemegaceros seems to replace Eucladoceros, at Comparisons and discussion - The genus Sus, of least in southwestern Europe, in the latest Villafranchian putative Asian origin, spread rapidly towards Europe since (Abbazzi, 2004). Unfortunately, the diagnostic characters the Pliocene and became a common element in the Early of all these forms reside mainly in the antlers, of which we Pleistocene faunal assemblages throughout the continent have no evidence in Vigna Nuova. No peculiar character (Cherin et al., 2020). The earliest member of this group can be identified in the upper deciduous or permanent is Sus arvernensis (Croizet & Jobert, 1828), occurring in dentition, neither in shape nor in size. For this reason, the Eurasia during the Ruscinian and Early Villafranchian. At large-sized deer of Vigna Nuova remains unidentifiable the beginning of the Early Pleistocene, this species was below the family level. replaced by the larger-sized S. strozzii. The latter survived in Europe at least until the end of the Early Pleistocene, although its presence during the latest Villafranchian (i.e., Family Suidae Gray, 1821 ca. 1.7-1.3 Ma) is yet not confirmed (see the so-called “suid gap” hypothesis by Martinez-Navarro et al., 2015) Genus Sus Linnaeus, 1758 (Cherin et al., 2018, 2020). Starting from the Middle Pleistocene, the wild boar Sus scrofa Linnaeus, 1758 replaced S. strozzii rapidly expanding throughout Eurasia Sus cf. strozzii Forsyth Major, 1881 up to the present day (Cherin et al., 2020; Iannucci et al., (Fig. 7i) 2020). The distinction between the closely-related S. Referred material - Fragmented right maxilla with arvernensis and S. strozzii on one side and S. scrofa on P4-M2 and broken M3 (SABAP_UMB 129847) and the other is easy when lower canines are available, even fragmented left maxilla with broken M3 (SABAP_UMB in the absence of complete skulls. This is because the first 129852). two species show a “verrucosic” morphology of lower canines, as opposed to the “scrofic” condition of S. scrofa Description - SABAP_UMB 129847 shows teeth (see Cherin et al., 2020). The lack of lower canines in the with an advanced stage of wear that prevents a detailed Vigna Nuova collection leads to rely only on biometric description of the occlusal pattern. However, wear makes considerations (also because the advanced state of wear of it possible to appreciate the considerable thickness of the teeth in SABAP_UMB 129847 prevents any detailed the enamel in all preserved teeth. The P4 has a squarish comparisons in dental morphology). The box-plots in Fig. occlusal outline, but appears slightly wider than long. A 9 clearly show that the Pliocene S. arvernensis can be mesial cingulum and a distal cingulum are visible. The distinguished from S. strozzii and S. scrofa by its smaller mesial one has a slight constriction in the middle. A hinted size (especially molar length). SABAP_UMB 129847 protoprestyle is visible. The protofossa is narrow, long has relatively large-sized teeth, which fall in the ranges and positioned lingually in the middle-mesial part of the of these two latter species. Although in the literature there tooth. The M1 is very worn (wear stage: 16, wear group: are several works in which an attempt has been made to 4; Zeder & Lemoine, 2020), so much that the main cusps recognize S. strozzii and S. scrofa on a dimensional basis are unrecognizable. The tooth has a figure-eight shape in (e.g., Ambrosetti et al., 1979; Faure & Guérin, 1983; occlusal view, with the mesial lobe smaller than the distal Van der Made, 1999; Van der Made et al., 2017), Cherin one and a labial constriction between them. The M2 is et al. (2020) showed that this approach alone cannot be similar in shape to the M1, but larger. Also in this molar, sufficient, especially when large samples are considered none of the four main cusps can be described in detail due both geographically and chronologically (the wild boar to the state of wear (wear stage: 12, wear group: 2; Zeder & shows enormous dimensional variation over time and Lemoine, 2020). However, the difference in width between space). As far as current knowledge is concerned, only the mesial and distal lobe is more marked than in the M1. the dimensions of the dp4 can be used for a statistically The mesial cingulum is flat and does not seem to bear a significant distinction between S. strozzii and S. scrofa protopreconule. The oval-shaped paracone is narrow and (Iannucci et al., 2020). In this context, the length of the in contact with the lingual cingulum. The residue of the teeth of SABAP_UMB 129847 alone is not decisive 238 Bollettino della Società Paleontologica Italiana, 61 (2), 2022 resembles S. strozzii rather than S. scrofa, which instead has a squarer P4. This evidence, associated with the very thick enamel possessed by the cheek teeth (i.e., one of the diagnostic characters of S. strozzii; Cherin et al., 2018), allows to refer the Vigna Nuova material to S. cf. strozzii. SYSTEMATIC PALEONTOLOGY (UPPER LAYER) Order Artiodactyla Owen, 1848 Family Bovidae Gray, 1821 Genus Leptobos Rütimeyer, 1877-1878 cf. Leptobos (Fig. 10) Referred material - Left hemimandible with p4-m3 and roots of p2-p3 (SABAP_UMB 129836); right metacarpal (SABAP_UMB 129838); atlas (SABAP_UMB 129839); intermediate phalanx (SABAP_UMB 129854); left P2 (SABAP_UMB 129855); left tibia, proximal epiphysis (SABAP_UMB 129858); left tibia diaphysis (SABAP_ UMB 129876); humerus distal diaphysis (SABAP_UMB 129878); horn core fragments (SABAP_UMB 22.3.248- 22.3.252, 22.3.267); distal phalanx fragments (SABAP_ UMB 22.3.256, SABAP_UMB 22.3.266); cranium fragment (SABAP_UMB 22.3.265); left medial malleolus fragment (SABAP_UMB 22.3.268). Description - An incomplete hemimandible and an isolated P2 were recovered from the Vigna Nuova upper layer. The mandible (SABAP_UMB 129836) includes the molars and the p4 with root fragments of p2-p3. The teeth are all heavily worn. In the p4, the parastylid and paraconid are separated by a shallow furrow, the protoconid is quite developed. The lingual valley between the paraconid and protoconid strongly indents mesially. The metaconid is large and has the shape of a blade, emerging lingually and strongly curving distally. The furrow separating the metaconid and entoconid is quite deep and narrow due to the distal shift of the former. The entostylid and entoconid are completely fused. The distal labial groove separating the protoconid from the hypoconid is strong. The molars are mesiodistally elongated with well-separated lobes. The parastylid is well developed and mesially projecting. The protoconid and hypoconid are morphologically similar except for a slight mesiodistal constriction of the latter. The ectostylid emerges mesially from the distal lobe and lies almost attached to the protoconid. The entostylid is smaller than the parastylid and projects lingually. The inner enamel Fig. 9 - Boxplots of upper tooth measurements (mm) comparing the suid from Vigna Nuova with Sus arvernensis, S. strozzii, and islets are crescent shaped. The mesial one is characterized S. scrofa. Boxplots show minimum, median, and maximum values by a constriction in the mid part. The islet of the distal with 25th and 75th percentile of each sample. L: length; W: width. lobe in m3 is marked by a bubaline fold (sensu Masini, 1989) in the distal portion. The cement is present in all the teeth, on both sides. The P2 SABAP_UMB 129855 has for the taxonomic identification of the specimen, while an advanced stage of wear. The paracone and parastyle considerations on their proportions can be. In particular, lie almost attached, separated by a shallow furrow that the P4 exhibits a relatively low length-to-width ratio (i.e., reaches the base of the tooth. The metacone is strongly it is more rectangular in occlusal view; Fig. 9). In this, it reduced. The metastyle projects distally. B. Azzarà et alii - Mammals from Vigna Nuova, Perugia (Italy) 239 Several horn core fragments are part of the collection. is divided by the sagittal crest into two facets for the Most of them present nutrient foramina and/or furrows on articulation with the proximal phalanx (abaxial and axial the outer side (e.g., SABAP_UMB 22.3.248 and SABAP_ glenoid cavities). Posteriorly to the crest there is a deep UMB 22.3.249) and clear sign of pneumatization in the U-shaped furrow. The abaxial cavity is slightly larger inner portion (e.g., SABAP_UMB 22.3.248 and SABAP_ than the axial. The abaxial tubercle and the axial pit are UMB 22.3.251). Despite their scanty preservation, it is visible although not well developed. The distal articular possible to state that all of the remains are characterized surface is subtriangular. The abaxial lobe has a pointed tip by a quite low degree of curvature and do not present developing toward the diaphysis of the phalanx. evidences of spiralization. The other available specimens are heavily fragmented The atlas SABAP_UMB 129839 preserves most of the and do not allow for any detailed description. body and the left wing. Dorsally, the atlas is characterized by two large openings, on each wing, shifted towards the Comparisons and discussion - The largest forms of anterior portion of the bone, in which two foramina are bovids are part of the subfamily Bovinae which, during located. The left wing projects posteriorly but not far from the earliest Pleistocene of Europe, was mainly represented the posterior zygapophyses. The dorsal tubercle and the by the genus Leptobos. This extinct taxon was populating dorsal portion of the neural arch are missing. The alar most of the subtropical and temperate areas of Eurasia, fossae, visible on the ventral side of the wings, are large spanning from the Iberian Peninsula to China, from and relatively deep depressions. The posterior tubercle is Northern India to, possibly, the British Islands (Cherin partially preserved and not well developed. The anterior et al., 2019b). Although Leptobos has been matter of articular surface is almost complete and extends on interest in numerous works (Merla, 1939; Pilgrim, 1939; the ventral side. The area connecting the two posterior Masini, 1989; Duvernois, 1990; Cherin et al., 2019b, zygapophyses is broad and flat. among others), there is still no consensus about its The only complete metacarpal (SABAP_UMB systematics and evolutionary history. Most of the scholars 129838) is relatively slender. It has an hourglass shape, divide European Leptobos into two lineages/groups or with the two epiphyses having approximately the same subgenera (Masini, 1989; Duvernois, 1990, 1992; Masini width, mediolaterally wider than the diaphysis. The et al., 2013; Cherin et al., 2019b; Sorbelli et al., 2021). proximal epiphysis, seen from above, is D-shaped; the The first includes the species Leptobos stenometopon anterior margin is sub-rounded whereas the posterior one (Sismonda, 1846), L. elatus (Pomel, 1853), L. merlai De is straight. The lateral articular facet is triangular and much Giuli (1987), and L. furtivus Duvernois (1989) (called smaller than the medial, which is sub-quadrangular with Leptobos ex gr. SEM hereinafter) constrained to the rounded anterior margin. The two facets are separated Early-Middle Villafranchian of Central and Western by a small crest perpendicular to the major axis of the Europe; the second includes L. etruscus (Falconer, 1868) articulation. The crest ends posteriorly in a deep, narrow and L. vallisarni Merla (1949) (called Leptobos ex gr. groove which hosts the proximal nutrient foramen. Another EV hereinafter), limited to the Late Villafranchian of foramen is present distally to the proximal end, connected Europe and, as concerning the latter species, Asia (Masini with the former through a very narrow furrow. All the et al., 2013; Tong et al., 2016; Sorbelli et al., 2021). anterior border of the proximal epiphysis is characterized Species of Leptobos ex gr. SEM are characterized by by a wrinkled tuberosity. The diaphysis has its minimum elongated neurocranium, “taurine” horns (i.e., laterally mediolateral width slightly above the mid-point. There emerging, with strong anticlockwise spiralization), are two shallow grooves in the proximo-anterior portion and slender limbs, whereas species of Leptobos ex gr. of the diaphysis. The anterior vascular groove at the distal EV share shorter postcornual portion of the cranium, shaft is narrow and shallow, getting deeper and marked posterolaterally emerging horns with single curvature, by sharp edges distally. The ovoidal foramen is located and stouter limb proportions. During the last stages of just above the distal end of the vascular groove, covered the Villafranchian, a new group of large bovids reached by sediment. The mediolateral width at the contact area Europe from Asia. These bovines represented the first between the diaphysis and the distal epiphysis is slightly western dispersal of Bison which, most probably, evolved smaller than that across the trochleae. The intertrochlear in mainland Asia from a derivate stock of Leptobos at the margins as well as the crests are subparallel. The lateral end of the Pliocene (Sorbelli et al., 2021 and references and medial trochlear pits are deep and marked by faint therein). These forms of large and stout bovids, grouped radial rugosities. The lateral abaxial hemicondyle is in the subgenus Bison (Eobison), were well adapted to anteroposteriorly thinner than the medial one. the harshened environmental condition that started to The tibia remains include a fragment of the diaphysis affect Europe during the Late Villafranchian and rapidly (SABAP_UMB 129876) and a partial proximal epiphysis replaced Leptobos in all the herbivore guilds of the (SABAP_UMB 129858). The articulation surface of the continent. With the faunal turnover of the Early-Middle latter is composed by the two large condyles. Neither of Pleistocene Transition (i.e., during the Epivillafranchian), these elements are complete. The proximally projected Eobison, in turn, was replaced by the “true” Bison, which intercondylar eminences are partially preserved. The will be the dominant large bovids in all the Holarctic diaphysis fragment represents the proximal half of the realm until the Holocene (Sorbelli et al., 2021). tibia due to the visible crest running along one side of Due to their general morphology and size, the sample the shaft. from Vigna Nuova is referable to a large bovid. As already The intermediate phalanx SABAP_UMB 129854 is remarked, during the Early Pleistocene, the only European complete and well preserved. The bone is stout, having representatives of this group are Leptobos and Bison a short and thick diaphysis. The proximal articulation (Eobison). The proportions of metapodials, in particular 240 Bollettino della Società Paleontologica Italiana, 61 (2), 2022 B. Azzarà et alii - Mammals from Vigna Nuova, Perugia (Italy) 241 Fig. 10 - cf. Leptobos from Vigna Nuova (Italy), upper layer. a) Left hemimandible with p4-m3 (SABAP_UMB 129836) in labial (a1) and occlusal (a2) views. b) Left P2 (SABAP_UMB 129855) in occlusal view. c) Atlas (SABAP_UMB 129839) in ventral (c1) and dorsal (c2) views. d) Right metacarpal (SABAP_UMB 129838) in proximal (d1) and anterior (d2) views. e) Horn core fragment (SABAP_UMB 22.3.248). f) Horn core fragment (SABAP_UMB 22.3.249). g) Intermediate phalanx (SABAP_UMB 129854) in abaxial (g1) and axial (g2) views. Scale bar: 5 cm. metacarpals, are a useful taxonomic tool in order to elements or a large sample of postcranial remains, it is discriminate between the various species belonging to virtually impossible to discern the two taxa. these genera (Maniakas & Kostopoulos, 2017; Sorbelli The lower dentition of Leptobos and Bison are quite et al., 2021). However, due to the high intraspecific similar, except for the evident increase of hypsodonty and variation that characterizes bovids, a cautious approach larger average size in the latter. The morphology of the is recommended when dealing with small samples, as in p4 has been used as tool in order to differentiate the two the Vigna Nuova case. The single metacarpal from this groups of Leptobos (Masini, 1989; Duvernois, 1990). The locality shows a relatively slender built, with a long and distally projected metaconid, the strong hypoconid, and thin diaphysis. In order to assess the robusticity of this the deep distolabial groove are characters shared between element, a biplot diagram comparing the total length of the Vigna Nuova and Leptobos ex gr. EV. Nonetheless, the metacarpal and the ratio between the transversal width several scholars underlined the strong variability affecting and the length is provided (Fig. 11). The Vigna Nuova the tooth morphology of large bovids, remarking how specimen falls within the range of the largest Leptobos, taxonomic inferences based on dentognathic remains in the area overlapping with Eobison and the slenderest should be taken cautiously (Skinner & Kaisen, 1945; specimens of Bison schoetensacki Freudenberg, 1914. Olsen, 1990; Sher, 1997, among others). The most imponent and robust species of Leptobos are The horn core morphology is one of the most useful those belonging to the EV group. While the postcranial tools for taxonomic purposes. Leptobos ex gr. SEM skeleton of L. etruscus is well known, that of L. vallisarni features long, spiralized horns emerging posterolaterally. is still poorly represented in the fossil record. Moreover, On the contrary, Leptobos ex gr. EV shows crescent moon- few limb bones referred to L. vallisarni are quite similar shaped cores with no or little torsion, emerging in a more to those of L. etruscus, thus, without complete cranial posterior position than in the former group (i.e., smaller Fig. 11 - Bivariate plot comparing the maximum length (Lmax; in mm) versus the ratio between the distal end width (DEW) and maximum length of the metacarpal in several samples of Eurasian bovines. The sample of Leptobos etruscus includes data from Faella/Matassino and Olivola (Italy), Senèze (France), and Fonelas P-1 (Spain). The sample of Leptobos vallisarni includes data from Upper Valdarno and Farneta (Italy). The sample of Leptobos ex gr. SEM includes the following sub-samples: Leptobos cf. stenometopon from Montopoli (Italy); Leptobos stenometopon/elatus from Upper Valdarno (Italy; several sites); Leptobos elatus from Les Etouaires (France) and Villarroya (Spain); Leptobos merlai from Saint Vallier (France); Leptobos cf. furtivus from Senèze and Olivola; Leptobos aff. furtivus from Upper Valdarno and Carsoli (Italy). The sample of Eobison includes the following sub-samples: Eobison sp. from Venta Micena (Spain), Eobison degiulii from Pirro Nord (Italy); Eobison cf. degiulii from Capena (Italy) and Mygdonia Basin (Greece); Eobison georgicus (Burchak-Abramovich & Vekua, 1994) from Dmanisi (Georgia); Eobison palaeosinensis Teilhard de Chardin & Piveteau, 1930 from Nihewan Basin (China). The sample of Bison menneri includes data from Untermassfeld (Germany). The sample of Bison schoetensacki includes data from Süssenborn and Mauer (Germany), Vallparadís composite section (Spain), Le Vallonnet, Le Vassirie, and Durfort (France). Source of data: Sorbelli et al. (2021), and references therein. 242 Bollettino della Società Paleontologica Italiana, 61 (2), 2022 angle between the sagittal plane and the core’s mid line) species in Vigna Nuova provides a new support of the and gently curving inwards. Eobison is characterized by dispersion of this species in the Italian Peninsula, which relatively small and stout horns, projecting laterally with may have been the source for the origin of E. stehlini strong upward bending and slight anticlockwise torsion. (Cirilli, 2022). The horn core fragments from Vigna Nuova show a low Some primitive characters of the sample of degree of bending and no sign of torsions, as in Leptobos Pseudodama from Vigna Nuova suggest that it may ex gr. EV. Other skeletal elements from Vigna Nuova be referred to a primitive species of the genus, thus do not bear/preserve diagnostic characters and are not supporting the attribution of the assemblage to the Middle discussed furtherly. Villafranchian. On the other hand, this hypothesis can Despite their scantiness, the analyzed elements only be confirmed by the discovery of more complete evidence that the bovid from the upper layer of Vigna fossils. Nuova has affinities with Leptobos ex gr. EV. However, Finally, the identification of the remains of Canis due to the limited number and preservation of the fossils sp. is noteworthy. Although these predators probably and the strong variation that characterizes this group of arrived in Europe already in the Early Villafranchian as large bovids, we prefer to cautiously refer the sample to evidenced by the Vialette record (Lacombat et al., 2008; cf. Leptobos pending for new material from the locality. Sotnikova & Rook, 2010), there is no trace of them in Italy before the second part of the Middle Villafranchian. Indeed, the record of Canis sp. from Coste San Giacomo CONCLUSIONS (Bellucci et al., 2021, 2014) marks the first occurrence of the genus Canis in Italy, which is confirmed here by the The fossil mammals from Vigna Nuova, collected Vigna Nuova fossils. more than thirty years ago, have revealed new extremely The recognition of a Middle Villafranchian assemblage intriguing data. First of all, the combination of taphonomic, in Umbria is novel information, given that the majority historical, and, to a lesser extent, geological data made of the other LFAs in the region, especially those from it possible to divide the sample into two different the Tiberino Basin, have been attributed to the Late assemblages. The one coming from what we have Villafranchian (see Cherin et al., 2019a, 2022 for a called the “upper layer” of Vigna Nuova (i.e., the only summary). The only exception is represented by the portion still visible in the outcrop), is made up of 18 small assemblage from the upper layers of Cava Toppetti identifiable remains, plus about 25 unidentifiable ones. near Todi (Perugia), which was referred to the Coste The identifiable material is all assigned to cf. Leptobos San Giacomo FU (Abbazzi et al., 1997). However, in in the light of morphological affinities with this genus of our opinion, the sample is too small and the fossils Villafranchian bovines. In particular, especially as regards too fragmentary to confirm such a biochronological the dimensions of the metacarpal (Fig. 11), the remains of conclusion. Vigna Nuova upper layer are similar to those of the most Our chronological results also open up stimulating derived forms of the group, namely L. etruscus and L. research prospects in the stratigraphic field. To date, vallisarni. Remains of the latter species are reported from the continental deposits of the Valdichiana Basin were the nearby locality of Selvella (De Giuli, 1987; Masini et generically referred to the Calabrian, especially on al., 2013) (Fig. 1a), whose referral to the Farneta FU (ca. the basis of biochronological data on mammals (e.g., 1.5 Ma) is supported by biochronological data on fossil Selvella, Farneta, Cava Liberatori LFAs, all dated to horse (Alberdi & Palombo, 2013; Cirilli et al., 2020). ca. 1.5 Ma; Masini & Sala, 2007). On the other hand, The assemblage from the “lower layer” (unfortunately recent studies focused on the paleoenvironmental history no longer cropping out at Vigna Nuova) includes at least of the marine portion of the Basin, which underwent ten taxa, some of which carrying crucial biochronological a complex depositional history at least throughout the information. The co-occurrence of C. ramosus and E. Early Pleistocene (Gelasian and Calabrian; Bizzarri & cf. senezensis allows an attribution of the fauna to the Baldanza, 2020). Further research is needed to identify in second part of the Middle Villafranchian, matching a the field the deposit that yielded the fossils of the “lower correlation with the Coste San Giacomo FU in the Italian layer” of Vigna Nuova, here dated to the Gelasian (ca. 2.2- biochronological framework. This result contrasts with 2.1 Ma), and to understand its stratigraphic relationships the first hypotheses on a younger age of the fauna (Tasso with the surrounding marine and continental units of the FU; Argenti, 2004), probably also failing to recognize the Valdichiana Basin. division into two assemblages. In Italy, scanty remains of Croizetoceros ramosus are reported only from Montopoli (ca. 2.6 Ma; De Giuli AUTHORS’ CONTRIBUTIONS & Heintz, 1974) and Coste San Giacomo (ca. 2.1 Ma; Bellucci et al., 2012, 2014). Therefore, the well-preserved M.C. and B.A. conceived the study. F.R., D.T., M.C., maxilla with upper dentition (SABAP_UMB 129863) and B.A. prepared the fossils. B.A., M.C., and F.R. from Vigna Nuova allows to confirm the presence of this collected the data. B.A., M.C., J.M.M., L.S., and O.C. enigmatic species in the Italian Middle Villafranchian. made the figures. M.C., B.A., M.B., J.M.M., L.S., and Similarly, the horse E. senezensis is rare in Italy, having O.C. developed the methods and performed the analyses. been reported with scanty remains only in Coste San B.A. and M.C. wrote the final version of the manuscript Giacomo and Montecarlo fossil sites, in contrast to the with input from the other authors. All authors contributed rich French sample from Senèze (Palombo et al., 2017; equally to develop the ideas, discussed the results, and Cirilli et al., 2021a; Cirilli, 2022). The occurrence of this reviewed the final version of the manuscript. B. Azzarà et alii - Mammals from Vigna Nuova, Perugia (Italy) 243 SUPPLEMENTARY ONLINE MATERIAL Azzaroli A. (1964). The two Villafranchian horses of the Upper Valdarno. Palaeontographia Italica, 59: 1-12. Supplementary data of this work are available on the Azzaroli A. (1983). Quaternary mammals and the “end- BSPI website at: https://www.paleoitalia.it/bollettino-spi/ Villafranchian” dispersal event - a turning point in the history of Eurasia. Palaeogeography, Palaeoclimatology, Palaeoecology, bspi-vol-612/ 44: 117-139. Azzaroli A. (1992). The cervid genus Pseudodama n. g. in the Villafranchian of Tuscany. Palaeontographia Italica, 79: 1-41. ACKNOWLEDGEMENTS Azzaroli A. (2000). On Equus livenzovensis Baigusheva 1978 and the “stenonid” lineage of equids. Palaeontographia Italica, We are grateful to Giorgio Carnevale and Alberto Collareta 87: 1-17. for having invited us to join this issue; Angela Baldanza, Roberto Azzaroli A., De Giuli C., Ficcarelli G. & Torre D. (1988). Late Bizzarri, Sergio Gentili, and Fausto Pazzaglia for the useful Pliocene to early mid-Pleistocene mammals in Eurasia: discussions on the stratigraphy of Vigna Nuova and information faunal succession and dispersal events. Palaeogeography, on the discovery of the fossils; Leonardo Brustenga, Cristiano Palaeoclimatology, Palaeoecology, 66: 77-100. Spilinga, and Giovanni Testaferri for their availability and help Azzaroli A. & Mazza P. (1992). On the possible origin of the Giant in accessing the outcrop; Marco Bagli for the translation of the Deer genus Megaceroides. Rendiconti Lincei, 3: 23-32. Capellini’s quote; Massimo Delfino, Marco P. Ferretti, Andrea Villa, Bajgusheva V.S. (1978). Krupnaja Loshad Khaprovskogo and Giorgio Carnevale for their opinion on the mustelid tooth; Luca Kompleksa is alluvija SeveroVostochnoto Priazovija. Invesija Pulcinelli and Paola Romi (SABAP_UMB) for constant assistance; Servo-Kavkazkogo Nauchnogo Zentra Vychei Shkoly, 1: 98-102. Dimitris Kostopoulos for the pictures of Greek Croizetoceros; Luca Baldanza A., Bizzarri R., Famiani F., Garassino A., Pasin G., Cherin Pandolfi for his thorough review; Luca Bellucci for his review M. & Rosatini F. (2018). The early Pleistocene whale-fall and for kindly providing additional data on canid vertebrae. O.C. community of Bargiano (Umbria, Central Italy): Paleoecological gratefully acknowledges Howard University, Washington D.C. for insights from benthic foraminifera and brachyuran crabs. space and resources for his postdoctoral research. L.S. is supported Palaeontologia Electronica, 21: 1-27. by the FI AGAUR fellowship (ref. 2020 FI_B1 00131) funded by the Ballesio R. (1963). Monographie d’un Machairodus du gisement Secretaria d’Universitats i Recerca de la Generalitat de Catalunya villafranchien de Senèze: Homotherium crenatidens. Travaux and the European Social Fund. et Documents des Laboratoires de Géologie de Lyon, 9: 1-129. Barchi M.R. & Marroni M. (2007). 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