Santimamiñe q i xxx (2013)

Quaternary International xxx (2013) 1e14

Contents lists available at SciVerse ScienceDirect

Quaternary International journal homepage: www.elsevier.com/locate/quaint

The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene Juan Rofes a, *, Xabier Murelaga a, Blanca Martínez-García a, Salvador Bailon b, Juan Carlos López-Quintana c, Amagoia Guenaga-Lizasu c, Luis Ángel Ortega d, María Cruz Zuluaga d, Ainhoa Alonso-Olazabal d, Jone Castaños a, Pedro Castaños e a Universidad del País Vasco UPV-EHU, Facultad de Ciencia y Tecnología, Departamento de Estratigrafía y Paleontología, Apartado 644, E-48080 Bilbao, Spain b UMR 7209-UMR 7149, CNRS Département Ecologie et Gestion de la Biodiversité (EGB), MNHN Bâtiment d’Anatomie Comparée CP55, 55 rue Buffon, 75005 Paris, France c AGIRI Arkeologia Kultura Elkartea, 208 Postakutxa, 48300 Gernika-Lumo, Spain d Universidad de País Vasco UPV-EHU, Facultad de Ciencia y Tecnología, Departamento de Mineralogía y Petrología, Apartado 644, E-48080 Bilbao, Spain e Sociedad de Ciencias Aranzadi, Zorroagagaina 11, 20014 Donostia-San Sebastián, Spain

a r t i c l e i n f o

a b s t r a c t

Article history: Available online xxx

The cave of Santimamiñe (Kortezubi, Bizkaia, Spain), on the southern slopes of the Ereñozar Mountain (Urdaibai biosphere reserve), is one of the most famous prehistoric localities of the Cantabrian range.

Between 2004 and 2006, a test trench revealed a 6 m-deep stratigraphic sequence in the inner vestibule of the cave covering w20,000 years, from the latest Late Pleistocene (MIS 2) to the middle Holocene (MIS 1). It comprises six chrono-cultural units: lower, middle and upper Magdalenian, Azilian, Neolithic, and Chalcolithic/Bronze Age, plus seven lower purely palaeontological levels. More than 47,000 microvertebrate elements (including mammals, birds, reptiles, amphibians and fishes) were recovered, of which 1587 were identified to the genus and/or species levels. Of these, small mammals were used for paleoenvironmental reconstruction since they are very sensitive to climatic conditions and their distributions over time, measured in terms of relative abundance, serve as reliable proxies of habitat and climatic change. The reconstruction of Santimamiñe past environments based on small mammals roughly coincides with other proxies, such as the amphibian/reptile, large mammal/bird, palynological, and cryoclastic records on the local scale; other paleoenvironmental reconstructions from north Iberia on the regional scale; and the oxygen isotopic curve of an ice core from central Greenland on the global scale. All Magdalenian levels from Santimamiñe were deposited during the Oldest Dryas, a mostly cold and humid period including the Heinrich event 1. Small mammalian, palynological and cryoclastic evidence likely document the shift from the cold Younger Dryas to the beginning of the Holocene Climatic Optimum during Azilian times. The uppermost level, deposited during a moderate re-opening of the landscape at the Subboreal, slightly postdates the Holocene Cooling Event 3. Ó 2013 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction

first cave paintings of the Basque Country were discovered by F. J. Bengoechea and other Gernika villagers in 1916 (FernándezEraso, 2011). Next year, H. Breuil discovered some new engravings and defined the main graphic units of the complex (LópezQuintana and Guenaga-Lizasu, 2011). The systematic excavation of Santimamiñe was the first one organized by a multidisciplinary team in the Basque Country, being conducted between 1918 and 1926 by the well-known archaeologists T. de Aranzadi, J. M. de Barandiarán and E. de Eguren (Aranzadi et al., 1925, 1931; Aranzadi

The cave of Santimamiñe (Kortezubi, Bizkaia) is one of the most famous prehistoric localities of the Cantabrian range, where the

* Corresponding author. Universidad del País Vasco UPV-EHU, Facultad de Ciencia y Tecnología, Departamento de Estratigrafía y Paleontología, Sarriena s/n, 48940 Leioa, Bizkaia, Spain.

E-mail address: juan.rofes@ehu.es (J. Rofes).

1040-6182/$ e see front matter Ó 2013 Elsevier Ltd and INQUA. All rights reserved. http://dx.doi.org/10.1016/j.quaint.2013.05.048

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

2

J. Rofes et al. / Quaternary International xxx (2013) 1e14

and Barandiarán, 1935). A second phase of excavation, directed by J. M. Barandiarán, took place between 1960 and 1962 (Barandiarán, 1962a, 1962b, 1962c).

In the 8 m deep archaeological trench excavated in the entrance of the cave by Aranzadi, Barandiarán and Eguren, a long stratigraphic sequence ranging from the Aurignacian to the roman period was unveiled (López-Quintana and Guenaga-Lizasu, 2011). A comprehensive synthesis of this sequence and its materials was published by I. Barandiarán (1988: 343e348).

Posed as a stratigraphic revision of all the previous work done in Santimamiñe, a new phase of excavation started in 2004 under the co-direction of J. C. López-Quintana and A. Guenaga-Lizasu (LópezQuintana and Guenaga-Lizasu, 2011). This third phase began with a test trench carried out between 2004 and 2006 (López-Quintana and Guenaga-Lizasu, 2007) and it continues up to the present with an excavation in zone taking the test trench profile as guide (López-Quintana and Guenaga-Lizasu, 2011).

The test trench was accomplished in the inner vestibule, some 20 m from the entrance of the cave, and it revealed a stratigraphic sequence, although not as complete as the one of the entrance, long enough to cover w20 millennia of history from the latest Late Pleistocene to the middle Holocene. Together with remains of large mammals (including humans), birds, molluscs, fishes, botanical evidence, and thousands of pieces of hand-made artefacts (lithic, bone, pottery), the sequence has yielded a rich sample of small vertebrates, with more than forty seven thousand items.

Digestion sub-products of birds of prey (e.g. owl pellets) and small carnivores are the main sources for small vertebrate deposition in archaeological sites (Andrews, 1990), and caves and rock shelters are particularly propitious for those animals to nest at the entrances, or to build burrows inside, respectively (Cuenca-Bescós et al., 2009; Rofes et al., 2013). Unlike large mammal remains (many of them product of human selection), microvertebrate accumulations reasonably well reflect local biocenosis, despite unavoidable biases due to specific predators (Andrews, 1990).

Moreover, small mammals are particularly sensitive to climatic and habitat changes, and their shifts along time in terms of species and

number of specimens can be successfully used for the reconstruction of past environments (e.g. Bertolini et al., 1996; Pokines, 1998; Repenning, 2001; Cuenca-Bescós et al., 2005, 2009, 2012; Sesé, 2005; López-García et al., 2010, 2011a,b, 2012; Rofes et al., 2013).

By distributing the different small mammalian species (in terms of relative abundance) according to their current habitat and climate requirements, and plotting them against time, the aim of this paper is to propose a reconstruction of past environments of the landscape around the cave of Santimamiñe, and to compare and correlate the results with other paleoenvironmental records on the local, regional and global scales.

2. The site The cave of Santimamiñe (Kortezubi, Bizkaia) is located in the eastern bank of the Oka basin (Urdaibai Biosphere Reserve), more specifically on the southern slopes of the Ereñozar Mountain, in a strategic position over the Mundaka estuary and marshland (Fig. 1).

Geologically, it opens on a karst reef limestone substrate of Aptian/ Albian (Lower Cretaceous) origin. The substrate includes corals and rudists conferring the rock a massive appearance (López-Quintana and Gunega-Lizasu, 2011).

The entrance of the cave, 137 m above present sea-level (a.s.l.), takes into a wide gallery that contains a long stratigraphic sequence covering nearly 20,000 years from the late Upper Palaeolithic to the Chalcolithic/Bronze Age transition. The deposit of the test trench is 6 m deep and it was divided into 27 archaeological levels which can be grouped into six chrono-cultural units: lower, middle and upper Magdalenian, Azilian, Neolithic, and Chalcolithic/Bronze Age, plus seven lowermost purely palaeontological levels and a thin hearthlayer of Mesolithic filiations (López-Quintana, 2011a).

As the lithology and archaeo-palaeontological contents of the different units have been extensively described elsewhere (LópezQuintana, 2011b), in Table 1 we present only a synopsis of the chronology and cultural filiations of those levels being quantitatively and qualitatively meaningful in terms of their small vertebrate contents, which are, from bottom to top: Arg-o, Csn-Camr,

Fig. 1. Location of the Santimamiñe cave in the Iberian Peninsula (up), and near the villages of Arteaga and Kortezubi in the Urdaibai Biosphere Reserve, Bizkaia, Spain (down).

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

J. Rofes et al. / Quaternary International xxx (2013) 1e14

Balm, Almp, Slnc, Arcp, Slm, and Lsm (Fig. 2). The cultural filiations of the successive levels were established on basis of their different lithic, bone and/or pottery industries.

3

Santimamiñe assemblage is not representative of the ecosystem in the surroundings of the cave at the time when the remains were deposited.

Table 1 List of radiocarbon ages from the Santimamiñe test trench including cultural periods, chrono-stratigraphic units, laboratory codes and the elements from where the samples were taken. Dates were calibrated at 95.4% confidence intervals (2s) using the IntCal09 data set (Reimer et al., 2009) and OxCal4.1.7 calibration software (Bronk Ramsey, 2009).

Mean probability calculated with the Calib6.1.1 (Stuvier et al., 2013). *Not cultural.

Cultural period

Level/sublevel

Radiocarbon age 14

C yr BP

Chalcolithic/Bronze Age Neolithic Azilian Upper Magdalenian Middle/upper Magdalenian? * Lower Magdalenian Bear den*

Lsm 4 Slm 6/7 Arcp 15 Slnc 20 Almp Balm Csn-Camr 41 Arg-o 75

3710 5010 10,100 12,790

40 40 60 70

14,670 70 20,530 110

3. Material and methods 3.1. Collecting techniques To obtain the small vertebrate sample, a total of 123 samples of sediment (0.33 m2 each) were alternatively taken from five 1 m2 excavation quadrats (Fig. 3) along the entire 6 m stratigraphic sequence of the test trench (Fig. 2). All stratigraphic units (arbitrary sublevels) were sampled with the exception of sublevels 11 and 13 (stalagmite flows: T3 and T2 in Fig. 2), and sublevel 12 (Mesolithic hearth: H-Sln in Fig. 2). A minimum of one quadrat by sublevel was sampled; being most of the 96 sublevels 5 cm deep.

The sediment was dried and weighted (to estimate the amount of remains by kg of sediment), and then washed and sieved using 4 mm- and 0.5 mm-mesh sieves. The small vertebrates were collected from residue coarser than 0.5 mm. Fossils were sorted, classified, counted, and studied with the aid of a binocular microscope (Nikon SMZ-U; 7 , 20 , and 40 magnifications). Most elements are teeth, isolated mandibles, skull fragments, and postcranial bones.

3.2. Taphonomic remarks The accumulation of small vertebrate remains in karstic cavities is usually due to biologic agents, commonly birds of prey which tend to accumulate their pellets near the entrance of the caves (Andrews, 1990). Those pellets are then transported inside by gravity or streams and deposited. Prior to deposition, the remains may suffer some degree of fragmentation which can be more or less intense depending on the size of the particles transported with them and the distance to the zone of deposition. Trampling (i.e. medium- to large-sized mammals, including humans) could also severely affect the remains. A detailed taphonomic study (Murelaga et al., 2011: 298e299), level by level, was carried out to evaluate the effects of all these factors in the samples. As a result, some samples presenting biases (e.g. anomalous quantities of remains) were not considered for the paleoenvironmental reconstructions.

The light to moderate digestion observed in the arvicoline teeth, indicates the bones were likely accumulated by an avian predator of category 1 (sensu Andrews, 1990) such as a barn owl (Tyto alba), which is an opportunistic rather than a selective hunter. We are not able to specify the exact predator responsible for the accumulation, but there are no signs of alteration suggesting that the

Lab code

Sample

4047 5746 11,695 15,202

Beta-240896 Beta-240897 Beta-240900 Beta-240902

Human mandible Charcoal Red deer bone Red deer bone

17,846 24,507

Beta-240904 Beta-240906

Red deer bone Red deer bone

Cal yr BP 2s

Mean prob.

4220e3926 5893e5653 11,979e11,396 15,856e14,785

18,443e17,528 24,956e24,170

The fossil remains of bats were likely accumulated from in situ mortality, although occasional predation by nocturnal birds of prey should not be discarded. In situ accumulation may also account for some amphibians and reptiles that at least temporarily frequent the karstic cavities (e.g. during wintering and/or aestivation).

3.3. Systematic attribution and quantification A total of 47,093 fossil items were recovered, of which 14,423 were identified to the order and/or family levels, and 1587 to the genus and/or species levels following the general criteria of smallvertebrate palaeontology. Specific attributions rest mainly on diagnostic elements: first lower molars for the Arvicolinae; mandibles, maxillas and isolated teeth for the Soricidae, Chiroptera and Lagomorpha; mandibles and post-cranial skeleton for the Talpidae; isolated teeth for the Gliridae and Murinae; skull elements for the Lacertidae; mandibles, vertebrae and osteoderms for the Anguidae; trunk vertebrae for the Colubridae, and cranial and post-cranial skeleton for the amphibians. The taxonomic classification for small mammals follows Wilson and Reeder (2005), and that for amphibians and reptiles follows Speybroeck and Crochet (2007).

Given the taphonomic nature and composition of the assemblage, the minimal number of individuals (MNI) per species should be a reasonably good quantitative measure to reconstruct past environments. The MNI of the small vertebrate record was calculated based on the amounts of a specific diagnostic tooth (e.g. first lower molar in arvicolines) or some equally identifiable cranial or postcranial element for small mammals (rodents, insectivores and chiropterans), reptiles, and amphibians. The low number and/or lack of precise taxonomical (i.e. to the species level) and habitat assignations of reptile and amphibian elements from Santimamiñe turns them unsuitable for long paleoenvironmental reconstructions, even though they can be used as complement in some cases.

There is classical debate on whether the NISP (number of identified specimens) or the MNI (minimal number of individuals) is a better method to quantify a given sample (Grayson, 1984). Both measures have advantages and drawbacks (Marshall and Pilgram, 1993), but the most frequently used among scholars dealing with small mammals and paleoenvironmental reconstructions is the MNI (Pokines, 1998; Cuenca-Bescós et al., 2005, 2009, 2011; LópezGarcía et al., 2010, 2011a, 2011b, 2012; Rofes et al., 2013), given the inability in attributing some cranial and post-cranial specimens to a specific taxon, i.e. the unrooted teeth different from m1 in mediumsized arvicolines.

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

4

J. Rofes et al. / Quaternary International xxx (2013) 1e14

Fig. 2. Stratigraphic sequence of Santimamiñe test trench (Kortezubi, Bizkaia, Spain).

3.4. Habitat types and climatic categories Following Cuenca-Bescós et al. (2009) and Rofes et al. (2013), we assigned the small mammals from Santimamiñe to five habitat types based on their environmental affinities. Being a site with a relatively recent chronology, all the Santimamiñe fauna is extant, which means that there is little doubt in the species-habitat correlations. Nevertheless, this classification lacks sharp boundaries, and, as with many other natural borders, transitions are gradual (ecotones). The habitat types are described below.

1. Forest: woodland, scrub forest and woodland edges. Mature forest including woodland margins and patchy forest with moderate ground cover.

2. Humid meadow: evergreen meadows with dense pastures and suitable topsoil. It indicates humid conditions.

3. Grassland: or open meadow; that is, meadows under seasonal climatic change. It indicates relatively dry conditions.

4. Rocky: highland and/or alpine, comprising species living in steppe grasslands with rocky substrates, usually above timberline.

5. Water: streams, lakes, ponds and marshes. It indicates abundant superficial water, either running or stagnant.

To evaluate the climatic requirements of the small mammals from Santimamiñe, we follow Rofes et al. (2013) and divide our taxa in two major categories, which are detailed as follows: 1. Cold: It ranges from the severe, steppe climatic conditions typical of the glacial advances (if dominant) to periods of cool or temperate weather (if not dominant). Rocky and humid meadow habitats are, respectively, characteristic of this category.

2. Warm: If dominant, it indicates warm, Mediterranean conditions. Wooded landscapes and grasslands are typical of this climatic category.

Data on the habitat and climatic preferences of extant species were taken from Janeau and Aulagnier (1997), Pokines (1998), Cuenca-Bescós et al. (2005, 2009), Palomo and Gisbert (2005), Sesé (2005), López-García et al. (2011a, 2012), and Rofes and Cuenca-Bescós (2011).

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

5

X X X X

X

57 255 422 8 6 a

19 6 8 16 3 44

5 5 2

NISP estimated by extrapolating the MNI proportions to the sample of indeterminate small-sized arvicoline teeth.

14 21 3 54 131 269 9 17 52 78 5 42

7 65 43 1 6 5 2 9 9 2 8 6 2 13 8

2 5 4 1 64 107 8

2 3 2 1

4 4 7 1 140 187 10

3 4 5 1

193 245 9

3 4 6 23 8

1 3 3 1 1 1 1

1 2 7 1 3 26 1 4 1 6 1 4 20 1 2

28

5 111 9

2 4 5 10 4 11 6

1 1 5 1

1 1 1 5 4 1 1 2 8 16

5 1

3 3

Arvicola amphibius 2 Arvicola cf. sapidus 1 M. (Terricola) lusitanicusa 1 Chionomys nivalisa M. (Alexandromys) oeconomusa 1 M. (Microtus) gr. agrestisearvalisa 3 Apodemus sylvaticus 106 Eliomys quercinus 1 Glis glis 6 Lepus sp.

Erinaceus europaeus Sorex minutus Sorex gr. araneusecoronatus 8 Talpa europaea 21 Myotis sp.

5 Microtines 61 Totals 216 N of species 11

2 1 1

3

3

2 3 6 2 4 11 2

3 4

2

2

1

1

3 2 4 29 4

2 24 3

2 2 3 18 3

5 5

1

16 17 1

1

1

1

2 3 1

1

1

X

X

X

X X

X

X

X X

X

X X X X

X X

X X X

Ro HM Gr Wa Fo Warm Cold Lsm-Sa Arb-o Arp-Sa Arg-o FC Csn-Camr Balm Almp Slnc Arcp Slm Lsm Level

NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI NISP MNI

Climate Habitat * * * Bear den* * L Mag * M/U Mag? U Mag

After calculating the NISP (1587) and MNI (275) of the different species of small mammals, the species were ordered by stratigraphic levels covering the lower, middle and upper Magdalenian, Azilian, Neolithic and Chalcolithic/Bronze periods (Csn-Camr to Lsm), plus the seven lowermost purely paleontological layers (LsmSa to Lsr-Ap). All these data plus the habitat and climatic preferences of the currently living species found in Santimamiñe are summarized in Table 2.

There are some levels both below and above the Arg-o unit that have not been considered for the paleoenvironmental reconstructions given their exiguous contents of small vertebrates in spite of their remarkable thickness (López-Quintana and GuenagaLizasu, 2011; Murelaga et al., 2011). Those levels are: Lsm-Sa, Arb-o and Arp, all three below Arg-o; and the “flooding complex” (comprising Avp-Sj, Arp-Sa, and Lsr-Ap), stratigraphically placed between Arg-o and Csn-Camr (Fig. 2). All these levels coincide in having no valid pollen samples available (Iriarte, 2011; LópezQuintana and Guenaga-Lizasu, 2011).

Five elements of Lepus sp. were identified in the sample, specifically in levels Csn-Camr, Almp and Arcp. Hares are not included in the habitat and climate reconstructions because, unlike other small mammals, they do not provide precise paleoenvironmental information and their presence at the site (or at least part of it) could be associated to human consumption. Due to their strong relation to the substrate, Cuenca-Bescós et al. (2009) do not include chiropterans in the habitat and climate reconstructions, but other authors use them (e.g. López-García et al., 2011a, 2011b, 2012). Here we leave bats aside mostly due to their scarcity and lack of precise taxonomic assignations.

There is a moderate proportion (30.6%) of identifiable microvertebrate elements in Santimamiñe. Even though, the preservation of small mammal bones (showing low digestion traces and moderate breakage) suggests that they accumulated naturally, mainly due to nocturnal birds of prey. Therefore, the small mammalian assemblage should be a feasible means of reconstructing at least part of the biocenosis around Santimamiñe, and small mammals serve as proxies of habitat and climate change due to their sensitivity to environmental conditions (Bertolini et al.,

Azilian

4.2. Small mammals and paleoenvironmental reconstructions

Neolithic

The small terrestrial vertebrate fauna from Santimamiñe and the ecological preferences of the different species were extensively documented elsewhere (Murelaga et al., 2011). Therefore, here we limit to synthesize the information besides adding some previously omitted species. The small vertebrates from Santimamiñe comprise 20 taxa: two soricids (Sorex minutus and Sorex gr. araneusecoronatus); one talpid (Talpa europaea); one erinaceid (Erinaceus europaeus); two glirids (Glis glis and Eliomys quercinus); one murid (Apodemus sylvaticus); six cricetids (Microtus (Microtus) gr. arvalis-agrestis, M. (Terricola) lusitanicus, Chionomys nivalis, Microtus (Alexandromys) oeconomus, Arvicola amphibius, and Arvicola cf. sapidus); one chiropteran (Myotis sp.); one leporid (Lepus sp.); two amphibians (Rana gr. temporariaeiberica and Epidalea calamita); and three reptiles (Lacertidae indet., Anguis fragilis, and Vipera sp.). Fig. 4 shows selected specimens of nearly all the small vertebrates recovered from Santimamiñe plus one large mammal (Cervus elaphus). In spite that in Fig. 4 we show two discernible molars of M. (M.) arvalis and M. (M.) agrestis (Fig. 4.6 and 4.7, respectively), most elements of the sample exhibit intermediate morphologies.

Chal/Bron

4.1. Small vertebrate assemblage

Cultural period

4. Results

Table 2 Number of Identified Specimens (NISP) and Minimum Number of Individuals (MNI) of the small mammal species from Santimamiñe ordered by cultural periods and chrono-stratigraphic units. Right: Small-mammal distribution by habitat and climate. Chal/Bron, Chalcolithic/Bronze Age; Mag, Magdalenian; U, Upper; M/U, Middle/Upper; L, Lower; FC, Flooding Complex; Ro, Rocky; HM, Humid Meadow; Gr, Grassland; Wa, Water; Fo, Forest. *Not cultural.

J. Rofes et al. / Quaternary International xxx (2013) 1e14

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

6

J. Rofes et al. / Quaternary International xxx (2013) 1e14

Fig. 3. Plan view of the entrance and vestibule of the Santimamiñe cave (Kortezubi, Bizkaia, Spain). Shaded quadrats are the area from where the small vertebrates were obtained.

1996; Pokines, 1998; Repenning, 2001; Cuenca-Bescós et al., 2005, 2009; Sesé, 2005; López-García et al., 2010, 2011a, 2011b, 2012; Rofes et al., 2013).

The number of taxa and the MNI percentage per species (relative abundance) are both measures of information abstracted from habitats, in that habitats with extreme climates and poor vegetation variety tend to be dominated by small numbers of species with high amounts of individuals, whereas more equable habitats with complex vegetation have larger number of species but no single species is dominant; thus, landscape complexity is correlated with high specific diversity and climatic equitability (Cuenca-Bescós et al., 2009, 2011).

4.3. Small mammalian community changes over time Fig. 5 (based on data from Table 2) shows the stratigraphic and quantitative distribution of the Santimamiñe small mammals.

Following the stratigraphic sequence from bottom to top, Level Lsm-Sa has only one identifiable item of A. sylvaticus on it, and the overlying Arb-o, only one of M. (M.) gr. agrestisearvalis. The next, Arp-Sa, shows no single identifiable element.

Level Arg-o is the first one with a good representation of small mammals. The association is composed, in descending order of relative abundance (as defined above), most of all by S. gr. araneuse coronatus and M. (M.) gr. agrestisearvalis, and, to a lesser extent, by M. (A.) oeconomus, A. sylvaticus, T. europaea, M. (T.) lusitanicus,

C. nivalis, and S. minutus. Overlying Arg-o is the flooding complex, comprising Levels Avp-Sj, Arp-Sa and Lsr-Ap. Only the uppermost (Lsr-Ap) has yielded identifiable small mammal remains which are derisory (NMI ¼ 6) for a 230e260 cm thick unit. The identified species, in order of abundance, are: S. gr. araneusecoronatus, M. (M.) gr. agrestisearvalis, and C. nivalis.

In Level Csn-Camr, the most abundant taxon is by far S. gr. araneusecoronatus, its relative abundance being similar to that of Arg-o. Second in abundance is T. europaea, closely followed by S. minutus and M. (M.) gr. agrestisearvalis, the latter one considerably dropping in representation respect to Arg-o, whereas T. europaea and S. minutus rise. Relative abundance of M. (T.) lusitanicus, M. (A.) oeconomus and A. sylvaticus is almost the same as in Arg-o. A. amphibius and Lepus sp. first appear in this level, and C. nivalis disappears.

Level Balm shows a general fall of both the number of species and the number of specimens present in the fossil record (Table 2).

Relative abundances of M. (M.) agrestisearvalis and T. europaea rises and those of S. minutus and S. gr. araneusecoronatus drop. Chionomys nivalis reappears but A. amphibius, M. (T.) lusitanicus, M. (A.) oecononus, A. sylvaticus, and Lepus disappear from the record. In Level Almp, the general number of taxa rises again: A. amphibius, M. (T.) lusitanicus, A. sylvaticus, and Lepus reappear. Despite most of the MNIs rise (Table 1), the relative abundances of C. nivalis, M. (M.) gr. agrestisearvalis, S. minutus, S. gr. araneusecoronatus, and T. europaea fall.

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

J. Rofes et al. / Quaternary International xxx (2013) 1e14

7

Fig. 4. Selected specimens of vertebrates from Santimamiñe. Eliomys quercinus 1 left m1; Glis glis 2 right m1 or m2; Apodemus sylvaticus 3 right M1eM2eM3; Arvicola amphibius 4 left m1; Microtus (Terricola) lusitanicus 5 right m1; Microtus arvalis 6 right m1; Microtus agrestis 7 right m1; Chionomys nivalis 8 right m1; Microtus (Alexandromys) oeconomus 9 left m1; Sorex gr. araneusecoronatus 10 right mandible in lateral view plus condyle in posterior view; Sorex minutus 11 incomplete left mandible with m1em2em3 in lateral view plus condyle in posterior view; Myotis sp. 12 incomplete left mandible with m2em3 in lateral view; Anguis fragilis 13 left dentary in medial view; 14 osteoderm; Vipera sp. 15 trunk vertebrae in lateral view; Lacertidae indet. 16 incomplete dentigerous in medial view; Rana gr. temporariaeiberica 17 sacral vertebrae in posterior view; 18 right ilium in lateral view; Epidalea calamita 19 left ilium in lateral view; Cervus elaphus 20 incomplete right maxilla. Specimens 1e9 are in occlusal view.

In Slnc, M. (A.) oeconomus reappears and the first identifiable element of Myotis sp. is recorded. The relative abundances of A. amphibius, C. nivalis, A. sylvaticus, S. minutus, and T. europaea rise, and that of M. (T.) lusitanicus, M. (M.) gr. agrestisearvalis, and S. gr. araneusecoronatus drop. Level Arcp exhibits a new decrease in the number of species, but not as pronounced as in Balm: A. amphibius, M. (T.) lusitanicus, and C. nivalis disappear but Lepus reappears. The record begins to be dominated by A. sylvaticus whose relative abundance rises together with those of M. (A.) oeconomus, S. gr. araneusecoronatus, and Myotis. Microtus (M.) gr. agrestisearvalis and T. europaea fall and S. minutus remains the same.

Especially meaningful is that six taxa disappear from the record in Level Slm: M. (A.) oeconomus, Lepus, S. minutus, S. gr. araneuse coronatus, T. europaea, and Myotis. The only item of E. europaeus from the entire sequence comes from this level which is clearly ruled by A. sylvaticus. In the uppermost level, Lsm, three species appear for the first time: Arvicola cf. sapidus, E. quercinus, and G. glis, and six more reappear: A. amphibius, M. (T.) lusitanicus, M. (A.) oeconomus, Sorex gr. araneusecoronatus, T. europaea, and Myotis.

The record is once again dominated by far by A. sylvaticus.

As shown in Fig. 5, the dominant species from Level Arg-o to Level Balm in terms of relative abundance is Sorex gr. araneuse

coronatus. Microtus (M.) gr. agrestisearvalis predominates in Levels Almp and Slnc, and A. sylvaticus in Levels Arp to Lsm. However, in terms of its presence throughout the sequence, the most stable taxon is without any doubt M. (M.) gr. agrestisearvalis, which is present from Arb-o to the uppermost level. Second species in stability is Sorex gr. araneusecoronatus, followed by T. europaea, A. sylvaticus, and S. minutus.

4.4. Habitat and climate evolution at Santimamiñe Fig. 6 (constructed by crossing data from Fig. 5 and Table 2) reconstructs Santimamiñe habitat and climate distribution based on changes in the small mammalian community over time. Both the three levels below Arg-o and the three above it (flooding complex) have not been included in the paleoenvironmental analysis given the scarcity of remains obtained for such a thick layers.

First to be noticed is the pre-eminence of the open landscape in the vicinity of the cave from Level Arg-o to Level Slnc, with a mix of dry and humid meadows, the latter habitat clearly dominating. This humid meadow predominance along with cold conditions, especially in Levels Arg-o, Csn-Camr, and Balm is determined by the

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

J. Rofes et al. / Quaternary International xxx (2013) 1e14

Arvicola amphibius

8

Fig. 5. Quantitative distribution of the small mammals from Santimamiñe (Kortezubi, Bizkaia, Spain) from the lower Magdalenian to the Chalcolithic/Bronze Age plus seven lowermost purely palaeontological levels. From left to right, columns represent the cultural periods defined by their archaeological contents, the stratigraphic levels, the arbitrary sublevels, and the fifteen taxa recorded at the site. Graphics represent the relative species abundances variations through time expressed in terms of the percentage of the minimal number of individuals (MNI) in each level (% of the MNI’s given at the top of each column). *Not cultural.

great representation of Sorex gr. araneusecoronatus in the record compared to other small mammals. Good representation of grasslands throughout the sequence is determined by the relative abundance of Microtus (M.) agrestisearvalis, which also contributes to the progressive climate amelioration observed from Level Balm upwards. This warming that parallels an increase in forestation of the surroundings (reaching its zenith at the uppermost levels), is mostly determined by the A. sylvaticus relative abundance.

Besides S. gr. araneusecoronatus, also significant for humid meadows representation are T. europaea and S. minutus. The relative abundance of these species together (but in a lesser extent) with those of M. (A.) oeconomus, C. nivalis, M. (T.) lusitanicus, and A. amphibius determine the humid meadow and cold peak of Level Csn-Camr. Aquatic (A. cf. sapidus and M. (A.) oeconomus) and rocky (C. nivalis) habitats are moderately to poorly represented throughout the sequence.

To summarize, we have a global panorama for the past environment of Santimamiñe (Fig. 6) that begins during the purely palaeontological Level Arg-o with a cold phase in which humid meadows prevail but grasslands were also important in the landscape; woodland, water and rocky habitats are poorly represented.

At the beginning of the Lower Magdalenian (Level Csn-Camr), conditions became even colder, with humid meadows reaching its peak to the detriment of grasslands and rocky habitats. During the not anthropic Level Balm, there was a moderate climate recovery mostly reflected in the advance of grasslands to the detriment of humid meadows, woodland, and aquatic habitats; the rocky habitat, by the contrary, reaches its highest upturn.

Towards the tentative transition between the middle and upper Magdalenian (Level Almp), there was a general warming which parallels a slight advance of grasslands and the reappearance of woodlands; the rocky component fall again. At the upper Magdalenian (Level Slnc), we may begin to speak of a patchy landscape, with the humid meadow habitat still prevailing, but with quite well represented forest and grassland components too, plus moderate presence of aquatic and rocky habitats. The progressive climatic amelioration continues during Azilian times (Level Arcp), with a patchy landscape surrounding the cave dominated by similar proportions of woodland and humid meadows together with moderate presence of grasslands and water courses.

Warm conditions characterize the Neolithic period in Santimamiñe (Level Slm) with mostly woodland but also a good proportion of grasslands dominating the landscape. The Chalcolithic/ Bronze Age transition (Level Lsm) is not as warm as the Neolithic but woodland still prevails over a moderately high humid meadow component with some grasslands and water courses completing the mosaic.

5. Discussion: comparisons and correlations 5.1. Amphibian and reptile record As stated before, we took the methodological decision of leaving the reptiles and amphibians out of the paleoenvironmental reconstructions. We are not able to extract habitat or climate inferences from the imprecise “Lacertidae indet.” or the widely

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

J. Rofes et al. / Quaternary International xxx (2013) 1e14

9

Fig. 6. Paleoenvironmental evolution of Santimamiñe (Kortezubi, Bizkaia, Spain) based on the distribution of habitat types and climatic oscillations along the stratigraphic sequence. From left to right, columns represent the cultural periods defined by their archaeological contents, the stratigraphic levels, the arbitrary sublevels, the species grouped by their habitat and climatic requirements, the d18O curve obtained from a deep ice core of central Greenland (GISP2, Grootes et al., 1993), some well-known north Atlantic and continental climatic episodes, and the ages expressed in cal ka BP. Dates for Levels Balm and Almp were estimated. H, Heinrich event; HCE, Holocene Cooling Event; OD, Oldest Dryas; B/A, Bölling/Alleröd; YD, Younger Dryas; Pb, Preboreal; Bo, Boreal; At, Atlantic; Sb, Subboreal; GS, Greenland Stadial; GI, Greenland Interstadial; LGM, Last Glacial Maximum; MIS, Marine Isotope Stages. Three phases of H1 sensu Stanford et al. (2011); HCE 3 sensu Bond et al. (1997, 2001); LGM sensu Mix et al. (2001). *Not cultural.

adaptable Vipera sp., but it is worth to mention the ecological requirements of the common/Iberian frogs (Rana gr. temporariae iberica), the slow worm (A. fragilis) and the natterjack toad (Epidalea calamita).

The common frog (R. temporaria) is a species commonly found at Quaternary deposits of Atlantic Europe; beech forests are its main habitat in the Basque Country, where it is continuously active throughout the year (Pleguezuelos et al., 2004). However, this species lives in a wide variety of habitats across Europe both open and forested, but always with high levels of humidity (Kuzmin et al., 2009). Rana iberica is an endemism of Portugal and northwestern and central Spain with environmental requirements similar to R. temporaria, but with a slightly higher preference for open landscapes (Tejedo et al., 2009). In Santimamiñe, remains of Rana gr. temporariaeiberica has been found in almost every level of the test trench, but the highest number of items come by far from Level Csn-Camr, not casually the one with the highest proportion of humid meadows of the entire sequence according to the small mammalian record (Fig. 6).

Epidalea calamita is a species currently found in open and unshaded light sandy soils of coastal dunes, lowland heaths, semi-

desert, high mountains, pine forest glades, gardens, parks, agricultural fields, sand and gravel quarries and meadows (Beja et al., 2009).

Remains of natterjack toads in Santimamiñe are only present at Levels Csn-Camr and Lsm, accounting for open-dry areas in the zone.

Anguis fragilis is a hygrophile, eurythermal reptile that, along with the common frog, currently inhabits the Eurosiberian region of Iberia. It is active from March to October, and is commonly found both in humid meadows and seasonal forests (Pleguezuelos et al., 2004). In Santimamiñe, there are few elements of slow worm in Levels Arg-o, Csn-Camr, Almp, and Slnc, but then an abrupt rising is recorded at the upper part of Level Arcp (1104 items among osteoderms and vertebrae). The amounts remain high at the uppermost levels (Slm and Lsm). This remarkable increase of A. fragilis items coincide with the notorious expansion of woodlands of the Holocene Climatic Optimum (Fig. 6).

5.2. Large mammal and bird record The large mammalian assemblage of Santimamiñe is composed by both wild and domestic taxa: Equus caballus, Bovini indet., Capra pyrenaica, Rupicapra rupicapra, C. elaphus (Fig. 4.20), Rangifer

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

10

J. Rofes et al. / Quaternary International xxx (2013) 1e14

tarandus, Capreolus capreolus, Sus scrofa, Bos taurus, Ovis/Capra, Sus sp., Ursus arctos, Vulpes vulpes, and Felis silvestris. The bird assemblage comprises: Gypaetus sp., Columba sp., Lagopus lagopus, Pyrrhocorax sp., Pica pica, Alectoris/Perdix, and Passeriformes indet.

In terms of the number of specimens and number of taxa, the ungulates clearly dominate the assemblage, wild species during the Upper Palaeolithic and both wild and domestic taxa during the Neolithic and Chalcolithic/Bronze Age transition. Carnivores and avifauna were also present but in a much lesser extent. As the cinegetic strategies along the sequence have been extensively treated elsewhere (Castaños and Castaños, 2011; López-Quintana, 2011a), we focus here in the large mammals and birds that are relevant for the paleoenvironmental discussion.

Most of the taxa from Santimamiñe are eurythermal, what means that they are tolerant to a wide range of temperatures and habitats. Nevertheless, there are some exceptions: the reindeer (R. tarandus), currently absent from the Iberian Peninsula, is a coldadapted species with preference for arctic and subarctic tundra, open montane and open woodland habitats (Henttonen and Tikhonov, 2008). In Santimamiñe, the few elements of R. tarandus recovered from the test trench come from Levels Arg-o, Csn-Camr and Almp, all three characterized as mostly cold and open by the small mammal record. In Almp, in particular, the presence of reindeer coincides with the only item of Willow Ptarmigan (Lagopus lagopus) found in the whole sequence, being this bird also a cold-tolerant taxon currently inhabiting northern Europe, Siberia, Alaska and northern Canada (BirdLife International, 2012).

Although the European wildcat (F. silvestris) can survive in quite open landscapes, it requires woodland masses in the vicinity (Palomo and Gisbert, 2005). In Santimamiñe, the sole remain of F. silvestris appears at Level Slm, the most densely forested and warmest of the sequence.

Finally, the wild boar (Sus scrofa) and the roe deer (Capreolus capreolus) have both forest affinities, especially the latter (Palomo and Gisbert, 2005). According to the paleoenvironmental reconstruction based on small mammals (Fig. 6), woodlands begin to rise during the Middle/Upper Magdalenian (Level Almp), and this is precisely the time when S. scrofa and C. capreolus appear for the first time. This forest advance is also detected in many other prehistoric deposits of the Cantabrian region, as El Pendo, Rascaño, Ekain, Erralla, El Juyo, and El Mirón (Leroi-Gourhan, 1980; Boyer-Klein, 1981; Dupré Ollivier, 1984; Pemán, 1985; Pokines, 1998; CuencaBescós et al., 2012), and even in caves from all the Iberian Peninsula, as Valdavara-1 in Galicia, and Les Cendres and Gorham’s in the Mediterranean region (Villaverde et al., 1999; López-García et al., 2011a, 2011b).

Some scholars claim that this increase of wooded environments would have favoured the proliferation of woodland-related ungulates, such as roe deer and wild boar, which would have increased their encounter times, leading to a greater consumption (Arias, 1992; García Moreno, 2010). However, other researchers, applying the Optimal Foraging Theory, consider that this is a simplistic point of view and argue that the Magdalenian people were following highly productive strategies in which the reforestation of the Middle Magdalenian and the rise in C. capreolus and S. scrofa exploitation were not necessarily related (Marín-Arroyo, 2009, 2010; Cuenca-Bescós et al., 2012).

dominated by open landscapes (high proportions of Poaceae in the herbaceous stratum, and of Pinus and Juniperus in the arboreal stratum); this scenario is quite similar to that inferred from the small mammals, with its predominance of humid meadows and grasslands, and cold general conditions. According to the pollen record, Level Csn-Camr (Lower Magdalenian) represents a colder and less humid period than Arg-o, also with open terrain dominating the landscape (i.e. rising of the Compositae in the herbaceous stratum and low arboreal covering comprising Pinus, Betula and Juniperus); the small mammalian record of this level reflects a similar open-land and cold pattern, with the difference of showing good levels of humidity inferred from the high numbers of, especially, Sorex and Talpa (among other humid meadow taxa) in the sample.

At the upper Magdalenian (Level Slnc), the preeminence of Compositae, the presence of Artemisia, the low proportion of fern spores, the scarce arboreal elements, and the reduced rate of taxa associated to humid environments, speaks of rigorous conditions and a mostly open-dry landscape; the small mammal association shows a retreat of humid meadows to the benefit of rocky, grassland and forest habitats, the latter one, nevertheless, still poorly represented (Apodemus). Level Arcp (Azilian) is warmer and more humid than Slnc, with more forested landscapes (i.e. appearance of Quercus robur and Corylus, retreat of Compositae, recovery of Ericaeae and Poaceae); the small mammals reflect a similar panorama, with the advance of woodland to the detriment of open landscapes (rising of Apodemus respect to other taxa become notorious). It is important here to remark that according to both pollen and small mammals the upper part of this level (treated in this paper as a whole) shows better climatic conditions than the lower (LópezQuintana and Guenaga-Lizasu, 2011; Murelaga et al., 2011).

During the Neolithic (Level Slm), the arboreal stratum reaches its maximum values due to a single taxon: Betula; Corylus and Q. robur keep their ascending trend, Poaceae dominates the herbaceous stratum, and the Compositae drop to its minimum values; towards the end of this period Betula drastically falls and Corylus and Quercus take its place; something very similar is inferred from the small mammals, with woodland and warm conditions reaching their peaks due to a sole species: A. sylvaticus; grasslands also advance paralleling Poaceae. Palynologically speaking, the uppermost level (Lsm) is very similar to Slm, although a slight retreat of Poaceae and fern spores is detected; the same happens from the small mammal point of view, with the retreat of grasslands and woodland to the benefit of humid meadow and water habitats.

There are two contradictions between the palynological and small mammalian records: a) peak of humid meadows (according to small mammals) versus moderate levels of humidity (according to pollen) at Level Csn-Camr; and, b) Coldest peak of the sequence at Level Csn-Camr (small mammals) versus coldest peak at Level Slnc (pollen). Nevertheless, these two discrepancies may be just apparent as far as all the sublevels of Csn-Camr and Slnc were sampled for the small mammal analysis (to obtain a compendium) whereas only one sublevel in each of these units was sampled for the palynological study. Hence, the paleoenvironmental scenario inferred from the pollen samples in each case could correspond to isolated periods within the general trend shown by the small mammals (see also López-Quintana and Guenaga-Lizasu, 2011: 36e39).

5.3. Palynological record

5.4. Cryoclastic record

Comparing our results (Fig. 6) with those obtained from the palynological study of Santimamiñe test trench (Iriarte, 2011), we find more coincidences than contradictions.

For the purely palaeontological level (Arg-o), the analysis of the only available sample shows a mostly cold and humid environment,

As in many other caves, with or without prehistoric deposits, some gelifraction events have been detected in the test trench of Santimamiñe (López-Quintana and Guenaga-Lizasu, 2011). In those events, the mechanical breakup and churning of the walls and/or roof of the cave (due to repeated freezing and thawing of water

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

J. Rofes et al. / Quaternary International xxx (2013) 1e14

within their cracks) causes rocks to fall, being the presence of these thick cryoclastic elements a proxy of past cold episodes. The counting of said large components, following the methods of the Analytic Stratigraphy and sampling the entire excavated surface, showed the existence of three cold phases that coincide with Levels Csn-Camr, Slnc and Arcp (lower part), being Csn-Camr the one with the highest number of thick cryoclasts, and, therefore, the coldest (López-Quintana and Guenaga-Lizasu, 2011: Figs. 40 and 50 therein). These results are not in discrepancy with the climate reconstruction based on small mammals (Fig. 6).

5.5. Other paleoenvironmental reconstructions with small mammals from north Iberia As the number of prehistoric deposits yielding fossil small mammals from the latest Late Pleistocene (MIS 2) and/or Holocene (MIS 1) times in north Iberia is considerably high, we focus here only in those with long archaeological sequences that have been analysed in terms of paleoenvironmental evolution.

In this sense, the first and most important site is, undoubtedly, El Mirón, a long cave deposit in the Asón river valley (Ramales de la Victoria, Cantabria) radiocarbon-dated to between 41 and 2 cal ka BP and chrono-culturally comprising the late Mousterian, initial Upper Palaeolithic, Solutrean, lower, middle and upper Magdalenian, Azilian, Mesolithic, Neolithic, Chalcolithic, and Bronze Age (Cuenca-Bescós et al., 2009). Comparing the paleoenvironmental reconstruction of Santimamiñe (Fig. 6) with the corresponding time span at the longer El Mirón sequence (CuencaBescós et al., 2009: Fig. 4), we find a very similar pattern of habitat evolution, with mostly open-humid landscapes for preMagdalenian, Magdalenian and Azilian times, and a remarkable reforestation during the Holocene. Both sequences coincide in the peak of grasslands detected during pre-Magdalenian times (Solutrean in El Mirón), the peak of humid meadows at the lower Magdalenian, the notorious woodland expansion of the Neolithic, and the moderate re-advance of humid meadows during the Chalcolithic/Bronze Age transition. The latter two episodes are also well recorded in the Neolithic-to-middle Bronze Age sequence of Peña Larga, a rock shelter in the southern slopes of the Cantabrian cordillera, in Álava (Rofes et al., 2013). The rocky habitat (Montane in El Mirón) is moderately better represented in El Mirón than in Santimamiñe, probably due to the former site is placed some 130 m higher than the second (260 m vs. 137 m a.s.l., Cuenca-Bescós et al., 2009; López-Quintana and Guenaga-Lizasu, 2011).

The small mammals from several other sites of the Cantabrian region with shorter chrono-stratigraphic sequences show similar distribution patterns to those found at Santimamiñe and El Mirón (see Pokines, 1998; Sesé, 2005 for compilations).

The cave of Valdavara-1 is located in the right bank of the river Narón, in the province of Lugo (westernmost Iberia), at roughly 600 m a.s.l. (López-García et al., 2011a). Excavations at the site revealed two sedimentary units: the upper, with a radiocarbon age of 4490 40 BP and chrono-culturally attributed to the Recent Prehistory; and the lower, dated between 13,770 70 and 15,120 70 BP, of Magdalenian filiations (López-García et al., 2011a). The paleoenvironmental reconstruction (based on small mammals) obtained for the lower unit (López-García et al., 2011a: Fig. 5) differs from that of the equivalent period in Santimamiñe (Fig. 6, dates of Valdavara-1 must be calibrated prior to comparison) in showing a much more forested realm with more water sources available. The rocky component is even higher than in El Mirón, what once again may be regarded to altitude. The upper unit of Valdavara-1, close in age to the Chalcolithic/Bronze Age level (Lsm) of Santimamiñe, exhibits a quite similar scenario to that level (also similar to the corresponding levels in El Mirón and Peña Larga); the

11

landscape dominated most of all by woodland and humid meadows.

On the northeastern edge of the North Plateau and the Duero Basin, not so far from the southern boundary of the Cantabrian Range, is placed the site of El Portalón, which represents the present-day entrance to Cueva Mayor karst system (Sierra de Atapuerca, Burgos) (López-García et al., 2010). The archaeological sequence of El Portalón has two units: the upper displays a Holocene chronology and the lower covers the latest part of the Pleistocene. The small vertebrates (mammals, reptiles and amphibians) of this Pleistocene unit, roughly dated to between 16 and 36 ka BP, have been studied and a paleoenvironmental reconstruction proposed (López-García et al., 2010). The upper portion of this reconstruction (i.e. w24e16 ka cal BP) can be correlated with the equivalent segment of the Santimamiñe sequence (Fig. 6). The mostly open-humid and cold panorama of El Portalón during this period, which includes the Last Glacial Maximum (LGM, 23e 19 cal ka BP sensu Mix et al., 2001), is very similar to that proposed for Santimamiñe and El Mirón.

5.6. GISP2 d18O curve Deep ice cores from central Greenland provide a long reliable climate record, which extends back to 105 ka (Grootes et al., 1993).

We have correlated our data on habitat and climate distribution from Santimamiñe with the d18O curve obtained from the Greenland Ice Sheet Project 2 (GISP2). We correlated the curve with the calibrated radiocarbon ages given in Table 1 (median probability). The GISP d18O curve in Fig. 6 shows a cold scenario for Levels Arg-o to Slnc, a transitory warming and a new cooling towards Level Arcp, and then a marked warming for Levels Slm and Lsm. This progression roughly coincides with our habitat and climate reconstructions, with the coldest (and open-land) peaks at Levels Arg-o and Csn-Camr, and the warmest (and forest) peaks at Levels Slm and Lsm. 5.7. Heinrich events Heinrich (H) events are characterized in North Atlantic sediments by horizons with increased Ice Rafted Debris (IRD) concentrations, high rates of the foraminifer Neogloboquadrina pachyderma sinistral, and light planktonic foraminiferal calcite d18O (meltwater dilution); they occurred quasi-periodically with a spacing of 5000e14,000 yrs (Heinrich, 1988; Hemming, 2004).

Records from around the North Atlantic and even throughout the Northern Hemisphere, indicate dramatic marine and terrestrial temperature reductions and increased aridity during H events (Hemming, 2004). The most widely accepted hypothesis holds that the low temperatures associated with these events resulted from reduced oceanic pole ward heat transport due to surface freshwater dilution in the North Atlantic and a consequent shutdown of the Atlantic meridional overturning circulation [AMOC] (Stanford et al., 2011).

Last two H events (H2 and H1) may be correlated with the coldest peaks recorded in the Santimamiñe sequence during the Late Pleistocene (MIS 2) (Fig. 6). H2 (w24 ka BP), in particular, roughly coincides with the cold episode of Level Arg-o (24,956e 24,170 cal BP), and H1 (17.5e16.7 ka BP) shortly postdates the very cold and humid episode of Level Csn-Camr (18,443e17,528 cal BP) and coincides with the estimated age for the also fresh, but less humid, Level Balm (the one with the highest peak of rocky habitats). However, Stanford et al. (2011) proposed a new concept for the sequence of events associated with the H1 iceberg/meltwater perturbation in the North Atlantic. They divided a much longer H1 event (some 4000 years) into three successive phases: Phase 1 (19e

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

12

J. Rofes et al. / Quaternary International xxx (2013) 1e14

17.5 BP) represents the onset of AMOC collapse, whereas Phase 2 (17.5e16.7 ka BP), defined as the “conventional” Heinrich event, represents the intense IRD deposition and freshening event in the IRD belt, at the time of the so-called “Heinrich layers” (Heinrich, 1988; Hemming, 2004). Phase 3 (16.7e14.6 ka BP) covers H1 in the Nordic Seas and at Eirik Drift and the termination of H1 cooling and AMOC resumption at the Bølling warm transition. This way, Level Csn-Camr can be tentatively correlated with the last part of H1-Phase 1, and Level Slnc (15,856e14,785 cal BP), and the preceding Level Almp, with cold pulses of H1-Phase 3.

5.8. Holocene Cooling Events During the Holocene (MIS 1), the moderately cold humid meadow expansion recorded at the uppermost level of Santimamiñe test trench (Lsm, 4220e3926 cal yr BP) (Fig. 6) slightly postdates a short “Ice Age” documented by Bond et al. (1997, 2001), as the Holocene Cooling Event 3 (HCE 3) by means of the record of lithic fragments (fresh volcanic glass from Iceland and hematite stained grains, both contained in drift ice) of European origin found at diverse sedimentary cores from the North Atlantic. The effects of this HCE 3 have been also recently detected at the nearby rock shelter of Peña Larga in Álava (Rofes et al., 2013).

5.9. North Atlantic and continental chronozones As a mean to correlate our results with other well-known continental and North Atlantic chronozones, we have included in Fig. 6, along with the paleoenvironmental reconstruction and the GISP d18O curve, the main climatic episodes of central Europe (pollen) and those recorded on ice cores from north and central Greenland (Andersen et al., 2006). The purely palaeontological Level Arg-o falls well within the very cold Würm III and the Greenland Stadial (GS) 3. All the Magdalenian levels, from Csn-Camr to Slnc, were deposited during the Oldest Dryas (equivalent to the upper half of GS 2), a mostly cold and humid period including the Heinrich event 1. The cold peak and humid meadow expansion of Level Csn-Camr came after the LGM and during the subsequent deglaciation, when central European and Iberian glaciers regained terrain around 18 cal ka BP (Knies et al., 2007; Jalut et al., 2010) as a result of the H1-Phase 1 (Stanford et al., 2011).

The Azilian occupation of Santimamiñe (Level Arcp) chronologically took place between the end of the markedly seasonal Younger Dryas (equivalent to the last part of GS 1) and the warm and humid Preboreal, the first climatic episode of the Holocene and the beginning of MIS 1. The Neolithic level (Slm) coincides with the warmest and most forested period recorded in the whole sequence, falling within the Atlantic chronozone. Level Lsm, attributable to the Chalcolithic/Bronze Age transition, was deposited during a moderate re-opening of the landscape at the warm Subboreal episode.

5.10. Occupational dynamics of the cave The information in this chapter (other than paleoenvironmental, which is extracted from Fig. 6) was taken and synthesized from Castaños and Castaños (2011), Gutierrez-Zugasti (2011), LópezQuintana and Gunega-Lizasu (2011), López-Quintana (2011a), and Roselló-Izquierdo and Morales-Muñíz (2011).

The lowermost level (Arg-o) of Santimamiñe test trench has no evidence of human occupation but a rich record of ungulate remains plus one carnivore (U. arctos) instead. The type of bone accumulation together with the finding of a long bone of red deer (C. elaphus) with gnawing marks point to the use of the cave as bear

den during this period which, paleoenvironmentally speaking, was cold, humid and with open areas dominating the landscape.

The archaeological record of Level Csn-Camr (lower Magdalenian), the coldest and most humid of the sequence (coetaneous with the end of H1-phase 1), shows that the cave was occupied by a human group mainly focused in the hunting and processing of red deer for both the procurement of meat and skin, and the elaboration of horn and bone tools.

The overlying level (Balm) and the lower part of the next (Almp) show a hiatus in human occupation, probably due to a flooding episode at least in the area of the test trench. In spite of the remobilization of materials from the overlying level (Slnc), there is evidence of occasional human presence in the upper part of Almp, which is tentatively assigned to a transition horizon between the middle and upper Magdalenian. People inhabiting the cave during this period probably witnessed the first steps towards a woodland recovery in the surroundings.

Level Slnc (upper Magdalenian) show evidence of longer and more intense human occupation of the cave by groups of hunters and collectors who exploited all the natural resources offered by a still predominantly open landscape subdued to humid and temperate conditions. They were mainly focused in the hunting of red deer and goats (Capra pyrenaica) complemented with roe deer, large bovids, and the fishing of salmon (Salmo sp.). According to González-Sainz and Ruiz-Idarraga (2010), all the parietal art of Santimamiñe was probably performed during this period.

The Azilian occupation of the cave (level Arcp) was not as intense as the previous. Subsistence was again based mainly in the capture of red deer whose number of remains exceeds by far those of goats, roe deer and large bovids. Fishing of salmon decreases and bone tools completely disappear from the record. Small mammalian, palynological and cryoclastic evidence (see above) document a remarkable climatic amelioration and forestation of the landscape at the upper part of this level, an evolution that, given the chronology of Arcp, well could be understood as the change from the cold Younger Dryas to the beginning of the Holocene Climatic Optimum.

During the middle Holocene, the cave was occupied, mostly in its external part, by eventual but recurring groups of farmers that also maintained a diversified exploitation of the landscape (ungulates hunting, essentially red deer, and intensive recollection of estuarine molluscs). Two levels deposited during this period: Slm, of Neolithic filiations, and Lsm, attributed to the Chalcolithic/ Bronze Age transition. The large mammal record of both levels is not surprisingly dominated by livestock (ovicaprids, cattle and pigs), the number of specimens increasing at the uppermost level (Lsm). Mollusc exploitation was especially meaningful during the Neolithic, most of it oriented to the extraction of the estuarine clam Scrobicularia plana.

During the Chalcolithic and Bronze Age, agriculture and stockbreeding spread drastically on the Iberian Peninsula (Fabián et al., 2006; Rofes et al., 2013). Santimamiñe is no exception, and to this could be regarded the decrease in woodland detected both in the pollen (Iriarte, 2011) and small mammalian (this paper) records of Level Lsm. An alternative and/or complementary explanation comes given by the already mentioned HCE 3 (Bond et al., 1997, 2001), which is dated precisely to the Chalcolithic/Bronze Age transition.

6. Summary and conclusions The cave of Santimamiñe (Kortezubi, Bizkaia) is one of the most famous prehistoric localities of the Cantabrian range. Between 2004 and 2006, a test trench revealed a long stratigraphic sequence in the inner vestibule of the cave. The 6 m-deep-deposit comprises

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

J. Rofes et al. / Quaternary International xxx (2013) 1e14

six chrono-cultural units: lower, middle and upper Magdalenian, Azilian, Neolithic, and Chalcolithic/Bronze Age, plus seven lowermost purely palaeontological levels (latest Late Pleistocene [MIS 2] to the middle Holocene [MIS 1]). More than forty-seven thousand microvertebrate elements were recovered from the test trench, of which 14,423 were identified to the order and/or family levels, and 1587 to the genus and/or species levels.

The small vertebrate assemblage is composed of mammals, birds, reptiles, amphibians and fishes, and it was naturally accumulated mainly by nocturnal birds of prey, although some degree of in situ deposition may account for at least part of the chiropteran, amphibian and reptile remains found at the cave.

Small mammals from the Santimamiñe test trench have been used for paleoenvironmental reconstructions: they are sensitive to climatic conditions and their distributions throughout the stratigraphic sequence, measured in terms of species relative abundance, serve as reliable proxies of habitat and climatic change.

Evolution of the environment in the vicinity of the cave begins with cold and humid conditions and a mostly open landscape from pre-Magdalenian times to the upper Magdalenian; it continues with mild conditions and a patchy landscape during Azilian times; and it ends up with a general warming and woodland expansion during the Neolithic, and a slight cooling and humid meadow recovery at the Chalcolithic/Bronze Age transition.

The paleoenvironmental reconstruction of Santimamiñe based on the small mammals roughly coincides with other habitat and climatic proxies, such as the amphibian/reptile record, the large mammal/bird record, the palynological record and the cryoclastic record on the local scale; other long paleoenvironmental reconstructions from north Iberia on the regional scale; and the oxygen isotopic curve of an ice core from central Greenland on the global scale.

The purely palaeontological Level Arg-o falls within the very cold Würm III and the GS 3, and roughly coincides with the Heinrich event 2. All Magdalenian levels (Csn-Camr to Slnc) were deposited during the Oldest Dryas (equivalent to the upper half of GS 2), a mostly cold and humid period including the Heinrich event 1, the latter chronologically coinciding with Level Balm, the one with the peak of rocky habitats. Small mammalian, palynological and cryoclastic evidence likely document the shift from the cold Younger Dryas to the beginning of the Holocene Climatic Optimum during the Azilian Level Arcp. The much forested Neolithic Level Slm falls within the warm Atlantic chronozone, and the uppermost Lsm (Chalcolithic/Bronze Age), deposited during a moderate reopening of the landscape at the Subboreal, slightly postdates the Holocene Cooling Event 3.

Acknowledgments Juan Rofes has a “Juan de la Cierva” postdoctoral fellowship (JCI2010-06148) of the Ministerio de Economía y Competitividad de España (MEC). We received financial support from the Servicio de Diputación Foral de Bizkaia and the Project GUI12/35 of the Universidad del País Vasco UPV-EHU.

References Andersen, K.K., Svensson, A., Johnsen, S.J., Rasmussen, S.O., Bigler, M., Röthlisberger, R., Ruth, U., Siggaard-Andersen, M.-L., Steffensen, J.P., DahlJensen, D., Vinther, B.M., Clausen, H.B., 2006. The Greenland ice core chronology 2005, 15e42 ka. Part 1: constructing the time scale. Quaternary Science Reviews 25, 3246e3257.

Andrews, P., 1990. Owls, Caves and Fossils. The Natural History Museum Publications, London.

Aranzadi, T., Barandiarán, J.M., 1935. Exploraciones de la caverna de Santimamiñe (Basondo: Cortézubi). Tercera Memoria e Yacimientos azilienses y paleolíticos.

Re-edited in Barandiarán, J.M. (1976). In: Obras completas, vol. IX, pp. 245e344.

13

Aranzadi, T., Barandiarán, J.M., Eguren, E., 1925. Exploraciones de la caverna de Santimamiñe (Basondo: Cortézubi). Primera Memoria e Figuras rupestres. Reedited in Barandiarán, J.M. (1976). In: Obras completas, vol. IX, pp. 11e89.

Aranzadi, T., Barandiarán, J.M., Eguren, E., 1931. Exploraciones de la caverna de Santimamiñe (Basondo: Cortézubi). Segunda Memoria e Los niveles con cerámica y el conchero. Re-edited in Barandiarán, J.M. (1976). In: Obras completas, vol. IX, pp. 91e243.

Arias, P., 1992. Estrategias económicas de las poblaciones del Epipaleolítico avanzado y el Neolítico en la región cantábrica. In: Moure, A. (Ed.), Elefantes, ciervos y ovicaprinos. Economía y aprovechamiento del medio en la Prehistoria de España y Portugal. Servicio de Publicaciones de la Universidad de Cantabria, Santander, pp. 163e184.

Barandiarán, J.M., 1962a. Exploraciones de la caverna de Santimamiñe (Basondo: Cortézubi). Cuarta Memoria e Campaña de 1960. Re-edited in Barandiarán, J.M. (1976). In: Obras completas, vol. IX, pp. 345e368.

Barandiarán, J.M., 1962b. Exploraciones de la caverna de Santimamiñe (Basondo: Cortézubi). Quinta Memoria e Campaña de 1961. Re-edited in Barandiarán, J.M. (1976). In: Obras completas, vol. IX, pp. 369e403.

Barandiarán, J.M., 1962c. Exploraciones de la caverna de Santimamiñe (Basondo: Cortézubi). Sexta Memoria e Campaña de 1962. Re-edited in Barandiarán, J.M. (1976). In: Obras completas, vol. IX, pp. 405e419.

Barandiarán, I., 1988. Historia General de Euskalerria. Prehistoria: Paleolítico. In: Enciclopedia General Ilustrada del País Vasco. Editorial Auñamendi, San Sebastián.

Beja, P., Kuzmin, S., Beebee, T., Denoël, M., Schmidt, B., Tarkhnishvili, D., Ananjeva, N., Orlov, N., Nyström, P., Ogrodowczyk, A., Ogielska, M., Bosch, J., Miaud, C., Tejedo, M., Lizana, M., Martínez-Solano, I., 2009. Epidalea calamita.

In: IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. www. iucnredlist.org (Downloaded on 14.03.13.).

Bertolini, M., Fedozzi, S., Martini, F., Sala, B., 1996. Late glacial and Holocene climatic oscillations inferred from the variations in the micromammal associations at Grotta Della Serratura (Marina di Camerota, Salerno, S. Italy). Il Quaternario.

Italian Journal of Quaternary Sciences 9, 561e566.

BirdLife International, 2012. Lagopus lagopus. In: IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. www.iucnredlist.org (Downloaded on 14.03.13.).

Bond, G., Showers, W., Cheseby, M., Lotti, R., Almasi, P., deMenocal, P., Priore, P., Cullen, H., Hajdas, I., Bonani, G., 1997. A pervasive millennial-scale cycle in North Atlantic Holocene and Glacial climates. Science 278, 1257e1266.

Bond, G., Kromer, B., Beer, J., Muscheler, R., Evans, M.N., Showers, W., Hoffmann, S., Lotti-Bond, R., Hajdas, I., Bonani, G., 2001. Persistent solar influence on North Atlantic climate during the Holocene. Science 294, 2130e2136.

Boyer-Klein, A., 1981. Análisis palinológico del Rascaño. In: González Echegaray, J., Barandiaran Maestu, I. (Eds.), El Paleolítico Superior de la Cueva del Rascaño (Santander). Centro de Investigación y Museo de Altamira, Santander, pp. 216e220.

Bronk Ramsey, C., 2009. Bayesian analysis of radiocarbon dates. Radiocarbon 51 (1), 337e360.

Castaños, P., Castaños, J., 2011. Estrategias de caza en la secuencia prehistórica de Santimamiñe. In: López-Quintana, J.C. (Ed.), La cueva de Santimamiñe: revisión y actualización (2004e2006), Excavaciones arqueológicas en Bizkaia, Kobie 1, pp. 197e206.

Cuenca-Bescós, G., Rofes, J., García-Pimienta, J.C., 2005. Early Europeans and environmental change across the EarlyeMiddle Pleistocene transition: small mammalian evidence from Trinchera Dolina cave, Atapuerca, Spain. In: Head, M.J., Gibbard, P.L. (Eds.), EarlyeMiddle Pleistocene Transitions: the LandOcean Evidence, Geological Society of London, Special Publications, vol. 247, pp. 277e286.

Cuenca-Bescós, G., Straus, L.G., González-Morales, M.R., García-Pimienta, J.C., 2009.

The reconstruction of past environments through small mammals: from the Mousterian to the Bronze Age in El Mirón Cave (Cantabria, Spain). Journal of Archaeological Science 36, 947e955.

Cuenca-Bescós, G., Melero-Rubio, M., Rofes, J., Martínez, J.L., Arsuaga, J.L., Blain, H.A., López-García, J.M., Carbonell, E., Bermúdez de Castro, J.M., 2011. The Earlye Middle Pleistocene environmental and climatic change and the human expansion in Western Europe: a case study with small vertebrates (Gran Dolina, Atapuerca, Spain). Journal of Human Evolution 60, 481e491.

Cuenca-Bescós, G., Marín-Arroyo, A.B., Martínez, I., González-Morales, M.R., Straus, L.G., 2012. Relationship between Magdalenian subsistence and environmental change: the mammalian evidence from El Mirón (Spain). Quaternary International 272e273, 125e137.

Dupré Ollivier, M., 1984. Palinología de los niveles VII a II. In: Altuna, J., Merino, J.M. (Eds.), El yacimiento prehistórico de la cueva de Ekain (Deba, Guipúzcoa). Eusko Ikaskuntza e Sociedad de Estudios Vascos, San Sebastián, pp. 61e63.

Fabián, J.F., Blanco-González, A., López-Sáez, J.A., 2006. La transición CalcolíticoBronce Antiguo desde una perspectiva arqueológica y ambiental: el Valle Amblés (Ávila) como referencia. Arqueología Espacial 26, 37e56.

Fernández-Eraso, J., 2011. Santimamiñe en la Prehistoria Vasca. In: LópezQuintana, J.C. (Ed.), La cueva de Santimamiñe: revisión y actualización (2004e 2006), Excavaciones arqueológicas en Bizkaia, Kobie 1, pp. 393e400.

García Moreno, A., 2010. Patrones de asentamiento y ocupación del territorio en el Cantábrico oriental al final del Pleistoceno. Una aproximación mediante SIG (PhD thesis). Universidad de Cantabria, Santander.

González-Sainz, C., Ruiz-Idarraga, R., 2010. Una nueva visita a Santimamiñe. Precisiones en el conocimiento del conjunto parietal paleolítico. Kobie, Serie Anejos 11, Diputación Foral de Bizkaia, Bilbao.

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048

14

J. Rofes et al. / Quaternary International xxx (2013) 1e14

Grayson, D., 1984. Quantitative Zooarchaeology. Academic Press, London.

Grootes, P.M., Stuvier, M., White, J.W.C., Johnsen, S., Jouzel, J., 1993. Comparison of oxygen isotope records from the GISP2 and GRIP Greenland ice cores. Nature 366, 552e554.

Gutierrez-Zugasti, F.I., 2011. Los moluscos alimenticios de la Cueva de Santimamiñe (Kortezubi, Bizkaia): campañas de excavación 2004e2006. In: López Quintana, J.C. (Ed.), La cueva de Santimamiñe: revisión y actualización (2004e 2006), Excavaciones arqueológicas en Bizkaia, Kobie 1, pp. 247e266.

Heinrich, H., 1988. Origin and consequences of cyclic ice rafting in the Northeast Atlantic Ocean during the past 13,000 years. Quaternary Research 29, 142e152.

Hemming, S.R., 2004. Heinrich events: massive Late Pleistocene detritus layers of the North Atlantic and their global imprint. Reviews of Geophysics 42, 1e43.

Henttonen, H., Tikhonov, A., 2008. Rangifer tarandus. In: IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. www.iucnredlist.org (Downloaded on 14.03.13.).

Iriarte, M.J., 2011. Polen y vegetación en la secuencia estratigráfica de Santimamiñe.

In: López Quintana, J.C. (Ed.), La cueva de Santimamiñe: revisión y actualización (2004e2006), Excavaciones arqueológicas en Bizkaia, Kobie 1, pp. 321e342.

Janeau, G., Aulagnier, S., 1997. Snow vole e Chionomys nivalis (Martins 1842).

Journal of Mountain Ecology 4, 1e11.

Jalut, G., Turu i Michels, V., Dedoubat, J.J., Otto, T., Ezquerra, J., Fontugne, M., Belet, J.C., Bonnet, L., García de Celis, A., Redondo-Vega, J.M., Vidal-Romaní, J.R., Santos, L., 2010. Palaeoenvironmental studies in NW Iberia (Cantabrian range): vegetation history and synthetic approach of the last deglaciation phases in the western Mediterranean. Palaeogeography, Palaeoclimatology, Palaeoecology 297, 330e350.

Knies, J., Vogt, C., Matthiessen, J., Nam, S.I., Ottesen, D., Rise, L., Bargel, T., Eilertsen, R.S., 2007. Re-advance of the Fennoscandian Ice Sheet during Heinrich event 1. Marine Geology 240, 1e18.

Kuzmin, S., Ishchenko, V., Tuniyev, B., Beebee, T., Andreone, F., Nyström, P., Anthony, B., Schmidt, B., Ogrodowczyk, A., Ogielska, M., Bosch, J., Miaud, C., Loman, J., Cogalniceanu, D., Kovács, T., Kiss, I., 2009. Rana temporaria. IUCN 2012. IUCN Red List of Threatened Species. Version 2012.2. www.iucnredlist.org (Downloaded on 14.03.13.).

Leroi-Gourhan, A., 1980. Análisis polínico de El Pendo. In: González Echegaray, J. (Ed.), El yacimiento de la cueva de El Pendo (Excavaciones 1953e57), Bibliotheca Praehistorica Hispana, vol. XVII. Instituto Español de Prehistoria. Consejo Superior de Investigaciones Científicas, Madrid, pp. 263e266.

López-García, J.M., Blain, H.-A., Cuenca-Bescós, G., Ruiz-Zapata, M.B., DoradoValiño, M., Gil-García, M.J., Valdeolmillos, A., Ortega, A.I., Carretero, J.M., Arsuaga, Bermúdez de Castro, J.M., Carbonell, E., 2010. Palaeoenvironmental and palaeoclimatic reconstruction of the Latest Pleistocene of El Portalón site, Sierra de Atapuerca, northwestern Spain. Palaeogeography, Palaeoclimatology, Palaeoecology 292, 453e464.

López-García, J.M., Blain, H.-A., Cuenca-Bescós, G., Alonso, C., Alonso, S., Vaquero, M., 2011a. Small vertebrates (Amphibia, Squamata, Mammalia) from the late PleistoceneeHolocene of the Valdavara-1 cave (Galicia, northwestern Spain). Geobios 44, 253e269.

López-García, J.M., Cuenca-Bescós, G., Finlayson, C., Brown, K., Giles-Pacheco, F., 2011b. Palaeoenvironmental and palaeoclimatic proxies of the Gorham’s cave small mammal sequence, Gibraltar, southern Iberia. Quaternary International, 137e142.

López-García, J.M., Blain, H.-A., Sanz, M., Daura, J., 2012. A coastal reservoir of terrestrial resources for Neanderthal populations in north-eastern Iberia: palaeoenvironmental data inferred from the small-vertebrate assemblage of Cova del Gegant, Sitges, Barcelona. Journal of Quaternary Science 27, 105e113.

López-Quintana, J.C., Guenaga-Lizasu, A., 2007. Avance a la secuencia estratigráfica de la cueva de Santimamiñe (Kortezubi), tras la revisión de su depósito arqueológico en las campañas de 2004 a 2006. Krei 9, 5e20.

López-Quintana, J.C., 2011a. La ocupación humana se Santimamiñe (Kortezubi): Paisaje, recursos y estrategias de explotación del medio desde el Magdaleniense inferior al Calcolítico-Edad del Bronce. In: López Quintana, J.C. (Ed.), La cueva de Santimamiñe: revisión y actualización (2004e2006), Excavaciones arqueológicas en Bizkaia, Kobie 1, pp. 421e446.

López-Quintana, J.C. (Ed.), 2011b. La cueva de Santimamiñe: revisión y actualización (2004e2006). Excavaciones arqueológicas en Bizkaia, Kobie 1, pp. 1e446.

López-Quintana, J.C., Guenaga-Lizasu, A., 2011. Revisión estratigráfica del depósito arqueológico de la cueva de Santimamiñe (Kortezubi, Bizkaia): campañas de

2004 a 2006. Cronoestratigrafía y paleoambiente. In: López Quintana, J.C. (Ed.), La cueva de Santimamiñe: revisión y actualización (2004e2006), Excavaciones arqueológicas en Bizkaia, Kobie 1, pp. 7e70.

Marín-Arroyo, A.B., 2009. The use of Optimal Foraging Theory to estimate Late Glacial site catchment areas from a central place. The case of eastern Cantabria, Spain. Journal of Anthropological Archaeology 28, 27e36.

Marín-Arroyo, A.B., 2010. Arqueozoología en el cantábrico oriental durante la transición Pleistoceno/Holoceno: La Cueva del Mirón. Servicio de Publicaciones Unican, Santander.

Marshall, F., Pilgram, T., 1993. NISP vs. MNI in quantification of body-part representation. American Antiquity 58, 261e269.

Mix, A.E., Bard, E., Schneider, R., 2001. Environmental processes of the ice age: land, ocean, glaciers (EPILOG). Quaternary Science Reviews 20, 627e657.

Murelaga, X., Bailon, S., Sáez de Lafuente, X., Castaños, P., López-Quintana, J.C., Guenaga-Lizasu, A., Ortega, L.A., Zuluaga, M.C., Alonso-Olazabal, A., 2011. La fauna de microvertebrados de Santimamiñe (Pleistoceno Superior-Holoceno) (Kortezubi, Bizkaia). In: López Quintana, J.C. (Ed.), La cueva de Santimamiñe: revisión y actualización (2004e2006), Excavaciones arqueológicas en Bizkaia, Kobie 1, pp. 197e206.

Palomo, J.L., Gisbert, J., 2005. Atlas de los mamíferos terrestres de España. Dirección General para la Biodiversidad, Madrid.

Pemán, E., 1985. Cazadores magdalenienses en la cueva de Erralla (Cestona, País Vasco). Capítulo 5. Aspectos climáticos y ecológicos de los micromamíferos del yacimiento de Erralla. Munibe 37, 49e57.

Pleguezuelos, J.M., Márquez, M., Lizana, M., 2004. Atlas y libro rojo de los Anfibios y Reptiles de España, third ed. Dirección General de Conservación de la Naturaleza-Asociación Herpetológica Española, Madrid.

Pokines, J.T., 1998. The Paleoecology of Lower Magdalenian Cantabrian Spain. In: British Archaeological Reports International Series 713. London.

Reimer, P.J., Baillie, M.G.L., Bard, E., Bayliss, A., Beck, J.W., Blackwell, P.G., Ramsey, C.B., Buck, C.E., Burr, G.S., Edwards, R.L., Friedrich, M., Grootes, P.M., Guilderson, T.P., Hajdas, I., Heaton, T.J., Hogg, A.G., Hughen, K.A., Kaiser, K.F., Kromer, B., McCormac, F.G., Manning, S.W., Reimer, R.W., Richards, D.A., Southon, J.R., Talamo, S., Turney, C.S.M., van der Plicht, J., Weyhenmeye, C.E., 2009. IntCal09 and Marine09 radiocarbon age calibration curves, 0e50,000 years cal BP. Radiocarbon 51 (4), 1111e1150.

Repenning, C.A., 2001. Beringian climate during intercontinental dispersal: a mouse eye view. Quaternary Science Reviews 20, 25e40.

Rofes, J., Cuenca-Bescós, G., 2011. Evolutionary history and biogeography of the genus Crocidura (Mammalia, Soricidae) in Europe, with emphasis on Crocidura kornfeldi. Mammalian Biology 76, 64e78.

Rofes, J., Zuluaga, M.C., Murelaga, X., Fernández-Eraso, J., Bailon, S., Iriarte, M.J., Ortega, L.A., Alonso-Olazabal, A., 2013. Palaeoenvironmental reconstruction of the early Neolithic to middle Bronze Age Peña Larga rock shelter (Álava, Spain) from the small mammal record. Quaternary Research 79, 158e167.

Roselló-Izquierdo, E., Morales-Muñíz, 2011. Evidencias de pesca en las ocupaciones de Santimamiñe. In: López Quintana, J.C. (Ed.), La cueva de Santimamiñe: revisión y actualización (2004e2006), Excavaciones arqueológicas en Bizkaia, Kobie 1, pp. 239e246.

Sesé, C., 2005. Aportación de los micromamíferos al conocimiento paleoambiental del Pleistoceno Superior en la región Cantábrica: nuevos datos y síntesis. In: Monografías, vol. 20. Museo y Centro de Investigación de Altamira, pp. 167e200.

Speybroeck, J., Crochet, P.-A., 2007. Species list of the European herpetofauna e a tentative update. Podarcis 8, 8e34.

Stanford, J.D., Rohling, E.J., Bacon, S., Roberts, A.P., Grousset, F.E., Bolshaw, M., 2011.

A new concept for the paleoceanographic evolution of Heinrich event 1 in the North Atlantic. Quaternary Science Reviews 30, 1047e1066.

Stuvier, M., Reimer, P.J., Reimer, R.W., 2013. CALIB 6.0. http://calib.qub.ac.uk/calib/.

Tejedo, M., Bosch, J., Martínez-Solano, I., Salvador, A., García-París, M., RecueroGil, E., 2009. Rana iberica. In: IUCN 2012. IUCN Red List of Threatened Species.

Version 2012.2. www.iucnredlist.org (Downloaded on 14.03.13.).

Villaverde, V., Martínez-Valle, R., Badal, E., Guillem, P.M., García, R., Menarques, J., 1999. El Paleolítico superior de la Cova de les Cendres (Teulada-Moraira, Alicante). Datos proporcionados por el sondeo efectuado en los cuadros AIB-17.

Archivo de Prehistoria Levantina 23, 9e65.

Wilson, D.E., Reeder, D.-A.M., 2005. Mammal Species of the World. A Taxonomic and Geographic Reference. The Johns Hopkins University Press, Baltimore.

Please cite this article in press as: Rofes, J., et al., The long paleoenvironmental sequence of Santimamiñe (Bizkaia, Spain): 20,000 years of small mammal record from the latest Late Pleistocene to the middle Holocene, Quaternary International (2013), http://dx.doi.org/10.1016/ j.quaint.2013.05.048