Gibert Etal-1999-Human Molar Barranco Leon

HUMAN EVOLUTION Vol. 14 - N.
1-2 (3-19)- 1999
J. Gibert Molar tooth fragment BL5-0: the oldest

LI. Gibert human remain found in the Plio-Pleistocene
of Orce (Granada province, Spain)
S. Albadalejo
A molar tooth fragment from the Plio-Pleistocene Barranco Le6n
F. Ribot site 5 at Orce is shown to belong to Homo by analyses of its
F. Sfinchez enamel in terms of the arrangement of the striae of Retzius and
Hunter-Shreger bands, presence of perikymata, and of the
E Gibert thickness of the enamel, when compared with teeth of similar-
lnstitut Paleontol6gic sized mammals of other taxa.
"M. Crusafont"
Diputaci6 de Barcelona
Carrer de l'Escola Industrial 23
08201 Sabadell, Barcelona
SPAIN
Keywords: tooth human remain,

Plio-Pleistocene, Orce Spain
Introductory background
Although well-known remains uncovered at Boxgrove in England (Roberts et al., 1994;
Stringer et al., 1998) and Atapuerca in northern Spain (Carbonell et al., 1995) show that Homo
was established in early Middle Pleistocene Europe, there are much older human remains and
palaeolithic artifacts from Orce in southern Spain that testify to human presence in Europe as
early as the Plio-Pleistocene boundary. Identification of the Olduvai magneto-subchron at site
7 in the Barranco de Oree (BO-7) (Agust~ et al., 1997) scarcely 150 m from site 5 in the
Barranco Le6n (BL-5) strongly supports attributing a similar antiquity to the latter and its
assemblage, which includes part of a human tooth crown and root (hominid BL5-0), palaeolithic
artifacts, and faunal remains.
In 1983 a human calvarial fragment (VM-0) was reported from a Plio-Pleistocene faunal
assemblage at the Venta Micena site at Orce (Gibert, Agusti & Solh-Moy~, 1983). Recent
research confirming its human juvenile form (Gibert, Campillo, Arqu6s et al., 1998) is well
corroborated by biochemical and immunological analyses (Borja, Garcfa-Pacheco, Garcfa-
Olivares et al., 1997; Borja, Garcfa-Pacheco, Ram irez-L6pez & Garcia-Olivares 1992; Gibert,
Campillo, Arqu6s et al., 1998), notwithstanding some ongoing controversy provoked by opinions
that VM-0 could have belonged to a young equid (Agusti & Moy~-Sol~_, 1987; Moy?~-Sol~ &
Agusti, 1989; Moy~-Sol?~ & K6hler, 1998; Palmqvist, 1998); the Venta Micena excavations
have also provided undoubtedly human humeral shafts - - a nearly complete young juvenile
specimen (VM-1960: Gibert, Sfinchez, Malgosa, Walker et al., 1992; Gibert, Sfinchez, Malgosa
& Martfnez, 1994; Sfinchez et al., this volume, and refs.) and a fragment of an adult one (VM-
3691, ibidem). A complete human phalangeal bone (CV-0) and two fragments of adult human
4 J. GIBERT, LL. GIBERT, ALBADALEJO, RIBOT, SANCHEZ, P. GIBERT
humeri (CV-1, CV-2) have also been described from the karstic Cueva Victoria near Cartagena
in Murcia province, 200 kilometres east of Orce, (Gibert & P6rez-P6rez, 1989; Gibert & Pons-
Moy~, 1984; Gibert, Pons-Moy~ & Ruz, 1985, 1989; Gibert, Sfinchez, Malgosa, Walker et al.,
1992; Sfinchez et al., this volume, and refs.; Santamaria & Gibert, 1992), which has a faunal
assemblage comparable to Venta Micena and is of uncontestably Lower Pleistocene age,
although no palaeolithic artifacts occurred. At Orce, by contrast, Lower Pleistocene many
palaeolithic artifacts have been excavated in sealed layers, most particularly at Barranco Le6n
site 5 and Fuentenueva site 3a (Gibert, Gibert, lglesias & Maestro, 1998; Gibert J., lglesias
A., Maillo, A. & Gibert, Li., 1992; Martinez et al., 1997; Roe, 1995; Tixier et al., 1995), and
cut-marks on animal bones, and intentional breakage of bones by percussion, are both
documented at the Venta Micena site (Gibert, Ferrfindez et al. 1992; Gibert & Jim6nez, 1991;
Jim6nez & Gibert, 1992).
Geological and geochronological background
The NE sector of the Guadix-Baza basin in SE Spain (Fig. 1) displays a continental

sedimentary sequence over 100 m thick, representing continual sedimentation from the Middle
Pliocene to the Upper Pleistocene. Plio-Pleistocene sedimentary outcrops correspond to five
unconformity-bounded depositional cycles (Fig. 2), each beginning with fluviatile sediments
and ending with lacustrine ones - - deposition of the one or the other depended on the relative
height of the lake level which, in turn, reflected global climatic oscillations. The BL-5 deposit
lies in a bed of fine sand that belonged to the distal part of a small alluvial system. Excavation
of 20 m 2 here in 1995 uncovered part of a human molar tooth (Fig. 3), and a Hippoptamus
amphibius antiquus mandible surrounded by over 100 palaeolithic artifacts; the fauna includes
Castillomys cf. crusafonti, Mimomys sp., Allophaiomys pliocaenicus and Equus granatensis.
The Venta Micena site (VM) was a den used by scavenging animals (Gibert & Caporicci 1989;
Martinez, 1992a,b). An intriguing feature of the assemblage is presence of cut-marks on some
bones and of signs of intentional breakage by percussion on others (Gibert, Ferrfindez et al.
1992; Gibert & Jim6nez, 1991; Jim6nez & Gibert, 1992) and, as it also includes apparent
manuports, there are implicit indications of sporadical human activity here, although the
human remains show evidence of carnivore action. Stratigraphical analysis situates Venta
Micena between Barranco Le6n-5 (BL-5) and Fuentenueva-3a (FN-3a) in Lower Pleistocene
times, in accordance with the VM fauna: Homo sp., Desmana sp., Allophaiomys pliocaenicus,
Apodemus aft. mystacinus, Castillomys crusafonti ssp., Eliomis intermedius, Hystrix major,
Prolagus capensis, Oryctolagus cf. lacosti, Ursus etruscus, Canis etruscus, Canis falconeri,
Vulpes praeglacialis, Homotherium latidens, Megantereon whitei, Lynx sp., Pachycrocuta
brevirostris, cf. Meles, Mammuthus meridionalis, Equus granatensis, Stephanorhinus etruscus,
Hippopotamus amphibius antiquus, Praemegaceros sp., Cervidae gen et sp. indet., Praeovibos
sp., Bovini indet. (aft. Bubalus), Soergelia minor, Hemitragus alba, Testudo sp., Lacerta sp.,
Opidia indet., Rana sp., and Charadriiforme indet. (aft. Laridae) (Fig. 3). Fuentenueva site 3a
(FN-3a) was situated near the edge of the basin and its sedimentary outcrop reflects the
environment of the lake margin, containing variable amounts of detrital and organic matter.
Systematic excavation in 1995 uncovered numerous palaeolithic artifacts associated with remains
of large mammals in a faunal context of Mimomys sp., Hystrix major, Allophaiomys sp.,
Hippopotamus amphibius antiquus, Mammuthus meridionalis, Stephanorhinus etruscus, Equus
granatensis, Praemegaceros sp., Cervus sp., Bovini indet., Hemitragus sp., Megantereon sp.
and Ursus sp. (Fig. 3).
MOLAR TOOTH FRAGMENT BL5-0 5
Palaeomagnetic determinations demonstrate presence of five normal-polarity episodes in a

geological section near Galera, just west of Orce, another one in the Barranco de Orce section,
and another probable one in the Fuentenueva section (Figs. 2, 3). The Olduvai magneto-
subchron has been identified at Bai:ranco de Orce site 7 (BO-7) and because, only 150 m to
the east, Barranco Le6n site 5 (BL-5) lies in the same stratum, and both sites have similar
fauna, it is eminently reasonable to assign BL-5 to the Olduvai magneto-subchron (Garc6s,
1993; Garc6s et al., 1997; Gibert, Arribas et al., 1994; cf. Agusti et al. 1997) and hence the
hominid molar fragment BL5-0 must also belong to this same early time.
Hominid molar fragment
Only mesial parts of the crown and root remained after the tooth (BL5-0) underwent an
ancient bucclolingual fracture that was not, however, parallel to its buccolingual axis (Fig. 4,
Fig. 5a, 5b). A large wear facet is present on the crown and dentine is exposed on the occlusal
surface. The fragment belonged to an upper, possibly left, adult molar. Crown height on the
mesial face is 4.6 mm and the length of the broken root is 2.9 mm. The maximum enamel
thickness is 1.2 mm.
Material and research methods
Advances in technology and methodology enable unequivocal identification of BLS-0 as

human, as follows:
(1) Striae of Retzius and Hunter-Shreger (H-S) bands were located and investigated with a
polarized-light analyzer coupled to a Wild M3Z binocular microscope equipped with a camera
and source of cold light illumination (ltralux 5000) with polarizers. In order to enhance
contrast during observation and photography BL5-0 was immersed in alcohol.
The last imbricate stria was located in the fractured part of the enamel, and the H-S bands
could be followed throughout nearly the entire crown fragment. The position of the bands, and
the angles they formed with the dentine, allowed us to separate ancient from modern forms of
Homo (cf. Benyon & Wood, 1987). The parallel pattern of the H-S bands permitted the human
pattern to be distinguished from the non-parallel pattern found in carnivores 90 as reported in
hyaenas by Stefen & Rensberger (1995) and extended here to several other carnivore molars
(particularly bear molars).
(2) The enamel prism patterns were examined b y scanning electron microscopy (SEM).
Patterns were grouped by Boyde (1964, 1965) into three types: 1, 2 and 3. Hominids display
type 3 patterns, together with carnivores, pinnipeds, and proboscideans (Boyde, 1965, 1971).
Gantt and colleagues differentiated pattern types 3a and 3b, identifying the former in hominoids
and the latter in Australopithecus and Homo (Gantt, 1982; Gantt & Cring, 1981; Gantt,
Pilbeam & Steward, 1977). The BL5-0 fragment was fixed on to a stud with a conductive
adhesive and photographed with the SEM at a low operating voltage, without further processing.
The SEM images also revealed perikymata of the enamel.
(3) Changes in enamel thickness were compared in different groups of mammals because
enamel thickness is a key feature in identifying specimens of Homo (Boyde, 1964, 1965, 1971,
1978), Gantt (1977, 1979), Gantt et al. (1977), Lavelle et al. (1977), von Koenigswald (1977).
BL5-0 was compared with different mammal groups (especially Ursus third molar sections).
Results
Low-power microscopy, following Benyon & Wood (1986~ 1987), enabled observation of
the striae of Retzius and H-S bands on the fractured surface of the inner enamel. The arrangement
of the H-S bands was also observed on the buccal surface (Fig. 6). The first stria of Retzius of
the imbricate part of the enamel was followed throughout the enamel, thereby permitting data
given by Beynon & Wood (1986), given first here, to be compared with our findings on BL5-
0, given second, as follows: stria of Retzius angle 30-33 ~ BL5-0 30~ H-S angle 50-68 ~
BL5-0 80~ enamel thickness 1.2-2.1 mm, BL5-0 1.2 mm. The position of the last imbricate
stria suggests BL5-0 belonged to an early Homo (cf. Benyon & Wood, 1987) (Figs. 7, 8).
Other important features were observed with the SEM. The enamel prism pattern was visible
near the base of the crown, and corresponds to type 3b (or keyhole) (Figs. 9a, 9b) (cf. Gantt &
Cring, 1981). Perikymata were observed on the tooth surface (Fig. 10). Enamel thickness in
BL5-0 is characteristic of Homo and separated the specimen from teeth of small artiodactyls,
pigs, carnivores (particularly bears), and cercopithecids (Fig. 11).
Discussion
BL5-0 is distinguishable from teeth of several other mammals of comparable body size. It
has already been mentioned that human teeth are distinguishable from those of hyaenas.
Similarly, BL5-0 can be distinguished from teeth of cercopithecids, carnivores, felids, canids,
ursids, suids, and small artiodactyls: these are distinguishable from BL5-0 in terms of enamel
thickness (Fig. 11), arrangement of the enamel in the crown, non-parallel as against parallel H-
S bands (Fig. 6), enamel prism pattern (Figs. 10a, 10b), and presence or absence of perikymata
(Fig. 12).
Conclusions
Analyses of the associated BL-5 fauna, stratigraphical position, and palaeomagnetic findings,
strongly suggest that BL5-0 is the oldest known human remain in Europe. Furthermore,
Barranco Le6n 5 (BL-5) and Fuentenueva 3a (FN-3a). This implies that the Orce Plio-
Pleistocene contains the oldest palaeoanthropological sites in our continent.
Acknowledgements - Professor B. Chiarelli and Dr. J. Moggi-Cecchi helped us and made constructive
suggestions that have improved this paper. The assistance of Dr. Fontarnau with the SEM and discussions
with Dr. A. Ruiz-Bustos about lhe biostratigraphy and geology are gratefully acknowledged. This work
has been supported by the research project DGICYT PB94-1222-CO2-01 of the Spanish Ministry of
Education and Science.
Fig. 1 Geological situation of the Guadix-Baza basin, SE Spain.

8 J. GIBERT, LL. GIBERT, ALBADALEJO, RIBOT, S,~NCHEZ, P. GIBERT
Fig. 2 General correlation of different stratigraphical sections in the Orce region, including the sections where
palaeomagnetic data exist: the stratigraphical sequence shows unconformity-bounded depositional cycles with high-
resolution time lines. Most of the palaeonWlogical sites in the eastern part of the region lie in a black detrital
member (7), fi~rmed by palustrine sedimentation within cycle 3 of the Orce lacustrine depositional sequence, and
their fauna belongs to a single group, belonging in time to the Normal magnetic episode defined at the Barranco de
Orce and Fuentenueva, interpreted as the Olduvai magneto-subchron by several authors.
Fig. 3 Simplified stratigraphical column showing the succession of sites, stone artifact.g and human remains, as
well as taxa of biostratigraphical significance.
OCCLUSAL VIEW
MESIAL
z: m
DISTAL
LINGUAL VIEW
BUCCAL
LINGUAL
DISTAL VIEW
~ M ESIAL
DISTAL
Fig. 4 Possible orientation of tooth fragment BL5-O.

Fig. 5a, b BL5-O: A= fi'actured face o f enamel and - B= mesiolingual view o f crown and root.
12 J. GIBERT, LL. GIBERT, ALBADALEJO, RIBOT, S.ANCHEZ, P. GIBERT
Fig. 6 BLS-O: polarized-light photograph to show parallel tfunter-Schreger bands.

Fig. 7 BL5-O: polarized-light photograph to show striae of Retzius and Hunter-Schreger bands.
MOLAR TOOTH FRAGMENT BLS-O 13
Fig. 8 Arrangement of last imbricate stria in BL5-O, early human and modern human teeth, according w the
criteria of Beynon & Wood (1986)
14 J. GIBERT, LL. GIBERT, ALBADALEJO, RIBOT, S,~NCHEZ, P. GIBERT
Fig. 9a, b BL5-O: scanning electron micrographs to show l)pe-3b prism enamel pattern.
Fig. lOa, l~ Scamziug electron micrograph of a cast of BLS-O to show (a) perikymata on mesiul sulface, and (b)
perikymata on the mesiolingual s'urface eroded by tran,sport.
FELISLEO URSUS ARCTOS SUS SP.

ENAMEL
HICKNESS
J~ ENAMEL 0,44 ENAMEL
~ TH~g4~Vss THICKNESS
0.54
M3 M1
CANISSP. CV SOERGELIAMINORV.M CERVIDAECV
~, 0.30 ~ S ENAMEL
THICKNESS
0.97
P4 ~ P3 \'Pa
"CERVUS"ELAPHUSV.M MACACASILVANA HOMO SP.
ENAMEL ENAMEL
THICKNESS I ~ ENAMEL
THICKNESS THICKNESS
0.69 MAX:0.66 1.2
M1 f M1 M
Fig. l l Schematic representation of enamel thickness in cross-sections of teeth of different mammals. Maximum
enamel thickness is indicated. CV = Cueva Victoria; VM = Venta Micena.
n
m
~e
Ills Ill
Fig 12 Presence or absence of perikymata

MOLAR TOOTH FRAGMENTBL5-0 17
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MOLAR TOOTH FRAGMENTBLS-0 19
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Journal of Human Evolution 65 (2013) 1e9
Contents lists available at SciVerse ScienceDirect
Journal of Human Evolution

journal homepage: www.elsevier.com/locate/jhevol
The oldest human fossil in Europe, from Orce (Spain)

Isidro Toro-Moyano a, *, Bienvenido Martínez-Navarro b, **, Jordi Agustí b,
Caroline Souday c, d, José María Bermúdez de Castro e, María Martinón-Torres e,
Beatriz Fajardo f, g, Mathieu Duval d, e, Christophe Falguères d, Oriol Oms h,
Josep Maria Parés e, Pere Anadón i, Ramón Julià i, José Manuel García-Aguilar j,
Anne-Marie Moigne d, k, María Patrocinio Espigares l, Sergio Ros-Montoya l,
Paul Palmqvist m
a
Museo Arqueológico de Granada, Carrera del Darro 41-43, 18010 Granada, Spain
b
ICREA, Institut de Paleoecologia Humana i Evolució Social, Àrea de Prehistoria, Universitat Rovira i Virgili, C/Marcel.lí Domingo s/n,
Campus Sescelades, 43007 Tarragona, Spain
c
Center for the Study of Human Origins e CSHO, 25 Waverly Place, New York, NY 10003-6790, USA
d
Museum National d’Histoire Naturelle, Département de Préhistoire, UMR 7194 du CNRS, 1 rue René Panhard, 75013 Paris, France
e
Centro Nacional de Investigación sobre la Evolución Humana (CENIEH), Paseo de Atapuerca, 09002 Burgos, Spain
f
Université Paul Valery, Montpellier III, UMR 5140 «Archéologie des sociétés méditerranéennes» Route de Mende, F-34199 Montpellier, France
g
Stone Age Institute, Indiana University, 1392 w. Ditterrore Rd., Gasport, IN 47433, USA
h
Departament de Geologia, Universitat Autónoma de Barcelona, Bellaterra, Spain
i
Institut de Ciències de la Terra ‘J. Almera’ (CSIC), C/L. Solé Sabarís sn, 08028 Barcelona, Spain
j
Departamento de Estratigrafía y Paleontología, Campus de Fuentenueva, Universidad de Granada, 18003 Granada, Spain
k
Centre européen de recherches préhistoriques, avenue Léon-Grégory, 66720 Tautavel, France
l
Museo de Prehistoria y Paleontología, Orce, Granada 18858, Spain
m
Departamento de Geología y Ecología, Universidad de Málaga, Campus de Teatinos, 29071 Málaga, Spain
a r t i c l e i n f o a b s t r a c t
Article history: The Orce region has one of the best late Pliocene and early Pleistocene continental paleobiological
Received 8 October 2011 records of Europe. It is situated in the northeastern sector of the intramontane Guadix-Baza Basin
Accepted 21 January 2013 (Granada, Andalusia, southern Spain). Here we describe a new fossil hominin tooth from the site of
Available online 05 March 2013
Barranco León, dated between 1.02 and 1.73 Ma (millions of years ago) by Electron Spin Resonance (ESR),
which, in combination with paleomagnetic and biochronologic data, is estimated to be close to 1.4 Ma.
Keywords:
While the range of dates obtained from these various methods overlaps with those published for the
Human tooth
Sima del Elefante hominin locality (1.2 Ma), the overwhelming majority of evidence points to an older
Early Pleistocene
Barranco León
age. Thus, at the moment, the Barranco León hominin is the oldest from Western Europe.
Ó 2013 Elsevier Ltd. All rights reserved.
Introduction evidence from both longitudinal extremes of the continent. The

Georgian site of Dmanisi has provided an important collection of
The date of the earliest human occupation of Europe has been Oldowan lithic artefacts and very significant fossils of early Homo,
recently challenged by new paleontological and archaeological ca. 1.8 Ma (millions of years ago) (Vekua et al., 2002; Lordkipanidze
et al., 2007; Ferring et al., 2011), the Spanish site of Sima del Ele-
* Corresponding author. fante in Atapuerca has yielded some lithic artefacts and a
** Corresponding author. mandibular fragment dated to w1.2 Ma (Carbonell et al., 2008), the
E-mail addresses: [email protected] (I. Toro-Moyano),
Italian site of Pirro Nord (1.3e1.7 Ma) (Arzarello et al., 2009), and
[email protected] (B. Martínez-Navarro), [email protected]
(J. Agustí), [email protected] (C. Souday), [email protected] (J.M. Bermúdez the French sites of Lézignan-la-Cèbe and Pont-de-Lavaud (1.57 and
de Castro), [email protected] (M. Martinón-Torres), [email protected] w1.1 Ma, respectively) (Crochet et al., 2009; Despriée et al., 2010)
(B. Fajardo), [email protected] (M. Duval), [email protected] (C. Falguères), have provided some lithic artefacts, and two early Pleistocene lo-
[email protected] (O. Oms), [email protected] (J.M. Parés), panadon@ calities placed in the northeastern sector of the Guadix-Baza basin
ija.csic.es (P. Anadón), [email protected] (R. Julià), [email protected]
in Spain, Fuente Nueva-3 and Barranco León, have yielded huge
(J.M. García-Aguilar), [email protected] (A.-M. Moigne), mpespigares@
gmail.com (M.P. Espigares), [email protected] (S. Ros-Montoya), [email protected] lithic assemblages and evidence of anthropic use of tools on large
(P. Palmqvist). mammal bones (Turq et al., 1996; Martínez-Navarro et al., 1997;
0047-2484/$ e see front matter Ó 2013 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.jhevol.2013.01.012
2 I. Toro-Moyano et al. / Journal of Human Evolution 65 (2013) 1e9
Gibert et al., 1998b; Toro-Moyano et al., 2003, 2009, 2011; thick enamel layer that matches the anatomy of BL5-0 tooth frag-
Palmqvist et al., 2005; Espigares et al., 2013). ment. Therefore, although the specimen has no clear anatomical
Here we report the finding of a first deciduous molar of early Homo resolution, the more parsimonious explanation (according to
dating back to an age older than 1.2 Ma, probably close to 1.4 Ma, from Occam’s razor) is that BL5-0 is probably part of a hippo tooth.
Orce, which was found in direct association with assemblages of lithic Thus, tooth specimen BL02-J54-100 from the level D of Barranco
artefacts and large mammal bones (Oms et al., 2000b; Toro-Moyano León is the first human remain found at the Orce sites.
et al., 2009; Martínez-Navarro et al., 2010).
The tooth comes from level BL D (previously referred to as BL 5; Paleoanthropology
i.e., Martínez-Navarro et al., 2005), square J54, of the section at Bar-
ranco León, a tributary creek of the Orce river, in the northeast of the The human specimen, BL02-J54-100 (Fig. 2), is a complete crown
Plio-Pleistocene lacustrine deposits of the Guadix-Baza Basin (Vera, of an isolated lower left first deciduous molar (dm1). It is heavily
1970; Sanz de Galdeano and Vera, 1992; Fernández et al., 1996) worn on the occlusal surface, category 5 according to Molnar (1971).
(Fig. 1). The Barranco León section spans the middle terrigenous Roots are missing due to resorption, suggesting that the tooth was
member (lowest part of section in Fig. 1) and the upper ‘Silty shed ante-mortem. A distal wear facet is present at the contact with
Calcareous Member’ of the Baza Formation (Vera et al., 1985; Oms dm2. The occlusal outline is oval and slightly asymmetrical. It pre-
et al., 2000a, 2011), which is dominated mainly by limestones, sents the five main cusps. The protoconid is the largest and it is
sandstones, carbonate silts and dark mudstones, deposited in a lake mesially displaced, followed by the metaconid, the hypoconid and
with an alternation of slightly saline and saline waters (Anadón et al., finally the entoconid. The distal bifurcation of the central groove sets
1994; Anadón and Gabàs, 2009). The excavated layers of the BL sec- the limits of a small hypoconulid. At the dentine and pulp cavity
tion show sediments associated with a swampy environment, except level, the hypoconulid is only hinted as a faint elevation (Fig. 2). A
level D, which shows fluvial features and encloses the archaeological well developed lingual groove and less developed buccal grooves are
level (see further details in Oms et al., 2011). Level BL D contains present. Mesial cusps represent the major part of the crown, and
abundant remains of early Pleistocene large and small mammals they are connected by a mid-trigonid crest. The tooth exhibits a
composed of Ursus sp., Canis mosbachensis, Lycaon lycaonoides, Vulpes strong mesial marginal ridge, prolonged by a vestigial paraconid,
cf. praeglacialis, Pachycrocuta brevirostris, Meles sp., Stephanorhinus separated from the metaconid by a deep V-shaped groove that opens
hundsheimensis, Equus altidens, Equus sussenbornensis, Hippopotamus toward the lingual face. This feature is present in Australopithecus
antiquus, Bison sp., Hemitragus cf. albus, Praemegaceros verticornis, and Homo, such as KNM-ER 820, KNM-ER 1507 and the Zhoukoudian
Metacervocerus rhenanus, Oryctolagus cf. lacosti, Mimomys savini, specimens (Weidenreich, 1937; Wood, 1991), and absent in Para-
Allophaoiomys aff. lavocati, Allophaiomys sp., Apodemus aff. mystaci- nthropus. BL02-J54-100 exhibits a tuberculum molare in the
nus, Histryx sp., Galemys sp., Erinacenae indet., Crocidura sp., Sorex mesiolingual angle, expanding onto the root trunk, but less devel-
minutus, and Sorex sp. (Agustí and Madurell, 2003; Furió-Bruno, oped than that found in the dm1 from Gran Dolina-TD6, Atapuerca
2003; Martínez-Navarro et al., 2010 and references there in), and (Bermúdez de Castro et al., 1999) (Fig. S1). The presence of the
stone artefacts (Oms et al., 2000b) that are broadly similar to other tuberculum molare, the relative expansion of the mesial cusps and
African and European lithic assemblages assigned to the Oldowan mesial marginal ridge, and the enlargement and relative position of
Industry Complex, or Mode I technology, characterized by a low the protoconid are classical diagnostic features in the genus Homo
degree of standardization (i.e., simple core forms and debitage). (Hillson, 1996; Keyser et al., 2000) (Fig. S1).
Although some controversial ‘hypothetical human remains’ Overall, BL02 is a large tooth although there is a great overlap with
from the Orce site of Venta Micena were published during the last the dm1 dimensions of other hominin groups (Table 1 and Fig. S2).
three decades (Gibert et al., 1983, 1998a, 2002; Campillo et al., The mesiodistal (MD) diameter is close to the mean of Paranthropus
2006; among others), including cranial fragment VM-0 and limb robustus and Australopithecus afarensis but it also falls within the
bone diaphyses VM-1960, VM-3691 and VM-12000, none of these range of variation of African, Asian and European early Pleistocene
specimens has enough anatomical resolution to be ascribed to the Homo as well as Homo heidelbergensis and the Middle Paleolithic
genus Homo and they were refuted by Moyà-Solà and Köhler Homo sapiens sample. The buccolingual (BL) diameter is also close to
(1997), Palmqvist (1997), Martínez-Navarro (2002), and Palmqvist the mean of Paranthropus robustus and Australopithecus afarensis and
et al. (2005), among others. Finally, there is also a tooth fragment larger than that of any of the studied groups except for Paranthropus
(BL5-0) found out of stratigraphic context in 1994 during a boisei and Homo antecessor. The crown computed area falls within the
geological survey at the Barranco León ravine by A. Arribas. This range of variation of Paranthropus robustus, Australopithecus afarensis
specimen, which was not unearthed from the excavation quarry of and H. heidelbergensis. The highest values of the Homo sample are
the site, was preliminarily identified as a putative hominin fossil by represented by the European early and middle Pleistocene fossils
Arribas. It is a small tooth fragment, which preserves the mesial such as Arago 66, ATD6-94 and Barranco León together with the
part of the crown and root (4.6 mm crown height, 2.9 mm root Neanderthal specimens from Teshik-Tash and Shanidar VII (Fig. S2).
length). The tooth has an advanced degree of wearing, showing the
exposed dentine and a reduced pulp cavity. Following the Archeology
comparative study of the striae of Retzius and Hunter-Schreger
bands, the shape of the enamel prism patterns and the enamel The lithic assemblage at Barranco León is composed of 1244
thickness (which is clearly thinner than in human teeth), Gibert artefacts (Fajardo, 2008), including 26 cores, 185 whole flakes, 78
et al. (1999) published the tooth fragment as part of an upper hu- flake fragments, 759 waste flakes or débris, 17 retouched pieces, 92
man molar of early Pleistocene age. However, such conclusion angular fragments, 12 modified cobbles (including hammerstones)
should not be accepted without caution, given the reservations on and 75 unmodified materials (cobbles and stones).
the stratigraphic position of the finding (Arribas and Palmqvist, Flint, limestone and quartzite compose the raw material of the
2002) and the possibility that it could belong to a deciduous lithic assemblage. Jurassic formations situated around 3 km south
tooth of H. antiquus, one of the large mammal species that are of the site, contain several primary sources of good raw material,
better represented in the faunal assemblage preserved in the while several flint sources in secondary deposits of alluvial and
swampy-lacustrine facies of Barranco León (Palmqvist et al., 2005). colluvial origin were also found near the site (Toro-Moyano et al.,
More specifically, the bunodont teeth of hippos show a relatively 2009, 2011).
I. Toro-Moyano et al. / Journal of Human Evolution 65 (2013) 1e9 3
Figure 1. Barranco León site. Left: Geological and geographical location. Right: Section with paleomagnetic results and ESR ages in the middle.
Humans knapped at the Barranco Leon D site. Two sets of present. The first stage shows the application of the orthogonal
refitted flint pieces were identified. The first set is composed of four technique with no preparation of the striking platform; cortical
pieces: one core and three flakes (see Fig. 3a). Although it is not surfaces and previous negative scars were used as striking plat-
complete, the first and the last stages of the reduction sequence are forms. The last stage is composed of the exhausted nucleus with
Figure 2. Upper: Tooth specimen BL02-J54-100, left dm1 from Barranco León D. a: Occlusal view; b: Buccal view; c: Distal view; d: Lingual view; and e: Mesial view. Lower:
Computed tomographic reconstruction of the enamel and dentine surfaces (left and center) and the pulp cavity (right) of the dm1. The arrow points to the presence of a small
hypoconulid.
ventral face exploitation of a flake showing the economy of good Chronology

quality raw materials. The second refitting set is formed by pieces
involuntarily detached during the knapping process. Electron spin resonance (ESR) dating was applied to optically
Two main techniques were used for knapping, hard-hammer bleached quartz grains extracted from sediments. This method is
percussion and direct unipolar and bipolar technique (axial and based on the detection of paramagnetic centres, e.g., Aluminum (Al)
non axial using an anvil). The choice of these techniques is related or Titanium (Ti), that are created by the interaction of the natural
to the texture and quality of the raw material. The cobbles and radioactivity with the quartz sample (Ikeya, 1993). Similar to
tabular fragments of flint were destined for the production of flake Optically Stimulated Luminescence (OSL) dating, ESR is an optical
cutting edges, while the less frequent limestone artefacts include dating method that relies on the zeroing of any previously present
cores, battered percutors and debitage. ESR signal by sunlight exposure at the time of deposition (see de-
The primary goal of the Barranco León knappers seems to have tails in Yokoyama et al., 1985). However, while OSL is usually
been obtaining small flakes, perhaps to fulfill immediate needs limited to late middle Pleistocene time range, ESR may potentially
including rapidly cutting meat from large herbivore carcasses. The go further back in time and cover the whole Quaternary time range
proportion of flakes, together with the high frequencies of débris (e.g., Voinchet et al., 2010), given the long term thermal stability of
(61.01%), broken and whole flakes, and angular fragments (28.54%), the ESR signal of the Al center (Toyoda and Ikeya, 1991) and its
which sum up to 89.55% of debitage elements in the whole absence of saturation at high irradiation doses (Lin et al., 2006).
assemblage, and the refitting pieces show that the debitage was in Five sediment samples for ESR dating (ESR BL-1 to ESR BL-5)
situ. Although all stages of core reduction are presented, the most were collected in situ in 2004, 2005 and 2006 from the sedimen-
abundant are from final stages of flaking. The proportion of tary sequence of the excavated area at BL (Fig. 1). This study focused
retouched artefacts (1.37%) is low and does not show stylistic on Al signal because no measurable Ti signal was detected in our
standardization. These are basically light-duty retouched pieces samples. Experimental conditions and age calculations are derived
(see Fig. 3b). from Duval (2008) and detailed in the Supplementary Online
The striation marks and polished areas (Toro-Moyano et al., Information.
2003) on the lithic material (Fig. 3c) are similar to those reported Electron spin resonance age estimates are overall consistent
in well known African and European assemblages such as Olduvai with the stratigraphy, i.e., following a general increase with depth:
Gorge, Koobi Fora or Monte Poggiolo (Keely and Toth, 1981; they range from 1.02 0.09 Ma to 1.88 0.19 Ma and are all in
Sussman, 1987; Peretto et al., 1998). These marks suggest that the agreement for attributing an early Pleistocene age to the deposits
tools were used on a variety of materials. (Table S2). ESR BL-1 was sampled at the base of the sequence, about
The lithic artefacts are found together with large mammal re- 1.5 m under the archaeological level D1, and yields a maximum age
mains showing evidences of anthropic activity, including true spiral of 1.73 0.17 Ma for the deposits. ESR BL-2 comes from D1 layer
or helical fractures, impact points, flake scars and bone flakes and provides an age of 1.46 0.17 Ma. ESR BL-3 and ESR BL-4
(Binford, 1981; Shipman, 1981a, b; Johnson, 1985; Blumenschine samples were collected from level D2, which overlies D1: their
and Selvaggio, 1988) (Fig. 3dei). ESR ages are 1.88 0.19 Ma and 1.23 0.12 Ma, respectively.
Table 1
Metric data of the specimen BL02-J54-100 from Barranco León, compared with other hominin samples.
Specimens n MD BL BL*MD (computed crown area) BL/MD*100 (crown shape index) Data source
X s.d. X s.d. X s.d. X s.d.
Paranthropus robustus
DNH 44, DNH 47, DNH 60, Kromdraai, SK61 (right), 7 10.28 0.66 8.38 0.63 86.55 11.70 81.53 3.15 Robinson, 1956;
SK3978, SK3979 Day, 1986;
Keyser et al., 2000
Paranthropus boisei
KNM-ER 1477, L704-2 2 11.65 e 9.30 e 108.41 e 80.03 e Howell and Coppens, 1973;
I. Toro-Moyano et al. / Journal of Human Evolution 65 (2013) 1e9

Wood, 1991
Australopithecus africanus
Stw104, Stw151, Stw296, Sts24, Taung 5 8.77 0.32 7.50 0.45 65.90 5.82 85.53 3.98 Robinson, 1956;
Moggi-Cecchi et al., 2006
Australopithecus afarensis
AL333-43 (left), AL333-30, LH3, LH2 4 9.77 1.48 8.35 0.77 82.41 20.44 86.030 6.41 White, 1980;
Johanson et al., 1982
Early Homo
DNH 35 1 9.10 e 7.50 e 68.25 e 82.42 e Keyser et al., 2000
Homo erectus
African Homo erectus: KNM-ER 820 (right), KNM-ER 1507 2 8.95 e 7.30 e 65.41 e 81.49 e Leakey and Wood, 1973,1974
Asian Homo erectus: Sinanthropus 123, 125 2 8.75 e 6.80 e 59.71 e 78.57 e Weidenreich, 1937
Homo antecessor
Gran Dolina-TD6 (ATD6-94)a 1 9.00 e 8.80 e 79.20 e 97.78 e
Homo heidelbergensis
Arago 34, 55, 66a 3 9.43 0.66 8.03 0.40 75.96 9.30 85.23 1.71
Homo neanderthalensis
Archi, Gibraltar II, Kulna III, Pech de l’Azé,a Shanidar III, 24 8.83 0.44 7.56 0.47 66.86 6.23 85.68 5.46 Tillier, 1979;
Shanidar VII, Chateauneuf I, Chateauneuf II, Krapina 67, Wolpoff, 1979;
Krapina A, Arcy 18, Arcy 25, Arcy 33, Roc de Marsal, Madre-Dupouy, 1985;
Dederiyeh, Kebara, Teshik-Tash, Combe-Grenal, Taubach, Bailey and Hublin, 2006
La Chaise H13, La Chaise H14, La Chaise Bourgeois
Delauney, Engis, La Ferrassie 8a
Homo sapiens
Middle Paleolithic: Skhül I, Skhül X, Qafzeh 4 3 9.43 0.60 7.50 0.30 70.63 1.76 79.86 8.35 Tillier, 1979
Upper Paleolithic: Miesslingtal, Isturitz 72, Badger’s Hole, 11 8.13 0.75 7.23 0.76 59.14 10.11 89.22 8.35 Frayer, 1978
Bruniquel 537, Bruniquel 540, Isturitz 4, Isturitz 9,
Laugerie Basse 905, Laugerie Basse 909,
La Madeleine 4, Le Figuiera
Contemporary: Musée de l’Homme collections (various 21 8.31 0.36 6.96 0.26 57.99 3.95 83.88 3.79
geographical origins)a
BL02-J54-100 1 10.05 e 8.48 e 85.22 e 84.4 e
a
Measurement taken by authors.
5
Figure 3. Archaeological evidences from level D of Barranco León. a: Refitted pieces; b: Retouched piece; c: Striation and one polished marks; def: Bone cut marks, BL03-L62-2, rib
fragment of megaherbivore (cf. Hippopotamus antiquus), it shows a long and curved cut mark oblique to the major edge of the rib in the central area of the bone, which is related to
the evisceration process; g: Bone impact point; hei: Bone flakes. Note: All lithic and fossil specimens shown in this figure were found during the excavations at Barranco León by the
members of the research team that coauthor this article, with the only exception of specimen BL95 C1 A3 n.15, the central piece of the refitting group of Figure 3A, which was
published in Gibert et al. (1998b).
Sample ESR BL-5, originating from level E1 at the top of the local encloses the archaeological layer at Barranco León. The quite large
sequence, yields a minimum age of 1.02 0.09 Ma for the deposits. final error may be interpreted as the consequence of the age scat-
Electron spin resonance age results obtained for layer D show tering and the limited number of samples. This age is in quite good
some apparent scatter. More specifically, the age of ESR BL-3 looks agreement with the ESR chronologies obtained for the nearby sites
somewhat overestimated in comparison with the other ages. This is of Fuente Nueva-3 and Venta Micena, of 1.19 þ 0.21 Ma and
probably due to the fitting of the dose response curve, which goes w1.4 Ma, respectively (Duval et al., 2011, 2012). However, one
above the natural point and thus might induce an overestimation of should be cautious in their interpretation, since the quite large
the equivalent dose value (Fig. S3). Nevertheless, this age may not error range does not allow any chronological distinction between
be discarded, regarding the small sample size (n ¼ 3). Consequently, the three sites from the Orce area. These results show the potential
based on the three ESR samples BL-2 to BL-4, a weighted mean ESR of ESR dating of quartz grains from early Pleistocene fluvio-
age of 1.43 0.38 Ma may be calculated for the layer D that lacustrine deposits. Future work will definitely need to be focused
on the improvement of the precision of this promising preliminary although it is commonly accepted that due to taphonomic reasons
ESR chronological framework based on quartz grains extracted the lack of record of a taxon in an assemblage is not an evidence of
from sediment. its absence from the original community, the nonappearance of
A new paleomagnetic study (Table S1), combined with previous suids in any of the Late Villafranchian Orce sites is a strong argu-
results from the same stratigraphic section (Oms et al., 2000b) ment that should not be avoided as a reliable biochronological
shows that the entire stratigraphic section of Barranco León has marker. Suids are found in Europe before and during the Olduvai
reverse polarity (Fig. 1). This fully reversed time interval for the subchron, including Fonelas P-1 site in the Guadix-Baza Basin,
whole silty calcareous member has been also confirmed in other dated to 2.0 Ma, which suid remains have been ascribed to Pota-
sections by Scott et al. (2007) [see also Gibert et al. (2006) and mochoerus magnus (Arribas et al., 2009), as well as in many other
comments in Agustí et al. (2007) and Oms et al. (2011)]. Conse- localities with the presence of Sus strozzii (see Rook and Martínez-
quently, the deposits may be correlated to the Matuyama chron Navarro, 2010). However, there are no pigs in the chronological
(0.78e2.58 Ma; Gradstein et al., 2005), and more likely between range comprised between the post Tasso Faunal Unit, in the base of
Olduvai and Jaramillo subchrons, as indicated by the ESR chronol- the Late Villafranchian (w1.8 Ma), and their arrival in Western
ogy and faunal evidence. Europe at the level TE9 from Sima del Elefante (w1.2 Ma), where
Both micro and macrofauna support the ESR ages and the pigs are recorded under the name of Sus sp. (Carbonell et al., 2008),
magnetostratigraphic interpretation. The combination of rodent and at the site of Untermassfeld (Germany), dated 1.0e1.1 Ma and
species at the site, including Mimomys savini, Allophaiomys aff. ascribed to Sus scrofa priscus (Gúerin and Faure, 1997). At sites like
lavocati, Castillomys crusafonti and Apodemus aff. mystacinus, Dmanisi (Georgia), as well as at Pirro Nord (Italy), Apollonia-1
further constrains its age. An age younger than the Olduvai sub- (Greece), Sainzelles (France) and, of course, at the Orce sites, no
chron (1.95e1.77 Ma) is inferred from the more derived record of pigs has never been documented. After 30 years of
morphology of Allophaiomys aff. lavocati compared with A. cf. excavation of 350 m2 at Venta Micena, 140 m2 at Barranco Léon, and
deucalion from the site of Kryzhanovka (Tesakov, 1998), which is 106 m2 at Fuente Nueva-3, and continuous research in the triangle
associated with the Olduvai subchron (Pevner et al., 1998). Allo- formed by the region of Orce-Fuente Nueva-Venta Micena, pig re-
phaiomys aff. lavocati is, in turn, more archaic than the microtine mains have never been found among the more than 25 000 large
species present at Vallonet (France) and Untermassfeld (Germany), mammal fossils unearthed from these sites, and this is probably the
two localities dated to the Jaramillo subchron (0.99e1.07 Ma; best record of the early Pleistocene fauna in Europe. When pigs are
Yokoyama et al., 1988; Wiegank, 1997). Therefore, Barranco León is in the ecosystem, they use to be abundant in the large mammal
placed within the reverse interval between the Jaramillo (0.99e community given their opportunistic feeding behavior and repro-
1.07 Ma) and Olduvai (1.95e1.77 Ma) subchrons. ductive success. For this reason, they are usually well represented
In the Guadix-Baza Basin, a biozonation based on rodents in- in the fossil assemblages. Suids are well recorded in the African and
cludes four biozones within this interval: Mimomys oswaldoreigi Asian archaeological and paleontological sites, including those
Zone, Allophaiomys ruffoi Zone, Allophaiomys aff. lavocati Zone and from the Levantine Corridor as ‘Ubeidiya (Geraads et al., 1986), or
Iberomys huescarensis Zone (Agustí et al., 2010; Oms et al., 2011). Evron Quarry (Tchernov et al., 1994)’, but they are not definitively
The average duration of each biozone is 200 thousand years. The present in Orce or in any other sites of the European continent
Mimomys oswaldoreigi Zone includes the sites of Barranco Conejos, during the period comprised between 1.7 and 1.2 Ma. Later, the
Orce 2 and Orce D. The Allophaiomys ruffoi Zone includes the genus Sus is recorded everywhere in Europe. For this reason, the
famous site of Venta Micena and other levels such as Cañada de absence of suids from level BL D constrains the upper limit of the
Murcia 1, Fuente Nueva 2, Orce 4 and Orce 7 (Agustí et al., 2010). chronological range estimated for this site to >1.2 Ma, in agreement
The Allophaiomys aff. lavocati Zone includes the sites of Barranco with the age estimated from the small mammal assemblage.
León D and Fuente Nueva 3. The Iberomys huescarensis Zone in-
cludes the sites of Huéscar 1, Puerto Lobo and Loma Quemada Conclusions
(Mazo et al., 1985; Agustí et al., 2010). The Last Occurrence (LO) of
Allophaiomys aff. lavocati can be established below the base of the The specimen BL02-J54-100 found at Barranco León site, a hu-
Iberomys huescarensis Zone, dated to the Jaramillo subchron at man tooth with an estimated age by ESR between 1.02 and 1.73 Ma,
1.07 Ma. The First Occurrence (FO) of Allophaiomys aff. lavocati is and by paleomagnetism between 1.07 and 1.77 Ma, but close to
more difficult to establish, since there is no section in the basin 1.4 Ma according to the biochronologic evidence, represents the
where the boundary between the Allophaiomys ruffoi Zone and the oldest anatomical evidence of human presence in Western Europe.
Allophaiomys aff. lavocati Zone has been recognized. However, a This finding, combined with the important lithic tool assemblage
recent dating of the level TE9 from Sima del Elefante, in the Ata- from the level D of Barranco León, confirms that Western Europe
puerca karstic complex (Cuenca-Bescós et al., 2001), contains a was colonized soon after the first expansion out of Africa, currently
more evolved Allophaiomys than A. aff. lavocati from Barranco León documented at the Dmanisi site.
D. The age of this level has been established at 1.22 0.16 Ma, based
on cosmogenic nuclids (Carbonell et al., 2008). Therefore, the age of Acknowledgments
the level BL D can be constrained between 1.77 Ma (top of Olduvai
subchron) and w1.2 Ma (age of Sima del Elefante). We thank all the members of the Orce research project involved
However, interpolation of metric parameters measured on the in the excavations and the study of the Barranco León paleoan-
lower first molar of diverse well dated early-middle Pleistocene thropological site. We thank Professor P. Rivas for his support and
microtine species provides a way to further constrain the age of the encouragement. Also we thank Professor H. de Lumley and his team
findings. The relative length of the anteroconid complex in micro- for their help. M.T. Alberdi, M.R. Palombo, L. Abbazzi, F. Lacombat,
tines (the A/L parameter) (Meulen, 1973) has proven to be a useful S. Bailón, H.A. Blain, J. Madurell, and M. Furió helped in the deter-
tool for dating Plio-Pleistocene sites (Maul et al., 1998). The A/L mination of the vertebrate fauna. C.F. and M.D. thank P. Voinchet
values of Allophaiomys aff. lavocati from level BL D suggest an age and J. Despriée for their help during the ESR sampling. R. Grün and
older than Sima del Elefante and close to 1.4 Ma (Table S3). J.-J. Bahain provided helpful comments on the ESR dating. We thank
The large mammals assemblage also points to an older chro- Ph. Mennecier, A. Fort and V. Laborde from the MNHN (Musée
nology for BL-D than level TE9 from Sima del Elefante. In addition, de l’Homme) for the access to fossil and extant collections.
Photographs and microscan of the hominin molar were carried at faune de vertébrés continentaux, associée à des artefacts dans le Pléistocène
inférieur del’Hérault (Sud de la France), vers 1,57 Ma. C.R. Palevol. 8, 725e736.
CENIEH of Burgos, Spain. We thank L. Martín-Francés and
Cuenca-Bescós, G., Canudo, J.I., Laplana, C., 2001. La séquence des rongeurs
M. Martínez de Pinillos for their help with the micro-CT analysis. (Mammalia) des sites du Pléistocène inférieur et moyen d’Atapuerca (Burgos,
ESR data were obtained at the Prehistory Department of MNHN Espagne). L’Anthropologie 105, 115e130.
(Paris, France). Paleomagnetic measurements were carried out at Day, M.H., 1986. Guide to Fossil Man, fourth ed. The University of Chicago Press,
Chicago.
the SCT of the Barcelona University. Scanning photographs of the Despriée, J., Voinchet, P., Tissoux, H., Moncel, M.-H., Arzarello, M., Robin, S.,
lithic artefacts were carried out at the CERP of Tautavel (France) Bahain, J.-J., Falguères, C., Courcimault, G., Dépont, J., Gageonnet, R., Marquer, L.,
by Mme. B. Deniaux. We thank Ms. Sheila Wglie for reviewing Messager, E., Abdessadok, S., Puaud, S., 2010. Lower and middle Pleistocene
human settlements in the middle Loire river basin, centre region, France.
the English version of this paper. D. Begun, E. Delson, E. Carbonell, Quatern. Int. 223e224, 345e359.
T. D. White, D. Lordkipanidze, S.C. Antón, M.M. Lahr, C. Stringer, Duval, M., 2008. Evaluation du potentiel de la méthode de datation par Résonance
M. Tappen, L. Rook, R. Ferring, the Associate Editor and three de Spin Electronique (ESR) appliquée aux gisements du Pléistocène inférieur:
étude des gisements d’Orce (bassin de Guadix-Baza, Espagne) et contribution à
anonymous reviewers revised a first draft of the manuscript. This la connaissance des premiers peuplements de l’Europe. Ph.D. Dissertation.
study has been sponsored by the Dirección General de Bienes Muséum National d’Histoire Naturelle, Paris.
Culturales (Consejería de Cultura, Junta de Andalucía, Spain), and it Duval, M., Falguères, C., Bahain, J.-J., Grün, R., Shao, Q., Aubert, M., Hellstrom, J.,
Dolo, J.-M., Agustí, J., Martinez-Navarro, B., Palmqvist, P., Toro-Moyano, I., 2011.
has been supported by projects CGL2010-15326, CGL2010-16821, The challenge of dating early Pleistocene fossil teeth by the combined uranium
CGL2012-38358, and CGL2008-04896 of the Spanish Ministry of serieseelectron spin resonance method: the Venta Micena palaeontological
Economy and Competitiveness. site. J. Quatern. Sci. 26, 603e615.
Duval, M., Falguères, C., Bahain, J.-J., Grün, R., Shao, Q., Aubert, M., Dolo, J.-M.,
Agusti, J., Martínez-Navarro, B., Palmqvist, P., Toro-Moyano, I., 2012. On the
limits of using combined U-series/ESR method to date fossil teeth from two
Appendix A. Supplementary data Early Pleistocene archaeological sites of the Orce area (Guadix-Baza basin,
Spain). Quatern. Res. 77, 481e482.
Supplementary data related to this article can be found online at Espigares, M.P., Martínez-Navarro, B., Palmqvist, P., Ros-Montoya, S., Toro, I.,
Agustí, J., Sala, R., 2013. Homo vs. Pachycrocuta: earliest evidence of competition
http://dx.doi.org/10.1016/j.jhevol.2013.01.012.
for an elephant carcass between scavengers at Fuente Nueva-3 (Orce, Spain).
Quatern. Int. 295, 113e125.
Fajardo, B., 2008. Les industries lithiques anciennes d’Orce: les sites de Barranco
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Two Deciduous Human Molars from the Early Pleistocene Deposits of Barranco León (Orce,
Spain)
Author(s): Francesc Ribot, Luis Gibert, Carles Ferràndez-Cañadell, Enrique García Olivares,
Florentina Sánchez, and María Lería
Source: Current Anthropology, Vol. 56, No. 1 (February 2015), pp. 134-142
Published by: The University of Chicago Press on behalf of Wenner-Gren Foundation for
Anthropological Research
Stable URL: http://www.jstor.org/stable/10.1086/679615 .
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134 Current Anthropology Volume 56, Number 1, February 2015
Two Deciduous Human Molars from the tools at the sites of Barranco León and Fuentenueva 3 (Gibert
et al. 1998b), and fragmentary human remains at Barranco
Early Pleistocene Deposits of Barranco León and at the stratigraphically lower site of Venta Micena—
León (Orce, Spain) the latter have been a matter of dispute, being supported by
Francesc Ribot, Luis Gibert, Carles Ferràndez-Cañadell, some authors (e.g., Aguirre 2008; Borja et al. 1997; Campillo
Enrique Garcı́a Olivares, Florentina Sánchez, and et al. 2003, 2006; Coppens 1992; Gibert and Palmqvist 1995;
Marı́a Lerı́a Gibert et al. 1989a, 1989b, 1998a, 2006a, 2006b; Lowenstein,
Borja, and Garcı́a-Olivares 1999; Martı́nez-Navarro 1996;
Museo de Prehistoria y Paleontologı́a José Gibert, Calle las Sánchez et al. 1999; Tobias 1998; Torres, Borja, and Garcı́a-
Tiendas, s/n.18858-Orce, Spain/Departament de Geoquı́m- Olivares 2002) and rejected by others (e.g., Martı́nez-Navarro
ica, Petrologia i Prospecció Geològica, Facultat de Geologia, 2002; Moyà and Agustı́ 1989; Moyà and Köhler 1997; Palmqv-
Universitat de Barcelona, Martı́ Franquès s/n, 08028 Barce- ist et al. 2005).
lona, Spain ([email protected])/Departament d’Estratigrafia, Recently Toro-Moyano et al. (2013) reported a second de-
Paleontologia i Geociències Marines, Facultat de Geologia, ciduous tooth from Barranco León and claimed it to be “the
Universitat de Barcelona, Martı́ Franquès s/n, 08028 Barce- oldest human fossil in Europe” (1, title), stating that a pre-
lona, Spain/Departamento de Bioquı́mica y Biologı́a Molec- vious human molar from the same site (BL5-0) had “no clear
ular e Inmunologı́a, Facultad de Medicina, Universidad de anatomical resolution” (2) and more likely belonged to a
Granada, 18012-Granada, Spain/Museo de Prehistoria y Pa- Hippopotamus antiquus deciduous tooth, based solely on the
leontologı́a José Gibert, Calle las Tiendas, s/n.18858-Orce, observation that H. antiquus is an abundant species in this
Spain/Departament de Dibuix, Facultat de Belles Arts, layer. In addition, they suggest that the fossil was found out
Universitat de Barcelona, Pau Gargallo, 4, 08028 Barcelona, of stratigraphic context. In this contribution we propose a
Spain. This paper was submitted 12 X 13, accepted 17 VII new interpretation of both finds after supplying information
14, and electronically published 27 I 15. on their stratigraphic location and anatomical data from de-
ciduous hippo teeth.
Recently Toro-Moyano et al. (2013) reported a deciduous
tooth from Barranco León (Spain; BL02-J54-100) and claimed Stratigraphic Context
it to be the oldest human fossil in Europe. In that paper, the
BL5-0 comes from a fossiliferous bed named BL5 (Arribas
authors suggest that a previously reported human molar frag-
and Palmqvist 2002). This bed is 15–30 cm thick and com-
ment from the same site (BL5-0) was not human but a de-
posed mainly of fine to medium sandstone, including Jurassic
ciduous molar of Hippopotamus found out of stratigraphic
marine and Pleistocene lacustrine carbonate pebbles, mammal
context. Here, we show the stratigraphic and spatial position
remains, and a large assemblage of lithic tools (Gibert et al.
of BL5-0, and we separate it from deciduous teeth of Hip-
1998b). This bed occurs within a lacustrine sequence and can
popotamus. We conclude that two human deciduous molars
be followed for more than 300 m on both sides of Barranco
have been discovered at the Barranco León site. Both teeth
León. It is the product of a lake-level fall that allowed erosion
were found 9 meters apart, have a similar size, are heavily
and resedimentation from the marginal area of a shallow lake
worn on the occlusal surface, have a nearly identical interstitial
(fig. 1A). A minimum age of 11.25 Ma was calculated for
contact facet, and in both cases the roots are practically miss-
this layer considering the stratigraphic distance to a paleo-
ing due to resorption. These similarities and the proximity
magnetic reversal interpreted as the top of the Olduvai sub-
of the finds suggest that both molars probably belonged to
chron (1.78 Ma; Scott, Gibert, and Gibert 2007). In 1994, A.
the same individual.
Arribas discovered a molar fragment after sieving a sediment
sample from the BL5 bed (Arribas and Palmqvist 2002). Later,
in 1995, J. Gibert opened a quarry on the area of the discovery
Introduction and initiated the excavation of this fossiliferous layer. New
paleontological excavations at BL5 were not permitted by the
With a continuous sedimentary record and archaeological
sites at different stratigraphic heights, the Orce region of administration until 1998 when a new team took the control
southern Spain is one of the candidate localities to uncover of the site and renamed the fossiliferous bed. As stated by
early humans in the European Pleistocene (Scott and Gibert Toro-Moyano et al. (2010, 2013), the level BL D, where they
2009). The Orce sites have yielded Early Pleistocene Oldowan found the tooth BL02-J54-100 in 2002, is also referred to as
BL5, so that the two teeth come from the same bed, which
䉷 2015 by The Wenner-Gren Foundation for Anthropological Research.
is the only one bearing tools and vertebrate fossils at Barranco
All rights reserved. 0011-3204/2015/5601-0007$10.00. DOI: 10.1086/ León.
679615 The spatial and stratigraphic location of BL5-0 was pub-

Ribot et al. Pleistocene Human Molars from Barranco León 135
Figure 1. Stratigraphic and archaeological location of BL5-0. (A) Plan of the excavation in the BL-5 bed made in 1995 indicating
the distribution of lithic artifacts, adult Hippopotamus remains, and the location of the tooth fragment BL5-0, recovered in 1994
at the edge of square A4. The curved line corresponds to the slope of the ravine. (B) Detail of the Black Detrital Unit (Gibert et
al. 1998b) in the Barranco León sedimentary sequence showing the stratigraphic position of BL5 bed where BL5-0 and other finds
were collected. In this stratigraphic series, large mammal fossils and lithic tools occur only at bed BL5. (C) Superposition of the
excavation plans from 2002 and 1995 (modified from Gibert et al. 1998b and Toro et al. 2010). BL5-0 was found at square A4 of
the 1995 plan and falls within the square P60 of later excavations by Toro-Moyano et al. (2010). The lateral distance between BL5-
0 and BL02-J54-100 was less than 9 m. Squares: 1 m.
lished in 1998 (figs. 3b and 4 in Gibert et al. 1998b), and its Archaeological Context
discovery and initial study were described in detail in Arribas
and Palmqvist (2002). A superposition of the excavation plans The site of Barranco León has archaeological relevance; five
from 1995 (Gibert et al. 1998b) and 2002 (Toro-Moyano et lithic artifacts were initially reported from the outcropping
al. 2010) shows that the two teeth were found less than 9 m bed BL-5 (Gibert et al. 1992). The first systematic excavation
apart from each other (fig. 1A, C). in 1995 revealed a mandible of Hippopotamus surrounded by

Figure 2. Examples of Oldowan technology from the BL5 site. These flint flakes were unburied during 1995 field season a short
distance from the BL5-0 tooth fragment location. The figure shows views of opposite faces of three flakes made with grey Jurassic
flint. Scale: centimeters.
more than a hundred lithic artifacts: 114 of flint, 1 of quartz- placed by chert. Blocks of quality flint (up to 10 cm) are
ite, and 1 of Jurassic limestone. The tools were associated with found in alluvial fan deposits down in the valley, only 2 km
early Pleistocene fauna: Castillomys cf. crusafonti, Mimomys southeast of the site. The origin of the quartzite should be
sp., Allophaiomys pliocaenicus, Equus granatensis, and Hip- found in Miocene fluvial deposits outcropping 8 km from the
popotamus antiquus. The flint cores and flakes are small site, where quartzite pebbles occur. The tools represent a lithic
(range: 20–61 mm; mean: 40.7 mm); the butts have a very technology with a very simple chaı̂ne opératoire, lacking any
variable morphology, and the flakes are not usually cortical. sign of bifacial flaking technique; accordingly it can be com-
Chopper-cores of flint and limestone are also present in BL- pared to the Oldowan (Gibert et al. 1998b; see fig. 2). The
5 (Gibert et al. 1998b). Subsequent excavations increased the presence in the same bed of cores and flakes associated with
number of artifacts to 11,200 (Toro-Moyano et al. 2013). The remains of large fauna suggests that the tools were occasionally
source of the flint and limestone was the Jurassic from Umbrı́a produced on the site and used to recover the available re-
and Periate ranges, where marine limestones are locally re- sources.

Figure 3. Morphological and enamel features of BL5-0. (A) Fractured face of enamel; note the reduced pulpar cavity. (B) Mesiolingual
view of the crown and root in polarized light to show striae of Retzius and Hunter-Schreger bands. (C) Polarized-light photograph
showing the parallel Hunter-Schreger bands. (D–F) SEM photographs showing the enamel prisms pattern of type 3b of Boyde
(1964). (G–H) Optical and SEM photographs of the insterstitial contact facet. (I–L) Comparison of BL02-J54-100 (I, J) and BL5-
0 (K, L), at the same scale, showing the similar interstitial contact facets (arrows). (I, J) Occlusal and mesial view of BL02-J54-100;
(K, L) Mesiolingual and nearly occlusal views of BL5-0. Images from BL02-J54-100 reproduced from Toro-Moyano et al. (2013)
with permission from Elsevier.
Figure 4. Comparison of BL5-0 enamel with human and hippo enamel. (A) Comparison of enamel thickness of BL5-0 (left) with
a modern human molar (right). Note the similar increase in thickness from the cervix to the crown. (B–D) Enamel thickness in a
deciduous mm2 of recent Hippopotamus amphibius (MZB-91-0214, 4–5 months old, Age Group I of Laws 1968). (B) Lingual view
with BL5-0 at the same scale. (C–D) Mesial view and detail showing the constant thickness of enamel from the cervix to the crown
and the reduced thickness at the cusp. (E) BL5-0 compared at the same scale with a broken adult tooth of H. antiquus from Venta
Micena (MNCN19273). Note the large and constant thickness of the enamel in H. antiquus.

Figure 5. Differences in enamel distribution and maximum thickness for different medium-sized mammals, Homo sp., and Hip-
popotamus antiquus from Venta Micena (modified from Gibert et al. 1999, not to scale). Both the distribution of the enamel and
the enamel thickness differentiate BL5-0 from hippos and other studied mammals. CV: Cueva Victoria; VM: Venta Micena.
The Molar Fragment BL5-0 Venta Micena, which shows an opposite pattern in the enamel
distribution (decreasing thickness toward the crown) than in
BL5-0 is a deciduous human molar fragment found in the
humans (increasing thickness toward the crown; fig. 5).
BL-5 bed. Only mesial parts of the crown and root remained
after an ancient buccolingual fracture. It is heavily worn, ex- Previous Work
posing the dentine on the occlusal surface, and has an inter-
stitial contact facet. The crown height on the mesial face is After the discovery, Arribas performed a comparative study
4.6 mm, the length of the broken root is 2.9 mm, and the and rejected the possibility that the tooth belonged to a non-
maximum enamel thickness is 1.2 mm (figs. 3A–C, G, H–L). human herbivorous or carnivorous mammal. He also un-
It was assigned to an early Homo based on common anatom- dertook a comparative study of the tooth fragment with ho-
ical features with Homo, including patterns on the micros- mologous sections of the upper and lower molariform teeth
tructure of the enamel (Gibert et al. 1999; see figs. 3C, E, F of omnivorous species (suids, ursids, and hominids), and ob-
and 4). Additionally, an anatomical study was performed, served that BL5-0 was analogous to the lower molars of Homo
based on enamel microstructure, distribution, and thickness, sapiens (Arribas and Palmqvist 2002:68). In consultation, Ber-
that differentiates BL5-0 from medium-size large mammals múdez de Castro agreed with a tentative human deciduous
present at the Early Pleistocene sites of Orce (Gibert et al. molar assignment (Arribas and Palmqvist 2002:68).
1999). The study analyzed enamel from Homo sp., Canis sp., The final assignment of BL5-0 to an early Homo was based
Ursus sp., Sus sp., Macaca sylvanus, Cervus elaphus, Felis leo, on the study of the microstructure of the enamel along the
and Soergelia minor, but did not consider hippos because of fracture surface, a character that plays a central role in in-
the large difference in size. Here we incorporate into this terpreting fossil hominin taxonomy (e.g., Lacruz et al. 2008).
previous study the enamel of Hippopotamus antiquus from This study shows the following human characteristics in

Figure 6. Morphology of deciduous teeth of recent juvenile Hippopotamus amphibius from the Museu de Zoologia of Barcelona
(MZB). (A–D) Individual MZB-91-0214, 4–5 months old. (A) Mandible with deciduous premolars dp2 and dp3, dp4 protruding
above bone level, and alveolus of dml open (Age Group I of Laws 1968). (B) Detail of right dp4 showing the well-developed cingulum
and the rugose enamel surface. (C–D) Details of the enamel in the cusp of the right dp3 (C) and left dp3 (D), showing the rugose
surface of the enamel. (E–H) Individual MZB-82-7007, about 1 year old. (E) Left hemimandible with wear evident on all three
cusps of dp4, Ml exposed above bone and alveolus of dm2 open (Age Group III of Laws 1968). (F–H) Worn left dp4 showing its
large size, and the rugose enamel surface.
BL5-0 (figs. 3A–L and 4A): the angle of the stria of Retzius of adult early Homo varies from 1.2 to 2.1 mm (Beynon and
(30º) and the Hunter-Schreger bands (80º), the lateral enamel Wood 1986), the lateral enamel thickness in the lower M1 of
thickness (1.2 mm), the position of the last imbricate stria, extant Homo from 0.96 to 2.19 (Mahoney 2010), and lateral
the enamel prism pattern of type 3b or “keyhole” defined by thickness of dm1 and dm2 in extant Homo from 0.32 to 1.27
Boyde (1964), the increased enamel thickness from the cervix mm (Mahoney 2010). Therefore, the thickness of the enamel
to the crown, and the presence of perikymata (figs. 3D–F and of BL5-0, 1.2 mm, falls within human variability (figs. 3 and
4A). 4).
Toro-Moyano et al. (2013:2) then assert: “More specifically,
Comparison with Hippo Deciduous Teeth and with Human
the bunodont teeth of hippos show a relatively thick enamel
Tooth BL02-J54-100
layer that matches the anatomy of BL5-0 tooth fragment”
Ignoring the enamel characters of BL5-0 provided by Gibert without showing any picture or measurements. To clarify this
et al. (1999), Toro-Moyano et al. (2013:2) claimed that BL5- point, we compared BL5-0 with deciduous molars of the re-
0 “has no clear anatomical resolution” and it is more likely cent H. amphibius, considering that the molar teeth in hip-
to belong to a H. antiquus deciduous tooth, based solely on popotamids are very conservative (Coryndon 1977).
the observation that H. antiquus is an abundant species in
this site. Toro-Moyano et al. (2013:2) claimed erroneously Differences between BL5-0 and Hippopotamus teeth. The dif-
that the enamel thickness of BL5-0 “is clearly thinner than ferences between BL5-0 and Hippopotamus teeth can be sum-
in human teeth.” The lateral enamel thickness in the molars marized as follows:

Comparison between BL5-0 and human specimen BL02-J54-

100. BL5-0 was described as a possible upper left adult molar
based on the heavily worn occlusal surface and the presence
of part of the root (Gibert et al. 1999). New observations on
BL5-0 and the comparison with specimen BL02-J54-100 re-
vealed that it may be also a deciduous tooth with a pulp cavity
similar to that in BL02-J54-100, as was previously suggested
by Bermúdez de Castro in 1994 (Arribas and Palmqvist 2002:
68).
In addition, BL5-0 has an interstitial contact facet strikingly
similar to the one in BL02-J54-100 (figs. 3G–L and 7).
Common characteristics in BL02-J54-100 and BL5-0 in-
clude:
1. Both are heavily worn on the occlusal surface, category
5 of Molnar (1971).
2. In both cases, the roots are practically missing due to
resorption. The root length preserved in BL02-J54-100 is 3̃.1
mm (estimated from the photos in Toro-Moyano et al. 2013),
and 2.9 mm in BL5-0.
3. Both fossils have a nearly identical interstitial contact
facet (figs. 3G–L and 7). In both teeth, these facets are very
marked and produce an almost flat interstitial teeth wall with
a U-shape.
4. Their size is very similar. The height of the crown in
BL02-J54-100 is about 4.4 mm (estimated from the photos
in Toro-Moyano et al. 2013) and 4.6 mm in BL5-0.
The comparison between BL5-0 and the newly discovered
tooth BL02-J54-100 indicates that BL5-0 has a thicker enamel
Figure 7. Teeth from Barranco León site compared with Homo (1.2 mm). Considering that dm2 has a thicker enamel than
sapiens lower left deciduous teeth. BL02-J54-100 would corre- dm1 (0.62–1.27 and 0.32–0.88 mm, respectively; Mahoney
spond to a left dm1 and BL5-0 to a proximal fragment of a left 2010), BL5-0 (1.2 mm) would be better classified as a dm2.
dm2, possibly from the same individual. Image from BL02-J54- Because of its fragmentary character, it is difficult to precisely
100 reproduced from Toro-Moyano et al. (2013) with permission locate its position, but possibly BL5-0 corresponds to the
from Elsevier. mesial part of a left dm2 (because dm2 are the last tooth, the
distal part lacks interstitial contact facets).
Therefore, it is possible that the two teeth, BL5-0 and BL02-
1. In Hippopotamus the enamel is thick and very rugose J54-100, belonged to the same individual, being contiguous
throughout the tooth surface (e.g., Boisserie 2005; Pavlakis teeth sharing an interstitial contact facet (fig. 7).
1990; see figs. 4C–E and 6B–H), whereas in BL5-0 and humans
it is completely smooth (fig. 3).
2. The thickness of the Hippopotamus enamel is uniform
Conclusion
from the cervix to the crown, whereas in BL5-0 the thickness BL02-J54-100 and BL5-0 are both deciduous human teeth
increases in this direction. Furthermore, in Hippopotamus the from the same site; they were located in the same layer at a
enamel is thinner over the cusps than elsewhere (Lucas et al. maximum distance of 9 m from each other. They have the
2008), whereas in humans the enamel is much thicker over same degree of wear and similar morphology and size, with
the cusps as in BL5-0 (figs. 3A–C, 4C–E, and 5). virtually the same crown height and root length, and with
3. Deciduous teeth of Hippopotamus do not have periky- the same degree of root resorption. Both teeth have a very
mata. similar interstitial contact facet affecting the enamel. The
4. Deciduous teeth of Hippopotamus have a cingulum at thickness of the enamel of BL5-0 fits better with a dm2 than
the base of the crown (fig. 6B) that is absent in BL5-0. with a dm1 like BL02-J54-100. The interstitial wear facet
5. The large dimensions of deciduous teeth of Hippopot- would indicate that BL5-0 corresponds to the mesial part of
amus do not permit an anatomical comparison with BL5-0. a left dm2.
Any of these features alone is enough to invalidate the Taken together, these observations lead to the conclusion
comparison of BL5-0 with Hippopotamus, and all of them that both teeth might possibly have belonged to the same
together clearly discard the assignment of BL5-0 to this genus. individual; they could possibly be contiguous teeth, in contact

through their wear facet. If so, they probably would corre- ueva-3: Three Archaeological Sites in the early Pleistocene Deposits of
Orce, south-east Spain. In The human evolution source book. Russell L.
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shed ante-mortem as interpreted for BL02-J54-100 by Toro- NJ: Prentice Hall.
Moyano et al. (2013). Given this new interpretation, the pos- Gibert, Josep, Lluı́s Gibert, Alfredo Iglesias, and Eudald Maestro. 1998. Two
‘Oldowan’ assemblages in the Plio-Pleistocene deposits of the Orce region,
sibility of finding more human remains at the site increases. Southeast Spain. Antiquity 72:17–25.
Gibert, Josep, Alfredo Iglesias, Alfons Maillo, and Luis Gibert. 1992. Industrias
lı́ticas en el Pleistoceno Inferior de la región de Orce. In Presencia humana
en el Pleistoceno Inferior de Granada y Murcia. Pp. 219–240. Orce, Spain:
Acknowledgments Museo de Prehistoria.
Gibert, Josep, and Paul Palmqvist. 1995. Fractal analysis of the Orce skull
This paper is a contribution to the Grup de Investigació Con- sutures. Journal of Human Evolution 28:561–575.
Gibert, Josep, Francesc Ribot, Carles Ferrández, Bienvenido Martı́nez, Roxana
solidat Geologia Sedimentària (2014 SGR 251 and the Ramón Caporicci, and Domènech Campillo. 1989a. Anatomical study: comparison
y Cajal Program of the Spanish government). The authors of the cranial fragment from Venta Micena (Orce, Spain) with fossil and
thank the staff of the Museu de Zoologia de Barcelona and extant mammals. Human Evolution 4:283–305.
———. 1989b. Caracterı́sticas diferenciales entre el fragmento de cráneo de
the Museo Nacional de Ciencias Naturales for their assistance Homo sp. de Venta Micena (Orce, Granada) y los équidos. Estudios Geo-
and to Robert A. Martin for helpful comments on the man- lógicos 45:121–138.
uscript. We dedicate this paper to the memory of Dr. Josep Gibert, Josep, Francesc Ribot, Patxu Gibert, and Lluı́s Gibert. 2006. Oblit-
eration study of lambdatic and obelionic region sutures in ruminant, car-
Gibert; thanks to his pioneer work, Orce appears in the sci- nivores and hominids. Estudios Geológicos 62:123–134.
entific records. Lacruz, Rodrigo S., M. Christopher Dean, Fernando Ramirez-Rozzi, and Tim-
othy G. Bromage. 2008. Megadontia, striae periodicity and patterns of
enamel secretion in Plio-Pleistocene fossil hominins. Journal of Anatomy
213:148–158.
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Gibert Etal-1999-Human Molar Barranco Leon

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HUMAN EVOLUTION Vol. 14 - N.

1-2 (3-19)- 1999

J. Gibert Molar tooth fragment BL5-0: the oldest

Keywords: tooth human remain,

Geological and geochronological background

The NE sector of the Guadix-Baza basin in SE Spain (Fig. 1) displays a continental

Palaeomagnetic determinations demonstrate presence of five normal-polarity episodes in a

Hominid molar fragment

Material and research methods

Advances in technology and methodology enable unequivocal identification of BLS-0 as

Fig. 1 Geological situation of the Guadix-Baza basin, SE Spain.

Fig. 4 Possible orientation of tooth fragment BL5-O.

Fig. 6 BLS-O: polarized-light photograph to show parallel tfunter-Schreger bands.

FELISLEO URSUS ARCTOS SUS SP.

Fig 12 Presence or absence of perikymata

du PlOistoc~ne infdrieur de la "rEgion d'Orce (Granada, Espagne). Comptes Rendues de l'Acad6mie

Contents lists available at SciVerse ScienceDirect

Journal of Human Evolution

The oldest human fossil in Europe, from Orce (Spain)

Introduction evidence from both longitudinal extremes of the continent. The

ventral face exploitation of a ﬂake showing the economy of good Chronology

X s.d. X s.d. X s.d. X s.d.

I. Toro-Moyano et al. / Journal of Human Evolution 65 (2013) 1e9

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Comparison between BL5-0 and human specimen BL02-J54-

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