Quaternary Science Reviews 121 (2015) 52e74

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Quaternary Science Reviews

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The Punta Lucero Quarry site (Zierbena, Bizkaia): a window into the Middle in the Northern Iberian Peninsula

* Asier Gomez-Olivencia a, b, c, d, , Nohemi Sala d, Diego Arceredillo e, f, g, Nuria García h, d, Virginia Martínez-Pillado i, j, Joseba Rios-Garaizar k, Diego Garate l, Gonzalo Solar m, Inaki~ Libano m a Dept. Estratigrafía y Paleontología, Facultad de Ciencia y Tecnología, Euskal Herriko Unibertsitatea, UPV-EHU. Apdo. 644, 48080 , b IKERBASQUE, Basque Foundation for Science, Spain c Equipe de Paleontologie Humaine, UMR 7194, CNRS, Departement de Prehistoire, Museum national d'Histoire naturelle, Musee de l'Homme, 17, Place du Trocadero, 75016 Paris, d Centro UCM-ISCIII de Investigacion sobre Evolucion y Comportamiento Humanos, Avda. Monforte de Lemos 5 (Pabellon 14), 28029 Madrid, Spain e Facultad de Humanidades y Ciencias Sociales, Universidad Internacional Isabel I de Castilla, C/ Fernan Gonzalez n 76, 09003 Burgos, Spain f Department of Geology, School of Science, University of Salamanca, 37008 Salamanca, Spain g Laboratorio de Prehistoria, Edificio IþDþi, Universidad de Burgos, Plaza Misael Banuelos~ s/n, 09001 Burgos, Spain h Department of Palaeontology, School of Geological Sciences, Universidad Complutense de Madrid, Ciudad Universitaria, 28040 Madrid, Spain i Department of Mineralogy and Petrology, School of Science and Technology, Universidad del País Vasco (UPV/EHU), 48940 , Bizkaia, Spain j ARANZADI Geo-Q, b/ Kortasenebarri s/n, 48940 Leioa, Bizkaia, Spain k CENIEH, Paseo Sierra de Atapuerca, 3, 09002 Burgos, Spain l Arkeologi Museoa, Calzadas de Mallona, 2, 48006 Bilbao, Spain m Edestiaurre Kultur Elkartea, Spain article info abstract

Article history: The period between the end of the Early Pleistocene and the mid-Middle Pleistocene (roughly between Received 2 February 2015 1.0 and 0.4 Ma BP) is of great interest in Western . It witnessed several climatic oscillations and Received in revised form changes in the fauna, the demise of a hominin species and the appearance of another, along with 21 April 2015 important cultural and technological changes. Thus, the few available sites with these chronologies is Accepted 5 May 2015 vital to the understanding of the tempo and mode of these changes. Middle Pleistocene sites in the Available online 31 May 2015 Northern Iberian Peninsula are very rare. Here we present the study of the site found at the Punta Lucero Quarry ( province, Northern Iberian Peninsula), which includes for the first time the complete Keywords: Galerian collection from the site. The fossil association from this site includes several ungulates, such as a Meg- latidens acerine , Cervus elaphus, large bovids (likely both primigenius and sp. are present), Ste- Canis mosbachensis phanorhinus sp., and carnivores, such as Homotherium latidens, gombaszoegensis, Canis Panthera gombaszoegensis mosbachensis and Vulpes sp. This association is typical of a middle Middle Pleistocene chronology and Natural trap would be the oldest macro- site in the Eastern Cantabrian region. This site would likely correspond to a chronology after Mode 1 technological complex and before the arrival of Mode 2 technology in this region. Thus, it offers a glimpse into the paleoecological conditions slightly prior to or contemporaneous with the first Acheulian makers in the northern fringe of the Iberian Peninsula. © 2015 Elsevier Ltd. All rights reserved.

1. Introduction

The period between the end of the Calabrian and the mid part of the Ionian (roughly between 1.0 and 0.4 Ma; MIS 30 to MIS 12) is a very interesting period from different perspectives. At the end of * Corresponding author. Dept. Estratigrafía y Paleontología, Facultad de Ciencia y the Calabrian (ca 0.78e1.0 Ma BP), Europe was inhabited by a Tecnología, Euskal Herriko Unibertsitatea, UPV-EHU. Apdo. 644, 48080 Bilbao, Spain. hominin that used Mode 1 technology. Sites that have yielded ev- E-mail address: [email protected] (A. Gomez-Olivencia). idence of the presence of this hominin can be found in the Iberian http://dx.doi.org/10.1016/j.quascirev.2015.05.001 0277-3791/© 2015 Elsevier Ltd. All rights reserved. A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74 53

Peninsula (Carbonell and Rodríguez, 1994; Rodríguez et al., 2011; 1.1. Objectives Vallverdú et al., 2014) and France (de Lumley et al., 1988), among others, reaching as far north as Pakefield (Parfitt et al., 2005). While The recent assessment of the paleontological remains from there are older hominin remains in Europe (Carbonell et al., 2008; Punta Lucero (PL) quarry, which included part of the collection Toro-Moyano et al., 2013), the fossil record of this hominin for the that was never published, has resulted in the identification of period between 0.8 and 1.0 is restricted to the TD6 layer from Gran new taxa and the re-identification of other taxa, which results in Dolina and has been used to name a new species, Homo antecessor a Galerian (Middle Pleistocene) association and is thereby older (Carbonell et al., 1995; Bermúdez de Castro et al., 1997). From a than the first assessment carried out by Castanos~ (1988),which faunal point of view, there was a total faunal turnover, which marks suggested it was from a time between Riss and Würm or at the the end of the Villafranchian and the beginning of the Galerian beginning of Würm. The objectives of this research are to pre- (ca 1.0e0.9 Ma; Azzaroli et al., 1988; Bellucci et al., in press, and sent for the first time a complete description of the faunal references therein). The period of co-existence between Villa- assemblage of the site, a taphonomic study of all the non-dental franchian faunas and Galerian elements (ca 1.2e0.8 Ma) is known remains, a geochemical analysis of the sediments attached to the as the Epivillafranchian by several authors (Kahlke, 2007; bones and a biochronological assessment of the assemblage. The Madurell-Malapeira et al., 2010; Kahlke et al., 2011; Bellucci et al., faunal association will be compared to other sites from the same in press) region. Around 600 kyrs BP, there is the oldest evidence available for the arrival of new macromammal taxa in Central and Western 1.2. Punta Lucero Quarry site Europe such as the (Bos primigenius). It is also roughly the earliest evidence in existence of a new hominin, generally known The construction and quarrying that was conducted in the as Homo heidelbergensis, and of the earliest appearance of Mode 2 building of the outer part of the drastically altered technology in Europe. The phylogenetic relationship between the the NE flank of the Punta Lucero mountain (307 m.a.s.l.; Zierbena; late Early Pleistocene and the Middle Pleistocene hominins is still Bizkaia) when a quarry was established there. The work a matter of debate (Bermúdez de Castro et al., 1997, 2003; Arsuaga completed at the quarry revealed a large that was visible et al., 1999; Martinon-Torres et al., 2007; Gomez-Robles et al., from the right side of the Ría de Bilbao. In September of 1987 one 2007, 2013). member of our team (IL) together with M.C. Salas, visited and In summary the period roughly between 1.0 and 0.4 Ma BP, prospected the different artificial terraces that were being left thus, saw not only several climatic oscillations and changes in during the works at the quarry. In the uppermost artificial terrace the fauna (Kahlke et al., 2011; Rodríguez et al., 2011), but also an old karstic deposit was discovered that yielded several macro- cultural changes and the replacement of a hominin species by mammal remains. The main excavation of the site was performed another.Therefore,thescarcenumberofsitesofthesechro- by Pedro Castanos~ (PC) and by IL at different times during 1988 nologies is vital to our understanding of the tempo and mode of (Castanos,~ 1988, 1989). An additional excavation was performed by these changes as well as of the context for the latest technolo- PC in 1992 due to imminent destruction of the site by the quarry gies based on flakes and the first human occupations with hand (Castanos,~ 1993; see Supplementary Fig. 1) followed by a final visit axes. to the site by IL who discovered additional carnivore remains from The Iberian Peninsula contains some sites from both the end of a sediment patch at the highest point of the sedimentary infilling the Early Pleistocene (e.g., Gran Dolina levels TD4-6; Vallparadís (Fig. 1). Section; Madurell-Malapeira et al., 2010, 2014; Rodríguez et al., The karst of Punta Lucero extends throughout in the Lower 2011) and the middle part of the Middle Pleistocene: Gran Dolina Albian () from the southern flank of the level TD10, Galería, Sima de los Huesos, Ambrona, Aridos, among synclinorium of Bizkaia. The original entry to the site was within others (Sese and Soto, 2005; Yravedra et al., 2010; Arsuaga et al., argiolitic limestones, which currently comprise the majority of 2014). Most of these sites are either from the continental and the mountain, below a calcareous crest that runs along the peak Mediterranean region of the Iberian Peninsula, and whether or not of this mountain (Castanos,~ 1988). The sedimentary sequence is they are representative of the whole peninsula remains uncertain. typical of an intermediate carbonate ramp, which occasionally The Cantabrian Mountain range that extends to the East along the would receive shallower resedimented products from an unstable Basque mountain range until it reaches the Pyrenees currently acts shallower talus, corresponding to grainstone-type calcarenites as an ecogeographic barrier between the Northern part (Euro-Si- (García-Mondejar et al., 2006). The site was located high in the berian) and the rest of the Iberian Peninsula (Carrion et al., 2010). quarry at about 200e220 m.a.s.l. (Castanos,~ 1988) in a wall, The past distribution of certain species has also been affected by approximately perpendicular to the main wall of the quarry at this barrier (e.g., the reindeer: Alvarez-Lao and García, 2011; that point, which corresponds to a joint plane (Fig. 2; Gomez-Olivencia et al., 2014). Moreover, certain Basque sites such Supplementary Fig. 2). as Lezetxiki II have yielded a record of European species (e.g., Sicista The partial destruction of the site at the time of discovery betulina; Mammalia; Dipodidae; Muscardinus avellanarius; Mam- together with the sediment's exposure to the elements, the degree malia; Gliridae) that have not been found elsewhere in the Iberian of diagenetic breakage (Supplementary Fig. 3) and fast and pre- Peninsula (Rofes et al., 2012; Garcia-Ibaibarriaga et al., 2015), carious excavation conditions have conditioned the study of the underscoring the connection that exists between the northern material. Although it is not possible to reconstruct the original fringe of this region and the rest of Europe. geometry of the site, the preservation of one of its sides, together The Early to Middle Pleistocene evidence found in the Iberian with the observations made in other nearby in the same northern fringe is less abundant and not as well preserved. This mountain, provide us with a general idea about it. It was initially record, with the exception of Mestas de Con (Crusafont, 1959)or interpreted as a pit by Castanos~ (1988). Based on our observations, Santa Isabel de Ranero (Torres et al., 2001), is mainly associated we can add that the site was likely limited by joint planes with the end of the Middle Pleistocene with sites such as La Parte, perpendicular to the main mining wall of the quarry. In fact one of Arlanpe or Lezetxiki and Lezetxiki II (Falgueres et al., 2005e2006; these vertical joints can be observed in Fig. 2. The sediment infilling Alvarez-Lao and García-García, 2006; Castanos~ et al., 2011; was composed of clays mixed with bones and blocks. Arrizabalaga and Rios-Garaizar, 2012; Rios Garaizar et al., 2013). When the site was discovered, there still remained a patch of 54 A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74

Fig. 1. Location of the Punta Lucero quarry site in the eastern Cantabrian region in the Northern Iberian Peninsula close to the city of Bilbao. The site is located at the end of the Nervion-Ibaizabal estuary. This map portrays other sites mentioned in the text. sediment of ca 2.5 m (vertically) by 1.5 m (horizontally). At the base 2.2. Geochemical analysis of the sequence the remains of two large feline taxa were found, while the top of the sequence yielded several canid fossils recov- The geochemical analysis of the sediments found at from the ered by IL after the last excavations. From the base of the sequence site was performed on two different samples. The first one (called to the current surface of the mountain, there was a minimum PL88-1) was taken by IL during the site excavation. The second one vertical distance of 4e4.2 m. (PL12-1) was sediment still attached to the fossils that was removed from them prior to their cleaning, at the end of 2012 e 2. Material and methods beginning of 2013 at the Arkeologi Museum of Bilbao. These two samples were observed by one member of our team (VMP) under a 2.1. Material stereoscopic zoom microscope in order to look for small verte- brates, a search that was unfruitful. After the identification and The material from Punta Lucero Quarry studied here comprises separation of the detritic matrix, a small subsample (3.3 g for PL88- the entire collection stored at the Arkeologi Museoa (Bilbao, 1 and 2.18 g for PL12-1) from each sample was separated in order to Bizkaia): both the elements excavated by P. Castanos~ and I. Libano perform an X-ray diffraction analysis. The analysis of the total rock (Castanos,~ 1988, 1989) and P. Castanos~ (Castanos,~ 1993) and ele- composition was carried out in the laboratories of the Parque ments found by I. Libano before and after these excavations which Científico Tecnologico de la Universidad de Burgos (Spain) using have remained unpublished. The collection yielded 782 individ- Bruker D8 Advance (Davinci Design) equipment. The diffracto- ually labeled items (PL.1ePL.782). One of these, an ovicaprid tooth grams obtained were processed with the Diffrac.EVA 2.1 (Bruker) (PL.759), was excluded from the study since it belongs to a software at the same university. The analysis of the clay mineralogy modern specimen based on its external appearance (different was conducted in the SgiKer laboratories of the Universidad del País external patina, with a more “fresh” appearance; it likely got Vasco/Euskal Herriko Unibertsitatea. Diffractograms were mixed in with the rest of the collection when the site was measured using a diffractometer PANalytical Xpert PRO equipped discovered). From the 781 Pleistocene fossils, it was possible to with a copper tube (1CuKa mean ¼ 1.5418 Å, 1CuKa1 ¼ 1.54060 Å refit 40 fragments which yielded a total of 741 specimens. From and 1CuKa2 ¼ 1.54439 Å), a vertical goniometer (Bragg-Brentano these, it was possible to taxonomically assess 301 elements. Many geometry), a programmable divergence slit, an automated sample indeterminate elements belonged to long bone shaft fragments, changer, a secondary graphite monochromator and a PixCel de- 134 of which were determined as belonging to a large size un- tector. In order to analyze the oriented aggregates of the <2 mm gulate (likely Bovini) and 29 to a medium size ungulate (likely fraction, three aggregates per sample were measured: 1) without Cervus). treatment; 2) after solvation with etilenglicol during 48 h at room A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74 55

Fig. 2. General (a) and closer (b, c) views of the Punta Lucero site during its excavation in 1988. The red shadow represents approximately the extent of the sediment excavated. 1 ¼ Approximate location of the Canis mosbachensis remains; 2 ¼ approximate location of the Homotherium latidens and Panthera gombaszoegensis. Remains; 3 ¼ Cone formed from the excavated sediment and blocks. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.)

temperature; 3) after solvatation with dimetilsulfoxide during Photographs were taken with the digital video microscope DINO- 72 h at 80 C. LITE AM-TFVW-A (DinoCapture 2.0 software). All the taxonomically identifiable remains, as well as those that 2.3. Paleobiological analysis only provide a size of the were studied. The body size was established based on weight and age of the ungulates (Bunn, 1986). The taxonomic assessment was conducted using standard In this work we consider: i) Medium-sized ungulate (MSU) as adult osteological atlases as well as modern and fossil samples housed in specimens of Cervus and megacerine deer and ii) Large sized- different research centers. We are well aware of different schools of ungulates (LSU) as adult specimens of Bos, Bison, and Stephano- thought in the naming of some taxa. In this work, we basically used rhinus genera. In the taxonomically unidentifiable remains, this the taxonomy used in Rodríguez et al. (2011). The fragmentary grouping was based on the thickness of the cortical bone. Those nature of the remains found at PL precluded any attempt to reach bones with cortical thicknesses around 1 cm were considered LSU. the subspecies level, especially in the red deer remains, where the The carnivore sample was studied independently of the ungulate absence of antler remains impeded its classification as “coronatus” body size. or “acoronatus”. In the case of the Homotherium remains they were The taphonomic observations were divided into physical alter- ascribed to Homotherium latidens due to the difficulty for some ations, biological alterations and fracturation type. The study of the authors to allow a species-level division within the Villa- physical alterations comprises presence/absence of crusting and franchianeGalerian within this genus (Anton et al., 2014). Distinc- the weathering stages following Behrensmeyer (1978). Regarding tion between Bos and Bison followed Lehmann (1949), Bibikova the biological alterations, several observations were made. The (1958), Stampfli (1963), Sala (1986), Slott-Moller (1990), Gee presence/absence of dissolution marks produced by roots, tram- (1993) and Sala et al. (2010). Age-at-death for red deer followed pling marks, and anthropogenic modifications including those Brown (1991). Age-at-death determination of the large bovid produced by fire and lithic industry were recorded. The last bio- dentition was hampered by the lack of associations. It has been logical alteration refers to that caused by carnivores. Tooth marks attempted only in M3s following Wegrzyn and Serwatka (1984) and on bone surfaces were classified into pits, punctures, furrowing, Niven and Hill (1998). The biometrical assessment follows the scores and dissolution due to gastric acids (Haynes, 1980, 1983; standards (Driesch, 1976). Maguire et al., 1980; Binford, 1981). Punctures, scores and pits were measured (length and width) in accordance with previous 2.4. Taphonomic analysis studies (Selvaggio and Wilder, 2001; Domínguez-Rodrigo and Piqueras, 2003; Delaney-Rivera et al., 2009). Metric data were A Nikon SMZ800 (stereoscopic zoom microscope) was used to compared to experimental data gathered from bones modified by examine surface modification on bone remains. The dental remains extant carnivores (Domínguez-Rodrigo and Piqueras, 2003; Andres were excluded and, thus, a total of 539 remains were studied. et al., 2012; Sala, 2012; Sala and Arsuaga, 2013; Saladie et al., 2013; 56 A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74

Sala et al., 2014). Finally, the fracturation study was carried out Table 2 pursuant to the criteria proposed by Villa and Mahieu (1991). Semi-quantitative estimation of the filosilicates in the studied samples. Sample PL12-1 PL88-1

2.5. Geochemical analysis %Illite 29 37 %Caolinite 39 40 Both samples are of a clayish nature and are partially cemented %Illite/Esmectite 32 23 with small limestone fragments. There are abundant macrofaunal bone fragments that measure smaller than 2 cm, but no micro- oldest adult (>50 months) would be represented by a left M3 faunal remain has been found. Within sample PL12-1 there is a (PL.100), which shows a fully functional M3. A younger adult would carbonate cemented limestone breccia fragment. be present with the erupted M3, but this individual's hypoconulid The total rock analysis using X-ray diffractometry (Table 1) has shows very little wear, and the P2 is still not functional yielded similar results in both samples. Clay minerals (especially (PL.97 þ 109 þ 116, PL.104, PL.105, PL.111, PL.127, PL.133). Wear in illite and with more than 40%) are the most abundant the hypoconulid starts around 26e33 months (Brown, 1991) and minerals in both samples, and both iron oxides (goethite) and thus an age-at-death of around 3 years would be reasonable for this calcium carbonates (calcite) show percentages close to or below 2%. individual. The third individual would be represented by a left d4 The groups of phylosilicate identified in the measured oriented (PL.123). aggregates were illite, caolinite and illite/smectite in- The length of the M3 is within the range of other Middle and Late terstratifications. Regarding the latter, the diffraction peaks were at Pleistocene samples (Table 4). Additional measures of the most 13 Å, at 9.40 Å and at 5.25 Å, and thus they are interpreted as complete postcranial elements (i.e., talus, first and second pha- interstratified illite/smectite minerals (Lynch et al., 2007) with a langes) can be found in Tables 5e7. While the talus is within a Late 30% of swelling smectitic layers. Finally the results of the content of Pleistocene sample from the nearby site of Labeko Koba (Table 5), different clay minerals of the <2 mm fraction using the “reflectant the length of the first phalanx surpasses the range of different powers” are presented in Table 2. samples (Table 6). Made et al. (2014) synthesized information about In summary, the similar mineralogical content is consistent with size fluctuations of C. elaphus throughout the Pleistocene. The size the hypothesis that both samples have the same origin. The mineral of the proximal epiphyses of the first phalanges from PL (Table 5)is association is typical of detritic sediments. The high percentage of similar to the MIS 11-14 samples and smaller than MIS 16-18 terrigenous elements, together with the presence of goethite sug- samples from Made et al. (2014). gests an allochtonous (i.e., exokarstic) origin of the sediment as well as the presence of vermiculite/kaolinite, whose formation is caused Megacerini indet. by the gradual transformation of illite and feldspars due to the increasing level of soil weathering (Arriolabengoa et al., 2015). There are only four specimens belonging to a megacerine cervid (described by Castanos~ as belonging to Megaloceros sp.): a complete 3. Systematic paleontology anterior second phalanx, two second phalanx proximal fragments and a very eroded pyramidal (Fig. 4). Castanos~ (1988) described the From the 741 specimens (after refitting), it was possible to distal radius epiphysis PL. 667 as also belonging to Megaloceros but taxonomically assess 301 elements (Table 3). Many indeterminate its general size more likely corresponds to C. elaphus elements belonged to long bone shaft fragments, 134 of which were (Supplementary Table 1). The measurements taken on the mega- determined as belonging to a large size ungulate (likely ) and cerine's second phalanges from PL are smaller than those from the 29 to a medium size ungulate (likely Cervus). Megaloceros sp. from Lezetxiki (Altuna, 1972, Table 8) and those from the Praemegaceros verticornis specimen from Bilhausen. In Order Artiodactyla Owen, 1848 regard the proximal breadth, they are also below the mean of the Family Cervidae Goldfuss, 1820 specimens presented by Lister (1994), which cluster between 33 Cervus elaphus Linnaeus, 1758 and 35 mm, though they fall within the variability range of his samples. With 25.58% of the NISP, the red deer is well represented by both cranial (mainly dentition) and postcranial elements such as long Table 3 bones, carpal and tarsal bones, and phalanges (Fig. 3). No antler Composition of the Punta Lucero Quarry faunal assemblage. remains were found and thus a subspecific assessment was not Taxa NISP %NISP MNI %MNI attempted. There are a minimum of three individuals (two adults and one immature) represented in PL based on different eruption Cervus elaphus spp. 77 25.58 3 16.66 and wear stages and the presence of three left first incisors. The Megacerini indet. 4 1.33 1 5.55 Bovini indet. 109 36.21 cf. Bos primigenius 25 8.31 4 22.22 cf. Bison sp. 38 12.62 5 27.77 Table 1 Stephanorhinus sp. 3 0.99 1 5.55 Composition (as a %) of the sediment from the Punta Lucero Quarry site based on X- Homotherium latidens 8 2.66 1 5.55 ray diffractometry. Panthera gombaszoegensis 11 3.65 1 5.55 Vulpes sp. 4 1.33 1 5.55 Mineral Sample PL12-1 Sample PL88-1 Canis mosbachensis 12 þ 3? 4.98 1 5.55 Illite 45.25 45.21 Carnivora indet. 7 2.34 Quartz 43.66 42.35 Total ungulates 256 85.04 14 77.78 Kaolinite/Vermiculite 8.34 8.47 Total carnivores 45 14.96 4 22.22 Goethite 2.00 1.93 Total 301 100 18 100 Calcite 0.75 2.04 NISP ¼ Number of identified specimens. MNI ¼ Minimum number of individuals. Sample PL12-1 corresponds to sediment attached to the fossils obtained when When the complete large bovid collection is taken as a whole, a minimum of 6 in- 12 fossils were cleaned at the end of 2012. dividuals is found based on the number of M and I2. When morphological dif- Sample PL88-1 corresponds to a sediment sample obtained by a member of our ferences that distinguish Bos and Bison are taken into consideration, the MNI is team (IL) during the excavation of the site in 1988. higher. A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74 57

Fig. 3. Selected cranial and postcranial Cervus elaphus remains from Punta Lucero.

Table 4

Length of the M3 of the Cervus elaphus compared to other sites.

Site Label Side Length Reference

Punta Lucero PL.111 r 32.5 This study PL.100 l 35.5 Labeko koba Mean (n) 32.93 (12) Altuna and Mariezkurrena, 2000 Range 31.0e35.0 Swanscombe Mean (n) 28.0 (3) Lister, 1986 Range 27.8e28.3 Arago-Upper Sequence (E-G) Mean ± SD (n) 32.16 ± 1.87 (30) Lister, 1990b Range 29.3e37.0 Arago-Lower Sequence (E-G Mean ± SD (n) 32.86 ± 1.44 (9) Lister, 1990b Range 20.0e35.3

Family (Gray, 1821) The large bovid remains constitute over 50% of the elements Tribe Bovini (Gray, 1821) (NISP) found in PL. Despite the absence of well-preserved horn cf. Bos primigenius remains, the morphology of the dental and postcranial elements cf. Bison sp. suggests the presence of two bovine taxa: cf. Bos primigenius and cf.

Table 5 Measurements (in mm) of the talus of Cervus elaphus from Punta Lucero compared to other samples.

Label/Site Side Lateral greatest Medial greatest Proximal Distal breadth Reference length (GLl) length (GLm) breadth (BP) (BD)

PL.087 l 61.3 57.1 38.4 37.8 This study Labeko koba Mean ± SD (n) 61.7 ± 3.2 (13) 57.1 ± 2.7 (11) 39.3 ± 2.1 (12) Altuna and Mariezkurrena, level IX (LP) 2000 Range 55.0e65.0 52.5 34.5e42.0 58 A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74

Table 6 Measurements (in mm) of the proximal phalanges of Cervus elaphus from PL compared to other Late and Middle Pleistocene samples.

Site/Species Label Position Abaxial length Proximal Shaft Maximum width of the References (GLpe) breadth (Bp) width (SD) distal epiphysis (Bd)

Punta Lucero PL.144 21.5 This study PL.132 65.5 22.9 20.8 20.9 PL.163 20.7 Labeko Koba (LP) Mean ± SD A (n ¼ 22) 56.32 ± 1.58 23.16 ± 0.88 18.16 ± 0.92 22.15 ± 1.06 Altuna and Mariezkurrena, Min-max A 54.0e60.0 22.0e25.0 16.9e20.5 20.5e24.5 2000 Mean ± SD P (n ¼ 14) 60.75 ± 1.14 23.36 ± 0.60 18.65 ± 0.54 22.69 ± 0.74 Min-max P 59.5e63.5 23.0e25.0 17.8e20.0 21.5e24.0 Camino-Pinilla Mean ± SD 57.4 ± 0.8 20.6 ± 1.1 Buitrago, 1992 del Valle (LP) Minemax 55.7e57.5 19.1e21.6 Mosbach (MP) Mean ± SD 22.39 ± 1.56 Lister, 1990a Minemax 18.7e28.2

A ¼ Anterior. P ¼ Posterior. LP ¼ Late Pleistocene. MP ¼ Middle Pleistocene.

Table 7 Measurements (in mm) of the medial phalanges of Cervus elaphus from PL compared to other Late Pleistocene samples.

Site Label Position Greatest Proximal Shaft width Maximum width of the References length (GL) breadth(Bp) (SD) distal epiphysis (Bd)

Punta Lucero PL.091 22.6 This study PL.081 A 42.6 21.7 17.8 18.5 This study PL.159 P 48.6 22.1 18.0 19.1 This study Labeko Koba (LP) Mean ± SD A (n ¼ 23) 42.89 ± 1.85 23.08 ± 0.59 16.78 ± 0.65 20.20 ± 0.97 Altuna and Mariezkurrena, Minemax A (n ¼ 23) 39e46.5 22.0e24.5 15.8e18.7 18.6e21.5 2000 Mean ± SD P (n ¼ 22) 44.74 ± 1.54 23.14 ± 0.87 17.94 ± 0.71 19.69 ± 0.94 Minemax P (n ¼ 22) 42.5e47.0 21.5e25.0 16.7e19.3 18.2e21.5

A ¼ Anterior. P ¼ Posterior. LP ¼ Late Pleistocene.

Bison sp., which concurs with the previous study by Castanos~ classified as Bovini indet. The co-occurrence of the genera Bos and (1988; this author attributed the Bison remains to Bison priscus). Bison is not unusual in both Middle and Late Pleistocene sites These attributions should be regarded as tentative, given the large (Auguste, 2009; Sala et al., 2010), as well as in different Upper degree of fragmentation and the aforementioned lack of well- Paleolithic cave-art sites regardless of being of Magdalenian (e.g., preserved horncore remains. There are, however, many fossils in Altamira, Altxerri, Tito Bustillo, etc.) or pre-Magdalenian (i.e., Arco which this distinction was not possible and, thus, they have been A, Lluera, Pasiega, etc.) chronology (Cacho Toca, 1999).

Fig. 4. Different views of two medial phalanges classified as belonging to a megacerine deer (Megacerini indet.). A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74 59

Table 8 Measurements (in mm) of the medial phalanges of the megacerine deer (Megacerini indet.) from PL compared to other samples.

Site/Species Label/Position Greatest length Proximal breadth Shaft width Maximum width of the Reference (GL) (Bp) (SD) distal epiphysis (Bd)

Punta Lucero PL.669 30.6 This study PL.666 56.2 30.8 24.8 26.1 Megaloceros sp. Lezetxiki (l.VI) Anterior 62.0 36.5 28.5 30.5 Altuna, 1972 Praemegaceros verticornis Bilhausen Anterior 61.0 33.5 26.5 Pfeiffer, 2002 Posterior 64.0 33.8 26.9

Table 9

Length of the M3 of the large bovids from Punta Lucero compared to other sites.

Site Taxon Label Side Length Reference

Punta Lucero cf. Bison sp. PL.446 r 44.9 This study PL.319 r 46.0 This study PL.327 l 45.9 This study PL.326 r 47.3 This study PL.447 l 45.1 This study cf. Bos primigenius PL.733 r 46.5 This study PL.744 r (42.5) This study (n ¼ 21) Bos primigenius Mean 48.8 Sala, 1986 Range 46.0e52.8 Charterhouse Warren Farm Bos primigenius l 45.6 Everton, 1975 La Borde Bos primigenius Mean ± SD (n) 46.34 ± 0.86 Slott-Moller, 1990 Minemax 42.7e51.2 Heidelberg (n ¼ 24) Bison schoetensacki Mean 42.8 Sala, 1986 Range 38.8e48.0 Isernia (n ¼ 30) Bison schoetensacki Mean 43.1 Sala, 1986 Range 40.0e49.0

The presence of another large sized species bovid (Hemibos record of this genus in the Iberian Peninsula is restricted to Hemibos galerianus) should be noted in the Early-to-Middle Pleistocene sp. aff. Hemibos gracilis from the early Pleistocene site of Venta transition of Ponte Galeria and Ponte Milvio (; Petronio and Micena (Orce) (Martínez-Navarro et al., 2011). Therefore, although Sardella, 1998; Martínez-Navarro and Palombo, 2004, 2007). The this genus is present in the Middle Pleistocene and the presence of

Table 10 Measurements (in mm) of the Bos primigenius metacarpal remains from PL compared to different samples.

Site Label Side Proximal breadth (Bp) Proximal depth (Dp) Distal breadth (Bd) Distal depth (Dd) Reference

Punta Lucero PL.309 R 86.3 50.3 This study PL.328 R 92.2 50.6 This study La Borde Mean ± SD (n) 70.5 (1) 43.5 (1) 72.32 ± 2.89 (5) Slott-Moller, 1990 Minemax 69.5e76.0

Table 11 Measurements (in mm) of the first proximal phalanx of large bovid from PL compared to different samples.

Label Position Abaxial Proximal Shaft width Maximum width of the Reference length (GLpe) breadth (Bp) (SD) distal epiphysis (Bd)

Punta lucero PL.329 Anterior 92.3 43.9 41.6 42.9 This study Bison priscus Cava Filo 20 Anterior 78.6 41.5 41.0 45.5 Sala 1986 18 Anterior 75.8 45.0 40.0 47.6 17 Anterior 78.0 45.7 42.0 43.0 Bison schoetensacki Isernia 1 Anterior 77.8 42.0 38.3 42.0 Sala, 1986 2 Anterior 76.3 37.0 32.4 35.0 Bison priscus Habarra Mean ± SD Anterior 76.4 ± 4.5 43.5 ± 4.4 38.5 ± 2.8 42.2 ± 4.1 Prat et al., 2003 Minemax Anterior 72.5e81.0 38.0e47.0 36.0e42.0 38.5e46.0 n4554 Liptovska Mara II Mean ± SD 81.11 ± 1.28 37.03 ± 1.07 32.07 ± 1.43 37.76 ± 1.98 Chroszcz et al., 2011 () n ¼ 7 Minemax 79.68e83.02 36.6e39.5 29.5e33.52 36.6e38.45 Charterhouse Warren Mean ± SD 83.4 ± 3.78 42.0 ± 1.41 Everton, 1975 Farm (n ¼ 5) Minemax 79.0e87.0 40.0e44.0

Values between brackets indicate preserved values. Everton (1975) provided maximum lengths of the phalanges. 60 A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74

Table 12 Measurements (in mm) of the medial anterior phalanx of large bovid from PL.

Site/Species Label Greatest Proximal Shaft Maximum width of the Distal depth Reference length (GL) breadth (Bp) width (SD) distal epiphysis (Bd) (Dd)

Punta Lucero PL.316 56.7 45.3 38.8 44.1 48.4 This study PL.028 [51.5] [45.9] 39.3 42.1 48.2 This study PL.318 62.9 46.6 36.5 37.0 44.9 This study Bison priscus Cava Filo (n ¼ 4) Mean ± SD 53.1 ± 0.84 42.65 ± 2.92 36.4 ± 3.35 39.22 ± 3.37 41.0 ± 3.23 Sala, 1986 Minemax 52.0e54.0 38.8e45.5 33.4e40.0 35.7e42.4 36.2e43.2 Bison schoetensacki Isernia Mean ± SD 54.6 ± 1.86 44.43 ± 1.10 34.50 ± 2.62 36.80 ± 2.87 40.37 ± 2.19 Sala, 1986 Minemax 51.8e57.2 42.0e45.8 32.2e41.0 32.0e42.0 37.5e43.4 n109 109 9 Bison priscus Habarra Mean ± SD 53.9 ± 3.2 46.7 ± 4.0 36.1 ± 3.5 40.4 ± 4.3 Prat et al., 2003 (anterior phalanges) Minemax 50.0e58.0 40.0e49.5 30.5e39.0 34.5e45.0 n6 5 6 6 Bos primigenius Charterhouse Warren Mean ± SD 50.5 ± 1.00 39.0 Everton, 1975 Farm (n ¼ 3) Minemax 54.0e56.0

Values between brackets indicate preserved values. this genus cannot be ruled out in PL, we consider its presence un- were hampered by the lack of associate dental rows. Based on the likely. In any case, new findings should help to establish the pale- M3 remains (NISP ¼ 6; MNI ¼ 3) one individual (PL.319, which ochronological and paleogeographical spread of this genus. could be the antimere of PL.475) would be between 2.7 and 3.9 Bos primigenius would be represented by four individuals based years of age. The other two individuals would be older than 5 years on the presence of seven upper M1-2. Age-at-death determinations old (M3 full wear and hypoconulid connected to hypoconid) but less were hampered by the lack of associate dental rows. Based on the than 12 (ectostylid worn but not connected). M3 remains (NISP ¼ 3; MNI ¼ 2) and the Bison dental eruption and The length of the lower M3 of both bovine taxa from Punta wear, one of the individuals (PL.744) would represent an individual Lucero are generally within the limits of similar taxa (Table 9). less than 2.5 years of age and the other (PL.594 þ PL.733, which are Regarding the postcrania, the metacarpal remains attributed to likely antimeres) would be older than 5 years old (M3 full wear and Bos primigenius from PL is larger than the Middle Palaeolithic hypoconulid connected to hypoconid) but less than 12 (ectostylid sample from La Borde (Table 10). The Bovini complete first phalanx worn but not connected). PL.329 also belonged to a large individual: its maximum length is Bison sp. would be represented by five individuals based on the above different Bison samples from Sala (1986) and closer to (but presence of five right I2. Once again, age-at-death determinations still above of) Bos primigenius samples (Table 11). The second

Fig. 5. Selected dentition attributed to cf. Bos primigenius from the Punta Lucero site. A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74 61

Fig. 6. Selected dentition attributed to cf. Bison sp. from the Punta Lucero site.

phalanges from PL are also in the upper limits or above the range of of the shaft and a posterior first phalanx which was previously different large bovid samples (Table 12)(Figs. 5 and 6). determined by Castanos~ (1988) to be a posterior second phalanx from the fourth row. There are four rhino taxa in the Pleistocene of Order Perissodactyla Owen, 1848 Europe: Stephanorhinus etruscus, Stephanorhinus kirchbergensis, the Family Rhinocerotidae Owen, 1845 Stephanorhinus hundsheimensis-hemitoechus line age and Coelo- Stephanorhinus sp. donta antiquitatis (Lacombat, 2009; Made, 2010). Castanos~ (1988) attributed the PL material to Dicerorhinus sp. (as a senior subjec- Punta Lucero has yielded four fragments that constitute three tive synonym). Firstly, he ruled out attributing it to C.antiquitatis Rhinocerotidae specimens: the fragment of a metapodial shaft based on the general gracility of the PL material. This author also (PL.90; maximum preserved length ¼ 48.6 mm), a left fourth suggested that the measurements were closer to Stephanorhinus metatarsal that lacks the proximal articulation (Table 13) and part hemitoechus rather than to S. kirchbergensis.

Table 13 Raw measurements (in mm) for the fourth metatarsal from Punta Lucero compared to other samples.

Site Label Length (L) Shaft width Distal transverse Distal antero-posterior Reference (SD) diameter (DTd) diameter (DAPd)

Punta lucero PL.89 þ 92 [155.5] 30.2 [35.1] 40.3 This study Stephanorhinus hundsmeihensis Different sites Mean (n) 161.31 (8) 27.33 (9) 32.0 (5) 37.86 (7) Guerin, 1980 Range 152.5e166.5 22.0e35.5 29.0e35.0 35.0e40.0 Stephanorhinus kirchbergensis Several sites Mean (n) 178.17 (3) 34.75 (4) 40.25 (4) 48.83 (3) Guerin, 1980 Range 170.0e182.5 33.5e36.5 27.0e43.0 44.4e51.5 Stephanorhinus hemitoechus Several sites Mean ± SD (n) 150.44 ± 11.10 (8) 27.31 ± 5.28 (8) 32.75 ± 3.81 (8) 37.06 ± 3.81 (8) Guerin, 1980 Range 136.0e169.5 20.0e34.5 27.0e39.0 32.5e43.0 Atapuerca-Galeria 150.6 28.3 34.2 Cerdeno~ and Sanchez, 1988 143.3 28.0 28.9 Alfaguara 25.8 24.0 Cerdeno,~ 1990 Pinilla del Valle (Camino) 32.6 Cerdeno,~ 1990 Pinilla del Valle (Camino) 32.4 Cerdeno,~ 1990 Congosto 32.8 33.8 Cerdeno,~ 1990 Neumark-Nord HK88:14, 21(l) 163.8 39.4 43.0 Made, 2010 HK88:14, 22(r) 163.9 39.1 43.6 Coelodonta antiquitatis Several sites Mean ± SD (n) 144.85 ± 6.15 (40) 30.56 ± 3.34 (40) 36.1 ± 2.43 40.79 ± 2.22 Guerin, 1980 Range 127.0e155.0 24.0e40.0 31.0e41.0 36.0e46.0

Values between brackets represent preserved values. 62 A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74

Fig. 7. The two most complete remains attributed to Stephanorhinus sp. from the Punta Lucero site. PL.89 þ 92 is a left fourth metatarsal and PL.96 is a phalanx.

In this study, a more in-depth comparative analysis has been presence of not only cranial but also of postcranial remains in PL is performed on the PL rhino material. The preserved length of the noteworthy, such a H. latidens pisiform that can be readily taxo- fourth metatarsal (155.5 mm) exceeds a large (n ¼ 40) sample of nomically distinguish from Panthera (Anton et al., 2005). From a Coelodonta antiquitatis. At the same time, the minimum width of metric point of view, the Homotherium remains from PL are within the diaphysis (SD) and the distal antero-posterior diameter (DAPd) the ranges of variation of sites such as Incarcal (Galobart et al., values fall below the range of a sample of S. kirchbergensis but are 2003, Table 14), among others. The pisiform is slightly longer within (or close to) the range for the samples of S. hundsheimensis than that of Seneze (Ballesio, 1963, Table 15). and S. hemitoechus published by Guerin (1980). Therefore, given the absence of dental material, we concur with previous studies that 3.1. Panthera gombaszoegensis the rhino material can be classified as Stephanorhinus sp. (Fig. 7). There are 11 remains, comprising both cranial and postcranial Order Carnivora Bowdich, 1821 remains (see Supplementary Tables 2 and 3), that have been Family Felidae Gray, 1821 attributed to a single (P. gombaszoegensis) individual (Fig. 9). Homotherium latidens (Owen, 1846) Some of these remains were first attributed by Castanos~ (1988) to Panthera leo. Our metrical analysis of the dentition demonstrates There are eight fossils attributed to this taxon comprising dental that the metrics of the Punta Lucero canine and P3 specimen are and one postcranial remain that have been attributed to a single closer to other P. gombaszoegensis specimens and below P. leo adult individual of H. latidens (Fig. 8). PL would be the second site (Fig. 10 and Tables 16 and 17; see also Fig. 5 by Marciszak (2014)) (after Mestas de Con in Asturias, see below) in the Northern fringe though some overlap occurs in the case of P4. Thus, the pantherine of the Iberian Peninsula that has yielded remains of this taxon. The felid from PL is a large-sized jaguar that could belong to a male A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74 63

Fig. 8. All the Homotherium latidens remains identified in Punta Lucero. All the dental remains are in buccal view except the left lower canine, which is also shown in lingual view.

individual. PL is the first site in the northern fringe of the Iberian northern fringe of the Iberian Peninsula that has yielded remains of Peninsula that has yielded remains of this taxon. this taxon. The second posterior cusplet of P4 is low and separated from the posterior cingulum (Fig. 12), which is a feature that dis- Family Canidae Fischer von Waldheim, 1817 tinguishes the C. mosbachensis line (García, 2003). The material Vulpes sp. from PL fits well within the range of variation of the large samples from Pirro Nord or CereCave( Table 19, Fig. 13). Four upper teeth: a canine, two P4s and an M1 have been recovered in PL (Fig. 11). From a metric point of view, the M1 is 3.2. Taphonomic analysis within the range of variation of both Vulpes praeglacialis and Vulpes vulpes (Table 11) and larger than Alopex lagopus. The transverse The taphonomic study was hampered by the difficulty of the diameter of the P4s (the only variable that can be measured on observation of the bone surfaces in most of the collection due to: these teeth) is below the range of variation of different samples of alterations and recent (excavation) breakaged and presence of V. vulpes (Tables 11 and 18). sediment/crust on the bone surface, among others. Table 20 dis- plays the results on the bone surface modifications (Fig. 14). Canis mosbachensis Soergel,€ 1925 There are a few alterations that may possibly be anthropogenic in origin (see discussion below). These are marks of intentional In the uppermost part of the site, Punta Lucero has yielded a breakage (percussion marks on long bone diaphysis) that are low in series of carnivore teeth (see above) most of which can be classified number (NISP ¼ 6; 1.1%). Some incisions were not considered here as canid and likely belong to the same individual of C. mosbachensis as they did not display the typical anthropogenic cut-mark (Fig. 12, Table 19; Supplementary Table 4). PL is the first site in the morphology. 64 A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74

Table 14 Raw measurements (in mm) for the Homotherium lower dentition from Punta Lucero compared to other samples.

Site Label/level (side) Lower I3 Lower C Lower P3 Lower P4 Lower M1 References

A-P Tr A-P Tr A-P Tr A-P Tr A-P Tr

Punta Lucero Label (side) PL.2 (l) PL.1(r)/PL.780(l) PL.781(l) PL.13(l) PL.19(r) This study Value 11.6 10.3 15.2/15.5 10.1/10.2 7.8 5.5 21.3 8.7 30.5 12.3 Gran Dolina-TD5 TDE5-T61-H16-10/ 13.1/13.0 9.2/9.4 García and Arsuaga, TDE5-T63-F18-1 1999 Incarcal IN-I 1057 (l) 12.2 10.8 15.5 11.7 10.8 4.6 21.1 9.6 36.2 13.5 Anton et al., 2014 IN-I 484(I3) (r)/346(C) 11.1a 9.0a 14.6a 9.4a 8.3a 5.7a 31.7 13.1 Galobart et al., 2003; (l)/8(P3)/322(M1) Galobart, pers. com. IN-I 549(I3) (l)/819(C) (r) 10.9a 9.2a 14.7a 9.6a IN-I 828(r)/830(r) 11.2a/10.7a 8.8a/8.6a IN-I 826 (l) 10.5 12.2 16.3 10.5 23.5 10.8 33.8 13.0 IN-II 5 14.8a 9.4a IN-V 1 (r) 14.9a 10.5a 19.0 9.2 27.2 11.7 IN-V 239 (I3) (l)/295(M1) (r) 11.2a 8.6a 33.4 12.6 IN-V 312 (r) 10.2 8.7 10.2 8.7 9.9 29.0 12.8 Perrier 11.0 9.0 15.0 11.0 6.5 5.0 18.2 9.8 32.5 13.5 Bonis, 1976 Sainzelles 24.0 10.5 34.0 12.5 Boule, 1901 (In Sardella and Iurino, 2012) Domegliara V 8911 23.7 10.0 28.0 Sardella and Iurino, 2012 Monte Peglia M.P.159-160 21.4 9.5 34.0 13.7 Sardella and Iurino, 2012 Seneze 11.0 11.0 8.0 5.5 22.0 11.0 32.0 13.0 Ballesio, 1963 Venta Micena 7.7 5.2 20.4 8.7 28.8 11.3 Pons-Moya, 1987 Untermassfeld 1993/24372 19.4 8.5 30.4 11.9 Hemmer, 2001 1997/25985 21.0 8.6 30.5 12.1 Semibalki-3 OP-827 8.1 5.7 20.0 29.9 Sotnikova and Titov, 2009 Liventsovka RSU/94 10.0 24.7 35.4 Sotnikova and Titov, 2009 Kuruksay 3120/344 8.0 21.5 29.9 Sotnikova and Titov, 2009 Fairbanks AM142492 (19.3) 8.6 Anton et al., 2014 AM142493 20.4 9.2 (32.0) 11.3 AM142494 9.6 10.4 18.2 8.2 AM142497 30.9 11.4

(r) and (l) refer to right and left sides. A-P ¼ antero-posterior diameter (mesio-distal). Tr ¼ transverse diameter (buco-lingual). Values between parentheses indicate estimated values. a Measurement taken at the base of the crown (Galobart et al., 2003).

Carnivore marks are clearly distinguished despite their low the fossil remains have been exposed to atmospheric agents. number (1.3% of the analyzed remains). These marks are pitting Around 17% of the remains are in stage 1 (according Behrensmeyer, (NISP ¼ 5), punctures (NISP ¼ 1) and scores (NISP ¼ 2). The size of 1978), 7.24% belongs to stage 2 and 2.78% are at stage 3. In some these tooth marks (Supplementary Table 5) are similar in size to cases signs of trampling were observed (2.97%) as well as signs of those produced by large-sized carnivores (lions, , ). dissolution by plant roots (2.04%). Thus, it is likely that the carnivore marks present in the PL collec- tion were made by any of the large felids (H. latidens or P. gom- 3.3. Biochronological assessment baszoegensis) present in the site. On the other hand, the study of fracture patterns on PL assem- PL has yielded several taxa that are useful for a biochronological blage indicates the presence of green bone breakage (curved purposes. First, the latest occurrence of C. mosbachensis is recorded orientation, smooth surfaces, oblique fracture angles and low pro- in L'Arago and L'Escale (France, MIS 12, ca 450 kyrs) (García and portion of complete diaphysis circumferences) in both, medium Arsuaga, 1999; Moigne et al., 2006) and this would provide a and large-sized ungulates. However, there are also transverse likely minimum age for PL. The earliest presence of an elaphine fractures represented by jagged and right edges, indicating fossil- deer in the Iberian Peninsula is found in the TD3eTD4 levels of diagenetic fracturing (dry bone) following the criteria of Villa and Trinchera Dolina (Sierra de Atapuerca, Burgos; Rodríguez et al., Mahieu (1991). 2011), located at the end of the Early Pleistocene and this would Regarding the non-biological modification, we observed that provide a maximum age for PL. The first record of Bos primigenius over 25% of the sample show signs of weathering, indicating that has been recently found in Tunisia with a chronology of ca 700 kyrs which shows that the genus Bos evolved in and then Table 15 dispersed out at the beginning of the Middle Pleistocene (Martínez- Raw measurements (in mm) for the Homotherium left pisiform from Punta Lucero compared to other samples. Navarro et al., 2014). In Europe the oldest occurrence of Bos pri- migenius is at the site of Venosa-Notarchirico, Italy, at a Punta Lucero (PL.14) Seneze ~500e600 kyrs (Piperno, 1999 in Martínez-Navarro et al., 2007), Maximum length 38.0 34.0 which, in case its presence could be completely certified in PL, Largest diameter of the articular head 21.0 23.0 would likely reduce the maximum age for PL. Based on the size a Ballesio, 1963. fluctuations provided by Made et al. (2014), the size of the A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74 65

Fig. 9. Selected Panthera gombaszoegensis cranial and postcranial remains from Punta Lucero.

C. elaphus from PL is closer to the MIS 11e15 sites of Hundsheim, derived phyllosilicates from existing soils and the presence of Mauer, Miesenheim than to those from MIS 16e18 sites of Süs- goethite which indicates an allochtonous precedence of the senborn, Voigtstedt, West Runton or Kozi Grzbiet. However, it sediment. should be noted that there is size overlap between the sites of these In PL, no remains were found. The abundance of bear chronologies. Thus, the size of the Cervus from PL would be remains and the presence of decidual dental remains are normally consistent with a younger age for the site between MIS 11e15 as associated with a hibernation den for this taxon, something that suggested by the likely presence of cf. Bos primigenius, but this can be excluded for PL. It should be noted, however, that bear cannot be used by itself as a chronological approximation. In remains can appear in sites interpreted as natural traps, and the Europe, Homotherium is already present at the beginning of the closest example of this is Kiputz IX (Castanos~ et al., 2012). The Pleistocene and becomes rare during the Middle Pleistocene scarcity of carnivore tooth marks, in addition to the absence of (Sardella and Iurino, 2012) after the entrance of P. leo, a likely coprolites and the absence of carnivore juveniles indicate that PL competitor, and there is only one record in the late Pleistocene was not a carnivore den (i.e., den) (Kuhn et al., 2010). (Reumer et al., 2003). The presence of Homotherium in PL is However, the aforementioned preservation issues should be taken consistent with the biochronological approach provided by other into consideration. A member of our team (IL) participated during taxa, but does not provide additional biochronological accuracy, the first excavations and neither in these nor in the latter no lithic which is also the case of P. gombaszoegensis. In summary, PL shows artifact was recovered. The absence of clear anthropogenic marks an association consistent with a Middle Pleistocene attribution. (see above) leads to believe that PL was not of anthropogenic origin. In summary, we favor the hypothesis that PL acted as a 4. Discussion natural trap. Nevertheless, due to the presence of some evidences of tooth and percussion marks in the assemblage (although very The partial destruction of the site at the time of discovery scarce in number), we cannot rule out the possible sporadic ac- together with the exposition of the sediment to the elements, and cess by hominins and large carnivores to the accumulated car- fast and precarious excavation conditions have conditioned the casses after their accumulation in the site by natural causes. study of the material. Whether PL was an open trap for a long or short period of time is a matter that is difficult to assess. In the case of Kiputx IX 4.1. Site formation (Castanos~ et al., 2014), dates throughout the sequence show that slightly more than 2 m of the sequence were accumulated in Given the geometry of the site and the knowledge that we around 10 kyrs, and taking the entire sequence into account, 3 m have of the karst formation in the Punta Lucero mountain, it is were accumulated in 20 kyrs. While it is difficult to extrapolate likely that PL was a karstic cavity that was developed through these numbers to PL, it is reasonable to assume that the filling of the joints and acted as a natural trap. Geochemical analyses the trap was relatively fast compared to the proposed chronology show a high percentage of terrigenous elements, the presence of of the site. 66 A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74

Fig. 10. Bivariate (upper canine and P3) and univariate (P4) comparison of the Punta Lucero Panthera gombaszoegensis remains with other samples of P. gombaszoegensis and Panthera leo.

4.2. Chronology With regards to the fauna present and the approximate chro- nology of the site, the only parallel in the Cantabrian region of the During the excavation, there was no stratigraphical information Iberian Peninsula is Mestas de Con (Crusafont, 1959). The macro- systematically recorded and we assume that all the faunal remains mammal species from this site include Homotherium crenatidens, recovered in the site could correspond to a single association. The Ursus cf. Ursus etruscus, Equus cf. E. sussenbornensis, Dicerorhinus taxonomic assessment of PL has not revealed any taxon not etruscus, Orthogonoceros? sp., Cervus cf. C. elaphus, Capreolus cap- consistent with the rest of the association. If there was indeed more reolus, B. priscus. The presence of Homotherium and an elaphine than one level in the site, the presence of C. mosbachensis in the deer indicate that this is an association roughly between 400 and uppermost part of the preserved sediment of the site directly 800 kyrs. Recent changes in the nomenclature would likely alter the provides a minimum chronology for the underlying faunal ele- taxonomy in this site. The megacerine deer (Orthogonoceros? sp.), ments. The absence of microfaunal remains in the collection im- represented by two upper molars would be classified within genus pedes further refinition of the chronological assessment based on Megaloceros or Praemegaceros, and the rhinoceros would be clas- the macro- remains. sified within genus Stephanorhinus. Unfortunately, the remains from this site have been lost (Alvarez-Lao, pers. comm.). 4.3. Parallels in the northern Iberian Peninsula and France Santa Isabel de Ranero (SIR), 25 km south west to PL as the crow flies, has yielded an important collection of Ursus deningeri that has It should be noted that all the taxa found in PL have been found been dated using bear dentine aspartic acid racemization (Torres in other sites from the Middle Pleistocene of the Iberian Peninsula. et al., 2001, 2002) and which has yielded a similar (or even A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74 67

Table 16 Raw measurements (in mm) for the Panthera gombaszoegensis dentition from Punta Lucero compared to extant and fossil jaguar samples.

Site Label/Other information Upper I3 Upper C Upper P3 Upper P4 References

A-P Tr A-P Tr A-P Tr A-P Tr

Punta lucero PL.18(l) PL.3(r)/PL.5(l) PL.17(r) PL.12(r) This study 13.2 10.2 21.8/22.0 17.6/16.9 23.8 11.2 33.0 [13.5] Panthera gombaszoegensis Huescar-1 HU1/86 A92 (Canine), 17.5 13.7 28.1 14.9 This study A49 (upper P4) Untermassfeld Mei 21775 (C); 20.8 16.9 22.0/22.1 11.0/11.2 32.0/32.2 18.4/18.0 Hemmer, 2001 Mei 20268 (P3eP4) L'Escale CD-795 22.7 12.2 32.0 16.4 Bonifay, 1971 CD-765(P3)/CD-764(P4) 23.1 11.8 32.3 CD-7634(P3)/CD-1142(P4) 23.1 11.5 31.5 18.7 Westbury-sub-Mendip Mean 21.4 16.9 21.7 10.7 18.5 Bishop, 1982 Range 18.5e24.1 15.0e19.3 29.7e33.0 n552 Gombaszog€ 32.9 19.0 Kretzoi, 1937/38 Petralona Range 23.6 11.9e13.0 Kurten and (n ¼ 1) (n ¼ 2) Pouliamos, 1977 Erpfingen 30.1 15.0 Lehmann, 1953 Panthera toscana Toscana Mean 19.6 14.7 21.3 9.1 30.0 15.8 Del Campana, Range 17.0e22.6 11.4e19.0 17.5e24.0 8.0e10.0 26.8e31.8 13.7e17.3 1915e1916 n 556566 Panthera onca augusta Extant Mean 21.1 17.6 19.9 10.6 31.3 16.1 Kurten, 1965, Range 19.5e23.0 16.1e20.3 16.9e22.3 9.2e12.2 26.8e35.0 13.1e18.7 1973 n 101110131311 Panthera onca Extant (Smithsonian Mean ± SD 18.2 ± 1.9 14.8 ± 1.6 17.5 ± 1.4 9.2 ± 0.8 26.9 ± 1.8 13.9 ± 1.4 This study Institution) Range 14.6e21.6 10.7e17.2 15.5e20.3 7.8e11.3 23.6e30.0 11.0e16.0 n 191920192020

The PL sample is composed of PL.18 (a left upper I3), PL.3 (a left upper canine), PL.17 (a right upper P3) and PL.12 (a right upper P4). Values between parentheses indicate estimated values. Values between brackets indicate preserved values. (r) and (l) refer to right and left sides.

slightly older) chronology of that of Sima de los Huesos (SH) would likely be older than Galeria due to the presence of Canis (ca 430 kyrs, Arsuaga et al., 2014; see below). The Santa Isabel lupus in the latter, and younger than TD8 and closer to TD10 and collection, however, lacks the most primitive morphotypes present Sima de los Huesos if the Bos primigenius presence could be verified. in Sima de los Huesos, which could be a sample bias due to the Gran Dolina is a well-known archaeo-paleontological site with a smaller collection from the former compared to the latter (Torres long sequence starting at the end of the Early Pleistocene and et al., 2001). Alternatively, it would mean that SIR represents a encompassing most of the Middle Pleistocene (Rodríguez et al., younger population than SH and thus, the differences in amino- 2011). Of special interest in this work are the levels TD8 and chronology would be the result of differences in fossilization con- TD10-3. All of these levels show a normal polarity and are therefore ditions (Torres et al., 2001). Due to the absence of U. deningeri in the younger than 780 kyrs (Pares and Perez-Gonz alez, 1999). PL as well PL assemblage, it is difficult to correlate these two sites, and we as SH, lies chronologically between the older TD8 and more recent could consider them as either roughly contemporaneous, or more TD10-3 and GIIb. likely, PL could be slightly older. The presence of Mimomys savini in TD8 gives a minimum age of There are additional sites in the Basque area that have yielded 500 kyrs for this level. Middle Pleistocene chronologies, but more towards its end: Goi- Combined ESR-U-series dating of ungulate tooth enamel has koetxe, Lezetxiki and Lezetxiki II, and Arlanpe. Direct dating of a yielded a mean value of 602 ± 52 kyrs (Falgueres et al., 1999) while rhinoceros tooth at Goikoetxe cave using dentine aspartic acid the Luminescence dating has yielded 820 ± 140 (Berger et al., racemization has yielded a date of 211.9 kyrs BP (Edeso et al., 2011). 2008). The carnivore association includes P. gombaszoegensis, Cro- Archaeopaleontological sites, such as Lezetxiki (levels VIeVII; cuta crocuta, C. mosbachensis, Hyaena sp. and Ursus sp. The un- Falgueres et al., 2005e2006; Alvarez and Arrizabalaga, 2012) and II gulates include Dama vallonnetensis, Equus altidens, Bison cf. (Castanos~ et al., 2011) and Arlanpe (Rios-Garaizar et al., in press), B. voigtstedtensis, Sus scrofa, Stephanorhinus etruscus, Hippopotamus which are more recent, would be located at the end of the Middle sp., Eucladoceros giulii, C. elaphus cf. C.e. acoronatus and Megaloceros Pleistocene. Finally, the site Azurtoki located in Ea (Bizkaia; Espejo solilhacus (Rodríguez et al., 2011). and de Torres, 1969), which has yielded only horse remains which The carnivore association of TD10-3 includes Homotherium sp., were used to describe a new subspecies, Equus caballus eaensis,was Ursus sp., V. vulpes and P. leo cf. P. l. fossilis (García and Arsuaga, suggested to have a pre-Riss chronology (de Torres, 1970). Some 2011). Ungulates include Hemitragus bonali, Dama dama clactoni- horse remains from this site were later directly dated using ami- ana, Equus ferus, C. elaphus priscus and maybe a small sized Bison noacid racemization which yielded a MIS5 chronology around (Rodríguez et al., 2011). 100 kyrs BP for this likely natural trap (Torres and Ortiz, 2006). GIIb level in Galeria has been proposed to be slightly younger Therefore, the best parallel to the faunal spectrum in Punta than TD10-3 and thermoluminescence dating was performed in the Lucero is found in the Middle Pleistocene sites of Gran Dolina, layers above (GIIIa: 466 ± 39) and below (GIIa-top: 422 ± 55) Galeria and Sima de los Huesos (all of which are located in the Si- (Berger et al., 2008; Rodríguez et al., 2011). The carnivore associa- erra de Atapuerca, 120 km south from PL as the crow flies). PL tion of GIIb comprises P. leo cf. P. l. fossilis, pardinus spelaeus, V. 68 A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74

Table 17 Raw measurements (in mm) for the Panthera gombaszoegensis dentition from Punta Lucero compared to extant and fossil lion samples.

Site Level/Label Upper C Upper P3 Upper P4 References

A-P Tr A-P Tr A-P Tr

Punta Luceroa PL.3(r)/PL.5(l) PL.17(r) PL.12(r) This study 21.8/22.0 17.6/16.9 23.8 11.2 33.0 [13.5] Panthera leo Petralona PEC 90 31.0 25.0 27.9 13.6 39.6 23.6 Baryshnikov and Tsoukala, 2010 Mokhnevskaya 41.2 30.1 32.6 19.1 41.8 22.9 Baryshnikov and Tsoukala, 2010 Gailenruth 26.2 19.4 36.0 18.6 Baryshnikov and Tsoukala, 2010 Ehringsdorf (MIS5e) EHR 8102 30.7 22.7 30.1 15.1 This study Taubach (MIS 7/5e) TAU 1532 28.9 14.9 This study Azoleta (Gorbea) 24.7(r) 17.4(r) 25.4(l)/25.5(r) 12.9(l)/12.9(r) 34.2(l)/34.8(r) 18.2(l)/18.0(r) Castanos,~ 2005 Lezetxiki Level VI 30.2 14.3 Altuna, 1972 Level VI 22.7 31.0 16.1 40.8 Extant Mean ± SD (n) 36.9 ± 2.3 (44) 19.4 ± 2.2 (44) García, 2003 MineMax 32.3e39.7 15.6e22.0 Panthera leo fossilis Zandobbio MCSNC 5127 24.2(r)/24.1(l) 13.7(r)/13.6(l) 36.6 17.0 Bona, 2006 Isernia 39.4 20.8 Sala, 1990 Sima de los Huesos 39.0 21.5 García, 2003; Arsuaga et al., 2014 P.leo fossilis 39.9 ± 3.8 Schütt and Hemmer, 1978 P.leo fossilis Mean (n) 26.7 (3) 16.0 (3) 39.9 (6) 19.6 (4) Schütt, 1969 Range 23.8e29.3 14.5e19.2 36.4e45.1 18.3e21.7 Panthera leo spelaea/Panthera spelaea (Western Europe) Panthera leo spelaea 38.4 ± 3.0 Schütt and Hemmer, 1978 Panthera leo spelaea/Panthera spelaea () Kondakovka K-1 28.7 19.5 28.0 16.3 40.0 20.8 Sotnikova and IPBPS-1 31.5 25.0 24.5 14.4 34.0 al Nikolskiy, 2006 Smolensk GIN-1123A 28.1 20.5 25.0 35.0 al Avdeevo GIN/AV86-12 41.1 21.0

Values between parentheses indicate approximate values. Values between brackets indicate preserved values. al ¼ alveolar. a For the labels of the PL material see Table 16. vulpes, Meles meles, C. lupus, Cuon alpinus europaeus, Mustela nivalis Pleistocene (Arsuaga et al., 1997). It also acted as a natural trap for and Mustela putorius (García and Arsuaga, 2011). Ungulates include carnivores and has a large carnivore diversity in which U. deningeri Megaloceros solilhacus, Hemitragus bonali, D. dama clactoniana, is the most common species, followed by Vulpes vulpes, with the Equus ferus, C. elaphus priscus, a small sized Bison, Stephanorhinus cf. presence also of P. leo fossilis, Panthera cf. P. gombaszoegensis, L. S. hemitoechus and Equus cf. E. hydruntinus (Rodríguez et al., 2011). pardinus spelaeus, Felis silvestris, M. meles, Canis sp., M. nivalis, M. Sima de los Huesos has not yielded ungulate remains and it putorius and Martes martes (García and Arsuaga, 2011; Arsuaga contains the largest accumulation of human fossils for the Middle et al., 2014).

Fig. 11. Vulpes sp. remains from PL. A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74 69

Table 18 Raw measurements (in mm) for the Vulpes sp. upper dentition from Punta Lucero compared to other samples.

Site/Region Upper C Upper P4 Upper M1 References

A-P Tr A-P Tr A-P Tr

Punta lucero Label (side) PL.333 (r) PL.330 (r)/PL.332 (l) PL.331 (r) This study Value 5.9 3.8 [11.9]/[10.3] 4.7/4.7 9.5 11.1 Vulpes praeglacilis Dolina-TD5 9.4 12.3 García, 2003 Il Tasso 7.6 10.3 García, 2003 Inferno 8.5 10.7 García, 2003 L'Escale 8.1; 8.4 9.8; 10.6 Bonifay, 1971 Saint Vallier 7.8 10.4 García, 2003 (original description in Viret, 1954) Alopex lagopus Crimea Mean ± SD (n) 7.60 ± 0.68 (9) 9.42 ± 0.57 (9) Baryshnikov, 2006 MineMax 7.0e8.6 8.9e10.3 Vulpes vulpes Sima de los Mean ± SD (n) 6.44 ± 0.57 (17) 4.43 ± 0.44 (17) 14.74 ± 0.68 (13) 6.27 ± 0.36 (13) 9.79 ± 0.53 (23) 11.74 ± 0.72 (23) García, 2003 Huesos MineMax 5.1e7.3 3.6e5.3 13.3e16.3 5.8e7.0 8.7e10.8 9.9e13.0 L'Escale Mean ± SD (n) 13.87 ± 0.26 (7) 6.68 ± 0.60 (6) 9.48 ± 1.33 (5) 11.62 ± 1.03 (5) Bonifay, 1971 Extant Mean ± SD (n) 6.56 ± 0.35 (26) 4.23 ± 0.32 (48) 13.87 ± 0.51 (48) 7.29 ± 0.44 (50) 9.42 ± 0.38 (50) 12.79 ± 0.55 (49) Gingerich and Winkler, 1979 MineMax 6.0e7.4 3.4e5.4 12.7e15.1 6.5e8.3 8.6e10.4 11.5e13.8

Values between brackets indicate preserved values.

In France, the “marne blanche” level of the Carpentier Quarry in 1997) also constitutes a parallel for PL. The faunal list for levels Abbeville (Auguste, 2009 and references therein) has yielded some FeHincludesLynx sp., Dinobastis sp. (¼Homotherium sp.), P. taxa comparable to those from PL: Bison sp., Bos primigenius, gombaszoegensis, Panthera spelaea, Panthera sp. ( size), Praemegaceros verticornis, Stephanorhinus hundsheimensis and H. Canis cf. C. etruscus, Ursus cf. U. deningeri, Cervus cf. latidens, together with other taxa not present in PL: Mammuthus C. simplicidens, Hemitragus sp., indet., Dicerorhinus meridionalis, Palaeoloxodon antiquus, Hippopotamus incognitus, S. (¼Stephanorhinus)cf.D. hemitoechus.The“Breches ” I-IV include scrofa mosbachensis, Equus mosbachensis and a stenonid Equus. This P. spelaea, Crocuta sp., M. putorius, M. meles cf. M.m. atavus, Canis site was compared to the MIS 15/17 faunas from Pakefield or cf. C. etruscus, Vulpes cf. V. praeglacialis, Ursus cf. U. deningeri, Mosbach and is suggested to belong to MIS 15 (Auguste, 2009) Cervus cf. C. simplicidens, Capreolus capreolus, Hemitragus sp., which is coincides with the radiometric ages (ESR on quartz) that Bovinae indet., Bison sp., Dicerorhinus (¼Stephanorhinus)cf. have yielded mean value of 600 ± 90 kyrs BP (Laurent, 1993 in D. hemitoechus. A dating places this site around 390 kyrs BP, and Auguste, 2009). the pollen indicates that it was not a cold period (and thus rules The layers that comprise the “Ensemble” moyen (Breches IeIV out MIS 12, 14 and 16). For this site Guadelli (1997) proposes a and levels FeH) of the Grotte XIV (Dordogne, France) (Guadelli, date around MIS 11.

Fig. 12. Selected remains of Canis mosbachensis from Punta Lucero. 70 A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74

Table 19 Raw measurements (in mm) for the Canis mosbachensis postcanine dentition from Punta Lucero compared to other samples.

Site Upper M2 Lower P2 Lower P4 Lower M3 References

A-P Tr A-P Tr A-P Tr A-P Tr

Canis mosbachensis Punta Lucero Label (side) PL.771(r) PL.776 (l) PL.763 (r) PL.775 This study Values 8.1 11.7 [7.7] 4.2 14.2 6.4 4.7 4.5 Huescar 1 HU1-66 A314 8.6 12.9 This study. Original description by Alcala and Morales (1989) Pirro Nord Mean (n) 7.8 (5) 11.1 (5) 9.9 (7) 4.5 (7) 13.7 (12) 6.5 (12) Sardella et al., 2015 Cere Cave Mean (n) 8.2 (16) 12.0 (16) 11.7 (55) 5.3 (56) 14.6 (98) 6.8 (98) 5.2 (13) 4.6 (12) Sardella et al., 2015 Untermassfeld Mean (n) 8.0 (6) 12.8 (6) 10.1 (13) 5.0 (12) 13.7 (17) 6.4 (17) 4.9 (5) 4.4 (5) Sotnikova, 2001 Gombaszog Mean (n) 8.3 (7) 11.0 (7) 11.3 (17) 5.1 (16) 14.4 (14) 6.5 (14) 4.8 (1) 4.2 (1) Sardella et al., 2015 Vertessz ol€ os€ Mean (n) 11.6 (4) 5.4 (4) 14.5 (2) 7.1 (2) Sardella et al., 2015 Soave 7.0 12.0 Sardella et al., 2015 Cal Guardiola 13.2 5.9 Madurell-Malapeira et al., 2009 L'Escale Mean ± SD 7.5 ± 0.4 11.5 ± 0.5 13.7 ± 0.8 4.7 ± 0.5 Bonifay, 1971 (n) (11) (11) (35) (19) Mosbach Mean ± SD 11.7 ± 0.0 5.5 ± 0.4 14.7 6.9 4.5 4.5 Baryshnikov and (n) (2) (3) Tsoukala, 2010 Westbury 12.1 5.6 14.8 7.1 4.7 4.4 Canis arnensis Petralona Mean ± SD (n) 6.2 10.9 10.14 ± 0.47 4.92 ± 0.36 13.10 ± 0.34 6.43 ± 0.43 4.5 ± 0.8 3.7 ± 0.6 Baryshnikov and (5) (5) (9) (9) (2) (2) Tsoukala, 2010 Canis lupus Extant Mean (n) 11.68 (14) 5.85 (14) 15.35 (14) 7.60 (14) Perez et al., 2010 Range 10.46e12.55 5.3e6.91 14.4e16.3 6.88e8.41

Values between parentheses indicate estimated values. Values between brackets indicate preserved values.

Fig. 13. Bivariate comparison of the Punta Lucero Canis mosbachensis remains with other European C. mosbachensis samples.

The lower levels (Ib-III) of Artenac, which were suggested to fall (Tournepiche, 1996). The younger levels (IVeV) have a faunal list between 500 and 700 kyrs have a faunal list that includes U. that includes U. deningeri, P. leo spelaea, Dinobastis (¼Homotherium) deningeri, P. gombaszoegensis, Dinobastis (¼Homotherium) latidens, latidens, C. lupus cf. C.l. lunellensis, Vulpes vulpes, Gulo gulo, E. mos- Ursus sp., Canis sp., Rangifer tarandus, Bovinae and Equus sp. bachensis, Coelodonta sp., Elephantidae indet., Bovinae indet., Cer- vus sp., R. tarandus, S. scrofa. The presence of reindeer, in addition to Table 20 the presence of the wolverine and woolly rhinoceros in the upper Surface alterations on the non-dental remains from Punta Lucero Quarry. levels point to less mild climatic conditions (Tournepiche, 1996). NISP %a In summary, compared to the best sequences in the North of the Percussion fractures (percussion cones and impact points) 6 1.11 Iberian Peninsula, i.e., those from the different sites of Atapuerca, PL Carnivorous activity marks 4 0.74 would be older than Gran Dolina-TD10-3 due to the presence of C. Trampling 16 2.97 lupus in this level but it could be younger than Gran Dolina-TD8 if Meteorization 146 27.09 the presence of cf. Bos primigenius in PL is verified (Rodríguez et al., Roots 11 2.04 2011). a Based on a total NISP of 539. A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74 71

Fig. 14. Surface modifications in the Punta Lucero site. a) Trampling (PL.238); b) Roots (PL.438); c) Depression made by a carnivore (PL.23); d) Score made by a carnivore (PL.532).

4.4. Middle Pleistocene human presence in the Eastern Cantabrian traits mixed with Middle Paleolithic technological traits, which is region characteristic of the Late Acheulean (regional Early Middle Paleo- lithic) in this region (Turq et al., 2010; Rios-Garaizar et al., in press), The eastern Cantabrian region or western Pyrenees, is also and probably do not date back further than 300 kyrs. Regardless of known as the Basque crossroads due to its important location whether PL represents a site that is older than the Basque Middle linking the Iberian Peninsula to the rest of Europe (Arrizabalaga and Pleistocene archeological sites or whether it is perhaps roughly Rios-Garaizar, 2012). This region offers different mountain passes contemporary with open-air sites such as Irikaitz or Mendieta, PL that connect the northern coastal region to the Ebro valley and the helps provide us with the knowledge of at least some of the taxa Meseta that were used by and humans during the present in the large mammal communities that the first human Pleistocene. occurrences in the northern fringe of the Iberian Peninsula would The Middle Pleistocene archeological record is biased by have met. different geomorphological, administrative and research factors. Cave sites containing Middle Pleistocene archeological deposits are 5. Summary and conclusions scarce, the most relevant ones are Lezetxiki and Arlanpe where human occurrences are presumably not older than 300 kyrs given In this research the first complete analyses of the faunal and the available absolute dates (Arrizabalaga and Rios-Garaizar, 2012). sediment remains from the Punta Lucero quarry site were made. The identification of Middle Pleistocene sites in open-air sedi- This site was discovered during the quarry construction in Punta mentary deposits, such as fluvial terraces, is no easy task either, due Lucero mountain and is currently completely destroyed. This site to the intense anthropization and general orography of the region has yielded remains of a megacerine deer, red deer (C. elaphus), (steep relief, short but relatively strong rivers). Only a few sites have large bovids (likely representing both Bos primigenius and Bison yielded archeological materials associated with Middle Pleistocene sp.), a rhinoceros (Stephanorhinus sp.), the scimitar-toothed cat H. terraces, and the most significant one is Irikaitz (Arrizabalaga and latidens, a jaguar (P. gombaszoegensis), a (Vulpes sp.) and the Iriarte, 2002). There are several open-air sites located in the right C. mosbachensis. This site has yielded the first evidence for C. margin of the Ibaizabal-Nebioi river that yielded Middle Pleisto- mosbachensis and P. gombaszoegensis and the second evidence of cene archeological remains (Rios-Garaizar et al., 2012, 2013). genus Homotherium in the northern fringe of the Iberian Peninsula. However, these are individual findings of lithic remains except in This association is typical of a middle Middle Pleistocene chronol- one site, Mendibarrena, where two teeth of a large bovid were also ogy and would be the oldest macro-mammal site found to date in found. Given that none of these sites has been directly dated it is the eastern Cantabrian region. difficult to specify a chronological framework for these human PL is interpreted as a natural trap into which both ungulates and occupations. Several sites, such as Irikaitz, Mendieta, Moreaga and carnivores fell, and at least a large-sized carnivore (likely a felid probably lower levels at Lezetxiki yielded assemblages without based on the faunal list) altered some of the bones. bifaces and without the characteristics of the first Early Middle This site likely corresponds to a date after the Mode 1 techno- Paleolithic sites of SW Europe (Santonja et al., 2014). Therefore logical complex and before (or contemporaneus) the arrival of industrial assemblages at these sites were interpreted as Lower Mode 2 technology in Western Europe. Thus, it offers a glimpse into Paleolithic without bifaces, thus non-Acheulean, and an age be- the paleoecological conditions slightly prior to or contempora- tween MIS5 and MIS11 was proposed (Arrizabalaga and Rios- neous with the first the first human occurrences in the northern Garaizar, 2012). The other aforementioned sites show Acheulean fringe of the Iberian Peninsula. 72 A. Gomez-Olivencia et al. / Quaternary Science Reviews 121 (2015) 52e74

Acknowledgments Arsuaga, J.L., Martínez, I., Gracia, A., Carretero, J.M., Lorenzo, C., Garcia, N., Ortega, A.I., 1997. Sima de los Huesos (Sierra de Atapuerca, Spain). The site. J. Hum. Evol. 33, 109e127. We are indebted to Mª Carmen Salas for the help and support in Arsuaga, J.L., Martínez, I., Gracia, A., Lorenzo, C., 1999. The Sima de los Huesos crania the discovery of Punta Lucero Quarry site. Further thanks to J. Reyes (Sierra de Atapuerca, Spain). A comparative study. J. Hum. Evol. 33, 219e281. Auguste, P., 2009. Evolution des peuplements mammaliens en Europe du nord- Aranda for his help. This work received the support from the ouest durant le Pleistocene moyen et superieur. Le cas de la France septen- Diputacion Foral de Bizkaia/Bizkaiko Foru Aldundia. We acknowl- trionale. Quaternaire 20, 527e550. edge P. Castanos~ for his work at the site and his preliminary studies. Azzaroli, A., De Giuli, C., Ficcarelli, G., Torre, D., 1988. Late pliocene to early mid- We would like to thank the following people and institutions for pleistocene mammals in Eurasia: faunal succession and dispersal events. Palaeogeogr. Palaeoclimatol. Palaeoecol. 66, 77e100. their help in the different phases of the research: I. García Camino, Ballesio, R., 1963. Monographie d'un Machairodus du gisement villafranchien de S. Anibarro, J.L. Ibarra and L. García (Arkeologi Museoa), M. Unzueta Seneze: Homotherium crenatidens Fabrini. Trabaux Lab. Geol. Fac. Sci. Lyon (DFB-BFA), M. Izquierdo (Eusko Jaurlaritza), D. Alvarez-Lao, A. Nouv. Ser. 9, 1e129. Baryshnikov, G., 2006. Late Pleistocene arctic fox (Alopex lagopus) from Crimea, Galobart, J. van der Made, S. Madelaine, A. Sanchís, N. García-Ibai- Ukraine. Quat. Int. 142e143, 208e217. barriaga, R. Carter, J. Burgos, N. Pedrosa, E. Iriarte, A. Aranburu, and Baryshnikov, G.F., Tsoukala, E., 2010. New analysis of the Pleistocene carnivores our colleagues and friends from EHU, MNHN, UCM-ISCIII and BBP. from (Macedonia, ) based on the Collection of the The- e Thanks to R. Sardella and J. Madurell-Malapeira for their helpful ssaloniki Aristotle University. Geobios 43, 389 402. Behrensmeyer, A.K., 1978. Taphonomic and ecologic information from bone comments that have improved this manuscript. Thanks to Lauren weathering. Paleobiology 4, 150e162. Ames for her help with English corrections. Thanks to J.M. Carretero Bellucci, L., Sardella, R., Rook, L., 2015. Large mammal biochronology framework in (LEH-UBU) for the kind access to the modern comparison materials Europe at Jaramillo: the Epivillafranchian as a formal biochron. Quat. Int. (in press). under his care. AGO was the recipient of a Marie Curie Intra- Berger, G.W., Perez-Gonz alez, A., Carbonell, E., Arsuaga, J.L., Bermúdez de European Fellowship during part of this work. NS received a Post- Castro, J.M., Ku, T.-L., 2008. 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