Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy) vol. 127(1): 179-187. March 2021 MIDDLE PLEISTOCENE FOX FROM THE VALLPARADÍS SECTION (VALLÈS- PENEDÈS BASIN, NE IBERIAN PENINSULA) AND THE EARLIEST RECORDS OF THE EXTANT RED FOX JOAN MADURELL-MALAPEIRA1*, SAVERIO BARTOLINI LUCENTI2,3 & DAVID M. ALBA1 1Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, c/ Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain. E-mails: [email protected]; [email protected] 2 Earth Science Department, Paleo[Fab]Lab, University of Florence, Via La Pira 4, 50121, Firenze, Italy. E-mail: [email protected] 3Natural History Museum, Geology and Paleontology Section, University of Florence, Via La Pira 4, 50121, Firenze, Italy. *Corresponding author. To cite this article: Madurell-Malapeira J., Bartolini Lucenti S. & Alba D.M. (2021) - Middle Pleistocene fox from the Vallparadís Section (Vallès- Penedès Basin, NE Iberian Peninsula) and the earliest records of the extant red fox. Riv. It. Paleontol. Strat., 127(1): 179-187. Keywords: Pleistocene; Iberia; Vallparadís Section; Carnivora; Canidae; Vulpes. Abstract. Dentognathic remains of European Middle Pleistocene Vulpini are scarce and fragmentary. They have classically been attributed to several species, but many taxonomic and phylogenetic uncertainties remain. Here we describe a fox well-preserved maxilla with associated mandible from the Middle Pleistocene layers of the Vallparadís Section (EVT3; ca. 0.6 Ma; Vallès-Penedès Basin, NE Iberian Peninsula) and compare them with the known fossil record of Vulpes from roughly contemporary European localities. The studied Vulpes remains described here differ from Early Pleistocene Vulpes alopecoides (including its junior synonyms V. praeglacialis and V. praecorsac) in several den- tognathic features (less individualized P4 protocone; M2 paracone larger than the metacone; more robust mandibular corpus, and stouter m1 with reduced metaconid) and most closely resemble extant Vulpes vulpes. The large size of the described remains also contrasts with the small dimensions of V. alopecoides and fits better within the variability of the extant red fox. We conclude that the studied specimens, like other available Middle Pleistocene fox remains from Europe, are attributable to the extant species V. vulpes. Therefore, the EVT3 remains represent the first well-dated appearance datum of the species in Europe. INTRODUCTION centi & Madurell-Malapeira 2020): Vulpes alopecoides (Del Campana, 1913), Vulpes praeglacialis (Kormos, The European Plio-Pleistocene fossil record 1932), and Vulpes praecorsac Kormos, 1932. of the genus Vulpes is fragmentary and patchy, and The absence of clear diagnostic features to available remains mostly come from old collections differentiate these three similar-sized species has led and are poorly dated. This hinders the reconstruc- to a confusing taxonomic allocation of several sam- tion of their taxonomic affinities and has led to the ples from Western Europe. Late Pliocene and ear- proliferation of a large number of species names liest Pleistocene remains have often been ascribed of doubtful taxonomic validity. The following three to V. alopecoides, whereas late Early to Middle Plei- species were traditionally recognized from Pliocene stocene fossils have been assigned to either V. prae- to Middle Pleistocene sites of Europe (Bartolini Lu- glacialis or V. praecorsac on the basis of size (large vs. small, respectively). Bartolini Lucenti & Madurell- Received: October 21, 2020; accepted: February 18, 2021 Malapeira (2020) recently allocated all Early Pleisto- 180 Madurell-Malapeira J., Bartolini Lucenti S. & Alba D.M. cene samples to V. alopecoides—thereby synonymi- colluvial processes (Madurell-Malapeira et al. 2010; zing with the latter multiple species distinguished by 2017). Magnetostratigraphic and biostratigraphic previously authors (Bonifay 1971; García et al. 1997; correlations indicate that the VCS spans from befo- Moigne et al. 2006)—based on their overall simila- re the Jaramillo paleomagnetic subchron (ca. 1.2-1.1 rities compared to the high intraspecific variation Ma) to the early Middle Pleistocene (layer EVT3, ca. typical of extant foxes (Daitch & Guralnick 2007; 0.6 Ma; Madurell-Malapeira et al. 2010, 2014, 2017; Szuma 2008a, 2008b, 2008c; Gimranov et al. 2015; Minwer-Barakat et al. 2011). Gimranov 2017) During the excavations carried out betwe- Despite the taxonomic confusion, V. alope- en 1997 and 2008 from CGR and EVT, more than coides was generally regarded as the ancestor of the 30,000 vertebrate fossil remains were recovered. extant red fox, Vulpes vulpes (e.g., Rabeder 1976; Mesowear and microwear analyses performed on a Bartolini Lucenti & Madurell-Malapeira 2020). The large sample of ungulate dental remains indicate that European Middle Pleistocene record of V. vulpes is around 0.9 Ma the environment of VCS shifted from utterly scarce. It is first recorded in the French locali- an open dry grassland with clear seasonality (layer ties of l’Escale (ca. 0.6 Ma; Bonifay 1971) and Caune EVT12; ca. 1.0 Ma; MIS 31) to more humid and pro- de l’Arago (ca. 0.6-0.45 Ma; Moigne et al. 2006). In bably even more seasonal woodlands (layer EVT7; the Iberian Peninsula, red foxes have been identified ca 0.86 Ma; MIS 21), in agreement with data from from Sima los Huesos (MIS12, ca. 0.45 Ma; García other Southern European roughly coeval sites (Stra- et al. 1997) and Galería (ca. 0.3 Ma; García & Arsua- ni et al. 2019). Specifically, the analysis of the EVT3 ga 1998) in Atapuerca. In Italy, it appears probably (ca. 0.6 Ma) remains suggests paleoenvironmental slightly later, at the MIS 12-11 transition (ca. 0.43 conditions characterized by heterogeneous landsca- Ma; Petronio et al. 2019), in sites such as Malagrot- pes, possibly including the predominance of open ta (Capasso Barbato & Minieri 1987) and Torre in habitats, as further indicated by the relatively high Pietra (Caloi & Palombo 1978). However, the pre- abundance of horse remains. This spread of open cise chronology of the appearance of V. vulpes and grasslands around 0.6 Ma is also attested at Isernia la its phylogenetic affinities with previous late Early Pineta (Italy), where herbaceous plants are the most Pleistocene V. alopecoides has never been subject to abundant in the pollen record (Strani et al. 2019). in-depth scrutiny. Here we describe the unpublished fox remains from the early Middle Pleistocene (ca. 0.6 Ma) Vallparadís Section and compare them with MATERIALS AND METHODS roughly coeval Vulpes remains from other European The described specimen from EVT3 (IPS120168) is housed in sites. the Institut Català de Paleontologia Miquel Crusafont, Sabadell, Spain. It is compared on qualitative morphological as well as morphometric grounds with other Early to Middle Pleistocene foxes belonging to GEOLOgiCAL AND PALEONTOLOgiCAL the extinct species V. alopecoides (sensu Bartolini Lucenti & Madurell- SETTING Malapeira 2020; see therein for the morphological and biometrical cri- teria used in this article) and fossil V. vulpes from selected European The described remains were collected from sites. In particular, the inspected comparative fossil material is housed in the following institutions (see abbreviations below): DST-UNIFI, layer EVT3 of the Vallparadís Estació (EVT) site, HNHM, and IGF. We also inspected all the relevant literature on Eu- which is part of the well-dated Vallparadís Compo- rasian fossil Vulpes (Odintzov 1965; Rabeder 1976; Qiu & Tedford site Section (VCS; Vallès-Penedès Basin, northea- 1990; Ginsburg 1998; Qiu et al. 2004; Koufos 2014; 2018; Wang et al. 2014). In turn, the extant comparative material includes specimens of stern Iberian Peninsula) together with the fossil site Vulpes corsac, Vulpes lagopus, and V. vulpes from the AMNH, HNHM, of Cal Guardiola (CGR; Madurell-Malapeira et al. and MZUF. 2010; 2017). The section has a total thickness of 20 m and multiple stratigraphic horizons. Specifically, Institutional abbreviations: ICP, Institut Català de Paleon- tologia Miquel Crusafont; DST-UNIFI, Earth Science Department the EVT site is a 14 m-thick succession composed of the University of Florence (Italy); IGF, Geological and Paleonto- by alternations of conglomerates and mudstones, di- logical Section of the Natural History Museum of the University of vided into two units separated by an erosive angular Florence (Italy); MZUF, La Specola, Zoological Section of the Na- unconformity (Madurell-Malapeira et al. 2010). All tural History Museum of the University of Florence (Italy); PP, site abbreviation of the Pirro Nord material housed in the collection of these sediments can be interpreted as the result of an Geological and Paleontological Section of the Natural History Mu- alluvial fan system paired by the influence of close seum of the University of Florence (Italy). Middle Pleistocene fox from the Iberian Peninsula 181 SYSTEMATIC PALEONTOLOGY protoconule is faint, and the metaconule is promi- nent but lower than the protocone. The hypocone Order Carnivora Bowdich, 1821 is separated from the lingual cingulum by a marked Family Canidae Fisher, 1817 furrow. The trigon basin is round and wide, and as Subfamily Caninae Fisher, 1817 deep as the talon basin, which is undulated but not Tribe Vulpini Hemprich & Ehrenberg, 1832 compressed. The M2 has a buccolingually elonga- Genus Vulpes Frisch, 1775 ted occlusal contour. The