Journal of Iberian Geology https://doi.org/10.1007/s41513-020-00127-y

RESEARCH PAPER

New remains of a primitive badger from Cueva de los Toriles (Carrizosa, Castilla‑La Mancha, Iberian Peninsula) suggest a new quaternary locality in the southern Iberian plateau

Daniel García‑Martínez1,2,3 · Alberto Valenciano4,5 · Aitziber Suárez‑Bilbao6 · Carlos A. Palancar2 · Irene Megía García7 · Davinia Moreno1 · Isidoro Campaña1 · Pedro R. Moya‑Maleno3,8

Received: 18 September 2019 / Accepted: 7 May 2020 © Universidad Complutense de Madrid 2020

Abstract The Cueva de Los Toriles belongs to a karstic system found in the southern Iberian plateau. Construction works in the (not related to paleontological activities) have allowed for the recovery of fossil remains of macromammals, including two lower molars of a mustelid that are presented in this work. They can be attributed to the species Meles cf. thorali based on morphological and morphometric comparisons. This extinct mustelid is known mostly from Villafranchian deposits of , , and spanning from the late Pliocene potentially, even reaching with some doubts the Early-Middle Pleistocene of France. Therefore, according to the presence of this badger in the cave, we could tentatively assume a chronological age ranging from the Late Pliocene to the Middle Pleistocene, chronologies that would be consistent with some lithic tools found in the cave. We highlight the importance of Cueva de Los Toriles for the Quaternary Paleontology of the southern Iberian plateau, one of the places from the Iberian Peninsula with a more limited fossil record from these chronologies.

Keywords Quaternary · Meles · Mustelidae · Iberian Peninsula · Castilla-La Mancha

Resumen La Cueva de los Toriles se encuentra en un sistema kárstico situado en la Submeseta sur de la Península Ibérica. A pesar de que actualmente no existe datación de los depósitos sedimentarios, algunos trabajos de acondicionamiento de la cueva (previos a las actividades paleontológicas) permitieron el hallazgo de restos fósiles de macromamíferos, entre los que se incluyen dos molares inferiores de un mustélido, principal foco de este trabajo. Estos molares pueden ser atribuidos a la especie Meles cf. thorali a través de comparaciones morfológicas y morfométricas. Este mustélido extinto se encuentra prin- cipalmente en depósitos Villafranquienses de Francia, España y Grecia entre el Plioceno tardío y el Pleistoceno temprano, llegando potencialmente, con algunas dudas, incluso al Pleistoceno Medio de Francia. Por lo tanto, de acuerdo con la pres- encia de esta especie de tejón en la cueva, podemos asumir tentativamente una edad cronológica que va desde el Plioceno Tardío hasta el Pleistoceno Medio, cronologías que serían coherentes con algunas piezas de insdustria lítica encontradas en la cueva. Destacamos la importancia de la Cueva de Los Toriles para la Paleontología Cuaternaria de la Meseta Ibérica meridional, uno de los lugares de la Península con registro fósiles más limitado de estas cronologías.

Palabras Clave Cuaternario · Meles · Mustelidae · Península Ibérica · Castilla-La Mancha

1 Introduction

The Pleistocene record of the Iberian Peninsula is very Daniel García-Martínez and Alberto Valenciano contributed rich. Jordá Pardo (2008) groups the Iberian Pleistocene equally to the manuscript. deposits according to their location in fve geomorphologi- * Daniel García‑Martínez cal subunits: (1) the Cantabrian Mountains and the Pyre- [email protected] nees, (2) the Central System, (3) the Iberian Range, (4) the Extended author information available on the last page of the article Coastal-Catalan Range, and (5) the Baetic Ranges and the

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Guadix-Baza Basin. Important Pleistocene fossiliferous sites (to the south). Geologically, the Paleozoic (Ordovician) sub- in the Cantabrian Mountains and the Pyrenees, in terms of strates extend across the western area of the region except amount and quality of the fossils preserved, are Cova Eirós for small outcrops of Precambrian rocks (Fig. 2). The eastern (Grandal-D’Anglade and Romaní 1997), El Sidrón (Fortea area is composed of Mesozoic sedimentary rocks (García et al. 2003; Rosas et al. 2013; García-Martínez et al., 2017) del Cura et al. 2000; Montero 2003; González et al. 2004), or Lezetxiki (Baldeón 1993), among others. In the Central represented by large limestone and dolomite bedrocks settled System and the Iberian Range, examples of Pleistocene fos- on highly folded Paleozoic substrates (Jiménez and Chaparro sil sites include Pinilla del Valle (Baquedano et al. 2016), 1983; Montero 2003) (Fig. 2). This can give rise to karstic TAFESA (Sesé 2010), Jarama VI (Jordá Pardo 2007) or the environments such as the lakes (Martínez Goytre worldwide-known Atapuerca (Aguirre and Lumley 1977; et al. 1988; González et al. 2004) or aquifers such as those Rosas et al. 2013; Bermúdez de Castro et al. 2011; Arsuaga of Las Tablas de (García Rodríguez and Almagro et al. 2015). In the Coastal-Catalan Range, sites include Costa 2004). Finally, the northern area is characterized by Vallparadís Section (Madurell-Malapeira et al. 2010), Bar- Cenozoic sediments (Jiménez and Chaparro 1983) (Fig. 3). ranc de la Boella (Vallverdú et al. 2014), Abric Romaní Based on these geological and geographical features, the (Fernández-García 2018), Cova Negra (Richard et al. 2019), Pleistocene deposits that can be found in this region are (1) Cova de Bolomor (Arsuaga et al. 2012) and Incarcal-I (Ros- fuvial sediments in the western and northern areas, related Montoya et al. 2012). Finally, in the Baetic Ranges and the to the paleo-basins of the Guadiana and Guadalquivir and Guadix-Baza Basin, important fossil sites include Quibas (2) karstic inflling sediments in the eastern part related to site (Piñero et al. 2016, 2020), Cueva Victoria (Gibert et al. the Mesozoic karstic systems. Few fossiliferous sites have 2016), Sima de las Palomas (Walker 2008), Orce (Agustí been described in La Mancha, and those that have been et al. 2015) and (Fynlanson et al. 2008). studied have not been explored in depth. Noteworthy are However, the list mentioned above highlights the fact Las Higueruelas (Badiola et al. 2007), Bonete y Piedrab- that some area of the Iberian Peninsula, such as the south- uena (Torres and Mazo 1991), and Valverde-2 (Alberdi ern Iberian plateau (also known as stable Meseta or Meseta et al. 1984) in the Campo de Calatrava, with Pliocene chro- Central) has few relevant sites, and in the region of Castilla- nologies. Early Pleistocene fossil sites include Valverde-1 La Mancha (including Guadalajara, , Cuenca, and Fuensanta del Júcar, with the presence of macromam- Toledo, and provinces) they are particularly scarce. mals such as Mammuthus meridionalis or Hippopotamus Several important factors contribute to this situation, such as antiquus, dated around 1.3–0.8 Ma (Aguirre 1989; Mazo the eminently agricultural use of the landscapes of the river 1999) and 1 Ma (Mazo et al. 1990), respectively. The El paleobasins that conforms most of this region, as well as the Provencio site is dated around 0.9 Ma and is coeval with focus of the regional academy in historical periods instead of Fuensanta del Júcar (Domínguez-Solera et al. 2020; Mazo pre-historical or paleontological periods (García-Martínez et al. 1990). There is no fossil evidence from the Middle- 2019). The presence of a potential previously unknown fos- Late Pleistocene in La Mancha. siliferous site in Castilla-La Mancha, Cueva de Los Toriles in the area called “La Mancha”, could shed light on this 1.2 The Cueva de los Toriles and its importance issue. in the southern Iberian plateau

1.1 Geographical and geological setting The Cueva de Los Toriles (Carrizosa, Ciudad Real, Campo de , Castilla-La Mancha, Central Iberian Penin- The area that is strictly called “La Mancha” is a not well- sula) (Fig. 1) is part of a large and still unexplored karstic delimited region located in the center east of the Castilla- system in Mesozoic (late Jurassic; Lias) bedrocks. It La Mancha autonomous community, with an area of about belongs to the Betico-Castellano foreland (Rincón et al. 30,000 ­km2 (Pérez and de la Peña 1986) that includes most of 2001) and may contain some pre-Holocene sedimentary Ciudad Real, as well as part of Cuenca, Toledo, and Albac- deposits (García-Martínez and Suárez-Bilbao 2018). Even ete (Madoz 1846). It is located in the southern Iberian pla- though the cave has not been properly excavated yet, con- teau (Fig. 1) and large plain terrains along the Guadiana and struction work in the cave (the removal of the natural sub- Guadalquivir basins typify its western and northern areas. strate of the cave access) led to the fortuitous discovery However, its eastern part and some other outcrops within the of a rich fossil assemblage (García-Martínez and Suárez- Campo de Montiel, are characterized by more rugged land- Bilbao 2018). The assemblage includes two mustelid teeth scapes. From a biogeographical point of view, the southern in excellent condition that may be important for taxonomic Iberian plateau is in a strategic position, since it connects assessment and provide new information on a possible the Central System (to the north) and the Iberian Range (to relative dating of the cave deposits. Even though the two the northeast) with the Sierra Morena and the Baetic Ranges fossils were found in reworked sediments (Fig. 4), they

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Fig. 1 The geographical location of the Cueva de Los Toriles in (c) Guadalajara province, (d) Toledo province, (e) Cuenca Province, Europe (top-left), Iberian Peninsula (top-centre) and in Castilla-La (f) Ciudad Real province, (g) Albacete province, (h) Campo de Mon- Mancha autonomous community (bottom-left), as well as a detail, in tiel (local term of a historical and administrative area) and (i) La Car- a color-map of the site position between the two closer populations. rasca (closest population to the cave) Legend: (a) Spain, (b) Castilla-La Mancha autonomous community, were both found in the same area of the cave and have a 2 Material and methods similar developmental status, so they may belong either to the same individual or to individuals of similar devel- 2.1 Materials opmental status. The teeth from the Cueva de los Toriles are temporar- 1.3 Aims of this study ily housed at the Museo Nacional de Ciencias Natu- rales (MNCN-CSIC) in Madrid (Spain). The m2 The fossil record in the southern Iberian plateau is scarce Tor’19·B·Sup·508 was first described by García-Martínez compared to other Iberian regions (García-Martínez et al. and Suarez-Bilbao (2018) as a potential p3 of a large 2019; Morales et al. 2015, 2018). In this context, the rodent. However, after a re-evaluation of the material, Cueva de los Toriles site, a yet unexplored fossil site, will it is reassigned herein to the genus Meles Brisson, 1762. contribute to flling this gap of knowledge. This study aims We compared the two teeth with samples of the extant to analyze and discuss the potential stratigraphic attribu- Eurasian badger Meles meles Linnaeus, 1758 from Swe- tion of the two mustelid fossil remains, and evaluate their den, , and Spain, housed at the Naturhistoriska taxonomical afnities through qualitative and quantitative Riskmuseet (NRM) in Stockholm (Sweden), the Naturhis- comparison with fossil and extant material, discussing the torisches Museum (NHMW) in Vienna (Austria) and the implications of the results for the paleontology in the Ibe- MNCN, respectively. Besides, the fossils were compared rian plateau. with samples of the extant hog badger Arctonyx colla- ris Cuvier, 1825 housed at the Field Museum of Natural

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Fig. 2 Location of the cave, marked with a star, on the regional geological map taken from the IGME, both at a 1: 1,000,000 (a) scale and at a 1: 50,000 (b) scale. The cave is located in the contact between the blue and red areas, cor- responding respectively to Jurassic and Upper Triassic (F. Keuper)

History (FMNH) in Chicago (USA), and the NHMW. In 3 Results addition to these modern species, selected Plio-Pleisto- cene badgers were used for comparisons by taking data 3.1 Potential assessment of the stratigraphic origin from the original publications, except for the holotype of the mustelid remains from Cueva de los of Arctomeles gennevauxi (Viret 1939) housed at Uni- Toriles versité Claude Bernard Lyon 1 (FSL) in Lyon (France), which was analyzed using pictures of the original speci- The two teeth of a mustelid appeared in reworked sediments men. Dental nomenclature follows Ginsburg (1999) and at the entrance of the cave (called Toriles-Boca and labeled Smith and Dodson (2003) (Fig. 5). Measurements were as Tor·B). The stratigraphic section of this area is formed made using Mitutoyo Absolute digital caliper to the near- mainly by two stratigraphic units, Tor I in the upper part est 0.1 mm. and Tor II in the lower part (Fig. 4). Tor I is formed by yel- lowish brown sandy silt with gravels. In this stratigraphy

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Fig. 4 Stratigraphic section of the Tor·B area

Fig. 3 Location of the specimens of Meles found in the cave as well as the section of the area close to the place where these specimens were found. a A general overview of the interior part of the cave. Specifcally, we show the cave area called “Boca” (Tor·B). The sym- bol * shows the place (out of context) where the teeth were found, and the symbol # shows the place where we explored the stratigra- phy of the “Boca”; b overall view of the two teeth of Meles found in the area marked as *; c frontal view of the stratigraphic section close to the teeth. Two diferentiated geological strata can be observed: the upper one (TOR·B·I) is rich in microfauna and present some remains of macrommamals (currently under study), whereas the lower one (TOR·B·II) does not contain fossils but presents Lower stone tools. This is currently under study but the lower stratum is older than the upper one unit, at least three layers can be identifed. Tor I has a dip towards the entry of the cave, suggesting that it was depos- Fig. 5 Terminology and measurements used for the description of ited by water fows from the interior of the cave towards its the lower dentition. a The terminology of the m1 and m2. b Meas- Cs En entrance. Tor I lies above an erosion surface of the top of urements of the lower dentition. Abbreviations: cuspulets, entoconid, Entcn entoconulid, Hp hypoconid, Hpn hypoconulid, L Tor II. Regarding the layers into Tor I, Tor I.1 is yellowish- maximum length, m1 lower frst molar, m2 lower second molar, Mt brown (10YR 5/4) sandy silt with sub-rounded gravels less metaconid, Pa paraconid, Pr protoconid, Tld talonid, Tr trigonid, W than 0.5 cm of the major axis, Tor I.2 is a yellowish-brown maximum width (10YR 6/4) sandy silt with sub-rounded gravels less than 2 cm of the major axis and Tor I.3 is a brown (10YR 5/3) sandy silt with sub-rounded gravels less than 0.5 cm of the cave too. This fact suggests that these layers flled this area major axis. and that the current cave is the result of the erosion of this Tor II consists of grey carbonate silt which can be difer- stratigraphic unit, but further work is required to clarify this entiated in three layers in the current stratigraphic section, issue. Tor II is slightly dipping to the entrance of the cave. although these carbonates are observed in the walls of the On the top of each layer, a yellowish partially cemented crust is observed. The layers of the section are named from 10

1 3 Journal of Iberian Geology onwards, based on the sedimentary layers observed on the Material: Tor’19·B·Sup·507, right m1 (Fig. 6a–d), and cave walls. This way, Tor II.10 is a brownish grey (10YR Tor’19·B·Sup·508, left m2 (Fig. 6e, f). 6/2) carbonate silt, Tor II.11 is a brownish grey (10YR 6/2) Description: The right m1 Tor’19·B·Sup·507 (maximum carbonate silt and Tor II.12 is also brownish grey (10YR length = 15.8 mm; maximum width = 7.7 mm; lingual length 6/2) carbonate silt. On the top, a cemented clay and silt of the trigonid = 8.9 mm; length of the talonid = 6.9 mm) sub-layer are found. This is a discontinuous sub-layer that is complete (Fig. 6a–d). It has a low crown, the protoco- is observed only in a limited area of the cavity. Tor II.12 is nid is the highest cusp of the trigonid, and the metaconid is currently the foor of the cave. Preliminary analysis of the tall and about the same height as the paraconid cusp. The sediments of Tor I and II suggest that the upper stratum talonid is relatively short in comparison with the trigonid. is a Holocene deposit rich in microfauna, containing also Both hypoconid and entoconid are well developed, and the some macromammal remains, whereas the lower stratum hypoconulid and entoconulid are less developed. There is a is a Pleistocene deposit that is poor in faunal remains but ring of distal cuspulets surrounding the talonid and a deep presents some Paleolithic stone tools (Garcia-Martinez et al., basin on it. There are strong buccal and lingual cingulids, in review). Since the fossils presented here were found in and the tooth has additional accessory roots. The left m2 reworked sediments coming from one of the two strata, it Tor’19·B·Sup·508 (maximum length = 5.3 mm; maximum cannot be assumed where the fossil teeth come from. How- width = 5.74 mm) has a single open root still in formation, ever, we know that the Holocene deposits would only be while the crown is complete and completely formed (Fig. 6e, characterized by the badger species M. meles whereas the f). The trigonid is buccolingually broader than the talonid Pleistocene deposits would be characterized by either the and shows two main cusps that are a protoconid and a shorter species M. meles or the species M. thorali. metaconid. There is a noticeable mesial and distal cingulid.

3.2 Systematic paleontology 4 Discussion Order CARNIVORA Bowdich, 1821 Suborder CANIFORMIA Kretzoi, 1943 4.1 The badger from Cueva de los Toriles Family MUSTELIDAE Fischer, 1817 in the context of European Plio‑Pleistocene Subfamily MELINAE Bonaparte, 1838 badgers Genus MELES Brisson, 1762 Type species: Meles meles Linnaeus, 1758 The Plio-Pleistocene badger record in Europe is fragmentary Meles cf. thorali but quite diverse in terms of the number of species (e.g., Locality: Cueva de los Toriles (Ciudad Real, Castilla La- Viret 1939; Teilhard and Leroy 1945; Stach 1951; Tedford Mancha, Spain). and Harington 2003; Madurell-Malapeira et al. 2009, 2011a, b; Baryshnikov 2009; Petrucci et al. 2013; Mecozzi et al.

Fig. 6 The lower dentition of Meles cf. thorali from Cueva de los Toriles. A-D. Tor’19·B·Sup·507, right m1. a Buccal view, b lingual view, c occlusal view, d distal view; e, f Tor’19·B·Sup·508, left m2. e Occlusal view, f distal view. Scale bar 2 cm

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2019). This issue led paleontologists to establish several new The badger from Cueva de los Toriles difers from the species based on very scarce material without taking into living M. meles in its mesiodistally larger m1 protoconid, account the intraspecifc variability of these forms and their in its shorter m1 talonid, and its less-developed m1 meta- living relatives. The systematics of Plio-Pleistocene badg- conid, hypoconulid, and entoconulid (Figs. 7, 8). It has a ers is intricate, partly due to the high polymorphism of the relatively wider and robust hypoconid and a shallower basin extant species M. meles (e.g., Baryshnikov 2009; Petrucci on the talonid. It also has less reduction in the m2 meta- et al. 2013; Mecozzi et al. 2019), and also because most conid compared to M. meles. All these diferences can be of the extinct species overlap in size with the extant one interpreted as more primitive traits than those of the living (Figs. 7, 8). Meles. Compared with M. meles and M. meles atavus from European Plio-Pleistocene badgers are included in the the Late Villafranchian (1.8–1.1 Ma) and Epivillafranchian genera Arctomeles (Stach 1951) and Meles (Brisson 1762). (1.1–0.8 Ma) of the Iberian Peninsula, such as Fuente Arctomeles is a large-sized Pliocene badger possibly related Nueva-3 or Vallparadís Estació (Madurell-Malapeira et al. to the living hog badger Arctonyx collaris according to 2011a, b), the badger from the Cueva de los Toriles also rep- Tedford and Harington (2003). This genus comprises the resents a more primitive form with a smaller overall size and European species Arctomeles pliocaenicus (Stach 1951) a smaller talonid with less-developed cuspids (Figs. 7, 8). It from , Arctomeles suillus (Teilhard and Leroy 1945) also difers from the Late Pleistocene M. meles from Grotta from Ukraine, , and Asia (Tedford and Haring- Laceduzza (Mecozzi et al. 2019) and Kudaro 1 and 3 ton 2003) and Arctomeles gennevauxi (Viret 1939) from (Baryshnikov 2009) in the same traits. The strong cingulid the Early Pliocene of Montpellier (France). The specimen and the accessory roots in the m1 from the Cueva de los Tor’19·B·Sup·507 difers from A. gennevauxi in its smaller Toriles are rarely observed in Meles, although some speci- size and less elongated m1 talonid (Fig. 9c, d). Moreover, mens of M. meles from Sweden analyzed in this study had A. gennevauxi has a double metaconid, a trait that is absent accessory roots (A.V. personal observation). As discussed in the extant Spanish specimen. Therefore, the badger from above, the intraspecifc dental variation of M. meles is very Cueva de los Toriles cannot be included in this genus. wide (Figs. 7, 8), but we can preliminarily rule out the inclu- The genus Meles comprises six species recorded in cen- sion of Meles from Cueva de los Toriles in M. meles based tral and western Europe ranging from the Late Pliocene to on morphological and morphometric diferences. However, present (Madurell-Malapeira et al. 2011a, and references only more fossil material from Cueva de los Toriles can therein): (1) Meles thorali (Viret 1951) from the Late clarify its taxonomic attribution. Pliocene to the Middle Pleistocene of France, Spain, and The size and proportions of the badger from the Cueva Greece (Bonifay 1971, 1981; Fosse 1996; Vos et al. 2002; de los Toriles are fully comparable with those of the Late Madurell-Malapeira et al. 2009); (2) M. iberica (Arribas Pliocene and Early Pleistocene M. thorali from Europe and Garrido 2007), from the Middle Villafranchian of Fon- (Viret 1951; Bonifay 1971, 1981; Fosse 1996; Vos et al. elas P-1 (Spain); (3) M. dimitrius (Koufos 1992), from the 2002; Madurell-Malapeira et al. 2009), especially to the Greek Villafranchian sites of Gerakarou and Apollonia-1; specimens from the type locality of Saint Vallier in France (4) M. hollitzeri (Rabeder 1976), from the Epivillafranchian (Early Pleistocene; Middle Villafranchian, MN17; Viret localities of Untermassfeld and Deutsch-Altenburg 2 (Ger- 1951). Meles thorali can be distinguished from M. meles many and Austria, respectively); (5) M. atavus (Kormos based on several cranial and dental characters (Viret 1951; 1914), from the Epivillafranchian of Beftia 5 (); Argant and Mallye 2005; Madurell-Malapeira et al. 2011a): and (6) the extant M. meles recorded from several Middle the relatively shorter muzzle, the more elongated auditory to Late Pleistocene sites throughout Europe (Madurell- bulla, the similar height of the M1 metacone and paracone, Malapeira et al. 2011a, b; Baryshnikov 2009; Petrucci et al. the presence of a small distal bulge-like cuspid in p2 and 2013; Mecozzi et al. 2019). Within this entire taxonomic p3, the higher p4, and the higher m1 paraconid, protoconid framework, Madurell-Malapeira et al. (2011a) in their and metaconid than those of the living M. meles. One of taxonomic review of the Villafranchian badgers of Europe, the key traits for distinguishing M. thorali from M. meles is synonymized these species and restricted them to M. tho- the longer length of the trigonid in relation with the talonid rali for the Early (3.6–2.6 Ma) and Middle Villafranchian in the lower carnassial of M. thorali. Madurell-Malapeira (2.6–1.8 Ma) forms and M. meles atavus for the fossils from et al. (2011a) stated that the Early and Middle Villafranchian the Late Villafranchian (1.8–1.1 Ma) and Epivillafranchian European badger possesses a longer m1 trigonid than talonid (1.1–0.8 Ma). However, Koufos (2018) still considers M. when compared with the late Villafranchian and Epivilla- dimitrius as a valid species, and according to Bonifay (1981) franchian forms. This is also observable in Fig. 8, where we and Fosse (1996), M. thorali is present in the Middle Pleis- plotted the maximum lingual length of the trigonid and the tocene of France (around 0.35 Ma). talonid. We observe that the Meles specimen from Cueva de los Toriles is located very close to the range of variability

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Fig. 7 Width (W) and length (L) in mm of the m1 of Meles cf. thorali the diferent badgers are shown on a real scale, so their absolute pro- from Cueva de Los Toriles, compared with Plio-Pleistocene badgers portions can be observed. Source: This work: 1; (Viret 1951): 2, 9; and living Meles meles and Arctonyx collaris. a = range of variation (Bonifay 1971): 3; (Mallye 2018): 4; (Gasparik and Pazonyi 2018): 5; of the extant M. meles analyzed in this work; b = range of variation (Viret 1939): 6; (Madurell-Malapeira et al. 2011a): 7; (Koufos 2018): of M. thorali according to Madurell-Malapeira et al. (2011a), includ- 8; (Arribas and Garrido 2007): 9; (Madurell-Malapeira et al. 2011b): ing the one from Cueva de Los Toriles; c = range of variation of the 10; (Petrucci et al. 2013): 11; and (Mecozzi et al. 2019): 12 extant A. collaris analyzed in this work. The pictures of the m1s of of M. thorali (e.g., Saint Vallier), and has very similar pro- cf. thorali from Cueva de los Toriles and M. iberica from portions to the Plio-Pleistocene Asiatic badgers Meles chiai Fonelas P-1 (Arribas and Garrido 2007) is not possible due Teilhard de Chardin, 1940, Meles magnus Jiangzuo et al. to the advanced wear of the lower dentition of the described 2018 and M. teihardi Qiu et al. 2004 (Fig. 8). In contrast, specimens from Fonelas P-1. It is noteworthy that one of the Middle and Late Pleistocene badgers such as M. meles the two described m1 of M. iberica is considerably smaller atavus, and M. meles (both Late Pleistocene and living popu- (maximum length = 13.3 mm; maximum width = 5.7 mm) lations), have relatively longer talonids (Fig. 8). Therefore, than that of the Cueva de los Toriles. In the Iberian Penin- based on morphological and morphometric comparisons, sula, several Middle Villafranchian species of carnivores, we preliminarily place the badger specimens from Cueva including M. thorali (Madurell-Malapeira et al. 2014), did de los Toriles in Meles cf. thorali. Meles thorali has been not survive the climatic shifts of the Late Villafranchian. previously described in the Iberian Peninsula by an M1 from However, it is also important to note that Fosse (1996) pro- Almenara-Casablanca 4 (Late Pliocene, MN16), which rep- posed that M. thorali was also present in the Middle Pleisto- resents the earliest record of Meles in Europe (Madurell- cene of France (around 0.35 Ma), by the taxon Meles thorali Malapeira et al. 2009). A direct comparison between Meles spelaeus from the locality of Lunel-Viel, and Mallye (2018)

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Cueva de los Toriles site could indicate a potential age rang- ing to the Late Villafranchian (Late Pliocene) to the Early- Middle Pleistocene, based on the known temporal range of this badger species (Viret 1951; Bonifay 1971, 1981; Fosse 1996; Vos et al. 2002; Madurell-Malapeira et al. 2009).

4.2 Importance of the Cueva de los Toriles for the fossil record of the southern Iberian plateau

Our fndings highlight the importance of a potential Late Plio- cene-Middle Pleistocene fossil site in the southern Iberian pla- teau and in the region called La Mancha specifcally, periods that are largely absent in this region (García-Martínez 2018). According to Rook and Martínez-Navarro (2010), the Villa- franchian spans the Late Pliocene to most of the Early Pleis- tocene (from around 3.5 Ma to about 1.0–1.1 Ma). Although other important fossil sites from the Pleistocene are found in the Castilla-La Mancha province, such as Cueva de Los Casares (Barandiarán and Beltrán 1973; Alcaraz-Castaño et al. 2017) or Jarama VI (Jordá Pardo 2007, 2008), these are not strictly located in the southern Iberian plateau but are located in the Central System and the Iberian Ranges respectively. A literature extensive review of the fossiliferous localities in La Mancha yields just a few references, such as Las Higuerue- las, Bonete y or Valverde-2 (Alberdi et al. 1984; Badiola et al. 2007; Torres and Mazo 1991), all of which are in the Campo de Calatrava (western region) in Pliocene chro- nologies. For the Early Pleistocene, the few fossil sites in the region are those from Valverde-1, Fuensanta del Júcar, and El Provencio (Aguirre 1989; Mazo 1990, 1999), dated around 0.9 Ma (Domínguez-Solera et al. 2020). Therefore, there is no paleontological evidence of the Middle-Late Pleistocene not only in La Mancha but in the entire southern Iberian pla- teau, and the evidence of the Pliocene and Early Pleistocene M thorali Fig. 8 Scatter plot of the maximum lingual length of the m1 trigo- is very scarce. The presence of . cf. from the Cueva nid (L tr) and maximum lingual length of the m1 talonid (L tld) in de los Toriles would highlight the potential of this site for the mm of Meles cf. thorali from Cueva de Los Toriles, compared with paleontology of the southern Iberian plateau. Future studies Plio-Pleistocene badgers and living Meles meles and Arctonyx colla- should confrm the results presented here with a more exten- ris. The pictures of the m1s of the diferent badgers are shown on a real scale, so their absolute proportions can be observed. The ellipse sive excavation of the fossil site. shows the range of variation of the considered specimens of Meles thorali published and the one from Cueva de Los Toriles. Source: (Mallye 2018): 1; (Arribas and Garrido 2007): 2, 3; (Vos et al. 2002): 5 Conclusions 3; This work: 4; (Viret 1939): 5; (Jiangzuo et al. 2018): 6; (Koufos 2018): 7; (Madurell-Malapeira et al. 2011a): 8; (Madurell-Malapeira et al. 2011b): 9; (Petrucci et al. 2013): 10; (Gasparik and Pazonyi The two new mustelid remains from Cueva de Los Toriles 2018): 11; (Baryshnikov 2009): 12 fossil site (Carrizosa, Castilla-La Mancha, Iberian Pen- insula) were found in reworked sediments but they could potentially be assessed as belonging to the stratum suggested the presence of Meles thorali from the locality of Tor·B·II, representing a Pleistocene deposit. The overall Unikoté (Late Pleistocene). However, the proportions of the morphology of the Cueva de los Toriles teeth, together m1 from Unikoté is closer to that of living M. meles, sug- with its proportions, indicate it difers from the living gesting a dubious asignation, clasifying it herein as M. cf. Euroasiatic badger (M. meles), presenting morphological thorali (Figs. 7, 8). The presence of Meles cf. thorali in the features that can be interpreted as more primitive traits

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Fig. 9 Lower carnassial (m1) of Meles cf. thorali from Cueva de los Toriles compared with other m1s of badgers (small diferences in the teeth orienta- tion should be considered when evaluating the difer- ent teeth depicted here). a, b Tor’19·B·Sup·507, right m1. a buccal view, b occlusal view; c, d FSL 40,188, right m1 of Arctomeles gennevauxi from Montpelier (France), Early Pliocene. c Buccal view, d occlusal view; e, f right m1 of an extant specimen of Meles meles housed at MNCN. e Buc- cal view, f occlusal view. Scale bar 2 cm

than those of the living Meles. These features include the excavation campaigns confrm this hypothesis, this would mesiodistally larger m1 protoconid and shorter m1 talonid, represent the second mention for this specie in the Iberian which in the lineage of the Eusoasiatic badgers, tend to Peninsula because it has only been previously found in increase the grinding area by an enlargement of the m1 Almenara-Casablanca 4 (Late Pliocene, MN16), represent- talonid and M1 talon (Ginsburg and Morales 2000). The ing the earliest record of Meles in Europe (Madurell-Mala- badger from Cueva de los Toriles also difers metrically peira et al. 2009). Besides, it is important to mention that the and morphologically to other extinct badgers from the Cueva de Los Toriles could represent one of the few fossil Iberian Peninsula and Europe. It difers from the Span- sites from the Late Villafranchian-Middle Pleistocene in the ish M. meles from Fuente Nueva-3 (Late Villafranchian, southern Iberian Plateau (García-Martínez 2019), along with Madurell-Malapeira et al. 2011a, b), M. meles atavus from those from Valverde-1, Fuensanta del Júcar and El Proven- Vallparadís Estació (Epivillafranchian, Madurell-Mala- cio (Aguirre 1989; Mazo 1990, 1999), dated around 0.9 Ma peira, et al. 2011a, b), the Italian M. meles from Grotta (Domínguez-Solera et al. 2020). Only future fndings on the Laceduzza (Late Pleistocene, Mecozzi, et al. 2019) and fauna and lithic industry can test that hypothesis. the Caucasian M. meles from Kudaro 1 and 3 caves (Late Pleistocene, Baryshnikov 2009). This is because the Meles Acknowledgements The authors thank Mr. Francisco León Garrido for kindly providing access to the cave to extract the fossils presented here, from Cueva de los Toriles has a more primitive dentition as well as Mr. Ramón Villa from the Junta de Comunidades de Castilla- with a smaller overall size and a smaller m1 talonid with La Mancha (JCCM) for permitting the study. We also thank Mr. Pedro less-developed cuspids to those of the former ones. Thus, Antonio Palomo, the Mayor of the Carrizosa locality, and his team the badger from the Cueva de los Toriles are compara- for kindly providing support while the stay of the paleontological and archaeological team in the village, J. Madurell-Malapeira (ICP) for the ble with those of the Late Pliocene and Early Pleistocene feedback about Plio-Pleistocene badgers from Europe, and S. Mayda M. thorali from Europe (Viret 1951; Bonifay 1971, 1981; (Ege University, ) for the pictures of Arctomeles gennevauxi Fosse 1996; Vos et al. 2002; Madurell-Malapeira et al. housed at FSL. We acknowledge Mr. Miguel García Martínez and Mrs. 2009), spanning cautiously to the Middle Pleistocene of Maria Navarro Navarro for contributing to the discovery of the cave and its preliminary exploration. The authors also acknowledge the fol- France (Fosse 1996; Mallye 2018). The badgers presented lowing curators for providing access to comparative material under here especially resemble the m1´s from the type locality their care: B. Petterson (FMNH), F. Zachos and A. Bibl (NHMW); D. of Saint Vallier in France (Early Pleistocene; Middle Vil- Kalthof (NRM), E. López Errasquin (MNCN). The “Juan de lafranchian, MN17; Viret 1951). Formación” program (FJCI-2017-32157), from the Spanish Ministry of Meles thorali Science, Innovation, and Universities funded also DGM. This research The presence of cf. in the Cueva de received support by A.V. from the SYNTHESYS3 Project https://www.​ los Toriles is here hypothesized. If future analyses and

1 3 Journal of Iberian Geology synth​esys.info/ (SYNTHESYS; AT-TAF-5457), which is fnanced by Arribas, A., & Garrido, G. (2007). Meles iberica n. sp., a new Eura- European Community Research Infrastructure Action under the FP7 sian badger (Mammalia, Carnivora, Mustelidae) from Fonelas “Capacities” Programme, and the European Union’s Seventh Frame- P-1 (Plio-Pleistoceneboundary, Guadix basin, Granada, Spain). work Programme (FP7/2007-2013) under Grant agreement no. 226506 Compts Rendus Palevol, 6, 545–555. (SYNTHESYS; SE-TAF-3637). A.V. was also co-funded by a Visit- Arroyo, A., & De la Torre, I. (2013). Acheulean large fake technol- ing Scholarship of Chicago Field Museum of Natural History (2016). ogy in Campo de Calatrava (Ciudad Real, Spain). , The support of the DST-NFR Centre of Excellence in Palaeosciences Ethnology, and Anthropology of Eurasia, 41(4), 2–10. https://doi.​ (CoE-Pal) toward this research for A.V. (COE2018-09POST and org/10.1016/j.aeae.2014.07.002. COE2019-PD07) is hereby acknowledged. Opinions expressed and Arsuaga, J. L., Peris, J. F., Gracia-Téllez, A., Quam, R., Carretero, J. conclusions arrived at, are those of the author and are not necessarily M., González, V. B., et al. (2012). Fossil human remains from to be attributed to the CoE-Pal. We acknowledge the Spanish Society Bolomor Cave (Valencia, Spain). Journal of Human Evolution, of Paleontology (SEP) for providing a research Grant (AJISEP 2018) 62(5), 629–639. https​://doi.org/10.1016/j.jhevo​l.2012.02.002. that contributed to carrying out the frst proper excavation campaign Arsuaga, J. L., Carretero, J. M., Lorenzo, C., Gómez-Olivencia, A., on the Cueva de Los Toriles. We acknowledge J. Madurell-Malapeira Pablos, A., Rodríguez, L., et al. (2015). Postcranial morphol- (ICP) and four anonymous reviewers for improving largely the quality ogy of the middle Pleistocene humans from Sima de los Huesos, of the text in previous versions of the manuscript. Spain. 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Afliations

Daniel García‑Martínez1,2,3 · Alberto Valenciano4,5 · Aitziber Suárez‑Bilbao6 · Carlos A. Palancar2 · Irene Megía García7 · Davinia Moreno1 · Isidoro Campaña1 · Pedro R. Moya‑Maleno3,8

Alberto Valenciano 4 Research and Exhibitions Department, Iziko Museums [email protected] of , PO Box 61, Cape Town 8000, South Africa Aitziber Suárez‑Bilbao 5 Department of Biological Sciences, Palaeobiological [email protected] Research Group, University of Cape Town, Private Bag X3, Rhodes Gift, Cape Town 7701, South Africa Pedro R. Moya‑Maleno [email protected] 6 Departamento de Estratigrafía Y Paleontología, Facultad de Ciencia Y Tecnología, Universidad del País Vasco, Leioa, 1 Centro Nacional Para El Estudio de La Evolución Humana Spain (CENIEH), Paseo Sierra de Atapuerca, 3, 09002 Burgos, 7 Departamento de Prehistoria Y Arqueología, Universidad Castilla‑León, Spain Autónoma de Madrid, Campus de Cantoblanco, 2 Paleoanthropology Group, Museo Nacional de Ciencias 28049 Madrid, Spain Naturales (CSIC), José Gutiérrez Abascal 2, 28006 Madrid, 8 Área de Prehistoria, Colaborador Honorífco, Facultad de Spain Geografía E Historia, Centro de Estudios del Campo de 3 Centro de Estudios del Campo de Montiel (CECM), Plaza Montiel (CECM), Universidad Complutense de Madrid, Mayor s/n, 13328 Almedina, Castilla‑La Mancha, Spain c/Profesor Aranguren S/N, 28040 Madrid, Spain

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