This is the accepted version of the journal article: López García, Juan Manuel; Piñero, Pedro; Agustí, Jordi; [et al.]. «Chronological context, species occurrence, and environmental remarks on the Gelasian site Pedr- era del Corral d’en Bruach (Barcelona, Spain) based on the small-mammal asso- ciations». Historical Biology, (April 2023). DOI 10.1080/08912963.2023.2180740 This version is available at https://ddd.uab.cat/record/272580 under the terms of the license 1 Chronological context, species occurrence, and environmental remarks on the Gelasian 2 site Pedrera del Corral d’en Bruach (Barcelona, Spain) based on the small-mammal 3 associations 4 Juan Manuel López-García1,2*, Pedro Piñero1,2, Jordi Agustí3,1,2, Marc Furió4,5, Julia 5 Galán6, Blanca Moncunill-Solé7, Francisco Javier Ruiz-Sánchez8, Hugues-Alexandre 6 Blain1,2, Montserrat Sánz9, Joan Daura9 1 7 IPHES-CERCA, Institut Català de Paleoecologia Humana i Evolució Social, Zona 8 Educacional 4, Campus Sescelades URV, Edifici W3, 43007, Tarragona, Spain. 9 jmlopez@iphes.cat; ppinero@iphes.cat; hablain@iphes.cat. 10 2 Universitat Rovira i Virgili, Departament d’Història i Història de l’Art, Avinguda de 11 Catalunya 35, 43002 Tarragona, Spain 3 12 Catalan Institution for Research and Advanced Studies (ICREA). 08010, Barcelona, 13 Spain. jordi.agusti@icrea.cat. 4 14 Serra Húnter Fellow. Facultat de Ciencies, Universitat Autònoma de Barcelona (UAB), 15 08193, Barcelona, Spain. marc.furio@uab.cat. 5 16 Institut Català de Paleontologia Miquel Crusafont. Universitat Autònoma de Barcelona 17 (UAB), 08193 Barcelona, Spain. 6 18 Departamento de Geología, Facultad de Ciencias y Tecnología. Universidad del País 19 Vasco/Euskal Herriko Unibertsitatea (UPV/EHU), Barrio Sarriena s/n, 48940, Leioa, 20 Spain, julia.galan@ehu.eus. 7 21 Grupo de Investigación en Cambio Ambiental (GRICA), Área de Paleontología, 22 Departamento de Física y Ciencias de la Tierra, Universidade da Coruña, 15008, A 23 Coruña, Spain, blanca.moncunill@udc.es. 8 24 PVC-GIUV (Palaeontology of Cenozoic Vertebrates Research Group). Àrea de 25 Palaeontologia, Universitat de València, Dr. Moliner 50, E-46100 Valencia, Spain. 26 Francisco.Ruiz@uv.es. 27 9 Grup de Recerca del Quaternari (GRQ-SERP), Departament d’Història i Arqueologia, 28 Universitat de Barcelona, Carrer Montalegre, 6, 08001 Barcelona, Spain. 29 montsesanzborras@ub.edu; jdaura_lujan@ub.edu 30 31 *Corresponding Author 32 Abstract 33 Here we present the results of a study of the small-mammal assemblage recovered from 34 a now-disappeared karstic fissure located in a quarry called “Pedrera del Corral d’en 35 Bruach”. The site was located at 330 m a.s.l. in the Garraf Massif, ca. 20 km south of 36 Barcelona. An archaeological rescue excavation was conducted in 2006 by the Grup de 37 Recerca del Quaternari, focusing on sieving the sediments that had accumulated at the 38 foot of the quarry face. Fortunately, some faunal remains and sediment were recovered, 39 which were later processed and sorted. Two hundred and seventy small-mammal remains 40 were identified at genus or species level, bringing to light great species richness (15 taxa). 41 The assemblage is typical of the Mammal Neogene 17 (MN17) biozone, placing this site 42 in the Gelasian stage of the Early Pleistocene. It is noteworthy that the first record of the 43 glirid Glirulus pusillus in Iberia is documented. On the basis of the small mammals 44 identified, the surrounding environment of the Garraf Massif is judged to have been 45 characterized by open forest landscape and more humid environmental conditions than 46 nowadays. Finally, the site provides a remarkable contribution to what is known of the 47 Iberian small mammals of this time period. 1 48 Keywords: Early Pleistocene, Eulipotyphla, Chiroptera, Lagomorpha, Rodentia, 49 Biochronology 50 1. Introduction 51 52 The Gelasian (ca. 2.5 – 1.8 Ma) is the first stage of the Pleistocene epoch (Cohen et al. 53 2020). This stage takes its name from the town of Gela in Sicily, which is near the Monte 54 San Nicola section, where the stage was defined (Río et al. 1994). The base of the 55 Gelasian stage coincides with Marine Isotope Stage 103 (MIS 103) and an increase in ice- 56 rafted detritus in the ocean sediment of the Northern Hemisphere (Raymo et al. 1989). 57 Furthermore, a series of changes in vegetation distribution patterns and migratory events 58 among large-mammal continental faunas took place at this time (Zagwijn 1974; Lona 59 1962; Lindsay et al. 1980; Azzaroli et al. 1988). It also coincides with the Gauss- 60 Matuyama magnetic polarity boundary, which occurred ca. 2.58 Ma (Suc et al. 1997). 61 This stage extends until ca. 1.8 Ma, more or less at the end of the Olduvai subchron, 62 matching the upper part of the Gelasian with the boundary falls between the highest 63 occurrence of Discoaster brouweri (below) and the lowest common occurrence of left- 64 coiling Neogloboquadrina pachyderma (above), and below the lowest occurrences of 65 medium-sized Gephyrocapsa (including G. oceanica) and Globigerinoides tenellus 66 (Gibbard and Head Martin 2012). Despite what is known of the Gelasian stage, there are 67 only a few sites in the Iberian Peninsula where the associated small-mammal fauna has 68 been recovered. Among these palaeontological sites are El Rincón-1 (Alberdi et al. 1997) 69 and Valdeganga 7 (Mein et al. 1978) in Albacete, Almenara-Casablanca-1 (Agustí et al. 70 2011) and Alto de la Cerdaña (Guillem-Calatayud 2013) in Castellón, Tollo de Chiclana 71 10 (Minwer-Barakat et al. 2004), Cortes de Baza- 1 (Peña 1975) and Galera G (Agustí et 72 al. 2001) in Granada, Alozaina in Málaga (Aguilar et al. 1993), Villarroya in La Rioja 73 (Laplana et al. 2016), and Islas Medes in Girona (Michaux 1971). This highlights the 74 importance of “Pedrera del Corral d’en Bruach” in the context of the ancient Early 75 Pleistocene, poorly represented in the Iberian Peninsula. 76 “Pedrera del Corral d’en Bruach” is a quarry situated 330 m a.s.l. in the Mesozoic Garraf 77 Massif, which is dominated by calcareous and dolomitic outcrops, both from the Lower 78 Cretaceous and the Jurassic (Fig. 1D-E). The site is located near the town of Gavà, ca. 20 79 km south of Barcelona (Fig. 1A-C). The site was discovered, together with other fissures, 80 by Joaquim Guillén, and these were published all together as Canal Negre 1 (Guillén 81 2010). The washing-sifting of the sediments in the 2006 rescure excavation campaing 82 (Fig. 1F-G), revelaed a considerable amount of small-mammal remains (Daura et al. 83 2007-2009). A first small-vertebrate identification was carried out and posted in the 84 excavation field report (Daura et al. 2007-2009). Among the large mammals recovered, 85 the presence of ursid, equid, rhinoceros, bovid and cercopithecine remains stands out. A 86 preliminary chronological estimate based on this large-mammal assemblage placed the 87 site within a range that could lie between MN16 and MN17 (ca. 2.5 Ma). In this context, 88 the main objective of our small-mammal study is to undertake a chronological approach 89 to the site and make some observations on the environment and the occurrence of species. 90 91 2. Material and methods 92 The small mammals of Pedrera del Corral d’en Bruach come from a now-disappeared 93 karstic fissure. They were recovered during the rescue field campaign conducted in 2006 2 94 by means of the water-screening of 18 tons of sediment from the fissure in question. Three 95 different meshes of 1.5, 0.8 and 0.5 mm were used for this procedure. The concentrated 96 sediment was then thoroughly sorted at the “La Guixera Laboratory” (Castelldefels City 97 Council) and the University of Barcelona (Daura et al. 2007-2009). The small-mammal 98 remains mainly consist of disarticulated bones and teeth. The insectivore and bat remains 99 are mainly composed of isolated teeth, fragmented mandibles, and humeri, whereas the 100 lagomorph and rodent remains are mainly composed of isolated teeth. The nomenclature 101 and measurements used for the identification of Soricidae follow Reumer (1984), and 102 those used for Talpidae follow Hutchinson (1974), for bats Sevilla (1988), for lagomorphs 103 Angelone (2005), for murids Pasquier (1974), for glirids Freudenthal (2004), for cricetids 104 Cuenca-Bescós (2003), and for arvicolines Van der Meulen (1973). 105 Our study of the small-mammal remains revealed 270 remains, corresponding to 55 106 individuals and comprising at least 15 species. Among these species, four insectivores 107 (Talpa cf. fossilis, Blarinoides mariae, Petenyia hungarica and Deinsdorfia doukasi), one 108 bat (Rhinolophus gr. ferrumequinum), one lagomorph (Prolagus cf. calpensis), and ten 109 rodents (Glis minor, Eliomys intermedius, Glirulus cf. pusillus, Muscardinus cf. 110 avellanarius, Apodemus atavus, Apodemus jeanteti, Castillomys rivas, Stephanomys 111 balcellsi, Allocricetus sp. and Mimomys medasensis) were identified (Table 1). The 112 previous preliminarily identified species (Daura et al. 2007-2009) of insectivores 113 (Asoriculus gibberodon), lagomorphs (Prolagus michaux, P. ibericus and Trischizolagus) 114 and rodents (Stephanomys minor, Apodemus gorafensis, A. mystacinus, Rhagapodemus, 115 Castillomys gracilis, C. crusafonti, Mimomys stehlini, Kislangia sp. and Allophaiomys 116 sp.) were not recognized in the assemblage from Pedrera del Corral d’en Bruach. 117 Therefore, the preliminary list should be discarded, considering as valid those identified 118 in the present work. 119 120 3. Systematic Paleontology 121 Order EULIPOTYPHLA Waddell, Okada and Hasegawa, 1999 122 Family TALPIDAE Fischer, 1814 123 Genus Talpa Linnaeus, 1758 124 Talpa cf. fossilis Petényi, 1864 125 Figure 2A 126 Material. Three humeri (PCB-Ta-1 [w13= 3.87 mm], PCB-Ta-2 [w13= 3.42 mm], PCB- 127 Ta-3 [w13= 3.51 mm]) (partially fragmented). 128 Description. The three elements recovered are poorly preserved, lacking all the important 129 parts of their epiphyses, such as the head of the humeri, the entepicondyles and the 130 ectepicondyles. The most important, clearly evident trait observed in these humeri is the 131 short, wide diaphysis. The olecranon fossa is rather deep, and the trochlea is well 132 delimited. The rest of the morphological characters are too weakly preserved to be 133 described. 3 134 Remarks. The morphology of the humerus in burrowing moles is unequivocal. The 135 extreme modifications of this element for efficient excavation result in many surfaces and 136 muscular attachment points with no equivalent among other vertebrates. Sadly, the 137 classification of the fossil burrowing talpids of Europe is unclear. Thousands of fossil 138 humeri from Plio-Pleistocene moles are stored in collections in all the European 139 institutions, tentatively labelled under several different names such as Talpa minor, Talpa 140 fossilis, Talpa europaea, Talpa gracilis, Talpa episcopalis or Talpa caeca, awaiting 141 definitive identification. Van Cleef-Roders and Van den Hoek Ostende (2001) revised the 142 mandibles and dentition of two extant species of Talpa, later discussing the implications 143 for the classification of the fossil elements of this genus. Unfortunately, their useful 144 observations and comments were not pursued any further by other authors. Thus, the 145 classification of fossil forms is still too weak and inconsistent to provide a solid specific 146 identification here. The size and the morphology of the three elements found fit within 147 the variation of Talpa fossilis, a species quite frequently cited within the specific literature 148 of Plio-Pleistocene small mammals. We are aware that Van Cleef-Roders and Van den 149 Hoek Ostende (2001) considered this form to be a likely junior synonym of T. europaea. 150 However, Sansalone et al. (2015) found significant differences in size and shape of their 151 corresponding humeri after a geometric morphometric analysis. In the light of this later 152 study, we feel it prudent to ascribe tentatively the material from Pedrera del Corral to the 153 fossil species. 154 155 Family SORICIDAE Fischer, 1817 156 Genus Blarinoides Sulimski, 1959 157 Blarinoides mariae Sulimski, 1959 158 Figure 2B 159 Material. One right m2 (PCB-Bl-1) (L=2.20; TRW=1.34; TAW=1.39) in a fragmented 160 hemimandible. 161 Description. The only tooth preserved is an m2. The tooth shows intense red pigmentation 162 covering the upper half of the labial face. The protoconid is the highest cuspid. Paraconid, 163 metaconid, entoconid and hypoconid are of similar height, all of them being lower than 164 the protoconid. There is no clear entoconid cristid. In the fragmented hemimandible, the 165 only useful character is the position of the mental foramen, which is placed between the 166 anterior and the posterior alveoli of the m1. 167 Remarks. The occurrences of the genus Blarinoides in the Iberian Peninsula have been 168 excellently analysed by Minwer-Barakat et al. (2007), who described B. aliciae, a new 169 endemic species. This Spanish form of Blarinoides is clearly smaller than its European 170 congeneric counterpart, B. mariae. The material from Pedrera del Corral d’en Bruach fits 171 perfectly within the range of the latter species. The large size of the only m2 recovered 172 precludes ascription to any other known Pliocene shrew, with the possible exception of 173 Beremendia fissidens. However, the rather irregular morphology of the labial cingulid of 174 the m2, the shape of the entoconid, and the alveoli of the m1 located more ventrally than 175 the base of the m2, support ascription to Blarinoides. The known stratigraphic range for 4 176 Blarinoides covers the Pliocene and the Early Pleistocene of Europe (Reumer 1984; 177 Rzebik-Kowalska 1998; Minwer-Barakat et al. 2007). However, according to the data 178 compiled by Furió et al. (2018), its occurrence in the Iberian Peninsula seems to have 179 been more short-lived, apparently constrained to the late Ruscinian (MN15) and the early 180 Villanyian (MN16). 181 Genus Petenyia Kormos, 1934 182 Petenyia hungarica Kormos, 1934 183 Figure 2C 184 Material. One right m2 (PCB-Pe-1) (L=1.26; TRW=0.75; TAW=0.81) in a fragment of 185 hemimandible. 186 Description. The upper zones on the labial side are distinctly covered with red 187 pigmentation. The talonid is slightly longer than the trigonid. In occlusal view, the outline 188 of the tooth is quadrangular. The base of the tooth is covered by a conspicuous cingulid, 189 especially at its mesial, labial, and distal margins. There is a rather stout entoconid in an 190 advanced position on the lingual side of the talonid. 191 Remarks. Although this is a rather frequent form in some European sites of Pliocene age, 192 Petenyia hungarica had only been reported in Spain three times before. Its occurrences 193 at Moreda (MN16; Ruiz Bustos 2002), Tollo de Chiclana 13 (MN16; Minwer-Barakat et 194 al. 2010) and Almenara-Casablanca 1 (MN17; Agustí et al. 2011) suggest that P. 195 hungarica is a form indicative of Villanyian ages in the Iberian Peninsula. 196 197 Genus Deinsdorfia Heller, 1963 198 Deinsdorfia doukasi Furió and Mein, 2008 199 Figure 2D, 3A-B 200 Material. One right m1 (PCB-De-1) (L=1.50; TRW=0.74; TAW=0.81), one right m2 201 (PCB-De-2) (L=1.37; TRW=0.74; TAW=0.74), one right m3 (PCB-De-3) (L=0.98; 202 W=0.59), one right I1 (PCB-De-4), one left i1 (PCB-De-5) (broken). 203 Description. All the dental elements recovered from Pedrera del Corral d’en Bruach are 204 stained with dark red pigmentation at the tips of their main cusps or cuspules. The most 205 significant diagnostic traits are the asymmetric and tilted cuspules of the i1, the non- 206 fissident I1 with an enlarged apex, the lower molars with rather short talonids and long 207 paralophids (especially in m1), and the m3 with a single-cusped talonid. 208 Remarks. The genus Deinsdorfia is a typical element in the Pliocene and Early 209 Pleistocene of Europe (Reumer 1984; Rzebik-Kowalska 1998). The Spanish finds of this 210 genus were revised and described by Furió and Mein (2008), resulting in the description 211 of an endemic Iberian species, namely Deinsdorfia doukasi. Nonetheless, the genus is 212 hitherto poorly documented in Spain, probably because many authors working on 213 Pliocene sites have confused its teeth with Sorex spp. remains. According to Furió et al. 214 (2018), in Spain this species is indicative of a Villanyian age (MN16-MN17). However, 5 215 this rough guideline could change in the future, once some Ruscinian or Early Pleistocene 216 sites have been revised. 217 218 Order CHIROPTERA Blumenbach, 1779 219 Family RHINOLOPHIDAE Gray, 1825 220 Genus Rhinolophus Lacépède, 1799 221 Rhinolophus gr. ferrumequinum (Schreber, 1774) 222 Figure 2E 223 Material. One left m1 (PCB-Rh-1) (L= 2.03 mm; W1=1.31 mm; W2=1.32 mm) 224 Description. The tooth recovered from Pedrera del Corral d’en Bruach shows in occlusal 225 view the entoconid connected with the hypoconulid in the talonid part of the tooth and in 226 labial view a cingulum that is thin and regular. These are characteristics of the genus 227 Rhinolophus. The molar measurements lie within the size range of the extant R. 228 ferrumequinum population, being larger than R. mehelyi (Sevilla 1988). 229 Remarks. There are two fossil species in the R. ferrumequinum group in Europe that are 230 described in the Gelasian and Calabrian Early Pleistocene: R. macrorhinus and R. 231 postdelphinensis (Topál 1979). Unfortunately, the diagnostic characters of these two 232 species pertain to the upper dental series, so with only one lower molar we can neither 233 rule out nor affirm that it is either of these species. In any case, in the Iberian Peninsula 234 there are no records of these two species to date, although R. ferrumequinum has been 235 identified in the Gelasian stage of Almenara-Casablanca 1 (Agustí et al. 2011). In light 236 of these data, and given the lack of diagnostic material, our specimen has been assigned 237 to the R. ferrumequinum group. 238 239 Order LAGOMORPHA Brandt, 1855 240 Family OCHOTONIDAE Thomas, 1897 241 Genus Prolagus Pomel, 1853 242 Prolagus cf. calpensis Major, 1905 243 Figure 2F 244 Material. One right p3 (PCB-Pr-1) (L=1.94 mm; W=1.57 mm) 245 Description. The only tooth recovered from Pedrera del Corral d’en Bruach pertaining to 246 the family Ochotonidae is a third lower premolar. Morphologically, this presents in 247 occlusal view a reduced entoconid, a “crochet” that is lost, and a thin protoconulid. These 248 morphological characteristics are also described in the specimens of P. cf. calpensis 249 recovered from the Gelasian sites of Almenara-Casablanca 1 (Gil and Sesé 1984), Moreda 250 and Islas Medes (López-Martínez 1989), and Fonelas P1 (Laplana and Blain 2008). The 6 251 measurements of our tooth also coincide with those of the P. cf. calpensis specimens from 252 the above-mentioned sites (Gil and Sesé 1984; López-Martínez 1989; Angelone 2005). 253 Remarks. Taking into account the sites referred to above together with El Rincón-1 254 (Alberdi et al. 1997), the presence of P. cf. calpensis in the Iberian sites seems to be 255 related with the Gelasian stage of the Early Pleistocene, lying within biozone MN17 and 256 indicating a chronology between 2.6 and 1.8 Ma (Sesé 2006). 257 258 Order RODENTIA Bowdich, 1821 259 Family GLIRIDAE Muirhead, 1819 260 Genus Glis Brisson 1762 261 Glis minor Kowalski, 1956 262 Figure 4A–G 263 Material. One right p4 (PCB-Gm-5) and one left p4 (PCB-Gm-32), two right m1 (PCB- 264 Gm-24; PCB-Gm-30), three left m1 (PCB-Gm-9; PCB-Gm-39; PCB-Gm-42), one left 265 m2 (PCB-Gm-38), one right m3 (PCB-Gm-7), one left m3 (PCB-Gm-46), one left P4 266 (PCB-Gm-3), two left M1 (PCB-Gm-47; PCB-Gm-14), one left M2 (PCB-Gm-30) and 267 one left M3 (PCB-Gm-18) 268 Description. p4: The occlusal pattern is composed of 6 ridges: anterolophid, metalophid, 269 centrolophid, mesolophid, posterotropid and posterolophid. In one specimen (PCB-Gm- 270 5) a small anterotropid can be recognized (Fig. 4E). Anterolophid and protolophid are 271 labially connected. The centrolophid is isolated. The posterolophid can be isolated or 272 lingually connected to the mesolophid. The posterotropid remains isolated; m1: The 273 occlusal pattern comprises 7 ridges: anterolophid, anterotropid, metalophid, centrolophid, 274 mesolophid, posterotropid and posterolophid. Anterolophid, anterotropid and metalophid 275 are lingually connected, although in one case the anterotropid remains isolated. 276 Centrolophid and metalophid are isolated. The posterolophid is connected lingually to the 277 posterotropid, although in two cases this ridge remains isolated; m2: The occlusal pattern 278 comprises 7 ridges.The anterolophid can be isolated or lingually connected to the 279 metalophid. The anterotropid remains. Centrolophid and mesolophid are isolated. The 280 posterotropid remains isolated; m3: As in the other lower molars, the occlusal pattern 281 comprises 7 ridges. The anterolophid is lingually connected to the metalophid. The 282 anterotropid can also be lingually connected to the anterolophid or remain isolated. The 283 centrolophid and mesolophid are isolated. Posterolophid and posterotropid are lingually 284 connected; P4: The occlusal pattern is composed of 6 main ridges: anteroloph, protoloph, 285 centroloph, metaloph, posterotrope and posteroloph. A tiny anterotrope can also be 286 distinguished. The centroloph is discontinuous and does not reach the labial wall of the 287 teeth. Centroloph and posterotrope are labially connected. The metaloph is also 288 discontinuous. The posterotrope is labially connected to the posteroloph; M1: The 289 occlusal pattern is composed of 7 ridges: anteroloph, anterotrope, protoloph, centroloph, 290 metaloph, posterotrope and posteroloph. All the ridges are isolated, with the exception of 291 the posterotrope and posteroloph, which are labially connected; M2: The pattern is the 292 same as in the M1. However, the posterotrope can reach the posteroloph; M3: The 7 293 occlusal pattern is composed of 7 main ridges.. All of these are lingually connected by a 294 continuous endoloph. Anteroloph, protoloph and centroloph are also labially connected, 295 though not fused. Metaloph and posteroloph are labially fused. 296 Remarks. Our specimens (Table 2) coincide with the measurements of G. minor from 297 central Europe (e.g. Striczky and Pazonyi 2014). Glis minor is a species described on the 298 basis of Pliocene material from Podlesice (Poland) by Kowalski (1959). This species is 299 well documented in the Early Pleistocene (Gelasian and Calabrian stages) in central and 300 southern Europe, at sites such as Osztramos 7 (Janossy 1986), Rivoli Veronese (Berto et 301 al. 2022), Le Vallonnet (Lumley et al. 1988), Betfia 13 (Terzea 1996) and Les Valerots 302 (Bonifay 1973), but until now there has been no known reference to the species in the 303 Iberian Peninsula. Thus, the presence of G. minor in Pedrera del Corral d’en Bruach 304 constitutes the first record of the species known to date in Iberia. 305 306 Genus Eliomys Wagner, 1849 307 Eliomys intermedius Friant, 1953 308 Figure 4H–K 309 Material. One right m2 (PCB-Ein-22), one right P4 (PCB-Ein-43), one right M1-M2 310 (PCB-Ein-11), two left M1-M2 (PCB-Ein-1; PCB-Ein-13), one right M3 (PCB-Ein-14), 311 four left M3 (PCB-Ein-17; PCB-Ein-30; PCB-Ein-29; PCB-Ein-19). 312 Description. m2: The occlusal pattern is composed of 5 ridges: anterolophid, metalophid, 313 centrolophid, mesolophid and posterolophid. The anterolophid and the centrolophid are 314 lingually connected. The metalophid is isolated. Mesolophid and posterolophid are also 315 lingually fused and with an accessory crest between them; P4: A small anteroloph is 316 present on the labial side of the anterior wall of the tooth. Paracone and metacone are 317 well-individualized cusps. The centroloph is reduced to a number of cuspules between 318 the protoloph and the metaloph. The posteroloph is long. Protoloph, metaloph, and 319 posteroloph are lingually connected by a continuous endoloph; M1-M2: There are four 320 main ridges: anteroloph, protoloph, metaloph and anteroloph, all of them lingually 321 connected by a continuous endoloph. Paracone and metacone are well marked. 322 Precentroloph and postcentroloph are present in one tooth, being lingually connected. In 323 the other two the postcentroloph is lacking, and only a short, isolated precentroloph is 324 present; M3: This is trapezoidal in shape. As in the case of the other upper molars, the 325 occlusal pattern is composed of 4 main ridges, all of them lingually connected by a 326 continuous endoloph. The anteroloph is the longest ridge, whereas the posteroloph is the 327 shortest. Paracone and metacone are not detached from protoloph and metaloph, 328 respectively. A short precentroloph is present in three teeth, whereas a short 329 postcentroloph is present in only one tooth. Both precentroloph and postcentroloph are 330 connected labially to the protoloph and the metaloph. All the morphological and metric 331 (Table 3) characters identified in our specimens coincide with the description provided 332 for Eliomys intermedius by García-Alix et al. (2008a). 333 Remarks. The first mentions of E. intermedius in the Iberian Peninsula are from Early 334 Pliocene (MN14) sites located in the Teruel Basin, the Cabriel and Alcoy Basins, and the 335 Granada and Guadix Basins (e.g. Masino et al. 2015). The species just reaches the Late 8 336 Pliocene (MN16) in sites such as Tollo Chiclana 3 and Almenara-Casablanca 4 (Agustí et 337 al. 2011) and the Early Pleistocene (Calabrian age) in the Sierra de Quibas (Montoya et 338 al. 2001), where it coexists with the current species E. quercinus (Piñero et al. 2015). 339 340 Genus Glirulus Thomas, 1906 341 Glirulus cf. pusillus (Heller, 1936) 342 Figure 4M 343 Material. One right m2 (PCB-Gpu-23) (L=0.99 mm; W=0.86 mm) 344 Description. The occlusal pattern is composed of 7 main ridges: anterolophid, 345 anterotropid, metalophid, centrolophid, mesolophid, posterotropid and posterolophid. A 346 small extra ridge is present between anterotropid and metalophid. In addition, two long 347 extra ridges are located on the two sides of the centrolophid. Anterolophid, metalophid, 348 centrolophid, mesolophid and posterolophid are lingually connected by a continuous 349 endolophid. The labial ends of the metalophid, mesolophid and posterolophid are 350 projected forwards. All these morphological characteristics are described in the species 351 Glirulus pusillus identified in central Europe (Van den Hoek-Oestende 2003). The 352 measurements of our specimens also coincide with those of the central European 353 specimens belonging to G. pusillus (Van den Hoek-Oestende 2003). Despite these factors, 354 due to the scarcity of the identified material, we have decided to assign our specimens to 355 G. cf. pusillus. 356 Remarks. G. pusillus is a rare component of Late Pliocene and Early Pleistocene faunas 357 in Europe (Nadachowski and Daoud 1995). G. pusillus specimens from the Late Pliocene 358 and Early Pleistocene differ only in size from the extant species, G. japonicus (which 359 currently only inhabits Japan), since the teeth of the current species are larger than those 360 of G. pusillus. Pedrera del Corral d’en Bruach is the first finding of G. pusillus in the 361 Iberian Peninsula. 362 Genus Muscardinus Kaup, 1829 363 Muscardinus cf. avellanarius (Lianneus, 1758) 364 Figure 4L 365 Material. One right m1 (PCB-Mu-35) (L=1.61 mm; W=1.35 mm) 366 Description. The tooth presents a subrectangular occlusal surface, with the posterior part 367 of the molar wider than the front part. In occlusal view, it presents 6 ridges: anterolophid, 368 anterotropid, metalophid, centrolophid, mesolophid and posterlophid. All of these are 369 isolated, with the exception of the anterolophid and the anterotropid, which are labially 370 and lingually fused. The above-mentioned morphological characters could be assigned to 371 the two Early Pleistocene Muscardinus species present in the Iberian Peninsula, M. 372 pliocaenicus and M. avellanarius (García-Alix et al. 2008b). According to Nadachowski 373 and Daoud (1995), the increase in size of the molars from M. pliocaenicus to M. 374 avellanarius could be a diagnostic feature separating the two species. Our single 375 specimen presents a length and width that lie within the measurement parameters of M. 9 376 avellanarius. It contrasts, for example, with the species identified in Somssich Hill 2 in 377 Hungary by Striczky and Pazonyi (2014) and Tegelen in the Netherlands by Van den Hoek 378 Ostende (2003), being larger than the specimens of M. pliocaenicus identified in these 379 deposits. Despite this and due to the scarcity of the identified material, we have decided 380 to assign our specimen to M. cf. avellanarius. 381 Remarks. According to García-Alix et al. (2008b), M. pliocaenicus and M. avellanarius 382 belong to the same lineage, and although these species are well documented in central 383 Europe (e.g. Nadachowski and Daoud 1995; Van den Hoek Ostende 2003; Striczky and 384 Pazonyi 2014) during the end of the Pliocene and the Early Pleistocene, the tooth in 385 Pedrera del Corral d’en Bruach is confirmed as belonging to MN17 by the result of the 386 small-mammal association as a whole. This could be the oldest citation known to date of 387 the species M. avellanarius in the Iberian Peninsula. 388 389 Family MURIDAE Illiger, 1811 390 Genus Apodemus Kaup, 1826 391 Description. Pedrera del Corral d’en Bruach has yielded 198 teeth displaying the common 392 features of the genus Apodemus: the presence of t7 and the t6-t9 connection in the M1 393 and M2, and a well-developed labial cingulum and the absence of a longitudinal crest in 394 the m1 and m2. 395 396 Apodemus atavus Heller, 1936 397 Figure 5A–L 398 Material. 12 left M1 (PCB-Aat-1, PCB-Aat-4, PCB-Aat-6, PCB-Aat-7, PCB-Aat-11, 399 PCB-Aat-12, PCB-Aat-13, PCB-Aat-14, PCB-Aat-22, PCB-Aat-25, PCB-Aat-26, PCB- 400 Aat-28), 19 right M1 (PCB-Aat-2, PCB-Aat-3, PCB-Aat-5, PCB-Aat-8, PCB-Aat-9, 401 PCB-Aat-10, PCB-Aat-15, PCB-Aat-16, PCB-Aat-17, PCB-Aat-18, PCB-Aat-19, PCB- 402 Aat-20, PCB-Aat-21, PCB-Aat-23, PCB-Aat-24, PCB-Aat-27, PCB-Aat-29, PCB-Aat- 403 30, PCB-Aat-31), 14 left M2 (PCB-Aat-32, PCB-Aat-33, PCB-Aat-35, PCB-Aat-36, 404 PCB-Aat-39, PCB-Aat-42, PCB-Aat-44, PCB-Aat-45, PCB-Aat-46, PCB-Aat-47, PCB- 405 Aat-48, PCB-Aat-49, PCB-Aat-50, PCB-Aat-51), six right M2 (PCB-Aat-34, PCB-Aat- 406 37, PCB-Aat-38, PCB-Aat-40, PCB-Aat-41, PCB-Aat-43), four left M3 (PCB-Aat-52, 407 PCB-Aat-53, PCB-Aat-57, PCB-Aat-58), three right M3 (PCB-Aat-54, PCB-Aat-55, 408 PCB-Aat-56), 16 left m1 (PCB-Aat-59, PCB-Aat-62, PCB-Aat-63, PCB-Aat-65, PCB- 409 Aat-66, PCB-Aat-69, PCB-Aat-71, PCB-Aat-73, PCB-Aat-76, PCB-Aat-80, PCB-Aat- 410 83, PCB-Aat-84, PCB-Aat-85, PCB-Aat-86, PCB-Aat-87, PCB-Aat-88), 14 right m1 411 (PCB-Aat-60, PCB-Aat-61, PCB-Aat-64, PCB-Aat-67, PCB-Aat-68, PCB-Aat-70, PCB- 412 Aat-72, PCB-Aat-74, PCB-Aat-75, PCB-Aat-77, PCB-Aat-78, PCB-Aat-79, PCB-Aat- 413 81, PCB-Aat-82), 13 left m2 (PCB-Aat-90, PCB-Aat-91, PCB-Aat-93, PCB-Aat-97, 414 PCB-Aat-100, PCB-Aat-102, PCB-Aat-103, PCB-Aat-107, PCB-Aat-110, PCB-Aat- 415 113, PCB-Aat-115, PCB-Aat-116, PCB-Aat-118), 17 right m2 (PCB-Aat-89, PCB-Aat- 416 92, PCB-Aat-94, PCB-Aat-95, PCB-Aat-96, PCB-Aat-98, PCB-Aat-99, PCB-Aat-101, 417 PCB-Aat-104, PCB-Aat-105, PCB-Aat-106, PCB-Aat-108, PCB-Aat-109, PCB-Aat- 418 111, PCB-Aat-112, PCB-Aat-114, PCB-Aat-117), six left m3 (PCB-Aat-119, PCB-Aat- 10 419 120, PCB-Aat-124, PCB-Aat-126, PCB-Aat-127, PCB-Aat-128), four right m3 (PCB- 420 Aat-121, PCB-Aat-122, PCB-Aat-123, PCB-Aat-125). 421 Description. The sample from Pedrera del Corral d'en Bruach includes 128 specimens of 422 a small-sized Apodemus species. The teeth show the typical traits of Apodemus atavus 423 (see Heller 1936): small size, the protoconid-metaconid pair connected to the lingual lobe 424 of the anteroconid, and the presence of accessory labial cuspids, a well-developed tma 425 and the presence of a small root, between the proximal and distal roots in some specimens 426 on the m1, a large posterior heel on m2 protruding beyond the outline, and a frequent 427 deep separation between the t4 and the t7 on the M1 and M2. The t12 is well developed, 428 and the t6 is joined to, or in contact with, the t9 in both the M1 and M2. Although 429 Rietschel and Storch (1974) proposed the presence of a large posterior accessory cuspid 430 (c1) as a diagnostic criterion of Apodemus atavus, Martín-Suárez and Mein (2004) ruled 431 this out as a diagnostic feature when they observed that the c1 is highly variable in size 432 among the m2s of this species from the localities of Schernfeld (Germany) and Balaruc 2 433 (France). We can corroborate this high variability: some specimens from Pedrera del 434 Corral d’en Bruach show a large c1 (33%), whereas in other specimens it can be small or 435 barely visible. It is even absent in five out of 30 cases. The size of the studied material 436 (Table 4) is very similar to that of the samples of Apodemus atavus from Tollo de 437 Chiclana 3 and 13 (Spain; Minwer-Barakat et al. 2005), Cañada del Castaño 1 (Spain; 438 Martín-Suárez 1988), Rambla Seca-A, Belmez-1 (Spain; Castillo Ruiz 1990), Alozaina 439 (Spain; Aguilar et al. 1993a), Calicasas 3 and 3B (Spain; García-Alix et al. 2008c), 440 Schernfeld (Germany; Pasquier 1974), Hambach (Germany; Mörs et al. 1998), Balaruc 441 6, Lo Fournas 4, Mas Rambault 2 (France; Bachelet 1990), Grand Serre (France; Aguilar 442 et al. 1993b), Monte Peglia (Italy; Meulen 1973), Monte la Mesa (Italy; Marchetti et al. 443 2000), and Notio 1 (Greece; Hordijk and de Bruijn 2009), among other sites. In 444 accordance with both morphological and biometric criteria, therefore, the studied molars 445 are assigned to Apodemus atavus. 446 Remarks. Several authors have considered Apodemus atavus and Apodemus dominans to 447 be extreme phenotypes of a single species, A. dominans being a junior synonym of A. 448 atavus (Fejfar and Storch 1990; Martín-Suárez and Mein 2004; Minwer-Barakat et al. 449 2005; García-Alix et al. 2008c; Colombero et al. 2014). The wide geographic distribution 450 of Apodemus atavus includes much of the Palearctic region, the species having been 451 recorded from Western Europe through Georgia to China (Cai and Qiu 1993; Martín- 452 Suárez and Mein 2004; Knitlová and Horáček 2017; Agustí et al. 2022). The time range 453 of this species extends from the latest Miocene to the Early Pleistocene (Rietschel and 454 Storch 1974; Fejfar and Storch 1990; Minwer-Barakat et al. 2005; Sala and Masini 2007; 455 García-Alix et al. 2008c; Colombero et al. 2014). The tooth morphology of Apodemus 456 atavus is relatively conservative, with little variation between the oldest and the youngest 457 populations (Martín-Suárez and Mein 2004; Minwer-Barakat et al. 2005). Apodemus 458 atavus is considered by some authors to be a direct ancestor of the extant Apodemus 459 sylvaticus (Rietschel and Storch 1974; Fejfar and Storch 1990; Martín-Suárez and Mein 460 1998; Piñero et al. 2022). 461 462 Apodemus jeanteti Michaux, 1967 463 Figure 5M–Y 464 Material. Eight left M1 (PCB-Aje-38, PCB-Aje-40, PCB-Aje-41, PCB-Aje-44, PCB- 465 Aje-45, PCB-Aje-49, PCB-Aje-51, PCB-Aje-52), nine right M1 (PCB-Aje-37, PCB-Aje- 11 466 39, PCB-Aje-42, PCB-Aje-43, PCB-Aje-46, PCB-Aje-47, PCB-Aje-48, PCB-Aje-50, 467 PCB-Aje-53), seven right M2 (PCB-Aje-54, PCB-Aje-57, PCB-Aje-59, PCB-Aje-61, 468 PCB-Aje-62, PCB-Aje-64, PCB-Aje-66), six left M2 (PCB-Aje-55, PCB-Aje-56, PCB- 469 Aje-58, PCB-Aje-60, PCB-Aje-63, PCB-Aje-65), one right M3 (PCB-Aje-68), three left 470 M3 (PCB-Aje-67, PCB-Aje-69, PCB-Aje-70), six right m1 (PCB-Aje-1, PCB-Aje-8, 471 PCB-Aje-9, PCB-Aje-12, PCB-Aje-14, PCB-Aje-15), nine left m1 (PCB-Aje-2, PCB- 472 Aje-3, PCB-Aje-4, PCB-Aje-5, PCB-Aje-6, PCB-Aje-7, PCB-Aje-10, PCB-Aje-11, 473 PCB-Aje-13), nine right m2 (PCB-Aje-17, PCB-Aje-18, PCB-Aje-19, PCB-Aje-20, 474 PCB-Aje-21, PCB-Aje-22, PCB-Aje-23, PCB-Aje-25, PCB-Aje-28), five left m2 (PCB- 475 Aje-16, PCB-Aje-24, PCB-Aje-26, PCB-Aje-27, PCB-Aje-29), five right m3 (PCB-Aje- 476 30, PCB-Aje-31, PCB-Aje-32, PCB-Aje-34, PCB-Aje-35), two left m3 (PCB-Aje-33, 477 PCB-Aje-36). 478 Description. The sample from Pedrera del Corral d'en Bruach includes 65 specimens of a 479 large-sized Apodemus species. The crown of these molars is higher than that of other 480 Apodemus species, which may lead to this sample being confused with Rhagapodemus. 481 However, the absence of tma in the studied m1 rules this option out in favour of the 482 species Apodemus jeanteti. The m1 and m2 have a well-developed labial cingulum with 483 high accessory labial cuspids. In the m1, the round or oval posterior heel is somewhat 484 smaller than in the m2. The m2 has a large labial anteroconid, whereas this is small or 485 absent in the m3. In the M1, the t1, t2 and t3 are almost aligned. The t12 is of medium 486 size in the M1 and small or absent in the M2. Overall, the t4 and t7 are separated, whereas 487 the t6 and t9 are connected in both the M1 and M2. In the M1, there is a posterior spur on 488 the t3, but this is absent on the t1 (except for PCB-Aje-39 and PCB-Aje-51), this latter 489 cusp being isolated in seven out of 15 cases. In the M2, the t9 is present in all but two 490 specimens, and the t1 and t3 are isolated. The t3 is absent in the M3. All these 491 morphological features are characteristic of Apodemus jeanteti (see Michaux 1967; 492 Pasquier 1974). The size of the specimens from Pedrera del Corral d'en Bruach (Table 5) 493 is close to that of Apodemus gorafensis. Nevertheless, the studied teeth can undoubtedly 494 be attributed to Apodemus jeanteti given the absence of tma in the m1. In addition, this 495 material coincides with the size of Apodemus jeanteti from its type locality Seynes, 496 France (Michaux 1967; Pasquier 1974), although the latter has somewhat higher mean 497 values. Similarly, the specimens from Pedrera del Corral d'en Bruach are close in size to 498 Apodemus jeanteti from Mont-Helene, Lo Fournas 13, Pla de la Ville, Nimes, Seynes, 499 Grand Serre, Mas Rambault 2 (southern France; Bachelet 1990; Pasquier 1974; Aguilar 500 et al. 2002), Moreda 1A and 1B, Belmez-1 and Alozaina (southern Spain; Castillo Ruiz 501 1990; Aguilar et al. 1993a). 502 Remarks. Apodemus jeanteti probably evolved from Apodemus gorafensis during the late 503 Ruscinian (MN15) (Martín-Suárez and Mein 1998). In the Early Pliocene, Apodemus 504 gorafensis also gave rise to Apodemus agustii, a species only recorded at its type locality, 505 Cañada del Castaño, Spain (Martín Suárez 1988). Apodemus jeanteti has been found in 506 both Spain and southern France (Pasquier 1974; Gil and Sesé 1985; Adrover 1986; 507 Bachelet 1990; Castillo Ruiz 1990; Aguilar et al. 1993a, 1993b, 2002). Its temporal 508 distribution extends as far as the Early Pleistocene, the species probably becoming extinct 509 before the onset of the Olduvai subchron (1.94–1.78 Ma). Apodemus jeanteti and 510 Apodemus atavus were replaced by Apodemus mystacinus, Apodemus sylvaticus and 511 Apodemus flavicollis in southern Europe in the course of the Early Pleistocene (Michaux 512 and Pasquier 1974; Martín-Suárez and Mein 1998; Piñero et al. 2022). 513 514 Genus Castillomys Michaux, 1969 12 515 Castillomys rivas Martín-Suárez and Mein, 1991 516 Figure 6A–G 517 Material. One right M2 (PCB-Cri-1), one left M2 (PCB-Cri-2), two left m1 (PCB-Cri-3, 518 PCB-Cri-4), two right m2 (PCB-Cri-5, PCB-Cri-7), one left m2 (PCB-Cri-6), one left m3 519 (PCB-Cri-8). 520 Description. The genus Castillomys is represented at Pedrera del Corral d'en Bruach by 521 only eight specimens. The small size, the absence of t7 and the development of a posterior 522 crest on the t1 in the M2, and the well-developed longitudinal crest on the m1 are 523 distinctive features of Castillomys. All the specimens have well-developed longitudinal 524 crests, completing the connection among the tubercles of the crown. The m1 and m2 have 525 a broad labial cingulum separated from the protoconid by a valley. These morphological 526 characters are present in the species Castillomys rivas. Moreover, these molars agree in 527 size (Table 6) with Castillomys rivas from Loma Quemada-1, its type locality (Spain; 528 Martín-Suárez and Mein 1991), as well as with Castillomys rivas from Quibas (Piñero et 529 al. 2015, 2022), Valdeganga 7 (Martín-Suárez and Mein 1991), Venta Micena 1, Orce 7 530 (Martín-Suárez 1988), Huétor Tájar 1 and 8 (García-Alix et al. 2009), and Tollo de 531 Chiclana 10B (Minwer-Barakat et al. 2005), among other sites. The studied specimens 532 can be distinguished from Castillomys gracilis and Castillomys crusafonti by their larger 533 size and the greater development of the longitudinal connections (Michaux 1969; Martín- 534 Suárez and Mein 1991). 535 Remarks. The first occurrence of Castillomys apparently coincides with the onset of the 536 Pliocene (van de Weerd 1976; Mein et al. 1990; Piñero and Agustí 2019; Piñero et al. 537 2018), whereas it became extinct at the Early-Middle Pleistocene boundary (Agustí et al. 538 1999). Martín-Suárez and Mein (1991) proposed the phyletic lineage Castillomys gracilis 539 – Castillomys crusafonti – Castillomys rivas, which underwent an increase in size and 540 greater development of the longitudinal connections over the course of the Pliocene and 541 Early Pleistocene. Castillomys rivas has been identified in a number of Early Pleistocene 542 Spanish and southern French localities. The oldest record of Castillomys rivas is reported 543 in earliest Pleistocene sites such as Tollo de Chiclana 10 and 10B (MN17; Minwer- 544 Barakat et al. 2005) and Valdeganga 7 (MN17; Mein et al. 1978), whereas its last record 545 is reported from the late Early Pleistocene level of Cúllar-Baza B (Agustí et al. 1999). 546 547 Genus Stephanomys Schaub, 1938 548 Stephanomys balcellsi Gmelig Meyling and Michaux, 1973 549 Figure 6H–O 550 Material. Two left M1 (PCB-Sba-9, PCB-Sba-11), one right M1 (PCB-Sba-10), one left 551 M2 (PCB-Sba-12), one right M2 (PCB-Sba-13), one left M3 (PCB-Sba-15), one right M3 552 (PCB-Sba-14), one right m1 (PCB-Sba-16), one left m2 (PCB-Sba-18), one right m2 553 (PCB-Sba-17), one left m3 (PCB-Sba-19). 554 Description. The eleven molars from Pedrera del Corral d'en Bruach share the main traits 555 of the genus Stephanomys: the high crown, the development of longitudinal crests on the 556 lower teeth, and the presence of posterior crests on the t1 and t3 in the upper teeth. The 557 M1 has high crests connecting the cusps. There is a small t12. The t1bis is smaller than 13 558 the t2bis. In the M2, the t6 is slightly larger than the t9, and the t12 is poorly developed. 559 In the M3, the t8 is separated from the t4. The lower molars have a high, longitudinal 560 crest. The m1 lacks a tma, the anteroconid being asymmetrical. Both the m1 and m2 have 561 the protoconid and hypoconid slightly displaced posteriorly with respect to the metaconid 562 and entoconid. They both have a low, crest-shaped posterior heel, and a high labial 563 cingulum. In the m3, the anterolabial cuspid is connected to the anterolabial part of the 564 protoconid. The Late Miocene and Early Pliocene species of Stephanomys, such as S. 565 stadii, S. ramblensis, S. dubari, S. cordii, S. margaritae, S. vandeweerdi, and S. 566 numidicus, are smaller and have a lower crown and less accentuated stephanodonty than 567 the studied molars (Mein and Michaux 1979; van de Weerd 1976; Cordy 1976; Adrover 568 1986; Aguilar et al. 1991; Coiffait et al. 1985; Ruiz Bustos et al. 1984; Piñero et al. 2017a, 569 2017b, 2018a; among others). The presence of a poorly-developed t1bis and t2bis, or even 570 their absence, in the M1 of S. prietaensis (see Aguilar et al. 1993a) is the main difference 571 with respect to the studied molars. Morphologically, S. donnezani, S. thaleri and S. calveti 572 can be distinguished from the teeth under study by the presence of a tubercular posterior 573 heel in the m1 and m2 (Depéret 1890; López-Martínez et al. 1998; Bachelet and Castillo- 574 Ruiz 1990). The presence of a crest-shaped posterior heel is shared with S. minor and S. 575 progressus. However, S. minor differs from the studied teeth in its smaller size (Gmelig 576 Meyling and Michaux 1973; van de Weerd 1976; Castillo Ruiz 1990; Minwer-Barakat et 577 al. 2005). The molars of Stephanomys progressus are slightly larger than those from 578 Pedrera del Corral d'en Bruach. In addition, the M2 of S. progressus shows a clearly less- 579 developed t9 than the studied M2 does (Cordy 1976; Minwer-Barakat et al. 2011). The 580 dimensions of the teeth fit the size range of Stephanomys balcellsi from its type locality 581 Islas Medas (Gmelig Meyling and Michaux 1973), Tollo de Chiclana 10 and 10B 582 (Minwer-Barakat et al. 2005), Valdeganga I (Mein et al. 1978) and Almenara-Casablanca 583 4 (originally published as Casablanca B; Gil and Sesé 1985; Agustí et al. 2011). 584 Moreover, the large size (Table 7), hypsodonty, the development of longitudinal 585 connections, the presence of a crest-shaped posterior heel in the m1 and m2, and the 586 presence of a well-developed t9 in the M2 are consistent with this species. Accordingly, 587 the specimens of Stephanomys from Pedrera del Corral d'en Bruach are identified as 588 Stephanomys balcellsi. 589 Remarks. The genus Stephanomys is a common murid in the Late Miocene to Early 590 Pleistocene rodent faunas of the Iberian Peninsula and France. The anagenetic 591 evolutionary line Stephanomys minor – Stephanomys balcellsi – Stephanomys progressus 592 underwent an increase in size, stephanodonty and crown height through the Pliocene and 593 earliest Pleistocene (Cordy 1976; Aguilar et al. 1993a; Renaud et al. 2005; Minwer- 594 Barakat et al. 2011). The descendant of S. balcellsi, i.e. S. progressus, is the most recent 595 representative of the genus (Minwer-Barakat et al. 2011). The oldest records of S. 596 balcellsi seem to be those from the Late Pliocene sites of Huélago-4 (MN16; Sesé 1989) 597 and Almenara-Casablanca 4 (Gil and Sesé 1985; Agustí et al. 2011). However, Sesé 598 (1989) had reservations in ascribing the material to this species, whereas Agustí et al. 599 (2011) assigned the material from Almenara-Casablanca 4 (Late Pliocene) to S. 600 progressus, which seems doubtful (no published descriptions, measurements or 601 illustrations are available). Likewise, Agustí et al. (2011) identified the Stephanomys 602 material from the somewhat younger site of Almenara-Casablanca 1 (Early Pleistocene) 603 as S. progressus. We have revised some specimens of Stephanomys from Almenara- 604 Casablanca 1 stored at the Institut de Paleoecologia Humana i Evolució Social 605 (Tarragona, Spain) in order to clarify their determination. We have found that this 606 material fits better with S. balcellsi than with S. progressus, as all the available M2s 607 develop a t9, a diagnostic character that differentiates S. balcellsi from S. progressus (see 14 608 Minwer-Barakat et al. 2011). There is therefore an inconsistency in the identification of 609 the descendant of S. balcellsi in the older site of Almenara-Casablanca 4. This 610 incongruence is best resolved if the material from the latter Late Pliocene locality 611 corresponds to S. balcellsi, as Gil and Sesé (1985) had determined. Indeed, these authors 612 reported the presence of a t9 in the M2s from Almenara-Casablanca 4. All the remaining 613 populations of S. balcellsi have been identified in earliest Pleistocene localities (MN17): 614 Islas Medas, Tollo de Chiclana 10 and 10B, Valdeganga I and Alfarerías (Gmelig 615 Meyling and Michaux 1973; Mein et al. 1978; Minwer-Barakat et al. 2005; Ruiz Bustos 616 1986). 617 618 Familly CRICETIDAE Fisher, 1817 619 620 Subfamily CRICETINAE Fisher, 1817 621 622 Genus Allocricetus Schaub, 1930 623 624 Allocricetus sp. 625 626 Figure 6P 627 628 Material. One left m1 (PCB-All-1) (L=1.8 mm; W=1.3 mm). 629 630 Description. Given the development of the protolophid and the metalophid, as well as the 631 mesalophid, the tooth in question can be ascribed to the genus Allocricetus in accordance 632 with Cuenca-Bescós (2003) and Cuenca-Bescós et al. (1997). According to the 633 Quaternary record of Western Europe, it could be one of the following three species: A. 634 bursae, A. ehiki or A. correzensis. The cingular stylid could be a diagnostic character that 635 separates the different species. The tooth under study has a well-developed cingular 636 stylid, as is present in all three species but more frequent in A. ehiki and A. correzensis 637 (Cuenca-Bescós, 2003). On the other hand, the measurements taken from the only 638 identified m1 are consistent with those taken from the Early Pleistocene A. bursae 639 specimens at the Hungarian sites of Osztramos 3 and Villány 3 (Hir 2002), differing from 640 the A. ehiki specimens from the same site and from the A. correzensis identified at the 641 Spanish Middle Pleistocene site of Sima de los Huesos (Cuenca-Béscos et al. 1997), 642 which are larger in size. In summary, our specimen is morphologically closer to A. ehiki 643 and A. correzensis, but metrically closer to A. bursae, preventing us from ascertaining the 644 species level with a single tooth. 645 646 Remarks. The first records of the genus Allocricetus in Europe are from the Hungarian 647 Gelasian sites (MN17) of Osztramos 3 and Villány 3 (Hir 1993). These were identified 648 as A. ehiki. However, the most ancient records of this genus in Iberia are known from the 649 Calabrian (post-Jaramillo) layers of the site of Gran Dolina (TD3-4) of the Sierra de 650 Atapuerca, identified as A. bursae (Cuenca-Bescós et al. 2010), and from Almenara- 651 Casablanca 3, Calabrian in age, identified as A. bursae duraciensis (Agustí et al. 2011). 652 Accordingly, if the age of Pedrera del Corral d’en Bruach is confirmed as MN17 by the 653 small-mammal association as a whole, this could be the oldest record known to date of 654 the genus Allocricetus in the Iberian Peninsula. 655 656 657 Subfamilly ARVICOLINAE Gray, 1821 15 658 Genus Mimomys Forsyth Major, 1910 659 Mimomys medasensis Michaux, 1971 660 Figure 7 661 Material. Six right m1 (PCB-Mm-1; PCB-Mm-5; PCB-Mm-8; PCB-Mm-9; PCB-Mm- 662 10; PCB-Mm-12) six left m1 (PCB-Mm-2; PCB-Mm-3, PCB-Mm-4; PCB-Mm-6; PCB- 663 Mm-7; PCB-Mm-11), one right M3 (PCB-Mn-13). 664 Description. In general, all the m1 are large in size (Table 8), comparable with the 665 specimens identified in Almenara-Casablanca I (Gil and Sesé 1984), El Rincón I (Alberdi 666 et al. 1997) and Tollo del Chiclana 10 (Minwer-Barakat et al. 2004). In occlusal view, the 667 m1 present a connected T1-T2, scarcely differentiated enamel (Mimomys-type) with SDQ 668 values between 156 and 166, comparable to that of other large-sized specimens from the 669 ancient Early Pleistocene of central Europe such as Mimomys ostramosensis (Kosciow 670 and Nadachowski 2002), relatively marked asymmetry between the labial and lingual 671 triangles, and the presence of little cement in the re-entrant angles. The ACC of the m1 672 in occlusal view is characterized by a T4 oriented backward, a T5 with a thick and 673 rounded shape, two labial re-entrant angles (“inselfalte” and “prismenfalte”) that are 674 shallow and relatively open, and in all cases the labial salient angle (“Mimomys-Kante”) 675 is differentiated and the LRA4 deep and marked. Of all the m1 identified, only two 676 specimens (PCB-Mm-1 and PCB-Mm-2) present an enamel islet, whereas in the rest of 677 the identified teeth the islet appears open, showing the interspecific variability revealed 678 in Sevilla et al. (1991). As in the lower molars, the only M3 is large in size (Table 8), 679 comparable with specimens identified in Almenara-Casablanca I (Gil and Sesé 1984), El 680 Rincón I (Alberdi et al. 1997) and Tollo del Chiclana 10 (Minwer-Barakat et al. 2004). 681 In occlusal view, this tooth presents a little cement in the re-entrant angles and scarcely 682 differentiated enamel. PL is isolated from T2, and T2 from T3; T4 is well delimited, and 683 T5 is small but differentiated; the BRA1 is deep, and the LRA2 and BRA3 are relatively 684 deep and opposed one another. It presents a large, ovoid enamel islet. 685 Remarks. As expounded above, the morphological traits of the Mimomys material 686 identified at Pedrera del Corral d’en Bruach are comparable with those of the species M. 687 medasensis of Islas Medas (Michaux 1971), Almenara-Casablanca 1 (Gil and Sesé 1984), 688 El Rincón I (Alberdi et al. 1997), Tollo de Chiclana 10 (Minwer-Barakat et al. 2004) and 689 Alto de la Ceradaña (Guillem-Calatayud et al. 2013). The presence of M. medasensis in 690 Pedrera del Corral d’en Bruach indicates a chronological range that could correspond, 691 according to Sesé (2006), to the MN17 biozone, within the Gelasian stage of the Early 692 Pleistocene, dated to between ca. 2.6 and 1.8 Ma. In addition to the sites mentioned above, 693 the Iberian Peninsula has other records of the species within the same chronological 694 range: Cortes de Baza-1 (Peña 1975), Galera G (Agustí et al. 2001), and Alozaina 695 (Aguilar et al. 1993a). The form Mimomys aff. medasensis has been identified at the latest 696 Pliocene site of Villarroya in La Rioja (Laplana et al. 2016). Given its fossil record, it 697 was thought to be an endemism restricted to the Mediterranean area of the Iberian 698 Peninsula. However, Girotti et al. (2003) identified M. medasensis at the Torre Picchio 699 site in Umbria, central Italy, showing, according to these authors, the possibility of 700 occasional migration to the Italian Peninsula. 16 701 702 4. Dicussion 703 704 4.1. Chronological remarks 705 706 From a chronological point of view, the small-mammal assemblage of Pedrera del Corral 707 d’en Bruach can be assigned to the MN17 biozone (e.g. Valli, 2004), the Gelasian stage 708 (ca. 2.5 - 1.8 Ma) (Rio et al. 1994). The most significant biochronological element is 709 Mimomys medasensis, an arvicoline restricted to the lower part of MN17 (Minwer- 710 Barakat et al. 2012; Agustí et al. 2015; among others). This rodent has also been found in 711 other Iberian localities correlated with the earliest Pleistocene, such as Islas Medas 712 (Michaux 1971), Almenara-Casablanca 1 (Agustí et al. 2011), Galera 2 (Agustí et al. 713 2001), Tollo de Chiclana 10 (Minwer-Barakat et al. 2004), El Rincón I (Alberdi et al. 714 1997), Alto de la Cerdaña (Guillem-Calatayud et al. 2013), Cortes de Baza-1 (Peña 1975), 715 Alozaina (Aguilar et al. 1993a) and Villarroya (Laplana et al. 2016). In addition to M. 716 medasensis, the simultaneous occurrence of the murids Castillomys rivas and 717 Stephanomys balcellsi is also characteristic of MN17 (Sesé 2006). The presence of the 718 latter murid confirms that Pedrera del Corral d’en Bruach is chronologically close to the 719 sites of Islas Medas, Tollo de Chiclana 10, Valdeganga I and Almenara-Casablanca 1 720 (Fig. 8). 721 722 The presence of M. medasensis is typical of the Kislangia gusii Zone (Agustí et al. 2015), 723 the earliest biozone established for the Pleistocene succession of the Guadix-Baza Basin 724 (ca. 2.6 - 2.2 Ma). This biozone is equivalent to the Mimomys medasensis Zone defined 725 by Minwer-Barakat et al. (2012) and immediately subsequent to the Kislangia ischus 726 Zone proposed by Piñero et al. (2018) (3.0 - 2.6 Ma). The Kislangia gusii Zone is 727 followed by the Mimomys cf. reidi Zone (2.2 - 1.85 Ma), which is correlated with the 728 upper part of the MN17 biozone (Agustí et al. 2015) and characterized by the occurrence 729 of a small, hypsodont arvicolid provisionally determined as Mimomys cf. reidi. In this 730 biozone K. gusii, M. medasensis and S. balcellsi are no longer present. As shown by the 731 Almenara-Casablanca 1 section, Kislangia gusii progressively disappeared through the 732 lower part of MN17, whereas M. medasensis was still present (Agustí et al. 2011). 733 Although the arvicoline sample from Pedrera del Corral d’en Bruach is not very 734 extensive, the continued presence of M. medasensis togheter the absence of other 735 arvicolines, as K. gusii, suggests that the site could be close in age to the transition 736 between the K. gusii and M. cf. reidi biozones, at roughly 2.2 Ma (Fig. 8), and is therefore 737 older than the lower boundary of the Olduvai geomagnetic subchron (1.94 - 1.78 Ma). 738 739 4.2. Species occurrence and environmental remarks 740 Given that the probable age of Pedrera del Corral d’en Bruach between 2.6 – 2.2 Ma and 741 problably close toca. 2.2 Ma, the presence of Glirulus pusillus constitutes the first record 742 of this species in the Iberian Peninsula. G. pusillus is a rare component of the Late 743 Pliocene and Early Pleistocene faunas of Europe (Nadachowski and Daoud 1995) and is 744 represented in only a few sites, such as Les Valerots Bed 2 (Bonifay 1973), Tegelen (Van 745 den Hoek Ostende 2003), Cascina Arondelli and the Fornace RDB Quarry (Kotsakis 17 746 1986), Podlesice (Kowalski 1956), Osztramos 7 (Janossy 1986) and Betfia 13 (Terzea 747 1996). It had not been found until now in the Iberian Peninsula. 748 The presence of Muscardinus avellanarius and the genus Allocricetus in Pedrera del 749 Corral d’en Bruach could likewise be the oldest mention of these taxa in the Iberian 750 Peninsula. M. avellanarius is an extant species that is relatively well documented in the 751 Late Pliocene and Early Pleistocene of central Europe (Nadachowski and Daoud 1995; 752 Van den Hoek Oestende 2003; Striczky and Pazonyi 2014). Until now, however, its 753 presence in the Iberian Peninsula has been restricted to the Late Pleistocene site of 754 Lezetxiki II (Garcia-Ibaibarriaga et al. 2015). The genus Allocricetus, appearing as A. 755 ehiki, has been identified in various Gelasian European sites, such as Osztramos 3 and 756 Villány 3 (Hir 1993). In the Iberian Peninsula, however, the genus has to date been 757 identified as A. bursae in the post-Jaramillo layers of Gran Dolina (TD3-4) in the Sierra 758 de Atapuerca (Cuenca-Bescós et al. 2010), and as A. bursae aff. balarucciensis and A. 759 bursae duraciensis in the likewise post-Jaramillo sites of Cueva Victoria and Almenara- 760 Casablanca 3, respectively (Agustí 1982; Agustí et al. 2011). Furthermore, the presence 761 of Blarinoides mariae in Pedrera del Corral d’en Bruach could be the most recent 762 occurrence of the genus in the Iberian Peninsula, since the chronological range of 763 Blarinoides is known to run from the Pliocene to the beginning of the Early Pleistocene 764 in Europe (Reumer 1984; Rzebik-Kowalska 1998; Minwer-Barakat et al. 2007), although 765 its presence in the Iberian Peninsula was until now apparently restricted to MN15 and 766 MN16 (Furió et al. 2018). 767 Currently, the environment around Pedrera del Corral d’en Bruach is characterized 768 mainly by scrubland with species such as kermes oak (Quercus coccifera) and the 769 Mediterranean dwarf palm (Chamaerops humilis), with dry conditions and a low diversity 770 in small mammals. The rodent-insectivore assemblage is now represented by Apodemus 771 sylvaticus, Mus spretus, Eliomys quercinus and Crocidura russula (Solís et al. 2000). In 772 contrast, the small-mammal assemblage of this area between 2.6 – 2.2 Ma was much more 773 diverse, as has here been demonstrated. The surrounding environment of the Garraf 774 Massif was probably more humid than nowadays, as suggested by the presence of 775 insectivores of the genus Talpa and the red-toothed shrews B. mariae, P. hungarica and 776 D. cf. doukasi. The landscape during the formation of Pedrera del Corral d’en Bruach was 777 characterized by the development of a forest denser than the present shrubland and open 778 woodland, as indicated by the occurrence of the murids A. atavus and A. jeanteti and the 779 highly diversified glirid association featuring G. minor, E. intermedius, G. cf. pusillus, 780 and M. cf. avellanarius. Such an interpretation is supported by the presence in the sample 781 of a rather diverse assemblage of amphibians and reptiles, also indicating more humid 782 local conditions than today in the area (Albanerpetontidae indet. and Blanus sp.), together 783 with taxa typically associated with open-dry areas and Mediterranean woodlands (cf. 784 Pelobates cultripes, Agamidae indet., Pseudopus sp., and Zamenis scalaris). 785 5. Conclusions 786 787 Our study of the small-mammal association of Pedrera del Corral d’en Bruach has led us 788 to the following conclusions: 789 18 790 A very rich, diverse small-mammal association has been revealed, with 15 taxa among 791 the 270 remains identified. 792 Chronological inferences from the small-mammal association show Pedrera del Corral 793 d’en Bruach to be Gelasian in age; the site can be placed at Kislangia gusii biozone (ca. 794 2.6 – 2.2 Ma) and probably could be close to the transition between the Kislangia gusii 795 and Mimomys cf. reidi biozones (ca. 2.2 Ma). 796 The presence of Glirulus cf. pusillus in Pedrera del Corral d’en Bruach is the first record 797 of the species known to date in Iberia. It could also be the oldest citation known to date 798 of the genus Allocricetus and the species Muscardinus avellanarius and the latest 799 occurrence of Blarionides mariae in the Iberian Peninsula. 800 Preliminary palaeoenvironmental observations show Pedrera del Corral d’en Bruach to 801 have had a much greater diversity of small-mammal species than what is currently 802 recorded in the area and an environment that was more humid than nowadays, featuring 803 a landscape dominated by forest formations. 804 805 Acknowledgments 806 This manuscript is part of a Knowledge Generation projects (PID2021-122533NB-100, 807 PID2021-123092NB-C21 and PID2020-117289GB-I00) of the Spanish Ministry of 808 Science and Innovation. The Institut Català de Paleoecologia Humana i Evolució Social 809 (IPHES-CERCA) has received financial support from the Spanish Ministry of Science 810 and Innovation through the ‘María de Maeztu’ program for Units of Excellence 811 (CEX2019-000945-M). J.M.L.-G, M.S. and J.D. were supported by a Ramón y Cajal 812 contract (RYC-2016-19386, RYC2021‐032999‐I and RYC-2015- 17667, respectively) 813 with financial sponsorship from the Spanish Ministry of Science and Innovation. P.P. is 814 supported by a ‘Juan de la Cierva-Incorporación’ contract (grant IJC2020-044108-I) 815 funded by MCIN/AEI/10.13039/501100011033 and the ‘European Union 816 NextGenerationEU/PRTR’. J.G. is the recipent of a postdoctoral contract 817 [ESPDOC20/83] from the University of the Basque Country (UPV/EHU) and her 818 work was also supported by the Eusko Jaurlaritza [IT1485-22]. B.M.-S. was funded by 819 the Consellería de Cultura, Educación e Universidade, Xunta de Galicia, Spain (ED481D- 820 2022-013). We want to thank Rupert Glasgow for reviewing the English language of the 821 manuscript. Thanks go to Castelldefels City Council (La Guixera Laboratory), where the 822 sieving process was carried out. 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La Piquera (central Iberian Peninsula): A new key vertebrate locality for the 1119 Early Pliocene of Western Europe. Acta Palaeontologica Polonica. 1120 Raymo, M.E, Ruddiman, W.F, Backman, J, Clement, B.M, Martinson, D.G. 1989. Late 1121 Pliocene variation in northern hemisphere ice sheets and 1122 North Atlantic Deep Water circulation: Paleoceanography, v. 4, pp. 1123 413–446. 1124 Renaud, S., Michaux, J., Schmidt, D. N., Aguilar, J. P., Mein, P., Auffray, J. C. 2005. 1125 Morphological evolution, ecological diversification and climate change in rodents. 1126 Proceedings of the Royal Society B: Biological Sciences, 272(1563), 609-617. 1127 Reumer, J.W.F., 1984. Ruscinian and early Pleistocene Soricidae (Insectivora, 1128 Mammalia) from Tegelen (The Netherlands) and Hungary. Scripta Geologica 73, 1–173. 1129 Rietschel, S., Storch, G., 1974. Aussergewöhnlich erhaltene Waldmäuse (Apodemus 1130 atavus Heller, 1936) aus dem Ober-Pliozän von Willershausen am Harz. Senckenbergiana 1131 Lethaea 54, 491–519. 1132 Rio, D, Sprovieri, R, Di Stefano, E, 1994, The Gelasian Stage: a proposal 1133 of a new chronostratigraphic unit of the Pliocene Series: Riv. It. Paleont. 1134 Strat., v. 100, pp. 103–124. 1135 Ruiz Bustos, A. 2002. Interpretación paleoambiental del yacimiento kárstico Plioceno de 1136 Moreda (Granada, España) a partir de los micromamíferos acumulados; pp. 419–428 in 1137 F. Carrasco, J. J. Durán, and B. Andreo (eds.), Karst and Environment. Fundación Cueva 1138 de Nerja, Málaga, Spain. 1139 Ruiz Bustos, A. 1986. Análisis del proceso evolutivo del género Stephanomys (Rodentia, 1140 Muridae). Paleomammalia 1(1):1–42. 1141 Ruiz Bustos, A., Sesé, C, Dabrio, C.J, Peña, J.A, Padial, J. 1984. Geología y fauna de 1142 micromamíferos del nuevo yacimiento del Plioceno inferior de Gorafe A (Depresión de 1143 Guadix-Baza, Granada) [Geology and micromammal fauna from the new early Pliocene 1144 site of Gorafe A (Guadix-Baza Basin, Granada)]. Estud Geol. 40:231–241. 1145 Rzebik-Kowalska, B. 1998. Fossil history of shrews in Europe; pp. 23–92 in J. M.Wójcik 1146 and M.Wolsan (eds.), Evolution of Shrews. Mammal Research Institute, Polish Academy 1147 of Sciences, Bialowieza. 27 1148 Sala, B., Masini, F., 2007. Late Pliocene and Pleistocene small mammal chronology in 1149 the Italian peninsula. Quaternary International 160, 4–16. 1150 Sansalone, G., Kotsakis, T., Piras, P. 2015. Talpa fossilis or Talpa europaea? Using 1151 geometric morphometrics and allometric trajectories of humeral moles remains from 1152 Hungary to answer a taxonomic debate. Palaeontologia Electronica 18.2.42A: 1-17 1153 Sesé, C. 1989. Micromamíferos del Mioceno, Plioceno y Pleistoceno de la cuenca de 1154 Guadix-Baza (Granada). Trabajos sobre el Neógeno-Cuaternario, 11, 185-213. 1155 Sesé, C. 2006. Los roedores y lagomorfos del Neógeno de España. Estudios Geológicos 1156 62, 429-480. 1157 Sevilla, P. 1988. 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"Towards the lowering of 1170 the Pliocene/Pleistocene boundary to the Gauss-Matuyama reversal". Quaternary 1171 International. 40: 37–42 1172 Terzea, E 1996. Biochronology of the Pleistocene deposits at Betfia (Bihor, Romania). 1173 Acta Zoologica Cracovensia 39, 531-540. 1174 Topál, G. 1979. Fossil bats of the Rhinolophus group in Hungary (Mammalia: 1175 Chiroptera). Fragmenta Mineralogica et Paleontologica 9, 61-101. 1176 Valli, A. 2004. Taphonomy of Saint-vallier (Drône, France), the reference locality for the 1177 biozone MN17 (Upper Pliocene). Lethaia 37, 337-350. 1178 Van Cleef-Roders, J.T., Van den Hoek Ostende, L.W., 2001. Dental morphology of Talpa 1179 europaea and Talpa occidentalis (Mammalia: Insectivora) with a discussion of fossil 1180 Talpa in the Pleistocene of Europe. Zoologische Mededelingen 75, 51–68 1181 Van de Weerd, A. 1979. Early Ruscinian rodents and lagomorphs (Mammalia) from the 1182 lignites near Ptolemais (Macedonia, Greece). Proc KNAW 82:127–170. 1183 Van den Hoek Oestende, L, W. 2003. Gliridae (Rodentia, Mammalia) from the Upper 1184 Pliocene of Tegelen (province of Limburg, The Netherlands). Scripta Geologica 126, 1185 203-205. 28 1186 Van der Meulen, A. J. 1973. Middle Pleistocene small mammals from the Monte Peglia 1187 (Orvieto, Italy) with special reference to the philogeny of Microtus (Arvicolidae, 1188 Rodentia). Quaternaria 16, 1-144. 1189 Zagwijn, W. H. 1974, The Pliocene-Pleistocene boundary in western and 1190 southern Europe: Boreas, v. 3, pp. 75–97. 1191 1192 1193 1194 1195 1196 1197 FIGURE CAPTIONS 1198 29 1199 Figure 1. Pedrera del Corral d’en Bruach (Gavà, Barcelona). A-C Location of the 1200 palaeontological site. The map was downloaded from http://maps-for-free.com/ (© 1201 OpenStreetMap contributors, accessed on 27 November 2022). The cartography in the 1202 OpenStreetMap map tiles is licensed under CC BY-SA 1203 (www.openstreetmap.org/copyright, accessed on 27 November 2022). The licence terms 1204 can be found on the following link: http://creativecommons.org/licenses/by-sa/2.0/ 1205 (accessed on 27 November 2022). D. Corral d’en Bruach quarry indicating the location 1206 of the site. E. Position of the fissure where the fossil material appears inside the quarry. 1207 F-G. Picture showing sediments and the water screening of the sediments recovered from 1208 the fissure. 1209 1210 Figure 2. ESEM images of insectivores, chiropters, and lagomorph from Pedrera del 1211 Corral d’en Bruach (northeastern Spain). (A) Talpa cf. fossilis: right humerus, PCB-Ta- 1212 1. (B) Blarinoides mariae: right mandibular fragment with m2, PCB-Bl-1. (C) Petenyia 1213 hungarica: right mandibular fragment with m2, PCB-Pe-1. (D) Deinsdorfia doukasi: right 1214 m1, PCB-De-1. (E) Rhinolophus gr. ferrumequinum: left m1, PCB-Rh-1. (F) Prolagus cf. 1215 calpensis: right p3, PCB-Pr-1. The upper scale bar represents 5 mm and goes with (A). 1216 The central scale bar represents 1 mm and goes with (B–E). The lower scale bar represents 1217 1 mm and goes with (F). 30 1218 1219 Figure 3. Images of Deinsdorfia doukasi from Pedrera del Corral d’en Bruach 1220 (northeastern Spain). (A) right I1 (PCB-De-4); (B) left i1 (PCB-De-5). Scale 1 mm. 31 1221 1222 Figure 4. ESEM images of glirids from Pedrera del Corral d’en Bruach (northeastern 1223 Spain). (A–G) Glis minor: (A) right P4, PCB-Gm-32; (B) left M1, PCB-Gm-47; (C) left 1224 M2, PCB-Gm-30; (D) left M3, PCB-Gm-18; (E) right p4, PCB-Gm-5; (F) left m1, PCB- 1225 Gm-9; (G) left m2, PCB-Gm-38. (H–K) Eliomys intermedius: (H) right P4, PCB-Ein-43; 1226 (I) left M1-2, PCB-Ein-1; (J) left M3, PCB-Ein-17; (K) right m2, PCB-Ein-22. (L) 1227 Muscardinus cf. avellanarius: right m1, PCB-Mu-35. (M) Glirulus cf. pusillus: right m2, 1228 PCB-Gpu-23. Scale bar represents 1 mm. The upper scale bar goes with (A–L), whereas 1229 the lower one goes with (M). 32 1230 1231 Figure 5. ESEM images of murids from Pedrera del Corral d’en Bruach (northeastern 1232 Spain). (A–L) Apodemus atavus (A) left M1, PCB-Aat-25; (B) right M1, PCB-Aat-31; 1233 (C) left M2, PCB-Aat-36; (D) right M2, PCB-Aat-43; (E) left M3, PCB-Aat-52; (F) left 1234 M3, PCB-Aat-53; (G) left m1, PCB-Aat-65; (H) right m1, PCB-Aat-77; (I) right m2, 1235 PCB-Aat-89; (J) right m2, PCB-Aat-108; (K) right m3, PCB-Aat-121; (L) right m3, PCB- 1236 Aat-125. (M–Y) Apodemus jeanteti: (M) left M1, PCB-Aje-38; (N) left M1, PCB-Aje- 1237 40; (O) left M1, PCB-Aje-44; (P) left M2, PCB-Aje-55; (Q) right M2, PCB-Aje-59; (R) 1238 right M3, PCB-Aje-68; (S) left M3, PCB-Aje-70; (T) left m1, PCB-Aje-2; (U) left m1, 1239 PCB-Aje-5; (V) right m2, PCB-Aje-17; (W) right m2, PCB-Aje-22; (X) right m3, PCB- 1240 Aje-31; (Y) right m3, PCB-Aje-32. Scale bar represents 1 mm. 33 1241 1242 Figure 6. ESEM images of murids and cricetids from Pedrera del Corral d’en Bruach 1243 (northeastern Spain). (A–G) Castillomys rivas: (A) right M2, PCB-Cri-1; (B) left M2, 1244 PCB-Cri-2; (C) left m1, PCB-Cri-3; (D) left m1, PCB-Cri-4; (E) right m2, PCB-Cri-5; 1245 (F) left m2, PCB-Cri-6; (G) left m3, PCB-Cri-8. (H–O) Stephanomys balcellsi: (H) left 1246 M1, PCB-Sba-9; (I) left M2, PCB-Sba-12; (J) right M2, PCB-Sba-13; (K) right M3, PCB- 1247 Sba-14; (L) right m1, PCB-Sba-16; (M) left m2, PCB-Sba-18; (N) right m2, PCB-Sba- 1248 17; (O) left m3, PCB-Sba-19. (P) Allocricetus sp.: left M1, PCB-All-1. Scale bar 1249 represents 1 mm. 34 1250 1251 Figure 7. ESEM images of Mimomys medasensis from Pedrera del Corral d’en Bruach 1252 (northeastern Spain). (A) right m1 in occlusal view, PCB-Mm-1. (B) right m1 in occlusal 1253 view, PCB-Mm-5. (C) left m1 in occlusal view, PCB-Mm-3. (D) left m1 in occlusal view, 1254 PCB-Mm-4. (E) left m1 in lateral view, left m1 in occlusal view, PCB-Mm-2. (F) right 1255 M3 in occlusal view, PCB-Mm-13. Scale bar represents 2 mm. 35 1256 1257 Figure. 8. Biostratigraphic correlation between Pedrera del Corral d’en Bruach and 1258 various Iberian localities, with the biochronologies proposed by Agustí et al. (2015) and 1259 Piñero et al. (2018), and the geomagnetic polarity time-scale (GPTS). 1260 1261 TABLE CAPTIONS NISP MNI %MNI Talpa cf. fossilis 3 2 3.64 Blarinoides mariae 1 1 1.82 Petenyia hungarica 1 1 1.82 Deinsdorfia doukasi 5 1 1.82 Rhinolophus cf. ferrumequinum 1 1 1.82 Prolagus cf. calpensis 1 1 1.82 Glis minor 15 3 5.45 Eiomys intermedius 10 4 7.27 Glirulus cf. pusillus 1 1 1.82 36 Muscardinus cf. avellanarius 1 1 1.82 Apodemus atavus 128 19 34.55 Apodemus jeanteti 71 9 16.36 Castillomys rivas 7 2 3.64 Stephanomys balcellsi 11 2 3.64 Allocricetus sp. 1 1 1.82 Mimomys medasensis 13 6 10.91 Total 270 55 100 1262 1263 Table 1. Representation of the small-mammal species identified at Pedrera del Corral 1264 d’en Bruach in terms of the number of identified specimens (NISP), the minimum number 1265 of individuals (MNI), and the MNI as a percentage. Length Width Element N min mean max σ N min mean max σ p4 2 1.09 1.10 1.10 0.01 2 1.12 1.13 1.14 0.01 m1 5 1.61 1.67 1.71 0.05 4 1.44 1.53 1.61 0.09 m2 1 1.73 1 1.73 m3 2 1.60 1.62 1.63 0.02 2 1.42 1.44 1.45 0.02 P4 1 1.31 1 1.44 M1 2 1.66 1.68 1.69 0.02 2 1.69 1.78 1.87 0.13 M2 1 1.43 1 1.65 M3 1 1.22 1 1.49 1266 1267 Table 2. Measurements of the identified elements (N) of Glis minor from Pedrera del 1268 Corral d’en Bruach in millimetres. Min: minimum, max: maximum; σ standard deviation. Length Width Element N min mean max σ N min mean max σ m2 1 1.44 1 1.66 P4 1 1.06 1 1.34 M1-M2 3 1.42 1.52 1.61 0.10 3 1,69 1.77 1.84 0.08 M3 5 1.26 1.33 1.48 0.10 5 1,50 1.70 1.89 0.16 1269 1270 Table 3. Measurements of the identified elements (N) of Eliomys intermedius from 1271 Pedrera del Corral d’en Bruach in millimetres. Min: minimum, max: maximum; σ 1272 standard deviation. Length Width Element N min mean max σ N min mean max σ M1 30 1.91 2.07 2.25 0.07 31 1.21 1.32 1.49 0.06 M2 19 1.26 1.41 1.54 0.07 19 1.13 1.26 1.36 0.07 M3 7 1.01 1.04 1.10 0.04 7 0.93 0.96 1.06 0.05 m1 24 1.74 1.92 2.04 0.08 26 1.04 1.15 1.22 0.05 m2 28 1.24 1.37 1.50 0.06 29 1.08 1.21 1.30 0.06 m3 9 1.06 1.12 1.23 0.07 10 0.90 1.01 1.16 0.08 37 1273 1274 Table 4. Measurements of the identified elements (N) of Apodemus atavus from Pedrera 1275 del Corral d’en Bruach in millimetres. Min: minimum, max: maximum; σ standard 1276 deviation. Length Width Element N min mean max σ N min mean max σ M1 15 2.18 2.37 2.62 0.11 17 1.39 1.51 1.65 0.06 M2 13 1.53 1.68 1.84 0.11 13 1.37 1.46 1.57 0.06 M3 4 1.21 1.24 1.29 0.04 4 1.00 1.10 1.16 0.07 m1 13 2.00 2.14 2.30 0.09 13 1.23 1.33 1.44 0.07 m2 14 1.51 1.63 1.80 0.08 14 1.25 1.40 1.52 0.07 m3 7 1.35 1.47 1.69 0.15 7 1.18 1.29 1.45 0.10 1277 1278 Table 5. Measurements of the identified elements (N) of Apodemus jeanteti from Pedrera 1279 del Corral d’en Bruach in millimetres. Min: minimum, max: maximum; σ standard 1280 deviation. Length Width Element N min mean max σ N min mean max σ M2 2 1.18 1.19 1.20 0.01 2 1.15 1.19 1.23 0.06 m1 2 1.53 1.62 1.70 0.12 2 1.03 1.08 1.12 0.06 m2 3 1.16 1.19 1.24 0.04 3 1.04 1.08 1.10 0.04 m3 1 - 0.91 - - 1 - 0.81 - - 1281 1282 Table 6. Measurements of the identified elements (N) of Castillomys rivas from Pedrera 1283 del Corral d’en Bruach in millimetres. Min: minimum, max: maximum; σ standard 1284 deviation. Length Width Element N min mean max σ N min mean max σ M1 1 - 3.19 - - 3 2.23 2.36 2.43 0.11 M2 2 2.38 2.45 2.52 0.10 2 2.22 2.24 2.25 0.02 M3 2 1.73 1.83 1.92 0.13 2 1.48 1.50 1.52 0.03 m1 1 - 2.52 - - 1 - 1.89 - - m2 2 2.21 2.37 2.52 0.22 2 2.14 2.31 2.47 0.23 m3 1 - 2.48 - - 1 - 1.86 - - 1285 1286 Table 7. Measurements of the identified elements (N) of Stephanomys balcellsi from 1287 Pedrera del Corral d’en Bruach in millimetres. Min: minimum, max: maximum; σ 1288 standard deviation. Length Width Element N min mean max σ N min mean max σ m1 5 3.39 3.49 3.62 0.09 5 1.61 1.68 1.73 0.04 M3 1 2.45 1 1.53 38 1289 1290 Table 8. Measurements of the identified elements (N) of Mimomys medasensis from 1291 Pedrera del Corral d’en Bruach in millimetres. Min: minimum, max: maximum; σ 1292 standard deviation. 1293 1294 1295 1296 1297 1298 1299 1300 39