One Million Years of Cultural Evolution in a Stable Environment at Atapuerca (Burgos, Spain)

Quaternary Science Reviews 30 (2011) 1396e1412

Contents lists available at ScienceDirect

Quaternary Science Reviews

journal homepage: www.elsevier.com/locate/quascirev

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 article info abstract

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 fi 16 February 2010 in llings 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 technology 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: [email protected] (J. Rodríguez), [email protected] (F. Burjachs), [email protected] (G. Cuenca-Bescós), [email protected] (N. García), [email protected] (J. Van der Made), [email protected] (A. Pérez González), [email protected] (I. Blain), [email protected] (I. Expósito), [email protected] (J.M. López-García), [email protected] (M. García Antón), [email protected] (E. Allué), [email protected] (I. Cáceres), [email protected] (R. Huguet), [email protected] (M. Mosquera), [email protected] (A. Ollé), [email protected] (J. Rosell), [email protected] (J.M. Parés), [email protected] (X.P. Rodríguez), [email protected] (C. Díez), [email protected] (J. Rofes), [email protected] (R. Sala), [email protected] (P. Saladié), [email protected] (J. Vallverdú), [email protected] (M.L. Bennasar), [email protected] (R. Blasco), [email protected] (J.M. Bermúdez de Castro), [email protected] (E. Carbonell). 1 Visiting Professor IVPP (Beijing).

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 . fi saw the development of two elements ( ). The rst of these was the Burgos ability to strike off relatively large flakes (.) suitable for a greater amount of invasive retouch. (.). The result was the bifacial tradition Madrid that is represented by the Acheulean and its variants. Mode 3 Portugal 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 T paleontological sites with a record of all these environmental r in c and cultural events at a local scale. Sima del Elefante, Gran Dolina h Galería e r and Galería, three sites located in the Trinchera del Ferrocarril a d (Railway Trench), provide evidence of both environmental (macro e l and microvertebrates and/or pollen remains) and cultural (stone F e tools and/or modified bones) changes for the 1.2 to 0.2 Ma period. r r o In addition, these three sites have yielded human fossils (Bermúdez c a

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

05m. 1025 102 forest landscapes to contract and the savannas to develop between 100010 2.8 and 2.5 Ma (De Menocal, 1995; Vrba et al., 1995). The opposite 00 5

m. view highlights the importance of social and demographic factors m. . as a motor of cultural changes (Carbonell et al., 1999a), and vice et al., 2008b). versa (Carbonell 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), corresponding 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 abundant 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 techniques (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 stratigraphic sequences. Fig. 2. Sections of Sima del Elefante (a), Gran Dolina (b) and Trinchera Galería (c) sites. Six Faunal Units (FU 1 to FU 6) based on the distribution of TZ (Trinchera Zarpazos) is a gallery only partially filled with sediments. The asterisk 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

1000

1200 1300 1400 1500 1600

1100

100 200 300 400 500 600 700 800 900 Table 2

0 Kyr. Lithostratigraphic subunits of Gran Dolina (Pérez-González et al., 2001) and corre- 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 omlpolarity Normal TD10-2 TD8-9 TD8b Cuenca-Bescós et al. (2001), López Antoñanzas and Cuenca-Bescós (2002) GIIIa 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 Jaramillo TD5 considered as indicative of a single habitat (probability 1 for the selected habitat and 0 for the rest, Table 4) based on the present TD5+TD3-TD4?

Reversed

Matuyama distribution of the genus reported by Palombo and Gisbert (2005).

Brunhes TD3-4 FU 2 We are aware that this is a deliberate underestimation of the adaptability of small mammals, but most small mammal species

polarity TE14 present at the Trinchera del Ferrocarril sites lack recent representatives 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 tailed eagle (Haliaetus albicilla)(Rosas et al., 2006). Conversely, the on micromammals as shown on Table 3 and “polarity” refers to the magnetization 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. even thermophilous species such as Hystrix refossa or Crocidura sp. All mammalian genera are considered indicators of a single habitat (see text). 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 1049 mm in the more humid period (Blain et al., 2008), whilst Bufo bufo 0.1 0.3 0.4 0.2 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 ssodactyla and Artiodactyla is shown in Tables 6 and 7. The envi- Natrix maura 1 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 preted as indicative of glacial conditions, as Praeovibos was asso- Castor 1 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

edge moist dry

woodland open water open drywoodland open moist water edgeopen dry conifersmesic treesopen moistwater edgePoaceaeMediterraneanopen UNITS 0 TE19

10 herpetofauna small mammals GIV 20

TD11 30 GIIIb 40

TD10-1 BRUNHES 50

TD10-2 60 GIII GIIb 70 TD10-3 GIIab GIIa TD8-9 TD10-4 80

sample TD8 90 TD7-3 and TD7-4 100 TD6-1 TD6-2 110 TD6-3

120 TD5

130 palynology TD5+TD3-TD4?

TUYAMA

140 MA 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 ee17 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 ee16 1552 than 300 pollen grains and spores and most fail to meet the ee GIIb 14 331 requirements to carry out statistical analyses because of their low TD10-3 15 1209 18 2327 diversity. The extremely low concentration of pollen and spores GIIa ee16 652 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 ee TD7 3/4 9 24 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 5to50 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 IIIeVofGarcía 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 e Pinus haploxylon type 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 to bone marrow extraction and including defleshing and visceral Quercus deciduous 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 stratigraphic 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

Agustí, J., 1991. The Allophaiomys complex in Southern Europe. Geobios 25, 7. Conclusions 133e144. Agustí, J., Castillo, C., Galobart, A., 1993. Heterochronic evolution in the Late Plio- e It is apparent that faunal and cultural changes are not related to cene Early Pleistocene arvicolids of the Mediterranean area. Quaternary International 19, 51e56. environmental change at Atapuerca. This result is especially Agustí, J., Oms, O., Remacha, E., 2001. Long Plio-Pleistocene terrestrial record of unexpected in the case of faunal changes, since they are usually climate change and mammal turnover in Southern Spain. Quaternary Research e interpreted in terms of climate change (Azanza et al., 2000; Agustí 56, 411 418. Andrews, P., 2006. Taphonomic effects of faunal impoverishment and faunal mix- et al., 2001). More data are needed from other similarly long ing. Palaeogeography, Palaeoclimatology, Palaeoecology 241, 572e589. temporal sequences in order to determine if this was just a local Arsuaga, J.L., Bermúdez de Castro, J.M., Carbonell, E. (Eds.), 1997. The Sima de los phenomenon or a continental scale pattern. Huesos hominid Site. Journal of Human Evolution 33 (special issue 2e3), 105e421. Two major local faunal changes have been detected: one around Arsuaga, J.L., Gracia, A., Lorenzo, C., Martínez, I., Pérez, P.J., 1999. Resto craneal de 1 Ma ago and the other one around 600e500 ka. The first transition Galería/Cueba de los Zarpazos (Sierra de Atapuerca, Burgos). In: Carbonell, E., occurred during Cultural Phase I, represented at TE9 and TD3-TD4 Rosas, A., Díez, J.C. (Eds.), Atapuerca: ocupaciones humanas y paleoecología del yacimiento de galería. Junta de Castilla y Leon, Valladolid, pp. 233e235. by Homo antecessor and Mode 1 technology, without any Azanza, B., Alberdi, M.T., Prado, J.L., 2000. Large mammal turnover pulses correlated apparent effect on hominid behavior. No faunal or environmental with latest Neogene glacial trends in the northwestern Mediterranean region. changes occurred during the transition towards the second cultural In: Hart, M.B. (Ed.), Climates: Past and Present. Geological Society, London, e phase represented at TD6 around 800 ka ago, and characterized by Special Publications, pp. 161 170. Bailon, S., 1991. Amphibiens et reptiles du Pliocène et du Quaternaire de France et new subsistence and technological strategies. The faunal change d'Espagne: mise en place et évolution des faunes. Thèse 3ème cycle, Université around 600e500 ka includes the replacement of Homo antecessor de Paris VII, France. by H. heidelbergensis and coincides with the transition from Bailon, S., 1999. Différenciation ostéologique des Anoures (Amphibia, Anura) de France. In: Desse, J., Desse-Berset, N. (Eds.), Fiches d'ostéologie animale pour Cultural Phase II to Cultural Phase III, although it is not related to l'archéologie, Série C: varia. Centre de Recherches Archéologiques e CNRS, pp. any significant environmental transition. This third cultural phase 3e41. ranges between 500 kae200 ka and is represented at Galería and Berger, A., 1988. Milankovitch theory and climate. Reviews of Geophysics 26 (4), 624e657. TD10 by Mode 2 technology and systematic and directional Berger, G.W., Pérez-González, A., Carbonell, E., Arsuaga, J.L., Bermúdez de Castro, J.- hunting. Finally, no faunal or environmental changes are related to M., Ku, T.-L., 2008. Luminescence chronology of cave sediments at the Ata- the transition to Mode 3 detected in TD10-1. puerca paleoanthropological site, Spain. Journal of Human Evolution 55, 300e311. From a technological point of view, there is no way at present to Bermúdez de Castro, J.M., Rosas, A., 1992. A human mandible fragment from the evaluate whether Mode 2 emerged from the local Mode 1 in Atapuerca Trench (Burgos, Spain). Journal of Human Evolution 22, 41e46. Atapuerca. Mode 1 from TD6-2 seems to be more evolved than that Bermúdez de Castro, J.M., Carbonell, E., Gómez, A., Mateos, A., Martinón-Torres, M., Muela, A., Rodríguez, J., Sarmiento, S., Varela, S., 2006. Paleodemografía del from TE9 and TD3-4, although both these latter levels yielded few hipodigma de fósiles de homínidos del nivel TD6 de Gran Dolina (Sierra de artifacts. The matter of whether a similar process of technological Atapuerca, Burgos): estudio preliminar. Estudios Geologicos 62, 145e154. change may have eventually produced Mode 2 cannot be answered Bermúdez de Castro, J.M., Pérez-González, A., Martinón-Torres, M., Gómez- as yet: a hiatus of approximately 200 ka without hominin activities, Robles, A., Rosell, J., Prado, L., Sarmiento, S., Carbonell, E., 2008. A new early Pleistocene hominin mandible from Atapuerca-TD6, Spain. Journal of Human represented by TD7 and TD8 levels, makes it impossible to link both Evolution 55, 729e735. technologies. At around 500 ka a different hominin species Bischoff, J.L., Williams, R.W., Rosenbauer, R.J., Aramburu, A., Arsuaga, J.L., García, N., appeared, and it is probably not a coincidence that at the same time Cuenca-Bescós, G., 2007. High-resolution U-series dates from the Sima de los Huesos hominids yields 600þ/-66 kyrs: implications for the evolution of the Mode 2 dispersed relatively quickly throughout Europe. By early Neanderthal lineage. Journal of Archaeological Science 34, 763e770. contrast, the TD10-1 lithic assemblage suggests that an incipient Blain, H.-A., 2005. Contribution de la Paléoherpétofaune (Amphibia et Squamata) à Mode 3 may have emerged at Atapuerca from the local Mode 2. la connaisance de l'évolution du climat et du paysage du Pliocène supérieur au Pléistocene moyen d'Espagne. Thèse de Doctorat du Muséum National d'His- toire Naturelle, Paris, France. Blain, H.-A., Bailon, S., Cuenca-Bescós, G., 2008. The Early-Middle Pleistocene Acknowledgements palaeoenvironmental change based on the squamate reptile and amphibian proxies at the Gran Dolina site, Atapuerca, Spain. Palaeogeography, Palae- The authors wish to thank J. Carrión, J. Rose and C. Stringer for oclimatology, Palaeoecology 261, 177e192. Blain, H.-A., Bailon, S., Cuenca-Bescós, G., Arsuaga, J.L., Bermúdez de Castro, J.M., their kind invitation to participate in this volume and R. Quam for Carbonell, E., 2009. Long-term climate record inferred from Early-Middle comments on the manuscript. This research has been funded by the Pleistocene amphibian and squamate reptile assemblages at the Gran Dolina MICINN projects CGL2009-12703-C03-01, CGL2009-12703-C03-02 cave, Atapuerca, Spain. Journal of Human Evolution 56, 55e65. Burjachs, F., 1990. Palinologia dels dòlmens de l'Alt Empordà i dels dipòsits qua- and CGL2009-12703-C03-03 and CGL2006-04548/BTE the JCyL GR ternaris de la cova de l'Arbreda (Serinyà, Pla de l'Estany) i del Pla de l'Estany 249-2008; and the MICINN project HAR2008-01984/HIST. We are (Olot, Garrotxa). Evolució del paisatge vegetal i del clima des de fa més de also indebted to Fundación Atapuerca, Grupos Consolidados del 140.000 anys al N.E de la Península Ibèrica. Publicacions de la Universitat Gobierno de Aragón and University of Zaragoza for the support Autònoma de Barcelona. Barcelona. Burjachs, F., López-Sáez, J.A., Iriarte, M.J., 2003. Metodología arqueopalinológica. In: provided to this work. Fieldwork at Atapuerca is funded by the Buxó, R., Piqué, R. (Eds.), La recogida de muestras en arqueobotánica: objetivos y Junta de Castilla y León. R. Blasco is beneficiary of a FI Grant from propuestas metodológicas. Museu d'Arqueologia de Catalunya, Barcelona, pp.11e18. Generalitat de Catalunya and financed by the European Social Carbonell, E., Bermúdez de Castro, J.M., Arsuaga, J.L., Díez, J.C., Rosas, A., Cuenca- Bescós, G., Sala, R., Mosquera, M., Rodríguez, X.P., 1995. Lower Pleistocene Found. P Saladié recieved a reaserch grant from Fundación Ata- hominids and artifacts from Atapuerca TD6 (Spain). Science 269, 826e830. puerca/Duques de Soria and R. Huguet from Fundación Atapuerca. Carbonell, E., Mosquera, M., Rodríguez, X.P., Sala, R., van der Made, J., 1999a. Out of J.M.L.-G. has been supported by a postdoctoral grant from Juan de la Africa: the dispersal of the earliest technical systems reconsidered. Journal of e fi Anthropological Archaeology 18, 119 136. Cierva Subprogram, with the nancial sponsorship of the Spanish Carbonell, E., Esteban, M., Martín Nájera, A., Mosquera, M., Rodríguez, X.P., Ollé, A., Ministry of Science and Innovation. Sala, R., Vergés, J.M., Bermúdez de Castro, J.M., Ortega Martínez, A.I., 1999b. The J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412 1411

Pleistocene site of Gran Dolina, Sierra de Atapuerca, Spain: a history of the Díez, J.C., Fernández-Jalvo, Y., Rosell, J., Cáceres, I., 1999. Zooarchaeology and archaeological investigations. Journal of Human Evolution 37, 313e324. taphonomy of Aurora stratum (Gran Dolina, Sierra de Atapuerca, Spain). Journal Carbonell, E., Rosas, A., Díez, J.C. (Eds.), 1999c. Atapuerca: ocupaciones humanas y of Human Evolution 37, 623e652. paleoecología del yacimiento de galería. Junta de Castilla y Leon, Valladolid. Eronen, J.T., Rook, L., 2004. The Mio-Pliocene European primate fossil record: Carbonell, E., García-Antón, M.D., Mallol, C., Mosquera, M., Ollé, A., Rodríguez, X.P., dynamics and habitat tracking. Journal of Human Evolution 47 (5), 323e341. Soahnouni, M., Sala, R., Vergés, J.M., 1999d. The TD6 level lithic industry from Evans, E.M.N., Van Couvering, J.A.H., Andrews, P., 1981. Palaeoecology of Miocene Gran Dolina, Atapuerca (Burgos, Spain): production and use. Journal of Human Sites in Western Kenya. Journal of Human Evolution 10, 99e116. Evolution 37, 653e693. Falguères, C., Bahain, J.-J., Yokoyama, Y., Arsuaga, J.L., Bermúdez de CAstro, J.M., Carbonell, E., Mosquera, M., Ollé, A., Rodríguez, X.P., Sahnouni, M., Sala, R., Carbonell, E., Bischofgf, J.L., Dolo, J.M., 1999. Earliest humans in Europe: the age Vergès, J.M., 2001. Structure morphotechnique de l'industrie lithique du Pléis- of TD6 Gran Dolina, Atapuerca, Spain. Journal of Human Evolution 37, 343e352. tocène infériur et moyen d'Atapuerca (Burgos, Espagne). L'Anthropologie 105, Falguères, C., Bahain, J.-J., Yokoyama, Y., Bischoff, J.L., Arsuaga, J.L., Bermúdez de 259e280. Castro, J.M., Carbonell, E., Dolo, J.M., 2001. Datations par RPE et U-Th des sites Carbonell, E., Bermúdez de Castro, J.M., Arsuaga, J.L., Allue, E., Bastir, M., Benito, A., Pleistocenes d'Atapuerca: Sima de los Huesos, Trinchera Dolina et Trinchera Cáceres, I., Canals, T., Díez, J.C., Van der Made, J., Mosquera, M., Ollé, A., Pérez- Galería. Bilan géochronologique. L'Anthropologie 105, 71e81. González, A., Rodríguez, J., Rodríguez, X.P., Rosas, A., Rosell, J., Sala, R., Fernández-Jalvo, Y., 1994. Small mammal taphonomy at la Trinchera de Atapuerca Vallverdú, J., Vergés, J.M., 2005. An Early Pleistocene hominin mandible from (Burgos Spain). A remarkable example of taphonomic criteria used for strati- Atapuerca-TD6, Spain. Proceedings of the National Academy of Sciences of the graphic correlations and paleoenvironment interpretations. Palaeogeography, United States of America 102, 5674e5678. Palaeoclimatology, Palaeoecology 114, 167e195. Carbonell, E., Bermúdez de Castro, J.M., Parés, J.M., Pérez-González, A., Cuenca- Fernández-Jalvo, Y., 1999. Tafonomía de micromamíferos del yacimiento de Galería Bescós, G., Olle, A., Mosquera, M., Huguet, R., van der Made, J., Rosas, A., Sala, R., de Atapuerca (Burgos). In: Carbonell, E., Rosas, A., Díez, J.C. (Eds.), Atapuerca: Vallverdú, J., García, N., Granger, D.E., Martinón-Torres, M., Rodríguez, X.P., ocupaciones humanas y paleoecología del yacimiento de Galería. Junta de Stock, G.M., Vergès, J.M., Allué, E., Burjachs, F., Cáceres, I., Canals, A., Benito, A., Castilla y León, Valladolid, pp. 95e128. Díez, C., Lozano, M., Mateos, A., Navazo, M., Rodríguez, J., Rosell, J., Arsuaga, J.L., Fernández-Jalvo, Y., Díez, J.C., Bermúdez de Castro, J.M., Carbonell, E., Arsuaga, J.L., 2008a. The first hominin of Europe. Nature 452, 465e470. 1996. Evidence of Early cannibalism. Science 271, 277e278. Carbonell, E., Mosquera, M., Rodríguez, X.P., Bermúdez de Castro, J.M., Burjachs, F., Fernández-Jalvo, Y., Díez, C., Cáceres, I., Rosell, J., 1999. Human cannibalism in the Rosell, J., Sala, R., Vallverdú, J., 2008b. Eurasian Gates: the earliest human Early Pleistocene of Europe (Gran Dolina, Sierra de Atapuerca, Burgos, Spain). dispersals. Journal of Anthropological Research 64, 195e228. Journal of Human Evolution 37, 591e622. Carretero, J.M., Ortega, A.I., Juez, L., Pérez González, A., Arsuaga, J.L., Pérez Foley, R., Mirazón-Lahr, M., 2003. On Stony Ground: Lithic technology, human evolu- Martínez, R., Ortega, M.C., 2008. A late Pleistocene-early Holocene archaeo- tion, and the emergence of culture. Evolutionary Anthropology 12, 109e122. logical sequence of Portalón de Cueva Mayor (Sierra de Atapuerca, Burgos, Franzen, J.L., Glizzoi, E., Jellinek, T., Scholger, R., Weidenfeller, M., 2000. Die spä- Spain). Munibe: Antropología y arqueología 59, 67e80. taltpleistozäne Fossilllagerstätte Dorn-Dürkheim 3 und ihre Bedeurtung für die Carrión, J.S., Scott, L., Arribas, A., Fuentes, N., Gil-Romera, G., Montoya, E., 2007. Rekonstruction der Entwicklung des rheinischen Flusssystems. Senck- Pleistocene landscapes in central Iberia inferred from pollen analysis of hyena enbergiana Lethaea 80, 305e353. coprolites. Journal of Quaternary Science 22, 191e202. Gabarró, J.M., García Antón, M.D., Gitralt, S., Mallol, C., Sala, R., 1999. Análisis de la Carrión, J.S., Fernández, S., González-Sampériz, P., Leroy, S.A.G., Bailey, G.N., López- captación de las materias primas líticas en el conjunto técnico del Modo 2 de Sáez, J.A., Burjachs, F., Gil-Romera, G., García-Antón, M., Gil-García, M.J., Parra, I., Galería. In: Carbonell, E., Rosas, A., Díez, J.C. (Eds.), Atapuerca: ocupaciones Santos, L., López-García, P., Yll, E.I., Dupré, M., 2009. Quaternary pollen analysis humanas y paleoecología del yacimiento de Galería. Junta de Castilla y León, in the Iberian Peninsula: the value of negative results. Internet Archaeology 25, Valladolid, pp. 83e297. 1e53. http://intarch.ac.uk/journal/issue25/5/toc.html. García Antón, M., Sainz Ollero, H., 1991. Pollen record from the middle Pleistocene Chaline, J., 1972. Les rongeurs du Pléistocène moyen et supérieur de France. Cahiers Atapuerca site (Burgos, Spain). Palaeoclimatology, Palaeogeography, Palae- de Paléontologie. C.N.R.S., Paris, France. oecology 85, 199e206. Clarke, D., 1968. Analytical archaeology. Methuen, London. García-Antón, M.D., Morant, N., Mallol, C., 2002. L'approvisionnement en matières Coltorti, M., Feraud, G., Marzoli, A., Peretto, C., Ton-That, T., Voinchet, P., Bahain, J.-J., premières lithiques au Pléistocène inférieur et moyen dans la Sierra de Ata- Minelli, A., Thun Hohenstein, U., 2005. New 40Ar/39Ar, stratigraphic and puerca, Burgos (Espagne). L'Anthropologie 106, 41e55. palaeoclimatic data on the Isernia La Pineta Lower Paleolithic site, Molise, Italy. García, N., Arsuaga, J.L., 2001a. Ursus dolinensis a new species of early Pleistocene Quaternary International 131, 11e22. ursid from Trinchera Dolina, Atapuerca (Spain). C.R. Academy of Science París Cuenca-Bescós, G., García, N., 2007. Biostratigraphic sucession of the Early and Science de la Terre et des Planetes 332, 717e725. Middle Pleistocene mammal faunas of the Atapuerca cave sites (Burgos, Spain). García, N., Arsuaga, J.L., 2001b. Les carnivores (Mammalia) des sites du pleistocène Courier Forschunginstitut Senckenberg 259, 99e110. ancien et moyen d'Atapuerca (Espagne). L'Anthropologie 105, 83e93. Cuenca-Bescós, G., Canudo, J.I., Laplana, C., 1995. Los Arvicólidos (Rodentia, Mam- García, N., Arsuaga, J.L., Bermúdez de Castro, J.M., Carbonell, E., Rosas, A., Huguet, R., malia) de los niveles inferiores de Gran Dolina (Pleistoceno Inferior, Atapuerca, 2008. The Epivillafranchian carnivore Pannonictis (Mammalia, Mustelidae) from Burgos, España). Revista Española de Paleontología 10, 202e218. Sima del Elefante (Sierra de Atapuerca, Spain) and a revision of the Eurasian Cuenca-Bescós, G., Laplana Conesa, C., Canudo, J.I., Arsuaga, J.L., 1997. Small occurrences from a taxonomic perspective. Quaternary International Quaternary mammals from Sima de los Huesos. Journal of Human Evolution 33, 175e190. of Baikalia: Stratigraphy, Paleontology and Paleoenvironments of the Plioce- Cuenca-Bescós, G., Laplana, C., Canudo, J.I., 1999. Biochronological implications of neePleistocene of Transbaikalia and Interregional Correlations 179, 42e52. the Arvicolidae (Rodentia, Mammalia) from the Lower Pleistocene hominid García, N., Arsuaga, J.L., 2011. The Sima de los Huesos (Burgos, northern Spain): bearing level of Trinchera Dolina 6 (TD6, Atapuerca, Spain). Journal of Human palaeoenvironment and habitats of Homo heidelbergensis during the Middle Evolution 37, 353e373. Pleistocene. Quaternary Science Reviews 30, 1413e1419. Cuenca-Bescós, G., Canudo, J.I., Laplana, C., 2001. La séquence des rongeurs García, N., Feranec, R.S., Arsuaga, J.L., Bermúdez de Castro, J.M., Carbonell, E., 2009. (Mammalia) des sites du Pléistocene inférieur et moyen d'Atapuerca (Burgos, Isotopic analysis of the ecology of herbivores and carnivores from the Middle Espagne). L'Antrhopologie 105, 115e130. Pleistocene deposits of the Sierra De Atapuerca, northern Spain. Journal of Cuenca-Bescós, G., Rofes, J., García-Pimienta, J.C., 2005. Early Europeans and envi- Archaeological Science 36, 1142e1151. ronmental change across the EarlyeMiddle Pleistocene transition: small Geist, V., 1998. Deer of the World. Stackpole Books, Mechanicsburg. mammalian evidence fromTrinchera Dolina cave, Atapuerca, Spain. In: Gil, E., Hoyos, M., 1987. Contexto estratigráfico. In: Aguirre, E., Carbonell, E., Ber- Head, M.J., Gibbard, P.L. (Eds.), EarlyeMiddle Pleistocene Transitions: the múdez de Castro, J.M. (Eds.), El hombre fósil de Ibeas y el Pleistoceno de la LandeOcean Evidence. Geological Society of London Special Publication, vol. Sierra de Atapuerca. Junta de Castilla y León. Consejería de Cultura y Bienestar 247, pp. 277e286. Social, Valladolid, pp. 47e55. Cuenca-Bescós, G., Rofes, J., López-García, J.M., Blain, H.-A., De Marfà, R., Galindo- Girard, M., Renault-Miskovsky, J., 1969. Nouvelles techniques de préparation en Pellicena, M.A., Bennàsar-Serra, M. Ll., Melero-Rubio, M., Arsuaga, J.L., Bermú- Palynologie appliqués à trois sediments du Quaternaire final de l'Abri Cornille dez de Castro, J.M., Carbonell, E., 2009a. Biochronology of Spanish Quaternary (Istres Bouches du Rhône). Bulletin A.F.E.Q 4, 275e284. small vertebrate faunas. Quaternary International. doi:10.1016/ Goeury, Cl., de Beaulieu, J.-L.,1979. À propos de la concentration du pollen à l'aide de la j.quaint.2009.06.007. liqueur de Thoulet dans les sédiments minéraux. Pollen et Spores 21, 239e251. Cuenca-Bescós, G., Straus, L.G., González Morales, M., García Pimienta, J.C., 2009b. Guthrie, R.D., 1990. Frozen fauna of the Mammoth steppe. The story of Blue Babe. The reconstruction of past environments through small mammals: from the The University of Chicago Press, Chicago & London. Mousterian to the Bronze Age in El Miron Cave (Cantabria, Spain). Journal of Huguet, R., 2007. Primeras ocupaciones humanas en la Península Ibérica: Paleo- Archaeological Science 36, 947e955. economía en la Sierra de Atapuerca (Burgos) y la cuenca de Guadix-Baza Delson, E., 1980. Fossil Macaques, Phyletic Relationships and a Scenario of (Granada) durante el Pleistoceno inferior. Ph.D. Thesis. Universidad Rovira i Deployment. In: Lindburg, D.G. (Ed.), The Macaques: Studies in Ecology, Virgili. Tarragona, Spain. Behavior and Evolution. Van Nostrand Reinhold Comp., New York, pp. 10e30. Huguet, R., Díez, C., Rosell, J., Cáceres, I., Moreno, M.V., Ibáñez, N., Saladié, P., 2001. Dennell, R., Roebroeks, W., 1996. The earliest colonization of Europe: the short Le gisement de Galería (Sierra de Atapuerca, Burgos, Espagne): un modèle chronology revisited. Antiquity 70, 535e542. archéozoologique de gestion du territoire au Pléistocene. L'Anthropologie 105, De Marfà, R., 2008. Oryctolagus giberti n. sp. (Lagomorpha, Mammalia) du Pléis- 237e257. tocène inférieur de Cueva Victoria (Murcia, Espagne). Comptes Rendus Palevol Kahlke, R.D., 1999. The history of the origin, evolution and dispersal of the Late 7, 305e313. Pleistocene Mammuthus-Colodonta faunal complex in Eurasia (large De Menocal, P.B., 1995. Plio-Pleistocene African Climate. Science 270, 53e58. mammals). Fenske Companies, Rapid City. 1412 J. Rodríguez et al. / Quaternary Science Reviews 30 (2011) 1396e1412

Laplana, C., Cuenca-Bescós, G., 2000. Una nueva especie de Microtus (Allophaiomys) Reumer, J.W.F., 1984. Ruscinian and early Pleistocene Soricidae (Insectivora, Mam- (Arvicolidae, Rodentia, Mammalia) en el Pleistoceno Inferior de la Sierra de malia) from Tegelen (The Netherlands) and Hungary. Scripta Geologica 73, Atapuerca (Burgos, España). Revista Española de Paleontología 15, 77e87. 1e173. López-Antoñanzas, Cuenca-Bescós, G., 2002. The Gran Dolina site (Lower to Middle Rial, J.A., 1999. Pacemaking the ice ages by frequency modulation of earth orbital Pleistocene, Atapuerca, Burgos, Spain): new palaeoenvironmental data based eccentricity. Science 285, 564e568. on the distribution of small mammals. Palaeogeography, Palaeoclimatology, Rodríguez, J., 2001. Structure de la communauté de mammifères pléistocenes de Palaeoecology 186, 311e334. Gran Dolina (Sierra de Atapuerca, Burgos, Espagne). L'Anthropologie 105, van der Made, J., 1999. Ungulates from Atapuerca TD6. Journal of Human Evolution 131e157. 37, 389e414. Rodríguez, X.P., 2004a. Technical systems of lithic production in the lower and Middle van der Made, J., 2001. Les Ongulés d'Atapuerca. Stratigraphie et biogéographie. Pleistocene of the Iberian Peninsula. B.A.R. International Series 1323, 1e199. L'Anthropologie 105, 95e113. Rodríguez, J., 2004b. Stability in Pleistocene Mediterranean mammalian commu- van der Made, J., 2005. La fauna del Pleistoceno europeo. In: Carbonell, E. (Ed.), nities. Palaeogeography, Palaeoclimatology, Palaeoecology 207, 1e22. Homínidos: las primeras ocupaciones de los continentes. Ariel, Barcelona, pp. Roebroeks, W., 2001. Hominid behaviour and the earliest occupation of Europe: an 394e432. exploration. Journal of Human Evolution 41, 437e461. van der Made, J., 2011. Biogeography and climatic change as a context to human Rofes, J., Cuenca-Bescós, G., 2006. First evidence of the Soricidae (Mammalia) dispersal out of Africa and within Eurasia. Quaternary Science Reviews 30, Asoriculus gibberodon (Petényi, 1864) in the Pleistocene of North Iberia. Rivista 1353e1367. Italiana di Paleontologia e Stratigrafia112,301e315. Márquez, B., Ollé, A., Sala, R., 1999. Análisis microscópico de los instrumentos de Rofes, J., Cuenca-Bescós, G., 2009a. A new genus of red-toothed shrew (Mammalia, Galería (Sierra de Atapuerca). Actividades desarrolladas y formas de uso. In: Soricidae) from the Early Pleistocene of Gran Dolina (Atapuerca, Burgos, Spain), Carbonell, E., Rosas, A., Díez, J.C. (Eds.), Atapuerca: ocupaciones humanas y and a phylogenetic approach to the Eurasiatic Soricinae. Zoological Journal of paleoecología del yacimiento de Galería. Junta de Castilla y León, Valladolid, pp. the Linnean Society 155, 904e925. 353e370. Rofes, J., Cuenca-Bescós, G., 2009b. First record of Beremendia fissidens (Mammalia, Márquez, B., Ollé, A., Sala, R., Vergès, J.M., 2001. Perspectives méthodologiques de Soricidae) in the Pleistocene of the Iberian Peninsula, with a review of the l'analyse fonctionnelle des ensembles lithiques du Pléistocène inférieur et biostratigraphy, biogeography and palaeoecology of the species. Comptes moyen d'Atapuerca (Burgos, Espagne). L'Anthropologie 105, 281e299. Rendus Palevol 8, 21e37. van der Meulen, A., 1973. Middle Pleistocene smaller mammals from the Monte Rosas, A., Bermúdez de Castro, J.M., 1998. The Mauer mandible and the significance Peglia (Orvieto, Italy), with special reference to the phylogeny of Microtus of Homo heidelbergensis. Geobios 31, 126e137. (Arvicolidae, Rodentia). Quaternaria 17, 1e144. Rosas, A., Artusi, E., Cuenca, G., Laplana, C., Rodríguez, J., 1999. Asociaciones de Minwer-Barakat, R., 2005. Roedores e insectívoros del Turoliense superior y el microvertebrados en el yacimiento mesopleistoceno de Galería, Sierra de Ata- Plioceno del sector central de la Cuenca de Guadix. PhD thesis, University of puerca. In: Carbonell, E., Rosas, A., Díez, J.C. (Eds.), Atapuerca: ocupaciones Granada, Granada, Spain. humanas y paleoecología del yacimiento de Galería. Junta de Castilla y León, Minwer-Barakat, R., García-Alix, A., Freudenthal, M., 2007. Blarinoides aliciae sp. Valladolid, pp. 129e134. nov., a new Soricidae (Mammalia, Lipotyphla) from the Pliocene of Spain. Rosas, A., Huguet, R., Pérez-González, A., Carbonell, E., Bermúdez de Castro, J.M., Comptes Rendus Palevol 6, 281e289. Vellverdú, J., Van der Made, J., Allué, E., García, N., Martínez-Pérez, R., Sala, R., Mitchell-Jones, A.J., Amori, G., Bogdanowicz, W., Krystufek, B., Reijnders, P.J.H., Saladié, P., Benito, A., Martínez-Maza, C., Bastir, M., Sánchez, A., Parés, J.M., Spitzenberger, F., Stubbe, M., Thissen, J.B.M., Vohralík, V., Zima, J., 1999. The atlas 2006. The “sima del Elefante” cave site at Atapuerca (Spain). Estudios Geo- of European mammals. Academic Press, London-San Diego. lógicos 62, 327e348. Mosquera, M., 1998. Differential raw material use in the Middle Pleistocene of Rosell, J., 1998. Les premières occupations humaines à la Sierra de Atapuerca Spain: the evidence from Sierra de Atapuerca, Torralba, Ambrona and Aridos. (Burgos, Espagne): les niveaux TDW-4 et TDW-4b. Économie Préhistorique: les Cambridge Archaeological Journal 8 (1), 15e28. comportements de subsistance au Paléolithique. XVIIIe. Rencontres Inter- Ollé, A., 2003. Variabilitat i patrons funcionals en els sistemes tècnics de Mode 2. nationales d'Archéologie et d'Histoire d'Antibes. Sophia Antipolis, France, pp. Anàlisi de les deformacions d'ús en els conjunts lítics del Riparo Esterno de 153e162. Grotta Paglicci (Rignano Garganico, Foggia), Áridos (Arganda, Madrid) i Galería- Rosell, J., 2001. Patrons d'Aprofitament de les Biomasses Animals durant el Pleistocè TN (Sierra de Atapuerca, Burgos). Ph.D. Thesis. Universidad Rovira i Virgili, Inferior i Mig (Sierra de Atapuerca, Burgos) i Superior (Abric Romaní, Tarragona, Spain. Barcelona). Ph.D. Thesis. Universitat Rovira i Virgili, Tarragona, Spain. Ollé, A., Cáceres, I., Vergès, J.M., 2005. Human occupations at Galeria site (Sierra de Ruiz Zapata, M.B., Dorado, M., Valdeolmillos, A., Gil García, M.J., Martín Arroyo, T., Atapuerca, Burgos, Spain) after the technological and taphonomical data. In: Pérez González, A., 2004. Registro paleoambiental y paleoclimático del Pleis- Molines, N., Moncel, M.-H., Monnier, J.-L. (Eds.), Les premiers peuplements en toceno medio y superior en depósitos fluviales del Valle del río Tajo (Toledo). In: Europe. Colloque international: Données récentes sur les modalités de Miscelanea en Homenaje a Emiliano Aguirre. Paleontología, Museo Arqueoló- peuplement et sur le cadre chronostratigraphique, géologique et paléo- gico Regional, Alcalá de Henares, vol. II, pp. 507e518. géographique des industries du Paléolithique ancien et moyen en Europe Saladié, P., 2009. Mordiscos de omnívoros. Aproximación experimental y aplicación British Archaeological Series, International Series, S1364. John and Erika Hedges zooarqueológica en la Sierra de Atapuerca. Ph. D. Thesis. Universidad Rovira i Ltd, Oxford, pp. 269e280. Virgili, Tarragona. Spain. Oms, O., Parés, J.M., Martínez-Navarro, B., Agustí, J., Toro, I., Martínez-Fernández, G., Sánchez-Marco, A., 1999. Implications of the avifauna for palaeoecology in the Turq, A., 2000. Early human occupation of Western Europe: paleomagnetic early Pleistocene of the Iberian Peninsula. Journal of Human Evolution 37, dates for two paleolithic sites in Spain. Proceedings of National Academy of 375e388. Sciences of United States of America 97, 10666e10670. Sittler, C., 1955. Méthodes et tecniques physicho-chimiques de préparation des Palombo, J.L., Gisbert, J., 2005. Atlas de los Mamíferos Terrestres de España. sédiments en vue de leur analyse pollinique. Rev. Inst. Fr. Petrol., Annales des Dirección General para la Biodiversidad-SECEM-SECEMU, Madrid. Comb. Liq. 10, 103e114. Parés, J.M., Pérez-González, A., 1995. Paleomagnetic Age for Hominid fossils at Velasco, J.C., Lianza, M., Román, J., Delibes de Castro, M., Fernández, J., 2005. Guía de Atapuerca Archaeological site. Nature 269, 830e832. los Peces, Anfibios, Reptiles y Mamíferos de Castilla y León. Náyade, Valladolid. Parés, J.M., Pérez-Gozález, A., 1999. Magnetochronology and stratigraphy at Gran Vergés, J.M., Allué, E., Angelucci, D., Burjachs, F., Carrancho, A., Cebrià, A., Dolina section, Atapuerca (Burgos, Spain). Journal of Human Evolution 37, Expósito, I., Fontanals, M., Moral, S., Rodríguez, A., Vaquero, M., 2008. Los 325e342. niveles neolíticos de la cueva de El Mirador (Sierra de Atapuerca, Burgos): Pérez-González, A., Parés, J.M., Gallardo, J., Aleixandre, T., Ortega, A.I., Pinilla, A., nuevos datos sobre la implantación y el desarrollo de la economía agropecuaria 1999. Geología y estratigrafía del relleno de Galería de la Sierra de Atapuerca, en la submeseta norte. Hernández, M., Soler, Actas del IV Congreso del Neolítico (Burgos). In: Carbonell, E., Rosas, A., Díez, J.C. (Eds.), Atapuerca: ocupaciones Peninsular. Museo Arqueológico de Alicante, Alicante, Spain. pp. 418e427. humanas y paleoecología del yacimiento de Galería. Junta de Castilla y Leon, Vrba, E.S., Denton, G.H., Partridge, T.C., Burckle, L.H. (Eds.), 1995. Paleoclimate and Valladolid, pp. 31e42. evolution, with emphasis on human origins. Yale University Press, New Haven, CT. Pérez-González, A., Parés, J.M., Carbonell, E., Aleixandre, T., Ortega, A.I., Benito, A., Whittaker, R.J., Nogues-Bravo, D., Araújo, M.B., 2007. Geographical gradients of Martín Merino, M.A., 2001. Géologie de la Sierra de Atapuerca et stratigraphie species richness: a test of the water-energy conjecture of hawkins et al. (2003) des remplissages karstiques de Galería et Dolina (Burgos, Spain). L'An- using European data for five taxa. Global Ecology & Biogeography 16, 76e89. thropologie 105, 27e44. Yokoyama, Y., Bibron, R., Falguères, C., 1988. Datation absolue desPlanchers sta- Pleguezuelos, J.M., Márquez, M., Lizana, M., 2002. Atlas y libro rojo de los Anfibios y lagmitiques de la grotte du Vallonnet à Roquebrune-Cap-martin (Alpes-Mari- Reptiles de España. Dirección General de Conservación de la Naturaleza e times) France, par larésonance de spin électronique (ESR). L'Anthropologie Asociación Herpetológica Española, Madrid. (Paris) 92, 429e436.