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Quaternary Science Reviews 30 (2011) 1396e1412

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Quaternary Science Reviews


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One million years of cultural evolution in a stable environment at Atapuerca


(Burgos, Spain)
J. Rodríguez a, *, F. Burjachs b, G. Cuenca-Bescós c, N. García d, e, J. Van der Made f, A. Pérez González a,
H.-A. Blain g, I. Expósito g, J.M. López-García g, M. García Antón h, E. Allué g, I. Cáceres g, R. Huguet g,
M. Mosquera g, A. Ollé g, J. Rosell g, J.M. Parés a, X.P. Rodríguez g, C. Díez i, J. Rofes d, R. Sala g, P. Saladié g,
J. Vallverdú g, M.L. Bennasar g, R. Blasco g, J.M. Bermúdez de Castro a, E. Carbonell g, j,1
a
Centro Nacional de Investigación sobre la Evolución Humana, Avenida de la Paz 28, 09004 Burgos, Spain
b
ICREA Research Professor at Institut Català de Paleoecologia Humana i Evolució Social, Plaça Imperial Tarraco 1, 43005 Tarragona, Spain
c
Area de Paleontología, Facultad de Ciencias, Universidad de Zaragoza, c/Pedro Cerbuna, 12, 50009 Zaragoza, Spain
d
Departamento de Paleontología, Facultad de Ciencias Geológicas, Universidad Complutense de Madrid, 28040 Madrid, Spain
e
Centro de Investigación (UCM-ISCIII) de Evolución y Comportamiento Humanos, c/Sinesio Delgado, 4 (Pabellón 14), 28029 Madrid, Spain
f
Departamento de Paleobiología, Museo Nacional de Ciencias Naturales, C.S.I.C., José G. Abascal 2, 28006 Madrid, Spain
g
IPHES (Institut Català de Paleoecologia Humana i Evolució Social). Área de Prehistòria, Universitat Rovira i Virgili, Plaça Imperial Tarraco 1, 43005 Tarragona, Spain
h
Departamento de Biología (Botánica), Facultad de Ciencias, Universidad Autónoma de Madrid, 28049 Madrid, Spain
i
Dpto. CC. Históricas. Laboratorio de Prehistoria. IþDþI, Plaza Misael Bañuelos, Universidad de Burgos 09001 Burgos, Spain
j
IPHES. Àrea de Prehistòria, Universitat Rovira i Virgili, Imperial Tarraco 1, 43005 Tarragona, Spain

a r t i c l e i n f o a b s t r a c t

Article history:
The present paper analyses the evidence provided by three sites (Sima del Elefante, Gran Dolina, and
Received 16 June 2009
Galería) located in the Trinchera del Ferrocarril of the Sierra de Atapuerca. These three sites are cave
Received in revised form
16 February 2010 infillings that contain sediments deposited from approximately 1.2 Ma to 200 kyr. Pollen, herpeto-
Accepted 16 February 2010 fauna, and small and large mammal remains are used as proxies to obtain a general picture of the
Available online 23 March 2010 environmental changes that occurred at the Sierra de Atapuerca throughout the one million-year
period represented at these sites. Similarly, cultural changes are tracked analyzing the evidence of
human behavior obtained from the study of several bone and lithic assemblages from these three sites.
At least three periods with different cultural features, involving technology, subsistence and behavior,
are determined from the available evidence. The first two periods correspond to the Mode 1 tech-
nology and Homo antecessor: the first is dated around 1.2 to 1.0 Ma and reflects opportunistic behavior
both in technology and subsistence. The second period is around 800 kyr BP. Mode 1 technology is still
maintained, but subsistence strategies include systematic hunting and the use of base camps. The third
period is dated between 500 ka and 200 ka and corresponds to the Mode 2 technology and the
acquisition of directional hunting and other organizational strategies by Homo heidelbergensis.
A transition from Mode 2 to Mode 3 seems to appear at the end of this time-range, and may reflect the
early phases of a fourth cultural change. With regard to the environment, our main conclusion is that
there was an absence of extremely harsh conditions at Atapuerca throughout this time period. The
presence of Mediterranean taxa was constant and the dominant landscape was a savannah-like open
environment, probably with small forest patches. An alternation of Mediterranean and mesic species

* Corresponding author. Tel.: þ34 947 255006.


E-mail addresses: jesus.rodriguez@cenieh.es (J. Rodríguez), francesc.burjachs@urv.cat (F. Burjachs), cuenca@posta.unizar.es (G. Cuenca-Bescós), ngarcia@isciii.es (N.
García), mcnjv538@mncn.csic.es (J. Van der Made), alfredo.perez@cenieh.es (A. Pérez González), hablain@prehistoria.urv.cat (I. Blain), isaeba@prehistoria.urv.cat (I.
Expósito), jlopez@prehistoria.urv.cat (J.M. López-García), mercedes.garcia@uam.es (M. García Antón), eallue@prehistoria.urv.cat (E. Allué), icaceres@prehistoria.urv.cat
(I. Cáceres), rhuguet@prehistoria.urv.cat (R. Huguet), marina.mosquera@urv.cat (M. Mosquera), andreu.olle@prehistoria.urv.cat (A. Ollé), jordi.rosell@urv.cat (J. Rosell),
josep.pares@cenieh.es (J.M. Parés), josepedro.rodriguez@urv.cat (X.P. Rodríguez), clomana@ubu.es (C. Díez), jrofes@unizar.es (J. Rofes), robert.sala@urv.cat
(R. Sala), palmira@prehistoria.urv.cat (P. Saladié), josep@prehistoria.urv.cat (J. Vallverdú), mlluc@prehistoria.urv.cat (M.L. Bennasar), rblasco@prehistoria.urv.cat
(R. Blasco), jm.ber@cenieh.es (J.M. Bermúdez de Castro), eudald.carbonell@urv.cat (E. Carbonell).
1
Visiting Professor IVPP (Beijing).

0277-3791/$ e see front matter Ó 2010 Elsevier Ltd. All rights reserved.
doi:10.1016/j.quascirev.2010.02.021
J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412 1397

as the dominant component of the tree storey was induced by the climatic cycles, and steppes spread
across the landscape during the drier periods. In any case, it is not possible to establish clear cut-off
points separating entirely different environmental episodes. Our results show no evidence of any
relationship between environmental change and cultural change at the Sierra de Atapuerca.
Ó 2010 Elsevier Ltd. All rights reserved.

1. Introduction evaluate the possible existence of correlations between cultural


and environmental changes.
The time period from the late Early Pleistocene to the end of the
Middle Pleistocene was an era of cyclic climatic changes (Berger, 2. The Trinchera del Ferrocarril sites
1988; Rial, 1999). This period also saw the human settlement of
Europe by Homo antecessor, (Carbonell et al., 2008a) and its even- The Sierra de Atapuerca is situated 15 km east of Burgos (Spain)
tual substitution in the fossil record by Homo heidelbergensis and contains several archaeological and palaeontological sites
(Arsuaga et al., 1997). From a cultural point of view, this period also dating from the Early Pleistocene to the Holocene (Arsuaga et al.,
records the appearance in Europe of Mode 1 and Mode 2 techno- 1997; Carbonell et al., 1999b,c, 2008a, Carretero et al., 2008;
logical complexes, and even the transition to Mode 3. Vergés et al., 2008; García and Arsuaga, 2011). Within the Sierra de
Technological Modes were defined by Clark in 1968 (Clarke, Atapuerca, the Trinchera del Ferrocarril includes three sites: Sima
1968). They provide a general framework to define the similarities del Elefante, Gran Dolina and Galería (Fig. 1). These sites are Pleis-
among the stone tool assemblages, grouping them into different tocene karstic deposits that were cut by a railway trench, outcrop-
production techniques. As a summarized by Foley and Mirazon Lahr ping their entire stratigraphic sequences at Gran Dolina and Galería,
(2003, pp 114) “Mode 1, comprising the Oldowan and Asian Pebble
Tool and Chopping Tool Traditions, constituted the simplest mode of
production, the striking of a flake off a core. The number of flakes
could vary, but what held this system of production together was
the simple platforms and lack of preparation involved. (.). Mode 2 Atapuerca
saw the development of two elements (.). The first of these was the Burgos
ability to strike off relatively large flakes (.) suitable for a greater

l
ga
Madrid
amount of invasive retouch. (.). The result was the bifacial tradition Po
rtu
that is represented by the Acheulean and its variants. Mode 3
represents a major shift in the output of lithic production, although Spain
it shares with Mode 2 elements of the way tools are produced. The
key difference is that the core is prepared (.). The outcome is
a much more diverse set of finished tools, and hence a greater
potential for variability and a greater emphasis on smaller items.”
(For an extensive explanation, see Foley and Mirazon Lahr, 2003).
We should clarify that preparation of cores already occurred during
Mode 2, but it was during Mode 3 that this preparation developed,
acquired complexity and became generalized.
The Sierra de Atapuerca includes several archaeological and Gran Dolina
paleontological sites with a record of all these environmental
Tri

and cultural events at a local scale. Sima del Elefante, Gran Dolina Galería
nc
h

and Galería, three sites located in the Trinchera del Ferrocarril


era

(Railway Trench), provide evidence of both environmental (macro


del

and microvertebrates and/or pollen remains) and cultural (stone


F

tools and/or modified bones) changes for the 1.2 to 0.2 Ma period.
erroc

In addition, these three sites have yielded human fossils (Bermúdez


arril

de Castro and Rosas, 1992; Carbonell et al., 1995, 2008a). Conse-


quently, the sites in the Atapuerca Trinchera del Ferrocarril provide Sima del Elefante
a unique opportunity to track the environmental and cultural
changes in an European locality over a one million-year period
covering several climatic cycles, and recording different techno-
logical complexes from Mode 1 to the transition to Mode 3.
A traditional premise states that environmental changes drive
faunal and cultural evolution, suggesting that the emergence of
technology is a result of environmental changes that led the African 42º 21’N
forest landscapes to contract and the savannas to develop between
1025 m..
1000

2.8 and 2.5 Ma (De Menocal, 1995; Vrba et al., 1995). The opposite
view highlights the importance of social and demographic factors
m.

as a motor of cultural changes (Carbonell et al., 1999a), and vice


versa (Carbonell et al., 2008b). 3º 31’ W
The aim of this paper is to summarize information from 0 50 100 m.
different sources to obtain a general picture of the environmental
and cultural changes throughout the Atapuerca sequence, and to Fig. 1. Map of the sites at the Trinchera del Ferrocarril (Atapuerca, Burgos, Spain).
1398 J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412

and most of it at Sima del Elefante. For a detailed description of the antecessor were found in 2008 and 2009 respectively in the TE9 unit
geology of Sierra de Atapuerca see Pérez-González et al. (2001) and associated to a number of stone tools (Carbonell et al., 2008a). These
Parés and Pérez-Gozález (1999). are the oldest human remains that have been found in Europe.
The Sima del Elefante (TE) site is a cave infill with a stratigraphic The Gran Dolina (TD) site is a cave infill 18 m thick located
succession 16 m thick. This site has been systematically excavated approximately 200 m from the Sima del Elefante site (see Fig. 1).
since 1996 (Rosas et al., 2006). The TE infill has been divided into 16 The Gran Dolina stratigraphic succession was divided into 11
stratigraphic units, named TE7 to TE21 from bottom to top (Rosas stratigraphic units named TD1 to TD11 from bottom to top by Gil
et al., 2006). The lower stratigraphic units TE7 to TE16 (Fig. 2a) are and Hoyos (1987) and reviewed by Parés and Pérez-González
dated as Early Pleistocene on the basis of their faunal content, (1999). We follow here the nomenclature published by PÉrez
reversed polarity and two dates based on the radioactive decay of González et al. (2001) for the Gran Dolina sequence, which is not
cosmogenic 26Al and 10Be (Carbonell et al., 2008a), whilst units TE17 entirely coincident with the nomenclature used by other authors
to TE21 date to the Middle and Late Pleistocene (Table 1 and Fig. 3). (Table 2 and supplementary material). A polarity reversal, inter-
A hominin mandible fragment, and a phalanx attributed to Homo preted as the Matuyama-Brunhes boundary (Parés and Pérez-
González, 1999), has been detected between the TD7 and TD8
units, allowing the division of the stratigraphic sequence into an
Early Pleistocene section (TD1-2 to TD7) and a Middle Pleistocene
section (TD8-TD11) coinciding with a gap in the fauna at TD7-TD8
as observed by Cuenca-Bescós et al. (2001). Several dates (Table 1
and Fig. 3) obtained by luminescence and combining electronic
spin resonance and uranium series (ESR/U-Th) techniques confirm
this interpretation (Falguères et al., 2001). The Gran Dolina TD6 unit
is famous for its so-called “Aurora stratum” (Table 2), correspond-
ing to the stratigraphic subunit TD6-2 (Bermúdez de Castro et al.,
2008), which yielded an impressive collection of human remains
attributed to Homo antecessor which were associated with abun-
dant stone tools and faunal remains (Carbonell et al., 1995, 2005).
The Galería (TG) site is located less than 50 m south-east of Gran
Dolina (Fig. 1). Galería is a cave infill with a stratigraphic succession
17 m thick, which has been divided into six lithostratigraphic units
named G I to G VI from bottom to top (Fig. 2c). A polarity reversal
attributed to the Matuyama-Brunhes boundary (Pérez-González
et al., 1999) has been identified inside the lowest unit (G I), which
contains no faunal nor archaeological items. The two middle units G II
and G III are subdivided into four subunits: G IIa, G IIb, G IIIa and G IIIb
from bottom to top. They have yielded abundant macro and micro-
vertebrate fossils and a rich collection of stone tools dating to the
500 ka to 250 ka time interval by ESR/U-Th and luminescence tech-
niques (Berger et al., 2008). Two human fossils attributed to Homo
heidelbergensis, corresponding to cranial and mandibular fragments
were found in these middle units (Bermúdez de Castro and Rosas,
1992; Arsuaga et al., 1999). The three uppermost units, G IV to G VI
are sterile in vertebrate remains and archaeological material: G V
being the last unit of infilling as G VI is an edaphic formation.

3. Chronology

As stated above, the oldest layer excavated to date in Trinchera


del Ferrocarril is TE9 with an estimated age of 1.2e1.1 Ma
(Carbonell et al., 2008a), whilst the youngest is G IV with an age of
approximately 200 ka (Table 1). The identification of the
Matuyama-Brunhes boundary in the three sites and the abundant
absolute dates available allow us to accurately correlate the three
sections so as to obtain a combined sequence for a time period
about one million years long (Fig. 3 and Table 1). It is important to
note that, although some of these dates are inconsistent, they are
coherent if both dating methods (ESR/U-Th and Luminescence) are
considered separately and the confidence intervals are taken into
account. In addition, ESR/U-Th tends to provide older dates than
luminescence for the more recent units and younger ones for the
older units (Fig. 2). Taking all these restrictions into consideration,
reliable correlations may be established between the three strati-
graphic sequences.
Fig. 2. Sections of Sima del Elefante (a), Gran Dolina (b) and Trinchera Galería (c) sites.
TZ (Trinchera Zarpazos) is a gallery only partially filled with sediments. The asterisk
Six Faunal Units (FU 1 to FU 6) based on the distribution of
marks the position of the Matuyama-Brunhes boundary. Heights are measured from vertebrate faunal assemblages have been established for the three
the railway trench floor. sites in Trinchera del Ferrocarril (Cuenca-Bescós and García, 2007;
J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412 1399

Table 1
Absolute dates for Galería, Gran Dolina and Sima del Elefante. ESR/U-Th: Electro Spin Resonance combined with U-series; IRSL: Infrared Stimulated Luminescence;
TL ¼ Thermoluminescence.

Site Unit Age Sample Method Reference


Galería GIV top 211þ/32 e ESR/U-Th
GIV bottom 185þ/26 TL97-14 IRSL Berger et al., 2008
GIIIb 256þ/23 Mean TL98-17 y TL97-24 TL & IRSL Berger et al., 2008
GIIIb 256þ/33 e ESR/U-Th Berger et al., 2008
GIIIa 466þ/39 Mean TL98-15 y TL97-19 Luminescence Berger et al., 2008
GIIa-top 422þ/55 TL98-12 Luminescence Berger et al., 2008
GIIa-bottom 503þ/95 TL97-06 Luminescence Berger et al., 2008

Gran Dolina TD11 240þ/44 OSL97-33 IRSL Berger et al., 2008


TD10-1a 337þ/29 Mean ATA9707. ATA9705, ATA9610 ESR/U-Th Falguères et al., 1999
TD10-1 bottom 379þ/57 ESR-ATA0606 ESR/U-Th Falguères et al., 1999
TD10-2 244þ/26 Mean 97-28, 97-45, 98-23 Luminescence Berger et al., 2008
TD10-2 337þ/51 ESR-ATA9703 ESR/U-Th Falguères et al., 1999
TD10-2 418þ/63 ESR-ATA9608 ESR/U-Th Falguères et al., 1999
TD10-3 430þ/59 TL98-20 Luminescence Berger et al., 2008
TD9 480þ/130 TL97-53 Luminescence Berger et al., 2008
TD8 602þ/52 Mean AT9604, AT9605, AT9702 ESR/U-Th Falguères et al., 1999
TD8 820þ/140 TL97-49 Luminescence Berger et al., 2008
TD7 bottom 960þ/120 TL97-41 Luminescence Berger et al., 2008
TD6-2 731þ/63 Mean AT9601, AT9602, AT9603 ESR/U-Th Falguères et al., 1999

Sima del Elefante TE9 1220þ/140 CTE9b CSM Carbonell et al., 2008
TE7 1130þ/180 CTE7 CSM Carbonell et al., 2008

Cuenca Bescós et al., 2009a). FU 1 is represented by the lower units of 4. Evidence of environmental change
Sima del Elefante (TE8-TE14) and includes species more primitive
than those present in post-Jaramillo faunal localities such as Allo- 4.1. Small vertebrates
phaiomys lavocati, Castillomys rivas, Asoriculus gibberodon and Cro-
cidura kornfeldi. Concerning large mammals, FU 1 is characterized by Most stratigraphic units from the sites in Trinchera del Ferro-
the presence of Villafranchian species like Pannonictis nestii (García carril contain remains of small vertebrates, whose distribution is
et al., 2008), cf. Baranogale antiqua, Mustela cf. palerminea/praeni- summarized in Table 3. The small-vertebrate fossil remains used for
valis, Vulpes cf. alopecoides and Lynx cf. issiodoerensis. As regards this study consist of disarticulated bone fragments collected by
ungulates, the remains of Bison sp. indicate an age younger than 1.2 screen-washing during the 1991 to 2007 excavation campaigns at
Ma, consistent with the available absolute date for TE9 (Fig. 3). Sierra de Atapuerca. Roughly 25 tons of sediment were water-
A major faunal turnover separates this Faunal Unit from FU 2, which screened in each campaign using superimposed 10, 5 and 0.5 mm
is represented at Gran Dolina TD3-TD4. Faunal Unit 2 is character- mesh screens (Cuenca-Bescós et al., 1995, 1997, 1999, 2001, 2005,
ized by the first appearance of advanced vole species such as Allo- 2009a; Blain et al., 2008, 2009; Rofes and Cuenca-Bescós,
phaiomys chalinei, Terricola arvalidens and Iberomys huescarensis, the 2009a,b). More than 70.000 fragments corresponding to
local appearance of Cervus elaphus, Sus scrofa, and the first European a minimum number of 13,314 small vertebrates, representing at
record of the spotted hyena (Crocuta crocuta) in TD3-TD4 (van der least 82 taxa, were recovered with this procedure.
Made, 2001; García and Arsuaga, 2001a,b). Both Cervus elaphus The sample analyzed in the present study is composed of
and Sus scrofa are recorded below the Matuyama-Brunhes boundary mandibles, maxillae and isolated teeth that are loose or in situ, and
only in Atapuerca and in the German site of Dorn Dürkheim (Franzen abundant postcranial elements. Identification at species level is
et al., 2000). The main characteristic of Faunal Units FU 3 and FU 4 is based on isolated mandibles and teeth and in exceptional cases on
the presence of the large red-toothed shrew Dolinasorex glyphodon the most diagnostic postcranial elements, such as the humerus of
(Rofes and Cuenca-Bescós, 2009a), whilst FU 5 is represented by the insectivores; amphibians and squamates are largely based on the
TD7 and TD8 assemblages and it is characterized by the last occur- post-cranial elements. The determinations are based on the works of
rences of primitive vole forms. A second major turnover is detected Chaline (1972), van der Meulen (1973), Reumer (1984), Agustí (1991),
in the transition from FU 5 to FU 6 coincident with the limit between Bailon (1991, 1999), Agustí et al. (1993), Cuenca-Bescós et al. (1995),
TD8 and TD8-9. FU 5 still includes species typical of the later part of Laplana and Cuenca-Bescós (2000), Blain (2005), Minwer-Barakat
the Early Pleistocene like Mimomys savini and many other taxa. TD8 (2005), Rofes and Cuenca-Bescós (2006, 2009a,b), Minwer-Barakat
also registers the last appearance of Stephanorhinus etruscus and et al. (2007), and De Marfà (2008), among many others.
Eucladoceros giulii in general, the last occurrence in Atapuerca of The method of habitat weighting (Evans et al., 1981; Andrews,
Canis mosbachensis, and Panthera gombaszoegensis and the first local 2006) was used to infer environmental information from the pres-
occurrence of Megaloceros solilhacus. The younger Faunal Unit at ence of small vertebrates (amphibians, squamates, and small
Trinchera del Ferrocarril (FU 6) includes the upper levels of Gran mammals) at the Atapuerca sites. This method assigns to each taxon
Dolina (TD8, TD10 and TD11), the middle units of Galería (GII and a probability of being found in each habitat type according to their
GIII), the upper levels of Sima del Elefante (TE18 and TE19) and Sima present distribution in the Iberian Peninsula. Habitats were classi-
de los Huesos (SH) sequence and it is characterized by the presence fied into four types in accordance with Cuenca-Bescós et al. (2005,
of species typical of the Middle Pleistocene like Terricola ata- 2009b) and Blain et al. (2008): 1) open dry: meadows under
puerquensis, Iberomys brecciensis, and Allocricetus correzensis, the seasonal climate regime; 2) open moist: evergreen meadow with
appearance of Dama clactoniana, an evolved form of Cervus elaphus, dense pastures and suitable topsoil; 3) woodland: mature forest
Stephanorhinus hemitoechus, Equus ferus, Equus hydruntinus, Bison including woodland margins and forest patches, with moderate
schoetensacki, Panthera leo, Canis lupus, Vulpes vulpes and the ground cover; 4) water: streams, lakes and ponds. The squamate
subspecies Cuon alpinus europaeus. and amphibian fossils from the Trinchera del Ferrocarril sites belong
1400 J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412

Table 2
1600
1500
1400
1300
1200

1000
1100

900
800
700
600
500
400
300
200
100
Kyr. Lithostratigraphic subunits of Gran Dolina (Pérez-González et al., 2001) and corre-

0
spondence with previously published names. TD11 in van der Made (2001) corre-
TE19 sponds only to the Top of TD10-1.

GIV Lithostratigraphic Lithostratigraphic Old References


Units subunits usage
TD11 TD11 TD11 van der Made (2001)
TD10 TD10-1 TD11
GIIIb TD10-2
TD10-3
TD10-1 TD10-4

TD8-9 TD8b Cuenca-Bescós et al.

Normal polarity
TD10-2
(2001), López Antoñanzas
GIIIa and Cuenca-Bescós (2002)
FU 6
TD8 TD8a López Antoñanzas and
GIIb
Cuenca-Bescós (2002)

TD10-3 TD7 TD7-1


TD7-2
GIIa TD7-3
TD7-4
TD9
TD6 TD6-1 TD6
TD8-9 TD6-2 TD6 “Aurora” Carbonell et al. (1995),
Pares and
TD8 Pérez-González (1995)
FU 5 TD6-3 TD6
TD7 TD5
TD5 þ TD3-TD4? TDE5 van der Made (2001)
TD6-1 TD3-TD4 TDW4b van der Made (2001),
FU 4 TDW4 Rodríguez (2001)
TD6-2

TD6-3
FU 3
TD5 considered as indicative of a single habitat (probability 1 for the
Jaramillo

selected habitat and 0 for the rest, Table 4) based on the present
TD5+TD3-TD4?
distribution of the genus reported by Palombo and Gisbert (2005).
Reversed polarity
Matuyama

We are aware that this is a deliberate underestimation of the


Brunhes

TD3-4 FU 2
adaptability of small mammals, but most small mammal species
TE14 present at the Trinchera del Ferrocarril sites lack recent represen-
tatives and this approach has the advantage of requiring fewer
TE13
assumptions than assigning probabilities of occurrence for different
TE12 habitats. The frequencies in Fig. 4 represent the percentage of the
FU 1 Minimum Number of Individuals indicative of the four habitats in
TE11
each sample. Table 5 shows the number of species and minimum
TE10 number of individuals (MNI) by stratigraphic unit.
The environmental information provided by small vertebrates is
TE9 summarized in Fig. 4. Small mammal assemblages from the lower
levels of Sima del Elefante (TE8-TE14) are characterized by the
TE7
dominance of species suggesting open moist habitats. The presence
of several taxa indicative of riparian or aquatic environments such
Fig. 3. Relative and absolute ages for the stratigraphic units of Sima del Elefante (TE),
Gran Dolina (TD) and Galería (G), according to different methods. The symbols indicate as a desman (Galemys cf. kormosi), shrew (Asoriculus gibberodon),
the mean and the bars two standard deviations above and below the mean for each and beaver (Castor fiber) is remarkable. The interpretation of
sample. Open squares: radioactive decay of cosmogenic 26Al and 10Be; vertical lines: a landscape with water courses and/or lagoon environments is
Luminescence; Open dots: Combined Electron Spin Resonance and Uranium Series; reinforced by the presence of waterfowl (Anas sp.) and a white
open triangle: Infrared Stimulated Luminescence. Faunal units FU 1 to FU 6 are based
on micromammals as shown on Table 3 and “polarity” refers to the magnetization
tailed eagle (Haliaetus albicilla) (Rosas et al., 2006). Conversely, the
direction recorded in the sediments. The chronology of the Jaramillo event and the percentage of open moist and water edge reptile and amphibian
Matuyama-Brunhes boundary are indicated as vertical lines for reference. (Data from species is moderate and the spectrum is dominated by species
Falguères et al., 1999; Berger et al., 2008; Carbonell et al., 2008a,b). indicative of an open dry environment (Fig. 4a).
The single sample from TD5þTD3-TD4 unit containing small
mammals is dominated by open moist species (Fig. 4). However, the
to living species, whose ecological requirements are reliably known. amphibian and reptile assemblage from this unit suggests a mosaic
This knowledge allows us to assign a different probability of environment and discards the existence of very extreme conditions
occurrence to each species for each habitat (Table 4) based on its (Fig. 4). This discrepancy may be partially explained by differences
current distribution (Pleguezuelos et al., 2002; Velasco et al., 2005. in the accumulation agent and partially by the relatively low rich-
As an example, and according to Table 4, the marble newt (Triturus ness of the mammalian assemblage (MNI ¼ 338) (Table 5). A slight
marmoratus) has a probability of 0.4 of being found in an open- increase of open dry and water edge mammals occurs at TD5. The
humid meadow, 0.2 of being found in a woodland and 0.4 of being herpetofauna indicates a slightly drier climate at TD6-2 (Fig. 4) in
found in a water edge. On the other hand, small mammals have been comparison with TD6-3, together with an increase of open
J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412 1401

Table 3
Stratigraphical distribution of small vertebrate species in the Trinchera del Ferrocarril sites. Faunal units FU1 to FU6 were defined by Cuenca Bescós and García (2007) and
Cuenca Bescós et al. (2009a). The letters “ja” refer to the subspecies Microtus agrestis jansoni.

(continued on next page)


1402 J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412

Table 3 (continued)

environments (Blain et al., 2008). However, the presence of Castor indicate that there were no harsh conditions during this period,
fiber at TD6-2 (“Aurora stratum”) is notable, being an indicator of contrary to what was suggested by Lopez-Antoñanzas and Cuenca-
the existence of a permanent water stream in the surroundings. Bescós (2002). Small mammal and herpetofauna spectra are both
Similarly, birds at TD6 are predominantly species of open country dominated by species indicative of open environments and the
and bushland habitats, whilst the presence of waterfowl (Anas sp.) presence of Galemys sp. and Castor fiber at TD8 proves the existence
and waders (Limosa limosa, Scolopax rusticola) constitute additional of permanent water streams in this period. Fig. 4 suggests a tran-
evidence supporting the existence of a large body of water sition inside TD8-9 unit from dry open woodlands to a more open
(Sánchez-Marco, 1999). TD7 is extremely poor in small vertebrate and humid environment. The small mammal assemblage from
remains, but the spectrum of squamate and amphibian species TD10-4 is absolutely dominated by open moist habitat species
indicates a high diversity of habitats (Fig. 4a). (Fig. 4) but, again, the amphibian and squamate assemblages
With reference to the Middle Pleistocene assemblages, the suggest a different and more diverse landscape (Fig. 4). The small
moderate abundance of woodland species at TD8 (Fig. 4), together mammal spectrum from the roughly contemporary GIIa unit of
with the presence of thermophilous amphibians and squamates Galería is also dominated by open habitat species, but with a higher
such as Blanus cinereus, Pelobates cultripes and Rhinechis scalaris percentage of open dry habitat species. Open moist habitat
J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412 1403

Table 4 gregaloides or Allocricetus bursae (Lopez Antoñanzas and Cuenca-


Habitat classification of amphibians, squamates and small mammals used to draw Bescós, 2002), they are always accompanied by temperate or
Fig. 4a. The numbers represent the probability of finding the taxon in each habitat.
All mammalian genera are considered indicators of a single habitat (see text).
even thermophilous species such as Hystrix refossa or Crocidura sp.
It is remarkable that the herpetofauna assemblage always includes
Open-dry Open-humid Woodland Water a significant woodland component. Furthermore, the paleo-
meadow and meadow and woodland edge
rocky habitats margin
temperatures estimated by Blain et al. (2008) for the Atapuerca area
from the Gran Dolina amphibian and squamate assemblages are
AMPHIBIA
Salamandra salamandra 0.4 0.5 0.1 always warmer compared with current temperatures in the Burgos
Triturus marmoratus 0.4 0.2 0.4 area, and the differences observed in estimated temperatures for
Alytes obstetricans 0.6 0.2 0.2 “cold” and “warm” periods are only about 2  C. Similarly, estimated
Pelobates cultripes 0.8 0.2 annual precipitations range from 750 mm in the drier period to
Pelodytes punctatus 0.6 0.2 0.2
Bufo bufo 0.1 0.3 0.4 0.2
1049 mm in the more humid period (Blain et al., 2008), whilst
Bufo calamita 1 current annual precipitation in Burgos is about 570 mm.
Hyla arborea 0.5 0.2 0.3
Rana sp. 0.4 0.4 0.2 4.2. Large mammals
Pelophylax sp. 1

SQUAMATA The stratigraphic distribution of large mammals, defined here as


Blanus cinereus 0.45 0.1 0.45 the species in the orders Primates, Proboscidea, Carnivora, Peri-
Anguis fragilis 0.25 0.75
Natrix maura 1
ssodactyla and Artiodactyla is shown in Tables 6 and 7. The envi-
Natrix natrix 0.5 0.25 0.25 ronmental information provided by the large mammal assemblage
Coronella austriaca 0.25 0.5 0.25 of the lower levels of Sima del Elefante (TE8-TE14) is limited. All
Coronella girondica 0.5 0.25 0.25 species represented are of temperate affinity or, at least, no species
Rhinechis scalaris 0.5 0.25 0.25
unequivocally indicative of arid or cold environments are present.
Vipera aspis 0.25 0.25 0.5
Vipera latasti 0.75 0.25 The presence of Pannonictis nestii in TE9 and TE10 suggests the
existence of a riparian environment in the vicinity, whilst the
MAMMALIA
presence of Hippopotamus in TE14 indicates the existence of either
Crocidura 1
Suncus 1 a deep water stream or a permanent lagoon.
Erinaceus 1 The assemblage from the lower levels of Gran Dolina (TD3-TD4
Sorex 1 to TD8) includes the presence of the spotted hyena, an opportu-
Talpa 1
nistic hunter/scavenger adapted to open landscapes, although the
Beremendia 1
Neomys 1
European jaguar (Panthera gombaszoegensis), an ambush hunter, is
Galemys 1 also present at Atapuerca in this period, suggesting the existence of
Asoriculus 1 forested areas. Again, this association does not include any species
Myotis 1 indicative of a cold environment. The overwhelming dominance of
Rhinolophus 1
Dama vallonnetensis, in TD8 and the presence of Hippopotamus sp.
Miniopterus 1
Marmota 1 suggest relatively humid and temperate conditions.
Pliomys 1 Large-sized lions (P. leo) first occur in Atapuerca at the Sima de
Allocricetus 1 los Huesos (w0.5ka; Bischoff et al., 2007), and successively occupied
Allophaiomys 1 the Middle Pleistocene ecosystems at the Sierra (Table 6). Lions are
Iberomys 1
Terricola 1
social predators, adapted to hunt in open landscapes, whilst dholes
Stenocranius 1 (Cuon alpinus) and wolves (Canis lupus), the other two large pred-
Hystrix 1 ators of this period at Atapuerca, are both social ambushing
Microtus 1 carnivores that may be found in a wide range of habitats.
Micromys 1
As in the case of small vertebrates, large mammal assemblages
Apodemus 1
Castillomys 1 lack any element unequivocally indicative of harsh conditions and
Clethrionomys 1 most species may be considered catholic or temperate. Although
Eliomys 1 Ovibos moschatus is a typical arctic inhabitant, the presence of its
Arvicola 1 related species cf. Praeovibos priscus at TD7 should not be inter-
Mimomys 1
Castor 1
preted as indicative of glacial conditions, as Praeovibos was asso-
Oryctolagus 1 ciated with temperate taxa, such as Hippopotamus, during the
Lepus 1 Pleistocene (van der Made, 2001). The genus Macaca, present in the
lower levels of Sima del Elefante and in the Gran Dolina TD8 unit, is
commonly found in Early Villafranchian woodlands associated with
mammals completely dominate the TD10-3 to TD10-1 spectra relatively humid areas, although from MN17 (2.0 Ma) it inhabited
(Fig. 4b). Woodland rodents and insectivores are virtually absent in a wide range of humidity (Delson, 1980; Eronen and Rook, 2004).
these assemblages but woodland amphibians and squamates are Furthermore, the continuous presence of megaherbivores in all the
well represented (Fig. 4a). Units GIIb and GIII from Galería, similar assemblages supports the persistent existence of open woodland
in age to TD10-3 and TD10-2, also indicate open landscapes but and/or grassland habitats throughout the entire sequence.
they contain a higher abundance of open dry habitat species. The results of isotopic stable analyses, an additional source of
A precise correlation of the upper levels of Sima del Elefante is not environmental information for FU 6, suggest the coexistence of trees
possible at present. In any case, the small mammal assemblage of and open landscapes. Red deer remains from Galería GII and GIII
TE19 indicates a remarkable diversity of habitats, with a large units demonstrate that this species lived in open forests or grass-
proportion of “open moist” and woodland habitat species. lands (García et al., 2009). Although red deer has traditionally been
In summary, although some small vertebrate assemblages considered a forest indicator, it is actually a mixed feeder able to live
include steppe or cold adapted species, for example Stenocranius in very different habitat types (Geist, 1998). On the other hand, other
1404 J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412

an
ge
t

ge
st

ne
s

es

ge
st
d

d
oi

oi
oo ry
ed

y
ed

rra
an

an

oi
tre

y
ed
m

dr
m

rs
d

ea

dr
m

ite
dl

dl
er

er

ife

ic
en

en

en

en

er

ac
en

en
oo

ed
es
at

at

at
op

op

Po
op

op

op

op
co
UNITS
w

M
w

w
0 TE19

10
herpetofauna GIV
small mammals
20

TD11

BRUNHES
30
GIIIb
40

TD10-1
50

TD10-2
60 G III
G IIb

70 TD10-3
G IIab
G IIa
sample

TD8-9 TD10-4
80
TD8
90
TD7-3 and TD7-4
100 TD6-1
TD6-2
110
TD6-3

MATUYAMA
120
TD5

130 palynology TD5+TD3-TD4?

140
TE8 to TE14
150
20 40 60 80 100 20 40 60 80 100 20 40 60 80 100
%NMI %NMI % taxa

Fig. 4. Environmental changes at Atapuerca from three different proxies: squamates and amphibians, small mammals and pollen analysis. Stratigraphic units from Trinchera
Elefante (TE), Gran Dolina (TD) and Galería (G) have been arranged in temporal sequence from the oldest to the youngest based on stratigraphic sequences (Fig. 2) and information
on Fig. 3. For small mammals and herpetofauna the x-axis represents the percentage of the minimum number of individuals, whilst for pollen samples it represents the percentage
of the total number of pollen grains. See text for additional information.

taxa from this stratigraphic unit, such as the herbivorous cave bears
(Ursus deningeri), and fallow deer, displayed more negative d13C
values, suggesting these taxa preferred more forested areas.
Table 5
Species richness and minimum number of individuals (MNI) in the stratigraphic 4.3. Vegetation
units analyzed in Fig. 4.

Herpetofauna Small Mammals Since the 1980s several sediment samples have been taken
Unit Sp. richness MNI Sp. richness MNI
along the exposed sections of the three sites at Trinchera del Fer-
rocarril (Sima del Elefante, Gran Dolina, and Galería). However, the
TE19 e e 17 52
TD10-1 9 151 16 540 pollen concentration of many of these samples is too poor to
TD10-2 11 338 17 756 warrant any environmental interpretation. Few samples have more
GIII e e 16 1552 than 300 pollen grains and spores and most fail to meet the
GIIb e e 14 331 requirements to carry out statistical analyses because of their low
TD10-3 15 1209 18 2327
GIIa e e 16 652
diversity. The extremely low concentration of pollen and spores
TD10-4 15 347 13 227 (0e664 grains/gram of dry sediment) is likely to be explained by
TD8-9 15 201 9 28 the highly oxidative sedimentary environment of the karstic cavi-
TD8 14 199 15 121 ties (Carrión et al., 2009). With regard to macroremains, the
TD7 3/4 9 24 e e
recovery of charcoal fragments from Early and Middle Pleistocene
TD6-1 14 747 13 297
TD6-2 12 234 8 32 deposits is exceptional, mainly because of postdepositional
TD6-3 16 1201 21 542 alterations. The important postdepositional processes suffered by
TD5 11 423 20 460 charcoals make it difficult to identify them taxonomically, since in
TD5 þ TD3-4 13 1098 18 338 most cases charcoal remains are preserved just as microcharcoals,
TE8-14 17 1532 21 1548
which can be quantified but not determined taxonomically.
J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412 1405

Table 6
Stratigraphic distribution of Carnivora at Sima del Elefante (TE), Gran Dolina (TD), Galería(G), and the nearby site of Sima de los Huesos (SH).

Although we are aware of all these caveats, we present here the Mediterranean taxa like Olea-Phillyrea. Acer and Quercus charcoals
available evidence with the aim of providing a “coarse-grained” that have been identified in the Sima del Elefante TE9 unit. The
picture of environmental conditions prevailing in the Atapuerca samples from the upper levels of Sima del Elefante (TE15-TE19) are
area during the one million-year time interval represented at the sterile in pollen content, although a number of charcoals have been
Trinchera del Ferrocarril sites. found at TE19. These show the presence of a Pinus sylvestris-type in
A composite palynological sequence for Gran Dolina and Galería this unit. The presence of Mediterranean taxa is continuous along
has been obtained by correlating the lithostratigraphic units of the the Gran Dolina and Trinchera Galería stratigraphic successions,
two sites as explained above (Fig. 4). Regular samples were taken although sometimes they are dominant and sometimes not (Fig.
every five centimeters from Galería GIII and GIV units (García Antón 4c). From a floristic point of view, the presence of tertiary relict
and Sainz Ollero, 1991) and these were treated using the traditional species at the lower levels of Gran Dolina (Taxodium-type, Pinus
chemical method (Sittler, 1955). The samples included in this anal- haploxylon-type, Carya and Cedrus) should be highlighted (see
ysis were chosen from the existing set, taken every 10, 20 or 30 cm Supplementary Material). Mediterranean taxa are scarce in TD5,
depending on the characteristic or interest of the units. They were where conifers and Poaceae dominate the spectrum. A moderate
treated according to the technique of Goeury and de Beaulieu (1979) increase in the presence of open dry taxa occurs in several samples
but slightly modified following the directives of Girard and Renault- from TD6-3 to TD6-1, although this should not be interpreted as
Miskovsky (1969) and the protocol developed in Burjachs (1990) and indicative of cold steppes but of steppe habitats in a mosaic
Burjachs et al. (2003). Only samples with more than 100 pollen environment. The high abundance of Celtis seeds at TD6-2 is
grains were included in Fig. 4 (see supplementary material). remarkable proof of Mediterranean conditions. A cold steppe
Charcoal remains were recovered manually and observed using would be incompatible with the abundance of Mediterranean
a metallographic microscope with reflected light with 5 to 50 species and the significant presence of mesic taxa in these samples.
magnifications. Each charcoal piece was fragmented by hand in Mediterranean taxa dominate in TD7 and TD8 at the expense of
order to observe the three anatomical sections that permit taxo- a decrease in the abundance of Poaceae (Fig. 4) but the relative
nomic identification. abundances of these two groups are again similar at TD8-9. The two
Fig. 4 summarizes the information provided by the pollen samples at the top of TD8 show a dominance of the mesic trees over
analyses using the ecological classification of taxa shown in Table 8. the Mediterranean taxa and a drastic reduction in conifers,
The extremely low concentration of pollen and spores in the suggesting a period of increased humidity. Interestingly enough the
samples from the lower levels of Sima del Elefante prevents their macrovertebrate fossils from TD8, that indicate a humid environ-
inclusion in the figure. The only point worthy of mention is that the ment, come from this part of the sequence. TD8 also records the last
floristic composition of these samples is similar to the rest of the appearance of relict taxa, a change in floristic composition that
sequence, with conifers, deciduous and evergreen Quercus as tree coincides with the major turnover event registered for both macro
species, and Asteraceae and Poaceae as herbaceous taxa (see and microvertebrates. The single sample from the bottom of TD10
Supplementary Material), together with the presence of (TD10-4) shows a significant increase of Poaceae at the expense of
1406 J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412

Table 7
Stratigraphic distribution of the Ungulates, Subungulates and Primates of Sima del Elefante (TE), Gran Dolina (TD) and Galería (G).

Mediterranean species and mesic trees, suggesting more 5. Cultural evidence


open landscapes. This pattern is more extreme in the samples from
TD10-3, where the abundance of Mediterranean species is extremely The palaeoanthropological and archaeological record of the TE9
low and Poaceae dominate the spectrum, although the presence of stratigraphic unit of Sima del Elefante constitutes the oldest-known
mesic trees in these samples does not suggest a really harsh site in Trinchera del Ferrocarril, and even in Europe, with hominin
environment. The three samples from TD10-2 indicate a climatic fossils and evidence of human activity. The presence of a mandible
improvement marked by a recovery of the Mediterranean compo- fragment attributed to Homo antecessor in TE9 (Carbonell et al.,
nent of the flora, which becomes dominant at TD10-1, matched by 2008a), allows us to consider this species as the hominin inhabit-
a higher abundance of mesic trees (Fig. 4). The temporal sequence is ing the Sierra de Atapuerca at that time. A Mode 1 lithic assemblage
completed with the samples taken at TD11 and Galería GIIIb and GIV composed of 32 artifacts made of Neogene and Cretaceous chert
units, since the samples from GII and GIIIa were extremely poor in has been recovered from TE9. Both these raw materials were
pollen and spores. Mediterranean taxa suffered a significant dimi- available within a radius of 2 km from Sima del Elefante and the
nution at GIIIb, accompanied by an increase of open-moist species presence of small waste flakes and two chert flakes apparently
and, in some samples, also of conifers and steppe species. García belonging to the same core suggest that hominins probably knap-
Antón and Sainz Ollero (1991) established five Pollen Zones for the ped inside the cave. Knapping strategies were simple and aimed at
GIIIb-GIV sequence. The samples from the GIIIb unit correspond to producing simple flakes using unidirectional knapping. Some long
their Pollen Zones I and II, and are dominated by open moist and bones of large herbivores from TE9 show evidence of hominin
mesic taxa, as are the samples from the bottom of the GIV unit, cor- processing in search of bone marrow or other nutritional resources.
responding to Pollen Zones IIIeV of García Antón and Sainz Ollero Several bones with cut marks and/or percussion marks have also
(1991). A group of samples at the middle of the GIV sequence, been recovered at other Early Pleistocene stratigraphic units of
which does not contain faunal nor archaeological remains, shows Sima del Elefante. Mammals of a wide range of body sizes were
a highly impoverished spectrum dominated by conifers and steppe consumed, from rabbits to large bovids, suggesting opportunistic
elements and almost devoid of Mediterranean taxa. These samples dietary habits (Huguet, 2007). Other evidence points to an early
are the only evidence of a harsh, cold and arid, environment in the access to the carcasses, although it is not possible to determine if
entire sequence of Trinchera del Ferrocarril and constitute the Pollen the animals were hunted or scavenged.
Zone IV of García Antón and Sainz Ollero (1991). The last Pollen Zone More recent evidence of human behavior at Atapuerca is located
(V) identified by García Antón and Sainz Ollero (1991) also has at the base of Gran Dolina, in the TD3-TD4 unit. The TD3-TD4 bone
a composition similar to that observed at the lower zone III, domi- assemblage is interpreted as the result of two independent
nated by Mediterranean taxa. processes: the accumulation of large mammals in a natural trap and
J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412 1407

Table 8 several groups of refittings have been identified in the TD6-2


Ecological classification of pollen taxa used to draw Fig. 4. assemblage, proving that hominins knapped inside the cave. The
Group Taxa increased complexity that characterize the technological behavior
Conifers Pinus of the Homo antecessor population represented at TD6-2 is paral-
Cupressaceae leled by a more complex trophic behavior, as evidenced by the
Taxodium-type bone assemblage recovered at the “Aurora stratum” (TD6-2). High
Pinus haploxylonetype
levels of anthropization characterize the assemblage, and all the
Picea
Cedrus. stages of carcass processing have been documented (Díez et al.,
1999; Rosell, 2001; Huguet, 2007; Saladié, 2009), from skinning
Mesic trees Betula
Quercus deciduous
to bone marrow extraction and including defleshing and visceral
Corylus removal. Primary and early accesses to animals by hominids is
Ligustrum reported by Díez et al. (1999), suggesting hunting activities.
Carya A different treatment of the carcasses consumed depending upon
Acer
the animal body size is also evident. Skeletal elements of small
Prunus
Tilia sized prey species (up to 100 kg) are evenly represented, sug-
Carpinus gesting they were transported to the site as complete carcasses. On
Castanea etype the other hand, axial elements of the largest mammals are scarce,
Fagus since they were probably butchered far away and only some parts
Juglans.
were taken to the cave. The high taxonomic diversity of the
Open moist Plantago assemblage, eight ungulate species and six carnivores plus H.
Ranunculaceae
antecessor is remarkable and points towards a non selective
Water edge Alnus hunting behavior. Nevertheless, the most striking feature of the
Fraxinus behavior of Homo antecessor documented at TD6-2 is probably the
Ulmus
practice of cannibalism (Fernández-Jalvo et al., 1996). More than
Salix
Viburnum-Sambucus 150 human fossil remains belonging to a minimum of 10 individ-
Populus uals including children, juveniles and young adults have been
Cyperaceae recovered (Bermúdez de Castro et al., 2008). The butchering
Typha-Sparganium. techniques observed on the hominin bones, aimed at meat and
Poaceae Poaceae. marrow extraction, and the pattern of post- processing discard of
Mediterranean Quercus evergreen these remains, identical to the treatment observed for animal
Olea-Phillyrea bones, is an argument against a ritual significance of this behavior.
Celtis In addition, the large amount of biomass represented by the TD6-2
Pistacia bone assemblage contradicts the hypothesis of an incidental
Coriaria
behavior in response to a starvation episode. Thus, this behavior
Myrtus.
has been interpreted as “nutritional” or “gastronomic” canni-
Open dry Cistaceae
balism associated with long periods in which humans were
Ephedra
Artemisia
feeding on other humans as part of their regular diet (Fernández-
ChenopodiaceaeeAmaranthaceae Jalvo et al., 1999; Bermúdez de Castro et al., 2006).
Cerealia-type Evidence of a different cultural behavior at Atapuerca in the
Asteraceae. Middle Pleistocene comes from the Galería GII and GIIIa strati-
graphic units. Galería is a horizontal cavity with a 7e9 m long
vertical duct to the surface, which acted as a pit fall trap (Huguet
the attritional accumulation of bears which died during hiberna- et al., 2001; Pérez-González et al., 2001). Ungulates, mainly juve-
tion (Rosell, 1998). Carnivore tooth marks, cut marks and evidence nile horses and deer, accidentally fell through the pit hole and both
of anthropic breakage are occasionally present in some elements, hominins and carnivores entered the cave in search of carrion.
proving that both carnivores and hominins sporadically exploited Carnivore activity is intense in the GII and GIII bone assemblages,
this opportunistic trophic resource (Huguet, 2007). Only five but carnivore access to the carcasses was secondary in most cases.
quartzite artifacts, two flakes and three flaked cobbles, have been Under-representation of the skeletal elements associated with large
recovered from TD3-4, although four unidentifiable chert frag- muscular masses and the predominance of axial and cranial
ments were also recovered. As in the TE9 assemblage they can be elements suggest transport of large meat portions outside of the
assigned to a Mode 1 technology. cavity for later consumption. As for the stone artifacts, they present
All these early occupations at Atapuerca seem to be sporadic, as the characteristics typical of Mode 2 techno-complex and, thus, they
characterized by their low intensity, an apparently opportunistic represent a major change when compared with the TD6 assemblage.
behavior and a lack of specialization. The lithic assemblages are Large shaped tools are well represented, many of them on flakes,
attributable to Mode 1 technology, and show low technological and such as handaxes and cleavers. Some of these tools lack the early
raw material diversity, although the scarcity of items argues for stages of the chaîne opératoire, which suggest that they were
caution in the interpretation of this pattern. produced outside the cave (Carbonell et al., 2001). Diversity of raw
A very different picture is shown at TD6-2 (“Aurora stratum”) materials is high, although all could be found inside a 2e5 km radius
where several episodes of high intensity occupation are around the site (Gabarró et al., 1999; García-Antón et al., 2002). That
represented. The lithic assemblage is also of Mode 1 technology different raw materials were selected to make different instruments
but a high diversity of knapping strategies has been observed and is evidence of complex behavior (Mosquera, 1998). Some knapping
seven different kinds of raw material have been identified, techniques that were common in the TD6-2 assemblage disappear
although almost half of the artifacts are made of Neogene chert and new ones are used for the first time at Atapuerca. Use-wear
(Carbonell et al., 1999d; Rodríguez, 2004a). All of the pieces analyses carried out on stone tools support the zooarchaeological
produced and used during the entire chaîne opératoire, as well as data (Ollé et al., 2005), and have documented mainly butchering
1408 J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412

activities, some hide working and the sporadic work on plant neanderthalensis, as proposed by Rosas and Bermúdez de Castro
material (Marquez et al., 1999, 2001; Ollé, 2003). Regarding the (1998).
hominin species that carried out these activities, the mandible
found in the lower levels has been attributed to Homo 6. Discussion
heidelbergensis by Bermúdez de Castro and Rosas (1992).
A different occupation is documented in the slightly more Climatic change around 1.2e1 Ma lead to a predominance of the
recent TD10-2 unit of Gran Dolina, adding to our understanding of 100 ka Milankovich cycles, which are related to the well known
the behavioral variability of this hominin population. The bone glacial cycles (De Menocal, 1995). This is reflected in a major faunal
assemblage is dominated by remains of a small species of bison turnover in Europe, with many extinctions and dispersals (e.g. van
(Bison sp.) and all the stages of the process of carcass exploitation der Made, 2005, Fig. 4.2; van der Made, 2011). In mid-latitude
are documented, evidencing intensive use of this trophic resource. Europe (north of the Pyrenees) or Eurasia this is noted in an
From a technological point of view, the artifact assemblage alternation of glacial and interglacial faunas. During glacials the
exhibits low raw material diversity and is dominated by chert closed environment of the taiga disappeared and the tundra and
artifacts. steppe united into what is called the “Mammoth steppe” (Guthrie,
The more recent cultural evidence at the Trinchera del Ferro- 1990), in which a fauna lived that includes mammoths, woolly
carril sites comes from the TD10-1 unit of Gran Dolina and GIIIb rhinoceroses and other mammals (Kahlke, 1999). This “glacial”
unit of Galería. These units are roughly coetaneous (Fig. 2), fauna derived mainly from the faunas that lived during interglacial
although GIIIb appears to be slightly younger. As in the former case, times in the tundra, steppe and mountains. The “interglacial” fauna
they document two different behaviors of allegedly the same survived during glacial times in the Southern peninsulas of Europe
population, since Galería was a natural trap and TD10-1 represents and parts of Asia, such as Anatolia - northern Iran in the west and
a base camp. Occupation was particularly intense at TD10-1, shown China in the east. In this context, the lack of evidence of harsh
by the fact that more than 80,000 bones and bone fragments and climatic conditions or extreme environments at Atapuerca during
20,000 artifacts and debris have been recovered from an 80 m2 the 1.2 Ma to 250 Ka time interval, shown in the previous sections,
excavation. This different occupational use of the two cavities reflects the prevalence of warm and equable conditions in Southern
probably explains the scarcity of artifacts and the absence of the Europe during this period. The only evidence suggesting a possible
early stages of the technological chaîne opértoire at Galería GIII, extremely cold period leading to a cold steppe landscape is found in
making it difficult to compare the two artifact assemblages. the composition of two pollen samples from Galería GIV unit, dated
Centripetal cores tending to Levallois characterize the TD10-1 around 200 kyr, a period without evidence of human presence at
assemblage, evidencing a transition towards a Mode 3 Atapuerca. Interestingly, steppe environments have been also
technology. Processing of animal carcasses continues to be intense, inferred for other sites in the center of the Iberian Peninsula of
although large bovids are substituted by medium sized ungulates similar or younger age (Ruiz Zapata et al., 2004; Carrión et al.,
(cervids and horses) as the main animal resource. 2007). However, apart from this episode, the rest of the pollen
The attribution of the lithic industries of Galería and TD10 to samples show, in varying proportions, a continuous presence of
Homo heidelbergensis is sustained by the presence of a mandible of Mediterranean taxa and mesic trees. In addition, the herpetofauna
this species at the lower levels of Galería. Here H. heidelbergensis is supports the permanent existence of woodland areas, although
considered a Middle Pleistocene European species ancestral to H. species linked to open (moist or dry) habitats always dominate the

Table 9
Summary of environmental and cultural changes in Atapuerca. Cultural Phases, Faunal Units (FU) and technological Modes are defined in the text. The environment is
a tentative interpretation based on the evidences summarized in Fig. 4.

Layer Technological Mode Human species Cultural Environment


Middle Pleistocene No Fauna GIV Open woodland and meadows
Steppe
Open wooland and meadows
FU6 GIIIb Mode 2 Phase III Open woodland and meadows
TD10-1 Mode 2/3 Phase III Mediterranean open woodland
TD10-2 Mode 2 Phase III Mediterranean open Woodland
GIIIa Mode 2 Phase III Mediterranean open woodland
TD10-3 Mode 2 Phase III Moist meadows with scattered trees
GIIb Mode 2 Phase III Open Woodland and meadows
GIIa Mode 2 H. heidelbergensis Phase III Open Woodland and meadows
TD10-4 Mode 2 Phase III Moist meadows with scattered trees
TD8-9 Open woodland and meadows
FU5 TD8 Humid temperate open woodland
Early Pleistocene TD7 Mediterranean open woodland
FU4 TD6-1 Mode 1 Phase II Open woodland and steppe
TD6-2 Mode 1 H. antecessor Phase II Open woodland and steppe
FU3 TD6-3 Open woodland and steppe
TD5 Open arid(?)
? TD3-TD4+TD5? Open woodland and moist meadows
FU2 TD3-TD4 Mode 1 Phase I Open woodland and moist meadows
FU1 TE14 Phase I Temperate open woodlands and meadows with lagoons.
TE13 Phase I Temperate open woodlands and meadows with lagoons.
TE12 Phase I Temperate open woodlands and meadows with lagoons.
TE11 Phase I Temperate open woodlands and meadows with lagoons.
TE10 Phase I Temperate open woodlands and meadows with lagoons.
TE9 Mode 1 H. antecessor Phase I Temperate open woodlands and meadows with lagoons.
J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412 1409

spectra. On the contrary, small mammals suggest an almost abso- documented at TD6-2 and cannibalism was apparently a common
lute dominance of open habitats, with a continuous predominance practice (Fernández-Jalvo et al., 1999). Finally, TD6-2 represents up
of “open moist” habitat species over “open dry” habitat ones. Here to now the earliest base camp known at Atapuerca. Open wood-
“open moist” refers to prairie environments, whilst the open dry lands also dominated the landscape coinciding with this cultural
category is equivalent to steppes. Thus, typical cold steppe species phase, although more Mediterranean in character than in the
are absent from the Atapuerca small mammals record and the same previous phase (Table 9). In addition, some periods within this
is true for the large mammals, since all the species present may be phase were probably dominated by steppes.
considered catholic or temperate. The third phase corresponds to the second half of the Middle
The apparent contradictions between the environments indi- Pleistocene (roughly 500e200 ka) and it is represented by the
cated by the herpetofauna and the small mammals may be explained Galería GII-GIII assemblages and the Gran Dolina TD10-4 to TD10-1
as the result of either the differential biases introduced by their assemblages. This phase represents a major cultural change,
accumulation agents or of methodological artifacts. It is conceivable marked by the appearance of a well developed Mode 2 technology.
that different agents sampling different habitats from a complex Both sites document two different and complementary behaviors
landscape, as is probably the case at Atapuerca (Fernández-Jalvo, of the Homo heidelbergensis populations: use of an opportunistic
1994, 1999; Rosas et al., 1999; Blain et al., 2008, 2009), would but well controlled resource at Galería and systematic hunting
produce assemblages that were apparently disharmonious. Con- reflected at the Gran Dolina TD10-2 and TD10-1 base camp, with
cerning methodological artifacts, herpetofauna and small mammals higher directionality as regards the TD6 base camp. Also, the
are environmental proxies with different resolution. Distribution presence in Galería of tools made outside the cave, suggests these
and diversity of reptiles and amphibians are tightly controlled by hominins were not entirely opportunistic, but that they controlled
climate (Whittaker et al., 2007) and, thus, we may regard them as and used their landscapes with a premeditated behavior to plan
reliable climate proxies. In addition all herpetofaunal remains their activities. Finally, the presence in the TD10-1 assemblage of
identified at Atapuerca belong to living species (Blain et al., 2008, some characteristics that suggest a shift towards a Mode 3 tech-
2009), whose habitat requirements are well known, and each of nology may be reflecting the appearance of a fourth episode of
these species may be assigned a probability of occurrence for each cultural change. Again, open although not tree-less landscapes
habitat. Conversely, most of the small mammals identified at the dominated throughout this cultural phase (Table 9). A particularly
Trinchera del Ferrocarril sites lack recent representatives and their open landscape may correspond to TD10-4 and, especially to TD10-
habitat requirements have been extrapolated from closely related 3, where Poaceae pollen is extremely abundant, Mediterranean
living species, assigning a single habitat type to each species. Few species are scarce in the pollen spectra and the small mammal
small European mammals may be classified as strictly woodland assemblages are devoid of woodland species. However, other
species, but most of the open country species may also be found in periods inside this phase were dominated by open woodlands
woodlands (Mitchell-Jones et al., 1999). This methodological artifact similar in character to those inferred for the two previous cultural
helps to explain the underrepresentation of woodlands in the phases.
landscapes inferred from the small mammal assemblages. It is important to note that the first two cultural phases corre-
Pollen spectra show the continuous presence of conifers, mesic spond to Homo antecessor, whilst the third one was developed by
and Mediterranean trees (evergreen Quercus, Olea-Phillyrea, Pista- a different hominin species, namely Homo heidelbergensis. Unfor-
cia, Myrtus) in varying proportions, together with Poaceae and open tunately, the lithostratigraphic units separating TD6 from TD10 (i.e.
dry species. These pollen spectra suggest two possible landscapes: TD7 and TD8) lack human remains and evidence of human activity.
a mosaic of woodland and open habitats or a savanna-like open Thus, with the evidence currently at hand, it is not possible to
woodland. The abundance of megaherbivoresand the scarcity of deduce if the third phase developed out of the local Early Pleisto-
forest adapted mammals are incompatible with the existence of cene culture or if it was an innovation produced elsewhere that
large forested areas, however a significant tree component is replaced the pre-existent local culture. What seems evident is that
always present in the pollen samples. In addition, the structures of these three cultural phases cannot be clearly related to any episode
the mammalian paleocommunities living at Atapuerca throughout of environmental change.
this period resemble in some aspects recent savanna communities It has been proposed that the early human presence in Europe
(Rodríguez, 2001), which supports the second interpretation. (e.g. before 0.5 Ma) was discontinuous (for instance, Dennell and
We have divided the sequence of cultural changes recorded at Roebroeks, 1996) and one of the reasons alleged for this disconti-
Atapuerca into three well differentiated phases (Table 9). The first nuity is glacial cyclicity. By now many localities, other than Ata-
phase includes the oldest occupations, found at the lower levels of puerca, testify to a human presence in Europe before 0.5 Ma ago,
Sima del Elefante and Gran Dolina TD3-TD4. These are low intensity including Barranco Léon 5, Fuente Nueva 3 (Oms et al., 2000),
occupations, characterized by very simple knapping strategies, very Vallonnet (Yokoyama et al., 1988), and Isernia (Coltorti et al., 2005).
low raw material diversity and an apparently opportunistic The evidence shown here supports the prevalence of temperate
behavior. Temperate open woodlands seem to have dominated the habitats at Atapuerca during much of the time between the earliest
landscape at this time. This was probably one of the most humid human arrival and about 0.5 Ma, when Homo heidelbergensis and
periods at Atapuerca, since several proxies point towards a signifi- Mode 2 appeared. This suggests that Southern Europe may have
cant presence of meadows and/or lagoons. been continuously inhabited during this period of about 0.7 Ma
A major cultural change is observed in the second phase, rep- duration and reflects the ecological preferences of Homo antecessor
resented by Dolina TD6-1 and TD6-2 assemblages and chronolog- for the warmer and more equitable conditions of Southern Europe
ically corresponding to the end of the Early Pleistocene. The lithic as suggested by several authors (Roebroeks, 2001; van der Made,
assemblage still presents the characteristics typical of Mode 1 1999). From a local point of view, the replacement of H. ante-
technology, but knapping strategies are more complex and diver- cessor by H. heidelbergensis may be considered as part of the faunal
sified, and a wide variety of raw materials is present. Although this turnover separating Faunal Unit FU 6 from FU 5. However, this
development may be a false reflection of lithic abundance at TD6, seems to be a purely taxonomic turnover, since the assemblages
regarding the scarcity in earlier phases, the rest of the hominin from FU 1 to FU 5 are not essentially different to those from FU 6 on
activities support a more complex behavior: primary access, the basis of their ecological characteristics. For instance: Stepha-
probably corresponding to opportunistic hunting, is well norhinus etruscus, Equus altidens, Dama vallonnetensis and Bison cf.
1410 J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412

voigtstedtensis are replaced by Stephanorhinus hemitoechus, Equus Appendix. Supplementary information


ferus, Dama dama clactoniana and Bison schoetensacki/Bison sp. The
lack of correlation between this turnover and any significant Supplementary material associated with this article can be
ecological change and the existence of a remarkable stability in the found in the online version at doi:10.1016/j.quascirev.2010.02.021.
ecological composition of the mammalian paleocommunities at
Atapuerca were previously highlighted by Rodríguez (2004b). References

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