Palaeoenvironmental and cultural dynamics of the coast of Málaga (Andalusia, Spain) during the Upper Pleistocene and early Holocene

Palaeoenvironmental and cultural dynamics of the coast of Málaga (Andalusia, Spain) during the Upper Pleistocene and early Holocene

Quaternary Science Reviews 27 (2008) 2176–2193 Contents lists available at ScienceDirect Quaternary Science Reviews journal homepage: www.elsevier.c...

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Quaternary Science Reviews 27 (2008) 2176–2193

Contents lists available at ScienceDirect

Quaternary Science Reviews journal homepage: www.elsevier.com/locate/quascirev

Palaeoenvironmental and cultural dynamics of the coast of Ma´laga (Andalusia, Spain) during the Upper Pleistocene and early Holocene ˜ iz b, Marı´a D. Simo´n-Vallejo c, M. Merce` Bergada`-Zapata d, Miguel Corte´s-Sa´nchez a, *, Arturo Morales-Mun Antonio Delgado-Huertas e, Pilar Lo´pez-Garcı´a f, Jose´ A. Lo´pez-Sa´ez f, M. Carmen Lozano-Francisco g, Jose´ A. Riquelme-Cantal h, Eufrasia Rosello´-Izquierdo b, Antonio Sa´nchez-Marco i, Jose´ L. Vera-Pela´ez g ´ rea de Prehistoria, Universidad de Co ´rdoba, Plaza Cardenal Salazar, 3, E 14071-Co ´ rdoba, Spain A ´noma de Madrid, E-28049 Madrid, Spain Laboratorio de Zooarqueologı´a, Departamento de Biologı´a, Universidad Auto c ´n Cueva de Nerja, Carretera de Maro, s/n. E 29787-Nerja, Malaga, Spain Fundacio d ` ria, Histo ` ria Antiga i Arqueologia, Facultat de Geografia i Histo `ria, Universitat de Barcelona, C/Montealegre 6, E-08001 Barcelona, Spain SERP, Departament de Prehisto e ´n Experimental del Zaidı´n, Profesor Albareda 1, E-18008 Granada, Spain Estacio f ´nica, Departamento de Prehistoria, Instituto de Historia, Duque de Medinaceli 6, E-28014 Madrid, Spain Laboratorio de Arqueobota g ´ laga, Spain ´gico de Estepona, Matı´as Prats, s/n. E-29680 Estepona, Ma Museo Municipal Paleontolo h Departamento de Prehistoria, Universidad de Granada, Campus Universitario de la Cartuja (CSIC-Granada), E-18071 Granada, Spain i Departamento de Paleobiologı´a, Museo Nacional de Ciencias Naturales, Jose´ Gutie´rrez Abascal 2, E 28006-Madrid, Spain a

b

a r t i c l e i n f o

a b s t r a c t

Article history: Received 24 January 2007 Received in revised form 19 March 2007 Accepted 31 March 2008

The impact of late Upper Pleistocene climate change in the western Mediterranean region has been mainly documented through marine records. Archaeological and geomorphological continental records now available for the coast of Ma´laga complement these records for the second part of the last major glacial episode and the early stages of the Holocene. This paper provides an overview of the archaeological, chronological and palaeoenvironmental data from the end of the Middle Palaeolithic period to the Epipalaeolithic. Sequences from the two major sites, Nerja and Bajondillo, indicate a mosaic-type response of ecosystems to the rapidly shifting conditions of the Last Glacial episode. Terrestrial mammals, for example, show no major variations from present-day communities. Plant remains in contrast, demonstrates the existence of localised refuge areas for species that would not otherwise have survived in the more widely prevailing climatic conditions of the time, whereas remains of fish and, secondarily, birds demonstrate the existence of communities without any present-day analogues combining Mediterranean and Boreal (i.e., northern Atlantic) taxa. From an archaeological standpoint, the major cultural shift is the onset of marine fishing, beginning in the Solutrean. This coincided with the end of the Last Glacial Maximum (LGM) episode. The faunal record also testifies to the rising importance of marine resources more generally. Although the data have not been used systematically to test the validity of the Broad Spectrum Revolution (BSR) hypothesis, the hints for it in the area seem compelling from the start of the Solutrean onwards. Ó 2008 Elsevier Ltd. All rights reserved.

1. Introduction The littoral fringe of Ma´laga (Fig. 1) features one of the most important archaeological records of the Southern Iberian * Corresponding author. E-mail addresses: [email protected] (M. Corte´s-Sa´nchez), arturo.morales@ ˜ iz), [email protected] (M.D. Simo´n-Vallejo), beruam.es (A. Morales-Mun [email protected] (M.M. Bergada`-Zapata), [email protected] (A. Delgado-Huertas), [email protected] (P. Lo´pez-Garcı´a), [email protected] (J.A. Lo´pez-Sa´ez), [email protected] (M.C. Lozano-Francisco), [email protected] (J.A. Riquelme-Cantal), [email protected] (E. Rosello´-Izquierdo), mcnac539@ mncn.csic.es (A. Sa´nchez-Marco). 0277-3791/$ – see front matter Ó 2008 Elsevier Ltd. All rights reserved. doi:10.1016/j.quascirev.2008.03.010

Peninsula. Thirteen prehistoric sites have been documented for the chronological interval ranging from OIS 5–1, the richest sequences being those retrieved at the caves of Bajondillo and Nerja (Table 1). The chronological, cultural and palaeoenvironmental information at these two sites, together with those from Hoyo de la Mina and Abrigo 6 of the Humo complex, allow insights into the impact of the last major glacial episode on this area as well as the mosaic of changes in ecosystems and taxa in response to shifting environmental conditions. It is within such a scenario of a shifting availability of resources that human populations adjusted their adaptive strategies during the Upper Palaeolithic and Early Holocene.

M. Corte´s-Sa´nchez et al. / Quaternary Science Reviews 27 (2008) 2176–2193

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Fig. 1. The Sea of Alboran and the coast of Ma´laga showing the location of sites mentioned in the text.

2. Geographical and geomorphological dynamics on the coast of Ma´laga The mountains of the coast of Ma´laga are part of the Internal Zone of the Be´tic mountain range and are rich in limestone massifs subjected to strong karstification. Caves and shelters are common and are occasionally filled with Upper Pleistocene sediments that record sequences of palaeoenvironmental change as well as human occupation. Additional geomorphological variables affecting landscape conditions include neotectonic activity and fluctuations of the karst water table, which promoted carbonated surficial deposits like travertines (Herna´ndez et al., 1996; Corte´s and Simo´n, 2000; Ferre et al., 2003; Guerra et al., 2003). Quaternary glacio-eustatic events repeatedly enlarged and reduced the width of the coastal platform, creating for long periods an extension of the coastal zone with established animal and vegetational communities. Such a dynamic geomorphology, coupled with a relatively mild climate during the coldest pulses of the Upper Pleistocene, created favourable conditions for the establishment of human populations, and this is consistent with the number of archaeological sites in the region. For the time period considered in this paper (c. 33–7 ky), and from a tectonic standpoint, the Bay of Ma´laga constitutes a stable zone or, at the most, one of moderate uplift (i.e., 1–2 cm/ky, Lario et al., 1998), whereas the easternmost sector of the region under study (Nerja) has undergone important episodes of uplift. Marine

regressions have been detected in various submerged erosional terraces lying at 120, 90, 73, 60, 47, 33 and 20 m below sea level (Herna´ndez et al., 1996). The 120 m isobath appears to correspond to the sea level of the coast during the Last Glacial Maximum (LGM, c. 20 ky) and the 90 m isobath with the level of the coast during the Tardiglacial (c. 14 ky). Such changes clearly affected the availability of terrestrial plant and animal resources and also the accessibility of the coast. At Nerja, the width of the coastal plain was reduced from 5.5 km during the LGM to 2.5 km during the Early Holocene (Table 2; today the cave lies a mere 1 km from the sea) (Aura et al., 2001). In the bay of Ma´laga, the coastal plan varied in width from 10.5 to 2 km (Corte´s and Simo´n, 2000). 3. Materials and methods The data presented derive mainly from two sites, Bajondillo and Nerja, with the occasional addition of data from two other deposits (Hoyo de la Mina and Abrigo 6/Humo). Both technological and typological criteria are used to create a chrono-cultural sequence (Aura, 1995; Corte´s, 2007). As such, the names given to the various periods are in general agreement with the current terminology for the Iberian Upper Palaeolithic and Epipalaeolithic. Bajondillo (Table 1) features the longest sequence as it covers OIS 5–1 (Corte´s and Simo´n, 1997; Corte´s, 2002; 2007). Human occupations have been documented for the Middle and Upper Palaeolithic,

2178

Table 1 Upper Pleistocene chrono-cultural sequences from archaeological sites of the coast of Ma´laga Cultural stage

Bajondilloa

Nerjab

Hoyo Minac

Epipalaeolithic, c. 7–10 ky

Bj/3 Bj/4 –





V/3 V/4

5b – – –

Magdalenian, c. 16–10 ky

Middle Solutrean, c. 19–21 ky Gravettian, c. 21–27 ky

Bj/6 Bj/7 Bj/8 –

Middle Palaeolithic, c. 34–114 ky

a b c d

M/ 16sup. M/16inf. – – – V/8

6 –

– – – –

– – – Bj/9

V/9 – V/10 –

– 7 – –



V/12





Aurignacian, c. 27–34 ky

T/16 T/17 V/5 V/6 V/7

Bj/10 Bj/11

V/13

– –



– –

Bj/12–13





Bj/14 Bj/15–Bj/19 Bj/20 Tufa

– –

– –

Corte´s and Simo´n, 1997. Aura et al., 2002. Ferrer et al., 2006. Corte´s et al., 1996.

C/AMS

CalPal2005 (calBP)

7.475  80 7.325  65 – 10.860  160 – 10.890  50 11.810  40 11.930  160 12.190  150 12.130  130 12.255  100 12.060  150

8.288  75 8.138  79 – 12.841  145 – 12.847  81 13.718  120 13.927  282 14.265  319 14.168  267 14.371  344 14.109  303

12.270  220 – – – 17.940  200 15.990  260 18.420  530 >9.000 – – – 19.990  480 25.600  4.800 23.400  2.300 21.760  970 24.300  1.400 24.480  1.100 – –

14.428  426 – – – 21.439  505 19.219  296 21.953  756 – – – – 23.862  694 30.012  5.347 25.692  2.532 26.146  1.244 28.865  1.434 29.481  394 – –

33.690  1.195 39.218  1.643 32.770  1.065 38.348  1.384 – – 37.005  1.790 41.617  1.623 >40.000 >43.000 – – 139.900 þ 33.000  22.000 (Th/U) 147.000 þ 9.200  8.500 (Th/U)

TL (ky BP) – – – – – –

Sample

Laboratory

Sites without datesd

Charcoal Charcoal – Charcoal

Ua-18269 Ua-21999 – UBAR-153

Victoria

Rock art (horizon) –

Level 8 (Abrigo 6/Humo) –





Victoria Higuero´n

– 16.438  1.497 17.582  1.521 –

Charcoal

Ua-19443

Carbonates Carbonates –

MAD-3927 MAD-3926 –

Level 9 (Abrigo 6/Humo)

Abrigo 3?/Humo

Navarro

Abrigo 4/Humo Victoria

– – – 18.701  2.154 –

– Bone Flint Charcoal Charcoal

Level 13 (Abrigo 6/Humo)

Higuero´n

Higuero´n

Nerja B–C

– AA-34710 MAD-2405 UBAR-343 UBAR-341 UBAR-342 UBAR-341 UBAR-340

24.344  2.653 26.013  2.777 28.019  2.334 –

Flint Flint Flint –

28.532  5.319 – –

– Charcoal Charcoal

MAD-2559 MAD-2482 Ua-17150 Ua-18050 MAD-2377 Ua-18270 Ua-16859

Stalagmite crust Travertine

CERAK–BAJ/6 CERAK–BAJ/5

Higuero´n? Nerja A Abrigo 3?/Humo Toro Abrigo 4?/Humo









Humo cave complex Abrigo 3/Humo Abrigo 4/Humo Caseta del Guardia Breccias of La Cala



M. Corte´s-Sa´nchez et al. / Quaternary Science Reviews 27 (2008) 2176–2193

Developed Solutrean, c. 16–19 ky

Bj/5? –

14

Table 2 Chrono-cultural and palaeoenvironmental sequence of the coast of Ma´laga during the Upper Pleistocene

SPECMAP

Period

Level of the sea1 (m. s l.)

Heinrich Event AMS MD 95-20433

Distance (km) to the coast Bay of Malaga4

Nerja5

Atlantic -

Holocene

-

-

Preboreal OIS-1 (c. 16-7 ky)

-50 (10.5 ky)

2.5

c. 10.8-7.4 ky Upper Magdalenian c. 11.9-10.8 ky c. 11.9-10.8 ky Upper Magdalenian c. 12.2-11.9 ky

-

-

-

-

Younger Dryas c. 12.5-11.5 ky

-73 (c. 12.5 ky)

3.1

3

Bölling

-90 (c. 14 ky)

5.1

4

16.4-12.2 ky

-

Developped Solutrean c. 17.5-16.4 ky

H1 (c. 16.8 ky)

Ancient Dryas OIS-2 (c. 24-16 ky)

2 -

Epipaleolitihic c. 7.4-7.2 ky

-

-

(Lascaux)

Bajondillo Layer

Phase

3

L

Cueva de Nerja Layer V3

4

Phase

(Dryas Ia) (Laugerie)

-120 (c. 19 ky)

10.5

5.5

(c. 27-16 ky) -

OIS-3.1 Early Cold Stage (c. 37-27 ky)

-

H2 (c. 24 ky)

-

H3 c. 29,1-31,1 ky3

-

-60-70 (50-40 ky)2

H4 c. 38,5-40 ky3 H5 c. 45,4-46,5 ky3

Nerja8 Charcoal

Atlantic (Humid) (

13

4

C, arid)

Erosive Episode -

-

M13-12 V4

Fauna

Phase8

Phase

Nerja 7

-

3

Warm & cold Humid 13 ( C, arid)

2

Mediterranean species (birds, mollusks)+ Boreal species (fishes)

Erosive Episode 5

K

M16 V7-6-5

Nerja 5

Mallaetes E

Erosive Episode 6

J

7

-

Phase G

Térmic Humide ( Mallaetes D

9

I

V8 V9 V10

Erosive Phase

Gravettian c. 27-21,7 ky

10

Aurignacian c. 34-27 ky

11

Transition (c. 37-34 ky)

12 13

F

Middle Paleolithic (c. >40 ky)

14

E

H

G

Nerja 1

-

-

Erosive Episode

-

Mallaetes C

Malaletes A

-

-

(

-

Cova Negra D

C, too)

13

C, too)

+ Refugia area +Cold Arid

-

V11 V12 V13

13

Cold Aride

Nerja 3

-

OIS-3.2 Transitional Stage (c. 44-37 ky)

-

Middle Solutrean c. 19,9-18,7 ky c. 21,7-19,9 ky

East Iberian7

Nerja 9

8 Last Glacial Maximum

Paleobotany Bajondillo 13 Pollen+( C)

(

13

C, too)

+ Refugia area Cold Arid 13 ( C, too) + milder conditions (refugia) -

Mediterranean species (Alectoris rufa Patella caerulea,etc.) + Boreal species (Somateria,Cygnus Clangula, gadids Modiolus,etc.)

1

-

-

-

-

-

-

-

-

-

-

M. Corte´s-Sa´nchez et al. / Quaternary Science Reviews 27 (2008) 2176–2193

Tardiglacial

Alleröd

Lithoestratigraphy Material Culture6

Thermic Humid milder conditions (refugia)

a

Herna´ndez et al., 1996. Shackleton et al., 1984. Sa´nchez et al., 2002. d Corte´s and Simo´n, 2000.. e Ferre et al., 2003. f Aura et al., 2002. g Corte´s, 2002. h Fumanal, 1995. i Badal, 1996. b c

2179

2180

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the Epipalaeolithic and the Neolithic. The sedimentary column incorporates 27 absolute dates (14C/AMS, TL and Th/U) and palaeoenvironmental changes have been analysed using stable carbon isotopes, sediment micromorphology and pollen data. Apart from molluscs, faunal data are almost absent. Nerja (Table 1) features a far shorter sequence (c. 25–4 ky) yet incorporates the richest archaeozoological samples thus far documented for the area, including terrestrial and marine faunas (Boessneck and Driesch, 1980; Pe´rez, 1986; Morales and Martı´n, 1995; Morales et al., 1998; Pe´rez and Raga, 1998; Jorda´ et al., 2003; Vera et al., 2003; Lozano et al., 2003, 2004; Riquelme, 2004; Riquelme et al., 2005/2006), as well as a rich record of human remains (Simo´n, 2003). No data from pollen isotopes or sediment micromorphology are available. Thus, Bajondillo and Nerja are complementary to a certain degree. Isotope measurements of sediment samples from Bajondillo were carried out at the Stable Isotope Laboratory of the Estacio´n Experimental del Zaidı´n (CSIC, Granada). All samples were treated with cold 1:1 HCl to remove carbonates. The carbon isotopic composition of residual organic matter was analysed by means of an EA–IRMS elemental analyser on line with a Delta Plus XL mass spectrometer. The overall precision of the analyses is 0.1& for d13C. The isotopic composition is reported as d values per mil [d ¼ (Rsample/Rstandard  1)  1000] where R ¼ 13C/12C for d13C values, using the PDB international standard. The chrono-cultural sequence has been defined on both technotypological and chronological (AMS, TL and Th/U) grounds and in terms of the relative stratigraphic position of archaeological material. The sedimentary–climatic sequence of Bajondillo is based on field descriptions and micromorphological analysis of 20 sedimentary levels identified in a 6-m deep profile. Pollen samples comprise, for each sample, a minimum of 200 pollen grains and 20 recorded taxa to enable statistically reliable inferences (McAndrews and King, 1976; Sa´nchez, 1993). The relative values for each taxon, whether arboreal, shrub or herbaceous, were obtained from the sum of the absolute values for each taxon referred to a pollen total. This total does not include hygrophilous taxa, cryptogam spores, undetermined pollen grains or Cichorioidea because of their hypothetical overrepresentation in sedimentary deposits due to their zoophyllic character (Carrio´n, 1992). Faunal remains were identified with the reference collections from the Natural History Museum at Madrid (birds), the Museum of Palaeontology at Estepona (molluscs) and the private collection of Arturo Morales housed at the Laboratorio de Arqueozoologı´a (Universidad Auto´noma, Madrid). The analytical protocols follow the conventions described in general works (e.g., Reitz and Wing, 1999). 4. Results 4.1. Chrono-cultural sequence The archaeological industries are comparable to those from other Mediterranean sites in the Iberian Peninsula. Artefacts associated with OIS 5–3 deposits, for example, correlate with those of the regional Middle Palaeolithic characterized by industries of typical Mousterian type (Corte´s, 2006). Deposits with Upper Palaeolithic industries are organized according to the following scheme (Aura, 1995; Corte´s, 2007, Table 1): (a) Aurignacian (c. 34–26 ky). Only present in one level at Bajondillo (Bj/11). (b) Gravettian (c. 26–21 ky). Present at Bajondillo and Nerja but possibly also in other caves in the Bay of Ma´laga area (e.g., Abrigo 4/Humo).

(c) Solutrean (c. 21–16 ky). Documented in no less than six sites featuring two different phases, an older one with industries of a full Solutrean type, and a younger phase corresponding to an evolved Solutrean. (d) Magdalenian (c. 16–10 ky). Five stratigraphic sequences incorporate an Upper Magdalenian. (e) Epipalaeolithic (c. 10–7 ky). Four sites show evidence of these last hunter-gatherers. In addition, cave art is recorded in five sites, three of them with human occupations from the Upper Pleistocene: Victoria, Higuero´n and Nerja, the latter with AMS dates; and Toro and Navarro, which have not yet been surveyed. Chrono-stylistic data indicate that all the cave art dates back to the LGM coinciding with the development of the Solutrean although its earliest manifestations could reach back into the late Gravettian (Villaverde, 2006). 4.2. Palaeoenvironmental records 4.2.1. Isotopes Values of d13C at Bajondillo range between 22.3 and 26.8&. Two relatively well-defined sections seem to account for this variability: samples older than 20 ky average 23.2&, whereas the younger samples give an average of 25.2&. More positive values (>24&) are related to a reduction of plant CO2. During arid periods plants close their stomas to avoid water loss through evapotranspiration thus increasing WUE (water use efficiency) values. Plant uptake of CO2 is lower, with less discrimination for 12C. Consequently, tissues from water-stressed plants show an increase in d13C values (Farquhar et al., 1982; Ehleringer, 1991; Pate, 2001). Thus, the relatively more positive values in the earlier part of the sequence, between 40 and 20 ky, indicate relatively arid conditions, whereas the more negative values in the later part of the sequence comprising the transition from the LGM to the Holocene indicates a reduction in aridity (Fig. 4). This is in agreement with global climatic trends. From level Bj/8 onwards there is a decline of d13C to 26.8 & in Bj/6, indicating lower water-stress and canopy effects, reinforced by an increased concentration of atmospheric CO2 from the end of the LMG by about 100 ppmv (Petit et al., 1999). Bajondillo levels Bj/5 and Bj/3 show a renewed increase in arid conditions though less arid than during the 40–20 ky maximum. These data indicate a general increase in humid conditions in the southern Iberian Peninsula resulting from the combined effects of a rise in temperatures or tempera-tures and sea level on a global scale (Dansgaard et al., 1993). 4.2.2. Sedimentary record Bajondillo (Bergada` et al., 2005) provide sequences of sediments that record sedimentary, pedological and anthropogenic processes. Sedimentary processes include: biochemical processes such as the accumulation of calcium carbonates and travertines; accumulation of detrital processes resulting from collapse of the roof and walls of the cave; remobilisation of material by pluvial and colluvial action; and aeolian inputs. Pedological processes are controlled by the physico-chemical nature of the environment and include dissolution, chemical and physical precipitation, structural deformation and eluviation caused by cycles of freezing and thawing, percolation of fine materials (silts, clays and charcoal particles) producing secondary accumulations; and biological activities. Anthropogenic processes incorporate allochthonous material, both inorganic materials (lithics, fragments of pottery, flints, etc.), organic animal (bones, shells, phosphatic masses) and organic botanical material (charcoal, phytoliths and ash). The preservation of such anthropogenic materials depends on the nature and intensity of the sedimentary and pedological processes to which they are subjected.

M. Corte´s-Sa´nchez et al. / Quaternary Science Reviews 27 (2008) 2176–2193

The sediments at Bajondillo were very homogeneous (Table 3). The coarse fraction was mostly of local origin resulting from the in situ fragmentation of travertine accumulations and carbonated deposits. In the Epipalaeolithic horizons there were allochthonous materials, including some igneous stone clearly brought in by human activity. The sandy fraction likewise exhibited a local origin caused by dissolution and fragmentation of materials by freeze-thaw cycles. The silt–sand fraction incorporated materials brought in by the action of rain and wind. The Late Upper Pleistocene to Early Holocene sequence is as follows (Table 3): Bajondillo P (Bj/14; >40 ky): morphogenesis dominated by travertine accumulations although these later become fragmented by freeze-thaw cycles. The upper part of this level shows some remobilisation caused by diffuse rain wash. Between 37 and 10 ky the growth of the travertine platform is interrupted and records the following episodes: Bajondillo Q (Bj/13 and Bj/12): rain wash, remobilisation of travertines and restricted incorporation of fine windblown materials. Bajondillo R (Bj/11): increased windblown material, indicating a harsher environment with higher aridity, lower temperatures and reduced tree cover. At some point after human occupation, gelifluction affected the sediments. Bajondillo S (Bj/10): freeze-thaw cycles accentuated causing a massive fall of boulders, retreat of the cave roof and a change in the morphology of the sediments, indicating a cold environment. Bajondillo T (Bj/9 and Bj/8): conditions deteriorated to those recorded for Bajondillo R. The reduction of tree cover facilitated aeolian inputs, though probably not as much as in the earlier stage. Once again, following cessation of anthropogenic activity, gelifluction processes set in, probably reaching a higher intensity than that recorded in Bj/11. Such conditions slowly ameliorated, already evident in Bj/8. Bajondillo U (Bj/7 and Bj/6): milder conditions with higher temperatures and humidity. This is seen in the formation of

2181

speleothems and in the epigenization of the limestones into phosphates, though winters remained cold. Bajondillo V (Bj/5): initially similar to the previous episode, but indicating subsequent deterioration and colder temperatures in comparison to Bajondillo U. Bajondillo W (Bj/4 and Bj/3; 7.3–7.4 ky): re-appearance of travertine formation. In the final stage there were inputs from low Tra-vertine or traver-tine to medium intensity rain wash. Conditions were temperate. From the above sequence and Nerja (Jorda´, 1986; Jorda´ et al., 1990; Aura et al., 2002) we can identify six major horizons, applicable to the Ma´laga coast area as a whole: (1) Middle/Upper Palaeolithic transition (c. <40–33.6 ky), subdivided into three stages: (1a) A temperate climate that later became colder with a slight decrease in humidity, corresponding to Bajondillo P (Bj/14). (1b) Renewal of milder conditions though with a slight increase in aridity, corresponding to Bajondillo Q (Bj/13, Bj/12). (1c) An erosional event detected in the contact zone between Bj/13–12 and Bj/11. (2) Aurignacian (c. 33.6–27/26 ky): a cold and arid climatic episode with frequent aeolian inputs, represented by Bajondillo R (Bj/11). Following the formation of this horizon conditions became more humid. (3) Gravettian (c. 27/26–21.7 ky): a cold phase with renewed aridity, corresponding to Bajondillo S (Bj/10). This horizon is also documented at Nerja, where it represents the beginning of the sequence (V/13–11 and M/19–17), with diffuse rain wash events caused by surface run off, as well as falls of boulders within a cold and humid climate setting. (4) Solutrean (c. 21.7–16.4 ky): subdivided into four stages: (4a) c. 21.7–19.9 ky: hiatus at both Bajondillo and Nerja. (4b) c. 19.9–18.7 ky: climate returned to cold and arid though not as extreme as for Bajondillo R, with conditions becoming moister at the end of this phase, corresponding to Bajondillo T (Bj/9–8).

Table 3 Sedimentological sequence of Bajondillo Cave, Thin sectioning and X-ray diffraction procedures evidence that the dominant minerals are quartz, calcite dolomite and aragonite

Level Bj/3 Bj/4 Bj/5

Episode

Facies

Bajondillo W

Carbonated accumulation (travertine) Gravel and sands

Bajondillo V

Bj/6

Colour 10 YR 5/2 10 YR 7/3 10 YR 3/1 10 YR 5/4

Bajondillo U Bj/7 Bj/8 Bajondillo T Bj/9

Cobbles, gravel and silty sands Silty sands with cobbles and blocks

Bj/10

Bajondillo S

Bj/11

Bajondillo R

Bj/13

Bajondillo Q

Gravel, cobbles and blocks with silty sand Sandy silt with and some cobbles Silty sands with gravel

Bj/14

Bajondillo P

Travertine cobbles

10 YR 3/2 10 YR 6/3 Bj/9a 10YR 4/2 Bj/9b 10YR 5/1 10 YR 6/3

Texture Bj/3a. Gravel and carbonate sandy Bj/3b. Carbonated sandy silt Carbonated accumulation Tabular gravel and sands Gravel , sands and some block Cobbles, gravel and sands Silty sand with gravel and some cobbles Sandy silt with gravels Silty sand with cobbles and blocks

10 YR 6/3

Gravel and cobbles with blocks and silty sands Sandy silt with gravels and cobbles Coarse sand with gravel

10YR 6/4

Travertine cobbles with sands

10 YR 3/1

Microstructure Granular/Fisure Massive/Granular Massive/Bridged grain Single grain/ Bridged grain Intergrain microaggregate/ Bridged grain Bridged grain/ Granular Granular/Fissure Granular/Spongy Granular/Bridged grain Granular/Platy Massive Granular/Bridged grain Single grain/ Intergrain microaggregate

Depositional conditions Medium intensity rain washes ( Bj/3a) Travertine Diffuse rain wash Frost shattering Non channelled rain wash Falls of blocks (+) Solifluction (gelifluction) (-) Eolian input/

Falls of blocks Frost shattering Solifluction (Gelifluction) Increased eolian input Diffuse rain wash Beginning eolian input Frost shattering Travertine

2182

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(4c) c. 18.4–17.9 ky: humid and temperate at the beginning with hints of colder and probably more arid conditions in the uppermost level, corresponding to Nerja stage 3 (V/10, V/9, V/8). (4d) c. 17.5–16.4 ky: progressively milder conditions culminating in the Bajondillo U episode (Bj/7 and Bj/6). (4e) c. 16.4–12.2 ky: hiatus at both Bajondillo and Nerja. (5) Magdalenian (c. 12.2–11.9 ky): this horizon is characterized by a cold pulse with evidence of modification by ice and diffuse rain wash. This horizon is documented only at Nerja (M/16 and V/7–5), and corresponds to Nerja phase 5. Although the Bajondillo V episode (Bj/5) lacks dates and a specific cultural assignment, it shows similar evidence of climatic deterioration, and this along with position within the overall sequence suggest a correspondence with this horizon. Between c. 11.9 and 10.8 ky, both Bajondillo and Nerja exhibit a hiatus. (6) Epipalaeolithic (c. 10.8–7.2 ky): subdivided into three stages: (6a) c. 10.8 ky: temperate or warm climate, and essentially dry with a regime of rainfall associated with storms. This horizon marks the beginning of the Holocene and is documented only at Nerja, corresponding to Nerja phase 7 (M/13–12 and V/4). (6b) c. 10.8–7.4 ky: an erosional an hiatus at both Bajondillo and Nerja. (6c) c. 7.4–7.2 ky: temperate conditions, corresponding to Bajondillo W (Bj/4–3) and Nerja 9 (M/11 and V/3). 4.3. Flora From a biogeographic standpoint the area is located in the thermo-Mediterranean zone of the Andalusian Province (i.e., the Malacitano-Alimjarense sector), with a corresponding vegetation of oak woods (Smilaci mauritanicae–Quercetum rotundifoliae), characterized by the predominance of the Holm oak (Quercus rotundifolia) along with an abundance of wild olive trees (Olea europaea subsp. sylvestris) and other thermophilous elements such as Chamaerops humilis, Pistacia lentiscus, Smilax aspera, Quercus coccifera and Rhamnus lycioides (Martı´nez and Peinado, 1987; Rivas, 1987). The botanical uniqueness of this zone is due to its recent Quaternary history, its calcareous landscape, the influence of the

coast, and the absence of deep soils (Cabezudo and Pe´rez, 2001). The history of the vegetation of the coast of Ma´laga was previously limited to a series of pollen analyses obtained from oceanic cores from the Sea of Alboran (D’Errico and Sa´nchez, 2003), and charcoal analyses from Nerja (Badal, 1996). Pollen analyses at Bajondillo have significantly augmented this record, particularly with reference to the Ma´laga coast during the end of the Pleistocene, and can be summarised in the following sequence (Fig. 2): Bajondillo Bj/19–14 (c. 110 ky (Th/U)–40 (AMS) ky), corresponding to the Middle Palaeolithic: extremely variable from the standpoint of climate, alternating warm and humid phases with colder and more arid episodes, leading to repeated fluctuations between forested and steppe conditions. However, even during the cold and arid phases thermophilous taxa persisted (Carrio´n et al., 2003). Bajondillo Bj/13–12, Middle–Upper Palaeolithic transition: recovery of mesic arboreal vegetation as well as Abies pinsapo, indicating moister interstadial conditions coincident with the end of OIS 3. Resilient thermophilous taxa survive as floral refugia. This alternation between warm/humid floras and colder/more arid ones continues during the Upper Palaeolithic. Bajondillo (Bj/11), Aurignacian: regression of forest cover, especially deciduous forest, even though both the evergreen formations of Quercus and the xerophilous maquis remain in the area, along with the thermophilous Whitania and Selaginella denticulata. From these data one can infer that the climate was essentially arid. Available dates from Bj/11 indicate a correlation with two cold pulses associated with Heinrich events 4 and 3 (c. 34–32 ky and 28–26 ky/AMS, respectively). The latter (i.e., Lazio VI–VII; 29.4–25.9 ky/AMS, Allen and Huntley, 2000) corresponds to a near-terminal stage of OIS 3 characterized by a warm steppe vegetation. Bajondillo Bj/10 (c. 26–21 ky, AMS and TL): the onset of a colder climate with a sharp increase in grass cover and a marked reduction in tree cover and maquis, but with continued presence of Whitania and Selaginella denticulata, indicating the persistence of isolated pockets of thermal floras. The TL dates from Bj/10 allow us to place this level

Fig. 2. Pollen diagram from Cueva Bajondillo (Bj/14–Bj/3).

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within the Pleniglacial (OIS 2), characterized by cold steppe vegetation. Bajondillo (Bj/9–7), Solutrean: similar to Bj/10, reinforcing the scenario of an arid and cold climate, though the slight recovery in the pollen values of Abies pinsapo and Alnus signals the onset of moister conditions. The presence of xerothermophilous elements, in particular S. denticulata, Lycium and Cosentinia, suggests that, although still cold and arid, the area of Bajondillo witnessed the continued persistence of isolated elements of a thermal flora. It is therefore possible to postulate the existence of local refugia for certain thermophilous species during the Upper Wu¨rm Pleniglacial (OIS 2). Bajondillo Bj/6 and Bj/5: recovery of arboreal taxa, especially Betula, of particular interest given its pioneer character in open areas, and A. pinsapo and Ilex, both probably reflecting conditions of increased rainfall. The pollen records also show a recovery of the thermophilous maquis and an important development of pine, mainly in the coastal areas near Bajondillo, but also in the foothills of the nearby mountains. Both levels may possibly reflect warmer and more humid conditions. Bj/6 exhibits an advanced Solutrean industry, and taking into account the TL chronology obtained for Bj/7, it is likely that Bj/6 was deposited toward the end of the Solutrean (c. 17–16 ky), coinciding with a phase of climatic amelioration that followed the LGM. The gap between Bj/6 and Bj/5 is marked by an erosional episode. Bajondillo Bj/5: higher Pinus values, tentatively assigned to the Bo¨lling/Allero¨d interstadial. Bajondillo Bj/4 and Bj/3 (c. 8 ky), Epipalaeolithic: a relatively dense Quercus forest landscape with a rich canopy (especially for Bj/4), developed under humid conditions, that correspond to the Atlantic Period. The coastal location of the site continues to allow for the presence of pine groves and a rich xerothermophilous maquis. Oak/kermes forest and oak/gall oak forest would have been the dominant vegetation in the area, the former in more arid areas, the latter on higher, north-facing slopes along watercourses. Charcoal analyses carried out at Nerja by Badal (1996), even allowing for the potential biases associated with the selective effects of human collection of firewood, essentially corroborate the pollen data from Bajondillo. Nerja V/13–V/8, Gravettian and Solutrean (c. 24.3–17.9 ky) shows very high percentages of Pinus nigra and Fabaceae, as well as the presence of thermal taxa like Rhamnus–Phillyrea and Cneorum tricoccum. The abundance of P. nigra is significant insofar as it is characteristic of the present-day coastal zone where Nerja and Bajondillo are located. For this reason Badal (1996, p. 172) argues that the palaeoclimatic conditions at the beginning of the Upper Palaeolithic would have been quite similar to those present in the area today. This means a dry or semi-arid climate corresponding with that of Bj/10 (Tables 1 and 2). In the Middle–Upper Solutrean of Nerja OIS 3, the vegetation, though still reflecting arid conditions, hints at warmer conditions, as indicated by the presence of both Pistacia and Cneorum. Levels V9 and V8 could be equivalent in palaeovegetational terms to the Solutrean of Bajondillo. In the Magdalenian levels (FA-5, Table 2), only V7 provides botanical information. This evidences the dominance of the Fabaceae along with a sparse presence of P. nigra and certain thermal elements like Quercus ilex–coccifera, Rhamnus–Phillyrea and C. tricoccum. No botanical information from this cultural period is available for Bajondillo. The Epipalaeolithic of Nerja (V4) is characterized by the appearance and relative abundance of Olea and Fabaceae, possibly a response to the climatic improvement that took place at the beginning of the Holocene. As such, this reinforces the picture that emerges from the pollen data at Bajondillo.

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4.4. Fauna From a palaeoenvironmental standpoint, as with charcoal, the archaeozoological record is subject to selective bias by human activities. This applies both to subsistence and to non-nutritional uses, for example, ornaments, where long-distance transport needs to be taken into account. There are two additional limitations to keep in mind when evaluating prehistoric faunas from the coast of Ma´laga: 1. The emergence of a coastal plain during periods of eustatic marine regression not only moved the shore away from the sites under discussion but also created an essentially sandy shore very different from the present-day rocky one. 2. Apart from Nerja, there is really no other local faunal record to compare with the palaeoenvironmental and botanical data. The uniqueness of the Nerja sample makes it difficult to judge to what extent the faunal patterns there are representative of the wider area or merely particularisms.

4.4.1. Molluscs During the earliest stages of the Upper Palaeolithic (i.e., Gravettian), mollusc remains appear exclusively in connection with ornamental uses, a practice that continues during the Upper Palaeolithic and Epipalaeolithic stages of the Nerja sequence (Corte´s et al., 1996). In addition to living taxa, ornamental shells at Hoyo de la Mina-6 include specimens of fossil scaphopods from the Pliocene outcrops of the Ma´laga basin (Vera et al., 2004). Although the oldest Upper Palaeolithic records of mollusc consumption on the coast of Ma´laga date back to the Gravettian of Nerja (V/11) and Bajondillo (Bj/10), such a practice has also been documented for the Middle Palaeolithic of the area, namely at Abrigo 3 y 4 of the Humo complex, at Bajondillo Bj/19 (OIS 5) and on Gibraltar in Level IV of Gorham’s cave (Barton, 2000; Aura et al., 2001; Vera et al., 2003; Fa, in press; personal communication). The data from Nerja show preferential harvesting of terrestrial molluscs during the Gravettian. In the Solutrean, marine molluscs become equally abundant, and eventually, in Magdalenian and Epipalaeolithic assemblages, the dominant group (Tables 7 and 8). Preferential gathering of marine molluscs appears typical of the Magdalenian more generally in the region, at the sites of Bajondillo, Abrigo 6 of the Humo complex and at Hoyo de la Mina. At Bajondillo, mollusc accumulations have been found in a chronostratigraphically problematic level (Bj/5 corresponding to the Bajondillo V episode), where pollen and archaeological features tentatively suggest a date within the Bo¨lling/Allero¨d interstadial (see above). The collection from Abrigo 6 of the Humo complex, on the other hand, comes from a test pit and incorporates two samples (Ramos et al., 2006; Corte´s et al., in preparation): a Magdalenian deposit, featuring 20 marine species, as well as barnacles (Balanus sp.) and sea urchins (Paracentrotus lividus); and an Epipalaeolithic sample, with 15 species of marine molluscs. In both instances, and except for the scallop Pecten maximus (Magdalenian), such evidence indicates a preferential harvesting of rocky shore taxa – Patellidae (limpets) represent 72% and 81% of the MNIs in the Magdalenian and Epipalaeolithic, respectively – in particular from the mesolittoral and upper infralittoral zones. At Nerja, the pattern is different. The richest assemblage, both in terms of diversity and abundance (NISPs), is that from the Epipalaeolithic (Tables 4–6). This ‘‘peak’’ is the culmination of a trend that started for molluscs in particular and for marine taxa in general at the end of the Solutrean (Tables 7 and 8). From the standpoint of harvested habitats at Nerja, rocky shores dominated during the Epipalaeolithic (42% of the species, >90% NISP) due to the dramatic rise of mussels. Yet during the Magdalenian, representation of

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Table 4 Faunal assemblages from the Last Glacial Maximum stage at Cueva de Nerja Last Maximum Glacial/Gravettian þ Solutrean (c. 24–17.5 ky BP) Mammals

NISP

Birds

NISP

Fishes

NISP

Molluscs

NISP

Equus sp. Bos primigenius Cervus elaphus Capreolus capreolus Capra pyrenaica Sus scropha Monachus monachus Lynx pardina Felis silvestris Crocutta sp. Oryctolagus cuniculus Erinaceus europaeus Apodemus sylvaticus Apodemus flavicollis Microtus cabrerae Arvicola sapidus Rodentia indet.

23 5 352 1 3.767 28 8 15 16 2 9.622 1 1 – – – 2

Sula bassana Calonectris diomedea Alectoris rufa Clangula hyemalis Somateria sp. Cygnus cygnus/olor Branta bernicla B. bernicla/rufficolis Anser albifrons Alca torda Pinguinus impennis Columba livia/oenas Athene noctua P. pyrrhocorax Corvus corax Pyrrhocorax/Corvus

7 1 1 1 1 1 6 2 1 1 5 7 1 2 1 4

Acipenseridae Salmonidae Congridae Gadidae Zeidae Moronidae Carangidae Sparidae Labridae Mugilidae Scombridae Triglidae

2 1 1 370 4 1 84 518 79 7 19 1

Antalis inaequicostatum Antalis vulgare Mytilus galloprovincialis Modiolus modiolus Ostrea edulis Pecten maximus Pecten jacobaeus Rudicardium tuberculatum Cerastoderma edule Venus verrucosa Callista chione Tapes decussatus Patella caerulea Patella ferruginea Patella intermedia Patella nigra Patella rustica Patella ulyssiponensis Patella sp. Gibbula richardi Gibbula sp. Monodonta turbinata Littorina sp. Melanopsis laevigata Melanopsis sp. Charonia lampas Nucella lapillus Stramonita haemastoma Cyclope pellucida Siphonaria pectinata Rumina decollata Sphinterochila hispanica Iberus alonensis Iberus marmoratus Otala lactea

5 1 271 5 4 21 1 6 88 1 1 58 105 3 41 14 9 27 25 71 1 5 1 2 12 1 5 1 1 2 25 3 2626 4 3

17 Taxa

13,843

16 Taxa

42

12 Families

1083

35 Taxa

3415

Combined data taken from Boessneck and Driesch, 1980; Pe´rez, 1986; Morales and Martı´n, 1995; Pe´rez and Raga, 1998; Jorda´ et al., 2003; Vera et al., 2003; Lozano et al., 2003, 2004; Arribas et al., 2004; Riquelme, 2004; Riquelme et al., 2005/2006 and from unpublished data (Rosello´, in preparation; Sa´nchez, in preparation). Data on fishes are preliminary.

molluscs from rocky and sandy-muddy shores was quite similar (37% and 34% in terms of taxa), although rocky shore species still dominated the bulk of the assemblage in terms of NISP (i.e., 70.5% vs. 12.5%). These characteristics are similar to those recorded for the Solutrean (Tables 3 and 4). A distinctive feature of all mollusc assemblages is the drastic decrease of terrestrial taxa with time (85% for the Solutrean, 20% for the Magdalenian and 7% for the Epipalaeolithic), indicating a more general pattern (Tables 3–5) (see below). Such pattern is to be seen, albeit less marked, within the Solutrean sequence from the Vestı´bulo sector (Table 8). Here, the increase in the number of marine taxa from the earliest Solutrean levels through to the Epipalaeolithic was explained in terms of the approaching coastline (Morales et al., 1994, 1998; Aura et al., 2001, 2002), whereas the shift from sandy to rocky shore taxa was taken to be a reflection of the changing character of the coastline as the existing coastal plain was gradually flooded (Serrano et al., 1995, 1997; Vera et al., 2003). A straightforward biogeographical interpretation of the Nerja mollusc assemblages is far from clear due to the impossibility of evaluating the biasing effects caused by human harvesting activities. Still, from a strictly biogeographical standpoint, the majority of the species were Mediterranean in character at all times. Noteworthy is the absence of the most Atlantic of the limpets (i.e., Patella vulgata), which constituted 40–80% of the patellids at nearby Gorham’s cave from the Middle Palaeolithic through Phoenician times (Fa, personal communication).

The complexities involved in interpreting the patellids in biogeographical terms are illustrated by the mollusc samples from Nerja. At all times, the more Mediterranean species (i.e., Patella caerulea) constitute the main component of the assemblages. Still, it is peculiar that the combined values of these taxa (i.e., P. caerulea or caerulea þ Patella ferruginea) decrease from the Solutrean (54.5% of the combined patellid NISP) to the Epipalaeolithic (40%) while the less sensitive species such as Patella rustica increase (4.5% LGM, 16% Tardiglacial, 17.5% Early Holocene). Given that the latest stages of the Tardiglacial at Nerja appear to coincide with the cold pulse of the Younger Dryas, it seems clear that the trend in the patellids neither reflects such an event nor the ensuing warming episode that took place during the Early Holocene. Of the cold adapted taxa, only some very few remains of the circumboreal mussel Modiolus modiolus are evidence of a sharp decline in sea surface temperatures (in fact, this species ‘‘peaks’’ during the milder Solutrean, not the Magdalenian) (Tables 4 and 5; Lozano et al., 2004). The presence in those same levels of warm water bioindicators such as Patella nigra and Siphonaria pectinata, prevalent throughout the sequence, indicates the time-averaged nature of the deposits. Under such circumstances, each one of the levels at Nerja should be taken as a palimpsest incorporating a multitude of shorter-term events, both cold and warm, which cannot be averaged to produce the sea temperature of a specific moment. Despite this it seems evident that, overall, warm water indicators were more prevalent than those of cold water at all times.

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Table 5 Faunal assemblages from the Tardiglacial stage at Cueva de Nerja Mammals

NISP

Birds

NISP

Fishes

NISP

Molluscs

NISP

Equus sp. Bos primigenius Cervus elaphus Capra pyrenaica Sus scropha Monachus monachus Delphinus delphis Cetacea indet. Lynx pardina Felis silvestris Meles meles Oryctolagus cuniculus Lepus sp. Eliomys quercinus Myotis myotis Miniopterus schreibersii Rodentia indet.

1 1 252 2.467 69 57 41 1 31 26 1 9.427 7 3 6 27 8

Gavia stellata Sula bassana Phalacrocorax carbo Phalacrocorax aristotelis Puffinus griseus Puffinus yelkouan Calonectris diomedea Alectoris rufa Melanita nigra Melanita fusca Tadorna tadorna Tadorna ferruginea Anas platyrhynchos Anas clypeata Anas crecca Aythya nyroca Aythya ferina/fuligula Branta bernicla Anser sp. Milvus milvus Buteo buteo Circaetus gallicus Falco tinnuculus Grus grus Fulica atra Larus canus Larus marinus Larus fuscus Sterna sandvicensis Uria aalge Alca torda Pinguinus impennis Columba livia/oenas Bubo bubo Hirundo rustica Alauda arvensis Monticola solitarius Turdus sp. Lanius excubitor C. coccothraustes Pyrrhocorax pyrrhocorax Corvus corax

1 35 3 4 14 2 10 8 1 2 2 2 11 1 2 1 1 3 1 2 1 2 1 1 3 3 2 2 1 9 2 8 24 1 1 1 6 2 2 1 2 2

Acipenseridae Clupeidae Belonidae Gadidae Serranidae Carangidae Sparidae Labridae Sphyraenidae Mugilidae Scombridae Triglidae Scorpaenidae

31 3 358 389 45 466 2.019 191 1 388 377 9 1

Mytilus galloprovincialis Modiolus adriaticus Ostrea edulis Pecten maximus Anomia ephippium Rudicardium tuberculatum Cerastoderma edule Cerastoderma glaucum Mactra stultorum Venus verrucosa Callista chione Tapes decussatus Patella caerulea Patella ferruginea Patella intermedia Patella nigra Patella rustica Patella ulyssiponensis Patella sp. Gibbula richardi Gibbula sp. Monodonta turbinata Jujubinus exasperatus Melanopsis laevigata Melanopsis sp. Turritella monterosatoi Charonia lampas Nucella lapillus Stramonita haemastoma Nassarius corniculus Siphonaria pectinata Rumina decollata Sphinterochila hispanica Iberus alonensis Iberus marmoratus Caracollina lenticula Otala lactea Helicidae

1219 1 4 48 1 7 270 1 1 1 1 1130 240 1 153 25 98 83 151 86 2 18 1 3 90 2 1 60 10 1 2 72 50 713 17 1 17 54

17 Taxa

12,425

42 Taxa

183

13 Families

4278

37 Taxa

4545

Combined data taken from Boessneck and Driesch, 1980; Pe´rez, 1986; Morales and Martı´n, 1995; Pe´rez and Raga, 1998; Jorda´ et al., 2003; Vera et al., 2003; Lozano et al., 2003, 2004; Riquelme, 2004; Riquelme et al., 2005/2006 and from unpublished data (Rosello´, in preparation; Sa´nchez, in preparation). Data on fishes are preliminary. In addition one remain of the European pond tortoise (Emys orbicularis) has been identified.

4.4.2. Fish The fish collection from Nerja, numbering in the thousands of specimens, is the richest ever found in the Iberian Peninsula, yet this data set is at present also the least studied compared to the other faunal remains of Nerja (Morales et al., 1994; Rosello´ et al., 1995). These samples from the Vestı´bulo sector, constituting some 90% of these collections, are either under study (Rosello´, in preparation) or else have been published in a most superficial way (Aura et al., 2001, 2002; Jorda´ et al., 2003). The first and most remarkable result is that marine fishing was already taking place during the Solutrean period. This pushes the onset of fishing activity in Europe, previously not recorded before the Tardiglacial (i.e., Magdalenian) (CleyetMerle, 1990; Rosello´, in preparation), back to the LGM. Equally importantly, fishing and shellfish harvesting show a gradual increase within the Solutrean (Table 8), a trend that constitutes part of a larger scale phenomenon of marine resource exploitation at Nerja (Tables 4–7). Although such trends may merely reflect circumstantial contingencies (i.e., an approaching coastline with a concomitantly greater chance of activities previously carried out on the beach now leaving their signature in the cave

deposits), the taxonomic composition of the fish assemblages is particularly informative also from a palaeoenvironmental standpoint. Of particular interest here is the relative abundance of cold water, often boreal, codfish species (Gadidae). Prominent among them are the Saithe (Pollachius virens), Cod (Gadus morhua), Haddock (Melanogrammus aeglefinus) and Ling (Molva molva) although the most common species at all times was the more temperate though still Atlantic Pollock (Pollachius pollachius) (Rodrigo, 1994; Rosello´, in preparation). Gadid abundances do not behave as shown in Tables 4–6, where their numbers drop to a mere 9% during the Tardiglacial. Our preliminary data instead indicate that gadid numbers comprise some 35% of fish NISPs in the LGM, then rise to 78% during the Tardiglacial, remaining stable during the Early Holocene (i.e., 79%; Rosello´, in preparation). The reasons for such ‘‘behaviour’’ are far from clear, since boreal taxa do not show any decline during this last stage (c. 10.8 ky), even though records from Bajondillo indicate a quite mild climate (see Section 4.2.2). One is therefore forced to consider less parsimonious explanations involving a preferential harvesting of codfish when sea temperatures dropped, a real

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Table 6 Faunal assemblages from the Early Holocene stage at Cueva de Nerja Mammals

NISP

Birds

NISP

Fishes

NISP

Molluscs

NISP

Bos primigenius Cervus elaphus Capra pyrenaica Sus scropha Monachus monachus Delphinus delphis Lynx pardina Felis sylvestris Vulpes vulpes Oryctolagus cuniculus Myotis myotis Miniopterus schreibersii Rhinolophus ferrumequinum Rodentia indet.

1 14 344 10 43 1 9 5 3 1632 8 1 3 5

Sula bassana Phalacrocorax aristotelis Puffinus griseus Calonectris diomedea Ardera cinerea Alectoris rufa Coturnix coturnix Melanita nigra Melanita fusca Mergus serrator Tadorna tadorna Anas platyrhynchos Aythya nyroca Aythya ferina Hieraaetus fasciatus Aquila adalberti Accipiter/Aquila Falco subbuteo Grus grus Fulica atra Catharacta skua Larus rudibundus Larus marinus L. cachinnans/fuscus Uria aalge Alca torda Pinguinus impennis Columba livia/oenas Hirundo rustica Luscinia megarhynchos Monticola solitarius Turdus merula Sturnus sp. P. pyrrhocorax Pyrrhocorax graculus Corvus corone Pyrrhocorax/Corvus

18 1 1 13 1 4 2 2 1 1 8 11 1 1 1 1 1 1 1 2 1 1 2 3 6 1 12 26 2 1 18 1 1 7 1 2 1

Acipenseridae Clupeidae Belonidae Gadidae Serranidae Moronidae Carangidae Sciaenidae Sparidae Labridae Mugilidae Scombridae Triglidae Scorpaenidae

5 175 11 1984 21 4 59 2 956 303 108 33 9 2

Antalis inaequicostatum Glycymeris sp. Mytilus galloprovincialis Ostrea edulis Pecten maximus Anomia ephippium Rudicardium tuberculatum Cerastoderma edule Ensis ensis Irus irus Tapes decussatus Patella caerulea Patella ferruginea Patella intermedia Patella nigra Patella rustica Patella ulyssiponensis Patella sp. Gibbula richardi Monodonta turbinata Monodonta sp. Melanopsis laevigata Melanopsis sp Turritella communis Charonia lampas Nucella lapillus Stramonita haemastoma Columbella rustica Mitra nigra Conus mediterraneus Siphonaria pectinata Rumina decollata Sphinterochila hispanica Iberus alonensis Iberus sp. Otala lactea Helicidae

2 5 9914 2 85 1 2 93 1 1 55 132 3 44 17 60 84 15 29 13 2 1 33 1 2 67 5 4 1 2 4 45 6 759 1 5 16

14 Taxa

2079

37 Taxa

157

14 Families

3672

37 Taxa

11,515

Combined data taken from Boessneck and Driesch, 1980; Pe´rez, 1986; Morales and Martı´n, 1995; Pe´rez and Raga, 1998; Jorda´ et al., 2003; Vera et al., 2003; Lozano et al., 2003, 2004; Riquelme, 2004; Riquelme et al., 2005/2006 and from unpublished data (Rosello´, in preparation; Sa´nchez, in preparation). Data on fishes are preliminary. In addition three remains of the Mediterranean terrapin (Mauremys caspica) have been identified.

possibility given the long-term nature of the deposits (see previous section) but impossible to verify unless these two parameters can be proved to correlate with each other. If all the taxa recorded for each of the levels did indeed coexist, then the Mediterranean fish add a new dimension to this peculiar fish community, for sea bream (Sparidae), the second most important group at Nerja, are nowadays scarce where gadids flourish and vice versa. Also peculiar is the fact that, as was the case with the limpets, sparids become scarcer with time (Tables 4–6), when one would, in fact, expect the opposite to hold. Obviously, there might exist both methodological and cultural biases affecting these abundances and producing misleading results. Among the former, sparids feature a far smaller number of vertebrae (24) than gadids (up to 40), but even a standardized quantification of remains, or incorporation of the minimum numbers of individuals as the abundance estimator, would not change the dominant

position held by codfish at all times, nor the trend of their increasing abundance with time in the Vestı´bulo area (Rosello´, in preparation). Preferential fishing of codfish may have served to exaggerate their presence but this is very hard to prove or to disprove with the data at hand. Another line of evidence that may reinforce the episodic character of the fish accumulations is that, in contrast with the molluscs and mammals, the heterogeneity of the fish assemblages is remarkable to the extent of being disturbing. Even within the Vestı´bulo assemblages, for example, the analyses by Aura et al. (2001, 2002) and Jorda´ et al. (2003) reveal an abundance of medium sized taxa such as the grey mullet (Mugilidae) and needlefish (Belonidae), which our studies have thus far been unable to detect (Rosello´, in preparation). It might be that some of this fishing also took place at different times of the year and from such a standpoint the needlefish as well as the jacks (Carangidae)

Table 7 Contributions (%NISP) of the main faunal groups to the assemblages from the three main stages of Nerja (Vestı´bulo sector)

Table 8 Contributions (%NISP) of the main faunal groups in the various Solutrean levels from Nerja (Vestı´bulo sector)

Faunal group

1 (%)

2 (%)

3 (%)

Faunal group

X (%)

XI (%)

VIII (%)

Terrestrial mammals Terrestrial molluscs Marine molluscs Fishes

75 9 9.5 6

57.5 4 17 20

12 4.5 61 21

Terrestrial mammals Terrestrial molluscs Marine molluscs Fishes

91.5 2.5 2 3.5

87.5 2.5 3 6.5

58 2 13.5 26.5

1: Last Glacial Maximum; 2: Tardiglacial; and 3: Early Holocene.

Last Glacial Maximum.

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and mackerels (Scombridae) are indicative of spring/summer fishing. Within this same context, wrasses (Labridae) have been taken as indicative of fishing close inshore, and thus as bioindicators of rough seas and winter fishing in Maori coastal settlements (Leach, 2006). If such coarse environmental bioindicators were to be taken into consideration at Nerja, then one would conclude that better sea conditions prevailed during the Tardiglacial (i.e., 4.5% wrasses vs. 7% for the LGM and 8% for the Early Holocene) or else that more of the fishing at this stage took place during the late spring and summer. The combined total for ‘‘needlefish þ jacks þ mackerels’’ amounts to 28% of the fish NISPs during the Tardiglacial, yet for the LGM and Early Holocene the values are only 9.5% and 3%, respectively (Tables 4–6). Such patterns, unfortunately, cannot be verified by the codfish, whose seasonal status remains undetermined. 4.4.3. Birds Birds constitute the richest faunal group at Nerja (65 taxa including 54 species) despite their contributions being orders of magnitude below those of the remaining animal assemblages (NISP ¼ 382) (Eastham, 1986; Herna´ndez, 1995a, b; Sa´nchez, in preparation). Though a large number of the taxa reflect a systematic hunting and transport of animals into the cave, an independent accumulation of remains undoubtedly took place through natural processes. Among the species belonging to this second taphonomic group are Little Owl (Athene noctua), Kestrel (Falco tinnunculus), various species of corvids including the raven (Corvus corax), choughs (Pyrrhocorax pyrrhocorax and Pyrrhocorax graculus) and even some or all of the rock pigeons (Columba livia). The LGM avifaunas retrieved in the Vestı´bulo sector reflect the highest proportion of cold adapted species (60%), and this is the only period featuring circumboreal taxa such as Eiders (Somateria sp.) and Harlequin Ducks (Clangula hyemalis) along with Swans (Cygnus sp.), Brent Goose (Branta bernicla) and White-faced Goose (Anser albifrons) (Sa´nchez, in preparation). Taking the percentages of the marine species as a proxy for the structure of a nesting community, we were able to correlate the

Fig. 3. Results from a correspondence analysis plotting the position of a series of Northeastern Atlantic and Mediterranean marine bird communities (R: breeding; I: wintering) according to latitude [ISL: Iceland; ESC: Scotland; MAN: English Channel; ATL: Galicia (NW Spain); MED: Mediterranean]. The location of the marine bird assemblages from Cueva de Nerja (dotted vertical lines) indicates a progressive ‘‘displacement’’ away from Mediterranean conditions during Neolithic times toward those presently found in Galicia during the Magdalenian. Although the Solutrean assemblages have not been included in this study, their boreal nature is more marked still (see Table 4) and corresponds to marine bird assemblages presently found in more northern latitudes (taken from Morales et al., 1998).

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Tardiglacial bird assemblage from the Mina þ Torca/Nerja sectors with a North Atlantic (Scotland) colony that evidenced a progressive latitudinal ‘‘descent’’ through the Early Holocene (Galicia) and the Middle Holocene (Neolithic), when it finally showed a strongly marked ‘‘Mediterranean’’ signature (Fig. 3; Herna´ndez, 1995a, b; Morales et al., 1998). Such data are now confirmed with our unpublished data from the Vestı´bulo/Nerja sector where the LGM marine bird assemblage would qualify as truly boreal (Sa´nchez, in preparation). This sharp ‘‘latitudinal shift’’ does not correlate with the data offered by the remaining marine faunas (see two previous sections). The reasons may lie in the mobility of birds and their marked migratory habits. These circumstances dictate that the group as a whole may equally well be indicative of local conditions at a particular site as of conditions prevailing in far away places (Sa´nchez, 1996, 2004; Morales, 1998). The presence at all times, albeit in small numbers, of taxa such as the red-legged partridge (Alectoris rufa), evidence that, as was the case for plants (see Section 4.3), the avian communities of Nerja never lost their Mediterranean character (Eastham, 1986; Herna´ndez, 1995a, b; Sa´nchez, 1996, 2004). For this reason, we believe that the changing proportions of cold adapted birds in this sequence more than anything else are reflecting the harsher conditions prevailing in the North Atlantic during the LGM. The terrestrial bird record indicates that the Mediterranean character of the avian assemblages of Nerja accentuated during the Early Holocene when both quail (Coturnix coturnix) and several species of forest passerines (e.g., nightingale Luscinia megarrhynchos and blackbird Turdus merula) appear for the first time (Table 6). Still, even during the Tardiglacial, species such as hawfinch (Coccothraustes coccothraustes) indicate that, despite lower temperatures and greater aridity, forests, whether as isolated pockets or as larger extensions, constituted an integral part of the local landscape. 4.4.4. Mammals Of all the faunal assemblages at Nerja, mammals constitute the least informative group from a palaeoclimatic standpoint (Pe´rez, 1986; Morales and Martı´n, 1995; Riquelme, 2004; Riquelme et al., 2005/2006). As can be seen in Tables 4–6 and 9, the bulk of the samples comprises two Iberian endemics, wild goat (Capra pyrenaica) and rabbit (Oryctolagus cuniculus), whose eclectic habits and wide spectrum of tolerances make them of little value as bioindicators (Morales et al., 1998). Likewise, both the small samples and low diversity of the micromammal remains (i.e., 0.03–0.8% of the mammalian NISP) preclude any meaningful comments along these lines. Hedgehog (Erinaceus europaeus), the field mouse (Apodemus sylvaticus) and the dormouse (Eliomys quercinus) are often considered forest animals but in Iberia they thrive equally well in more open landscapes. These, as the various species of bats, are insensitive to climate, meaning that they are able to tolerate a wide range of temperatures. Hibernating and migratory practices set in when conditions deteriorate and all of them qualify as temperate species in a broad sense. Only wild cattle (Bos primigenius), wild boar (Sus scropha), and wood mouse (Apodemus flavicollis) are taken to be

Table 9 Contributions (%NISP) of the main mammal groups to the mammal assemblages from the three main stages of Nerja (Vestı´bulo sector) Taxon

1 (%)

2 (%)

3 (%)

Large herbivores Wild goat Rabbit

3 27 69.5

2.5 20 76

1 16.5 78.5

1: Last Glacial Maximum; 2: Tardiglacial; and 3: Early Holocene.

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Table 10 Relative contributions, in terms of edible meat represented by the wild goat, rabbit and molluscs at Cueva de Nerja assuming that each remain represents one individual Stage

LGM (%)

Tardiglacial (%)

Early Holocene (%)

Wild goat Rabbit Molluscs

88.5 11.5 0.05

84 16 0.02

80.5 19 0.4

Estimated average weights as follows: wild goat (30 kg), rabbit (1.5 kg), and mollusc (0.005 kg). NISP data taken from Tables 3–5.

forest indicators. The presence of all three of them in the LGM assemblages and of the first two during the Tardiglacial lends support to the pollen data from Bajondillo and the avian record from Nerja indicating the existence of at least pockets of forest even during the coldest pulses of the Upper Palaeolithic. Except for the now extinct wild cattle, all of the species shown in Tables 4–6 still thrive in the Iberian Peninsula. The clearest pattern is a cultural one, linked with the decline of high-ranked prey taxa (i.e., ungulates) and an increase in the lowranked rabbit (Aura and Pe´rez, 1992). Of equal relevance is the sharp decline of mammal remains during the Early Holocene (Table 6). These trends do not change the role of the wild goat as the main meat provider throughout the sequence as the percentages of the estimated biomasses from the three stages reveal (Table 10; Morales et al., 1998). Marine mammals fall into two categories: dolphins and monk seal (Monachus monachus), a temperate water species, becomes more abundant with time, its percentage contribution from the Tardiglacial being an order of magnitude higher than from the LGM (i.e., 0.5% vs. 0.05%) yet barely four times more than that in the Early Holocene (2%; Tables 4–6; Pe´rez and Raga, 1998). This

trend follows a general one at Nerja of marine resources increasing with time and is coupled with a concomitant increase in the number of young and subadult individuals within each sample, which may be taken as indicative of an intensification and eventual depletion of this resource with time (monk seal can be considered, for all practical purposes, extinct in the Sea of Alboran today). Alternatively, as was mentioned for other marine resources, the trend may be spurious, reflecting the fact that as the coastline approached more seal carcasses were brought into the cave. Dolphins are absent from the LGM assemblage and almost so in the Early Holocene. Their relative abundance during the Tardiglacial may simply reflect a stochastic phenomenon or else a higher number of strandings as a result of rougher seas (see the comment on wrasses in Section 4.4.2). 5. Discussion The integration of the various lines of evidence into a coherent picture of the Upper Pleistocene environment along the Ma´laga coast is far from straightforward, a major problem being the diverse origin of the deposits. Pollen and sediments essentially accumulated through natural processes whereas most of the fauna and the charcoal assemblages were generated by human activities. For such reasons, the latter two records reflect a regional dimension dictated by the site catchment area for humans at any time and also by longdistance interchanges that, in our case, appear to have been restricted to the barest minimum. A second problem has to do with the conflicting evidence generated by different sets of data. Our preliminary synthesis of the palaeoenvironmental dynamics and human activities along the Ma´laga coastal lowlands indicates the existence of four major stages (Fig. 4).

Fig. 4. Palaeoenvironmental sequence of the Ma´laga coast during the Upper Quaternary based on sedimentary, pollen, isotope and chronological data from Cueva Bajondillo.

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5.1. Second half of OIS 3/transition from the Middle to the Upper Palaeolithic (c. 40–26 ky) Integrating sedimento–climatic (Bergada` et al., 2005) and pollen data with AMS dates from the Bj/14–Bj/11 levels at Bajondillo, it appears that this ‘‘stratigraphical moment’’ can be placed, grosso modo, at an advanced, though not final, moment of OIS 3 (Fig. 4). It would likewise be possible to correlate it with episodes located within Heinrich events 4 (Bj/14) and 3 (Bj/11), also detected in the Sea of Alboran (Sa´nchez et al., 2002), although the Bajondillo sedimentary sequence has a slightly longer chronological range (Table 2). The deterioration of the climate during OIS 3 has been proposed as one of the triggers of the extinction of the Neanderthals (D’Errico and Sa´nchez, 2003). Such deterioration, evident in the data from the oceanic records (Sa´nchez et al., 2002), is also seen in the Bajondillo sequence (e.g., Bj/14–11) as the pollen, isotopes and sedimentary micromorphological records show (Fig. 4). The undoubtedly complex process of Neanderthal extinction, as well as the final record of Middle Palaeolithic cultural manifestations from Southern Iberian peninsula, require a wider geographical and ecological scale and more detailed dates than the ones thus far available in order to be properly addressed (Finlayson, 2004; Finlayson et al., 2006). As things presently stand, climate might have merely acted as yet another de-stabilizing factor but it is debatable whether major or minor. Notwithstanding culturallyimposed limitations (e.g., strong territoriality), Neanderthals emerged and prospered in a permanently fluctuating environment that included many previous episodes of climatic deterioration before their final demise. Also, one should keep in mind that thus far there is no recorded instance where climate can be claimed as the sole cause of total (vs. local) extinction of a mammal species, whether large or small. Within this stage, the beginning of the Upper Palaeolithic is marked by the appearance of Aurignacian lithic technocomplexes. For the moment these have only been documented at Bajondillo (Bj/11; Corte´s and Simo´n, 1997), where four AMS and TL dates are now complemented by a collection of some 2000 artefacts (Corte´s, 2007; Table 1). Such scarcity of Aurignacian deposits seems puzzling and may have been caused by at least four kinds of phenomena: (a) A systematic lack of surveys documenting the transition from the Middle to the Upper Palaeolithic in Southern Iberia. (b) A low demography of the Aurignacian populations that translated into a feeble archaeological signature in the sedimentary deposits. (c) A limited amount of work on sites featuring Gravettian occupations. Of the four thus far known from Southern Iberia (e.g., Vale Boi, Bajondillo, Nerja and Higueral), the excavations of the Gravettian layers from the latter two have been quite restricted despite the fact that at Higueral the sequence reaches down into the Mousterian. (d) The existence of a mosaic of synchronous occurrences of technocomplexes (i.e., Mousterian in the Bay of Gibraltar sector and Upper Palaeolithic in the Bay of Ma´laga) blurring clear-cut patterns of artefact distribution at a regional scale. None of these explanations proceed beyond the stage of educated guesses, and only future studies may eventually provide an answer to this issue. 5.2. Last Glacial Maximum (c. 26–16 ky), Gravettian–Solutrean The Upper Palaeolithic spread throughout the coastal region of Ma´laga with the Gravettian technocomplex, detected at both

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Bajondillo and Nerja and probably also in some of the Humo complex sites (Tables 1 and 2). It is quite possible that Palaeolithic art records (i.e., Cueva del Toro, and possibly Nerja) originated during this same period (Villaverde, 2006). The second cultural tradition, exclusively within the LGM, is the Solutrean, recorded in 11 levels from six sites and fixed by six datings (AMS and TL; Tables 1 and 2). All these Solutrean deposits correspond to the Iberian facies of the Solutrean (Corte´s, 2002). Multiproxy data in oceanic cores from the Sea of Alboran show the strong impact of the LGM in this sea (Cacho et al., 2000, 2001; Sa´nchez et al., 2002; Colmenero et al., 2005). Coinciding with this global scenario, the Bajondillo LGM levels evidence a cold and arid climate (Bj/10–8). The main faunal and anthracological record from Nerja derives from a later moment (Nerja 3) but the scarce data available from charcoal as well as the lithostratigraphy of the Nerja 1 occupation lend support to such a harsh climatic hypothesis. The multiproxy data from the oceanic cores likewise indicate the onset of rapid cooling episodes (Heinrich events and Dansgaard–Oeschger stadials) throughout the LGM and not only during its initial phase (i.e., Pleniglacial). Alquenones and coccolithophorids from cores in the Sea of Alboran and the Gulf of Ca´diz demonstrate that during these cold pulses there existed a temperature gradient in the superficial water mass such that MST (mean surface temperature) was, on the average lower than that of the surrounding regions, an amplification effect of the general lowering of the MST during glacial episodes (Va´zquez et al., 1991; Cacho et al., 2000, 2001; Colmenero et al., 2004). Data from planktonic foraminifera in the OPF 976 core sample from the Sea of Alboran show that MST at c. 21 ky was below 10  C at all times (Pujol and Vergnaud-Grazzini, 1989). All these data indicate that during the LGM the oceanographic repercussions of climate changes were far more intense in the Mediterranean Sea than at mid-latitude in the Atlantic, creating favourable conditions for cold adapted marine taxa (Colmenero et al., 2004, 2005). The faunal evidence from the Solutrean does not lend itself to a straightforward interpretation within the framework of the previous interpretation. For birds, the coldest assemblages are recorded during this period and feature a majority of taxa that now thrive up to circumboreal latitudes. One problem is that these avian assemblages show the largest number of boreal taxa at the beginning of the sequence (Sa´nchez, in preparation), and all these taxa could be taken to represent harsh conditions in the far away northern Atlantic lands and not necessarily in Alboran, given their migratory habits. Mammals and land snails are essentially worthless as climatic bioindicators, since most of the taxa recorded at Nerja still thrive in the Iberian Peninsula today. Within the marine sector, molluscs were essentially Mediterranean or indifferent to climatic limitations, and of the cold adapted taxa only five remains of Modiolus, also recorded at Bajondillo, qualify as boreal (Table 4). The resilience of the Mediterranean species is therefore striking and calls for an explanation. As for the fish, the increase of boreal elements represented by codfish seems gradual throughout the Solutrean and only in the latest level (V/8) does one see numbers rise significantly (Rosello´, in preparation). Of all the faunal groups at Nerja, these seem to be the best indicators that local sea temperatures dropped drastically during the Solutrean but why this is not also reflected by the molluscs remains unclear. Disturbing in particular is the abundance of gadids at the end of the Solutrean, when conditions apparently became milder in the area. This could either reflect a harvesting bias, or else harsh conditions somewhere else, since all these gadids are also migratory. If the latter scenario held, why then is such a trend not seen also in the birds? One way or the other, the most important aspect revealed by the fauna during this stage relates to shifts in the harvesting of particular groups. Mollusc records in the area date back to the Middle Palaeolithic and, although the Gravettian only records molluscs being

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used as ornaments, littoral adaptations seem beyond question. One reason for the low frequencies of marine faunas during these initial stages of the Upper Pleistocene may relate to the distance of the archaeological sites to the coast at that time (Corte´s and Simo´n, 2000; Aura et al., 2002). Yet during the Solutrean these distances remained essentially the same as during the previous period. For such reason, the gradual increase of marine taxa documented within the Solutrean at Nerja (Table 8) may variously represent either an enlargement of the site catchment area, shifts in exchange networks with the shore, technological developments in the preservation and transport of foodstuffs, or a real shift in human subsistence practices. The uniqueness of the Nerja faunal record prevents one from choosing any of these alternatives, or a combination of them, as the most likely cause for such a shift. The Solutrean nevertheless marks the onset of an activity (marine fishing), previous evidence for which in Europe reached back only to the Magdalenian (i.e., Tardiglacial; Morales et al., 1998). Likewise, the intensification of the harvesting of marine resources in general, whether fishing or gathering, appears to coincide with the final stages of the LGM and the onset of a climatic amelioration event, at least in terms of humidity data as the isotopic analysis of Bj/8 and the pollen data from Bj/9–8 (Bajondillo T episode) suggest (Fig. 4; Table 3). During the LGM some sites in the area still evidence sporadic human occupation alternating with hyena coprolites (e.g., Nerja and Abrigo 6 of the Humo complex). Correlating with the disappearance of this carnivore, the evidence of other large predators vanishes from the record whereas faunal assemblages of human origin increase both in size and in diversity. This leads one to suggest that it is during the Solutrean period that the patterns of mobility and subsistence of the Upper Palaeolithic become definitively established on the coast of Ma´laga. Such phenomena probably led to permanent occupations that would explain why, among other things, we have a superposition of the Magdalenian over the Solutrean in no less than five settlements (Table 1). Finally, one should remark that it is during the LGM that humans symbolically ‘take hold’ of the coastal zone as exemplified by rock art in five cavities located on the southern façade of the littoral mountains (Fig. 1). 5.3. Tardiglacial (c. 12–10 ky), Upper Magdalenian Erosive processes similar to those recorded in other areas of the Iberian Mediterranean (Aura, 1995) have removed sediments from the 16 to 13 ky interval (Table 2). For such reason the regional sequence of the coast of Ma´laga resumes only during the Tardiglacial, slightly after the last cold pulse of some intensity is detected in southern Iberia. This pulse is evident in the development of glaciers in the Sierra Nevada (c. 14 ky) and of particularly cold conditions in the Sea of Alboran coinciding, grosso modo, with the Younger Dryas episode (c. 12.5–11.5 ky) (Go´mez et al., 1992; Cacho et al., 2001, 2002; Colmenero et al., 2004, 2005). At Bajondillo, sedimentological and isotopic data indicate that Bj/5 probably corresponds to this moment, though the scarce pollen data currently available for that level remain inconclusive (Figs. 2 and 4). This constitutes yet another instance where the different kinds of records are difficult to reconcile. The latest stages of the Nerja 5 occupation phase (i.e., V/7–5; M/16) appear to coincide with this cold pulse although it now seems clear that the precise location of the Younger Dryas within the regional sequence of the Ma´laga coast is one of those issues that will require further work before being settled. Except for the birds, whose diversity trebles, the Tardiglacial faunas are quite similar in terms of diversity and number of remains to those of the LGM, the proportional increase of marine resources being their most noteworthy feature. Although the data on fish are

preliminary and the contributions of the boreal codfish will be far larger than the 9% shown in Table 5, one cannot detect during the Magdalenian any significant change in the variety or proportional contribution of the boreal taxa from those of the last Solutrean level (Rosello´, in preparation). Tardiglacial birds are lacking most of the truly boreal ducks (e.g., eiders) that were present during the LGM although the small-size of most samples does not allow one to determine to what extent these absences are merely stochastic in nature. By comparison with the LGM assemblage, the main differences of the Tardiglacial avifaunas are the presence of the lower latitude, climatically insensitive, ducks, preybirds, gulls and passerines (Table 5). In the case of molluscs, one does detect a decline of the Mediterranean component by comparison with the LGM assemblages but this is more than compensated by an increase in the number of climatically insensitive taxa rather than by that of the boreal taxa, essentially restricted to one single specimen of Modiolus (i.e., 0.02% of the mollusc NISP). Given that mammals maintain the same temperate species throughout the sequence, their Tardiglacial assemblages could hardly qualify as more boreal than those from the LGM. The de-coupling of the fish assemblages from the remaining faunal records at Nerja thus calls for an explanation. During the Tardiglacial the coast of Ma´laga featured the incorporation of new technologies (e.g., harpoons, small blanks, tools of small-size with abundant backed bladelets). The appearance of tools at Nerja such as the so-called straight hooks (Aura and Pe´rez, 1992) may reflect the development of a local tradition of fishing. This tradition, part of a larger phenomenon of coastal adaptations, is probably of a more regional than local nature. Indeed, although the faunal record from Nerja is unique and exceptional, recent finds of molluscs at Bajondillo, Hoyo de la Mina, Abrigo 6 of the Humo complex and Victoria cave evidence the generalization of such kinds of harvesting activities during the Tardiglacial. 5.4. Early Holocene (c. <10–7 ky), Epipalaeolithic This last stage of the sequence from the coast of Ma´laga is a prolongation of the cultural and economic adaptations recorded for the previous period under milder environmental conditions. From a palaeoenvironmental standpoint, although the inorganic and pollen records are absent from Bajondillo for the earliest moments of this stage, data from Nerja (V/4) indicate a climatic amelioration seen in the rising contributions of wild olive (Olea sp.) and Fabacea. In the marine domain, the disappearance of the cold SST (sea surface temperature) foraminifer Emiliania huxleyi from the MD 95 to 2043 core highlights a major shift for the Sea of Alboran (Colmenero et al., 2004, 2005). A second important factor is the approaching coastline, which in the case of Nerja, essentially replaces previously predominant sandy beaches with rocky shorelines (though not in the Bay of Ma´laga zone where sandy beaches continued throughout the Holocene). When one turns to the fauna, the de-coupling discussed for previous periods continues to be evident. Once again, the fish assemblages witness the rise of codfish to a dominant position at Nerja (V/4) although the main taxon (i.e., pollack, Pollachius pollachius) is more temperate than boreal in character (Rodrigo, 1994; Rosello´, in preparation). Birds, on the other hand, exhibit an increase in forest passerines and terrestrial taxa to the detriment of marine birds. Thus the bird assemblages become better indicators of local environments than any from previous stages (Table 6). The only boreal mollusc on record (Modiolus) disappears but the tendency for the Mediterranean taxa (e.g., P. caerulea, P. ferruginea, etc.) to decline at the expense of their more climatically insensitive vicariants (e.g., P. rustica, Patella ulyssiponensis) continues. Only the anecdotal replacement of the colder adapted European pond tortoise (Emys orbicularis) by the Mediterranean terrapin

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(Mauremys caspica) might be taken to reflect a climatically induced ‘‘turnover’’ (Tables 4 and 6). The faunal record of Nerja has higher value as an indicator of changing subsistence strategies. The prominent features are a collapse in the number of mammals as well as a dramatic increase in the number of mussels. The latter phenomenon testifies to the availability of mussels as a combined result of the development of a rocky shore and rising sea levels that brought the shore closer to the cave. The former phenomenon might be taken as yet another case of resource depletion, the truly informative trends here being the increase of rabbits and the decrease of ungulates (Table 9). Still, wild goat kept its dominant position within the animal economy as the main meat provider (Table 10; Morales et al., 1998). Lack of reliable information concerning the demography of these human populations forces such hints of intensification to remain at a necessarily descriptive (i.e., qualitative) level of analysis. 6. Conclusions Though preliminary and often much too restricted to proceed beyond coarse generalizations, the integration of these analyses with previous archaeological records and multiproxy core data from the Sea of Alboran allow a tentative model for the palaeoenvironmental and cultural dynamics of the coast of Ma´laga during the Upper Pleistocene and Early Holocene to be established (Fig. 4). The data thus far gathered indicate that this zone constituted a strategic place for human populations during the 200–7 ky interval despite the fact that detailed evidence of such circumstance is presently only available for the 40–7 ky period (Corte´s et al., 2005). Both Homo sapiens neanderthalensis and Homo sapiens were thus involved in this occupation, making the region a target zone for the study of Neanderthal–modern human interactions and of their evolutionary implications. Among the factors determining such strategic relevance of the area one should mention an abundance and reliability of water sources linked with the presence of aquifers, a diversity of biotopes determined by the topography, fluctuating sea levels and the productivity of the Sea of Alboran, along with a climate that must have been at all times milder than that existing in the nearby regions of the hinterland. It appears that the environmental fluctuations of the second half of the last major glacial episode accentuated the mosaic of biotopes in this region. In this way, despite the general and sharp decline in precipitation indicated by the sedimentary, pollen and isotope records from Bajondillo, along with the marine core data from the Sea of Alboran, one detects the presence of both more humid and temperate refugia throughout the whole sequence at Bajondillo, as well as evidence for the resilience of mesic and thermic plant taxa in microclimatic pockets (Figs. 2 and 4). One major problem now is that the main sequences of the coast of Ma´laga, Bajondillo and Nerja, are not duplicates in terms of either chronostratigraphy or available materials. The problem is all the more pressing due to an absence of comparable sequences in nearby areas, stressing the uniqueness of the data thus far presented. Such uniqueness implies that the patterns that we have been able to infer must of necessity remain difficult to verify through the analysis of alternative data sets. As such, they should be considered as a testable framework for future studies. A second problem has to do with the occasional de-coupling of the different data sets that compromise the elaboration of a coherent, straightforward and unitary sequence of chronostratigraphical and palaeoenvironmental events in the area. In the case of Nerja this is reflected in the conflicting palaeoenvironmental information provided by the different faunal groups, birds signalling coldest conditions at the beginning of the Solutrean, while fish

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suggest coldest conditions at the end of this period and up to Epipalaeolithic times when the other indicators show that the mildest conditions prevailed. Such discrepancies may simply reflect methodological (e.g., sampling) or cultural biases given that most of the taxa have been accumulated as a result of human activities. Still, we feel that there might exist idiosyncrasies dictating such a decoupling, some of them having to do with the differential mobility of each group. These differential responses conventional faunal analyses are normally unable to calibrate. Such de-coupling of the faunal data sets, also seen when different kinds of records are confronted, in no way invalidates the premise that the diversity of the animal and botanical taxa reveals an ecologically complex mosaic of the terrestrial and marine domains on the Ma´laga coast at all times. It is often difficult to proceed beyond a coarse level of interpretation given that these long-term deposits mostly represent palimpsests. The overall picture emerging from our analyses highlights the resilience of Mediterranean ecosystems to relatively harsh climatic conditions in the southern Iberian Upper Palaeolithic, reflecting the biogeographical relevance of the coastal fringe of Ma´laga and the refugial character that this zone repeatedly displayed. Multiproxy data also hint that the temperature decline was far more marked and took place at a later time in the Sea of Alboran than in the terrestrial ecosystems. The response of the fauna to these changes was far more heterogeneous and eclectic than that of the plants and, in the case of the birds and fishes, appears to have been dictated also by conditions that took place in the far away ecosystems of the North Atlantic. For all these reasons we believe that future studies of the southern Iberian littoral, rather than concentrating on the contrasts revealed within each particular data set, should try to reconcile the qualitative and quantitative aspects within and among the various records in an integrated, multivariate manner. Acknowledgements The authors would like to thank the following funding agencies: Consejerı´a de Cultura de la Junta de Andalucia (Project: ‘‘Estudio y contextualizacio´n cronoestratigra´fica de las antiguas excavaciones del Patronato de la Cueva de Nerja’’, ‘‘Muestreo de Cueva Bajondillo’’, ‘‘Prospeccio´n Arqueolo´gica superficial de Torremolinos’’; Miguel Corte´s); Patronato de la Cueva de Nerja (Project: ‘‘Correlacio´n de la secuencia estratigra´fica de las cuevas de Nerja y Bajondillo’’; Miguel Corte´s); and the Spanish Ministry of Science and ˜ iz). Clive Technology (Grant CGL2004-00891; Arturo Morales-Mun Finlayson and Darren Fa (Gibraltar Museum) and three anonymous referees are gratefully acknowledged for their critical review of the paper and for improving the English. References Allen, J.R.M., Huntley, B., 2000. Weichselian palynological records from southern Europe: correlation and chronology. Quaternary International 73/74, 111–125. Arribas, A., Aura, J.E., Carrio´n, J.S., Jorda´, J.F., Pe´rez, M., 2004. Presencia de hiena manchada en los depo´sitos basales (Pleistoceno Superior Final) del yacimiento ˜ a). Revista Espan ˜ ola de arqueolo´gico de la Cueva de Nerja (Ma´laga, Espan Paleontologı´a 19/1, 109–121. Aura, J.E., 1995. El Magdaleniense mediterra´neo: la Cova del Parpallo´ (Gandı´a, Valencia). Diputacio´n Provincial de Valencia, Valencia. Serie de Trabajos Varios 91. Aura, J.E., Jorda´, J.F., Pe´rez, M., Rodrigo, M.J., 2001. Sobre dunas, playas y calas. Los pescadores prehisto´ricos de la Cueva de Nerja (Ma´laga) y su expresio´n arqueolo´gica en el tra´nsito Pleistoceno–Holoceno. Archivo de Prehistoria Levantina 24, 9–39. Aura, J.E., Jorda´, J.F., Pe´rez, M., Rodrigo, M.J., 2002. The far south: the Pleistocene– Holocene transition in Nerja Cave (Andalucia, Spain). Quaternary International 93/94, 19–30. Aura, E., Pe´rez, M., 1992. Tardiglaciar y Postglaciar en la regio´n mediterra´nea de la Penı´nsula Ibe´rica (13.500–8.500 B.P.): Transformaciones industriales y econo´micas. Saguntum 25, 25–47.

2192

M. Corte´s-Sa´nchez et al. / Quaternary Science Reviews 27 (2008) 2176–2193

Badal, E., 1996. La ve´ge´tation du Pale´olithique Supe´rieur et de l’E´pipale´olithique aux alentours de la Cueva de Nerja (Ma´laga, Espagne). Revue d’Arche´ometrie, Supple´ment Actes du colloque de Pe´rigueux 199, 171–176. Barton, N., 2000. Mousterian hearths and shellfish: late Neanderthal activities on Gibraltar. In: Stringer, C.B., Barton, R.N.E., Finlayson, J.C. (Eds.), Neanderthals on the Edge. Oxbow, Oxford, pp. 211–220. Bergada`, M.M., Corte´s, M., Simo´n, M.D., 2005. Secuencia geoarqueolo´gica del Pleistoceno superior y Holoceno inicial en el sur de la Penı´nsula Ibe´rica: Cueva Bajondillo (Torremolinos, Ma´laga). In: Rodrı´guez, J., Finlayson, C., Giles Pacheco, F. (Eds.), Cuaternario Mediterra´neo y poblamiento de homı´nidos. Gibraltar Museum, Gibraltar, pp. 74–75. Boessneck, J., Driesch, A.V.D., 1980. Tierknochenfunde aus vier su¨dspanischen Ho¨hlen. Studien u¨ber fru¨he Tierknochenfunde von der Iberischen Halbinsel 7, 1–83. Cabezudo, B., Pe´rez, A., 2001. Datos sobre la vegetacio´n termo´fila del litoral oriental ˜ a). Acta Botanica Malacitana 26, 229–240. de Ma´laga (Espan Cacho, I., Grimalt, J.O., Canals, M., Sbaffi, L., Shackleton, N., Scho¨nfeld, J., Zahn, R., 2001. Variability of the Western Mediterranean Sea surface temperature during the last 30,000 years and its connection with the northern hemisphere climatic changes. Paleoceanography 16, 40–52. Cacho, I., Grimalt, J.O., Sierro, F.J., Shackleton, N., Canals, M., 2000. Evidence for enhanced thermohaline circulation during rapid climatic coolings. Earth and Planetary Science Letters 183, 417–429. Cleyet-Merle, J.J., 1990. La Prehistoire de la Peˆche. Editions Errance, Paris. Carrio´n, J.S., 1992. Late Quaternary pollen sequence from Carihuela Cave, southeastern Spain. Review of Palaeobotany and Palynology 71, 37–77. Carrio´n, J.S., Yll, E.I., Walker, M.J., Legaz, A., Chaı´n, C., Lo´pez, A., 2003. Glacial refugia of temperate, Mediterranean and Ibero North African flora in south-eastern Spain: new evidence from cave pollen at two Neanderthal man sites. Global Ecology and Biogeography 12, 119–129. Colmenero, H., Flores, J.A., Sierro, F.J., Ba´rcena, M.A., Lo¨wemarkb, L., Scho¨nfeld, J., Grimalt, J.O., 2004. Ocean surface water response to short-term climate changes revealed by coccolithophores from the Gulf of Cadiz (NE Atlantic) and Alboran Sea (W Mediterranean). Palaeogeography, Palaeoclimatology, Palaeoecology 205, 317–336. Colmenero, E., Flores, J.A., Sierro, F.J., Grimalt, J.O., 2005. Reconstruccio´n de las temperaturas superficiales marinas del margen ibe´rico sudoriental durante el u´ltimo periodo glacial mediante el empleo de cocolitofo´ridos. Geogaceta 38, 195–198. Corte´s, M., 2002. El Paleolı´tico Medio y Superior en el sector central de Andalucı´a (Co´rdoba y Ma´laga). Unpublished Ph.D. Universidad de Co´rdoba, Co´rdoba, Spain. Corte´s, M., 2006. El extremo occidente neandertal. El Paleolı´tico Medio en el sur de la Penı´nsula Ibe´rica. In: Reunio´n Cientı´fica-Mesa de Trabajo Neandertales Canta´bricos. Estado de la Cuestio´n. El Paleolı´tico Medio canta´brico: hacia una revisio´n actualizada de su problema´tica. Museo de Altamira, Monografı´as 20, Santander, pp. 55–74. Corte´s, M. (Ed.), 2007. Cueva Bajondillo (Torremolinos). Secuencia cronocultural y paleoambiental del Cuaternario reciente en la bahı´a de Ma´laga. Diputacio´n Provincial de Ma´laga, Ma´laga. Corte´s, M., Ferrer, J.E., Marque´s, I., Baldomero, A., Simo´n, M.D., 2005. Aportaciones cronoestratigra´ficas de Cueva Bajondillo al tra´nsito Paleolı´tico Medio-Superior en el sur de la Penı´nsula Ibe´rica. In: Santonja, M., Pe´rez-Gonza´lez, A., Machado, M.J. (Eds.), Geoarqueologı´a y patrimonio en la Penı´nsula Ibe´rica y el entorno mediterra´neo. Adema, Soria, pp. 181–196. ˜ oz, V.E., Sanchidria´n, J.L., Simo´n, M.D., 1996. El Paleolı´tico en Corte´s, M., Mun Andalucı´a. La dina´mica de los grupos predadores en la prehistoria andaluza. ˜ os de investigacio´n (1770– Ensayo de Sı´ntesis. Repertorio bibliogra´fico de 225 an 1995), Co´rdoba. Corte´s, M., Simo´n, M.D., 1997. Cueva Bajondillo (Torremolinos, Ma´laga). Aportaciones al Paleolı´tico en Andalucı´a. In: El mo´n Mediterrani despre´s del Pleniglacial (18.000–12.000 BP). Centre d´Investigacions arqueolo`giques de Girona, Gerona, pp. 275–290. Se´rie Monogra`fica 17. Corte´s, M., Simo´n, M.D., 2000. Bahı´a de Ma´laga: algunos aspectos fisiogra´ficos y su incidencia sobre los yacimientos arqueolo´gicos pleistocenos en medio ka´rstico de su a´mbito de influencia. In: Actas 1 Congreso Andaluz de Espeleologı´a, Sevilla, pp. 217–224. D’Errico, F., Sa´nchez, M.F., 2003. Neanderthal extinction and the millennial scale climatic variability of OIS 3. Quaternary Science Reviews 22, 769–788. Dansgaard, W., Johnsen, S.J., Clauson, H.B., Dahl-Jensen, D., Gundestrup, N.S., Hammer, C.U., Hvidberg, C.S., Steffensen, J.P., Sveinbjornsdottir, A.E., Jouzel, J., Bond, G., 1993. Evidence for general instability in past climate from a 250 kyr ice-core record. Nature 364, 218–220. Eastham, A., 1986. The birds of the cueva de Nerja. In: Trabajos sobre la Cueva de Nerja, vol. 1. Cave of Nerja Foundation, Malaga, pp. 109–131. Ehleringer, J.R., 1991. 13C/12C fractionation and its utility in terrestrial plant studies. In: Coleman, D., Fry, B. (Eds.), Carbon Isotope Techniques. Academic Press, New York, pp. 187–200. Fa, D.A., in press. A report on the marine mollusca from the Gorham’s cave excavations 1998–2005: preliminary results and interpretation. In: Finlayson, J.C., et al. (Eds.), Where The Last Neanderthals Lived. Gorham’s Cave Monographs, Oxbow Books, Oxford. Farquhar, G.D., O’Leary, M.H., Berry, J.A., 1982. On the relationship between carbon isotope discrimination and the intercellular carbon dioxide concentration in leaves. Australian Journal of Plant Physiology 9, 121–137.

Ferre, E., Corte´s, M., Ramos, J., Senciales, J.M., Lozano-Francisco, M.C., Vera-Pela´ez, J.L., Auilera, R., Navarrete, I., 2003. El Cuaternario reciente en el sector oriental de la Bahı´a de Ma´laga. Rasas y depo´sitos marinos, continentales y arqueolo´gicos. Cuaternario y Geomorfologı´a 18, 73–93. Ferrer, J.E., Marque´s, I., Corte´s, M., Baldomero, A., Ramos, J., 2006. Excavaciones en ˜ a). Contrastacio´n de una Cueva del Hoyo de la Mina (Ma´laga, Andalucı´a, Espan secuencia arqueolo´gica cla´sica para el estudio del Tardiglaciar–Holoceno antiguo en el sur de la Penı´nsula Ibe´rica. In: Sanchidria´n, J.L., Ma´rquez, A.M., Fullola, J.M. (Eds.), La cuenca mediterra´nea durante el Paleolı´tico Superior 38.000–10.000. IV Simposio de Prehistoria Cueva de Nerja y Reunio´n de la VIII Comisio´n de Paleolı´tico Superior U.I.S.P.P. Cave of Nerja Foundation, Malaga, pp. 316–325. Finlayson, C., 2004. Neanderthals and Modern Humans. An Ecological and Evolutionary Perspective. Cambridge University Press, Cambridge. Finlayson, C., Giles, F., Rodrı´guez, J., Fa, D.A., Gutierrez, J.M., Santiago, A., Finlayson, G., Allue´, E., Baena, J., Ca´ceres, I., Carrio´n, J.S., Ferna´ndez, Y., Gleed, C.P., Jimenez, F.J., Lo´pez, P., Lo´pez, J.A., Riquelme, J.A., Sa´nchez, A., Giles, F., Brown, K., Fuentes, N., Valarino, C.A., Villalpando, A., Stringer, C.B., Martinez, F., Sakamoto, T., 2006. Late survival of Neanderthals at the southernmost extreme of Europe. Nature 443, 850–853. Fumanal, M.P., 1995. Los depo´sitos cuaternarios en cuevas y abrigos. Implicaciones sedimentoclima´ticas. In: El Cuaternario del Paı´s Valenciano. Universidad de Valencia: 115–124. Go´mez, A, Sa´nchez, S., Simo´n, M., Salvador, F., Esteban, A., 1992. Sı´ntesis de la morfologı´a glaciar y periglaciar de Sierra Nevada. In: Lo´pez, F., Conesa, C., ˜ a. Soc. Esp. Romero, M.A. (Eds.), Estudios de Geomorfologı´a en Espan Geomorfologı´a I, Murcia, pp. 379–392. Guerra, A., Serrano, F., Ramallo, D., 2003. Geomorphic and sedimentary Plio-Pleistocene evolution of the Nerja area (northern Alboran basin, Spain). Geomorphology 60, 89–105. ˜ as de 1980 y Herna´ndez, F., 1995a. Cueva de Nerja (Ma´laga): las aves de las campan 1982. In: Trabajos sobre la Cueva de Nerja, vol. 5. Cave of Nerja Foundation, Malaga, pp. 221–293. Herna´ndez, F., 1995b. El papel de las aves en la reconstruccio´n paleoecolo´gica. Su aplicacio´n al Cuaternario del Sur de la Penı´nsula Ibe´rica. Unpublished Ph.D., Universidad Auto´noma Madrid, Madrid, Spain. Herna´ndez, F.J., Somoza, L., Rey, J., 1996. Pleistocene–Holocene high-resolution sequence analysis on the Alboran Sea’s continental shelf. In: De Batist, M., Jacobs, P. (Eds.), Geology of Siliciclastic Shelf Seas. Geological Society Special Publication, vol. 117, pp. 137–154. Jorda´, J.F., 1986. Estratigrafı´a y sedimentologı´a de la cueva de Nerja (Salas de la mina y del vestı´bulo). In: Trabajos de la Cueva de Nerja, vol. 1. Cave of Nerja Foundation, Malaga, pp. 41–97. Jorda´, J.F., Aura, J.E., Jorda´, F., 1990. El lı´mite Pleistoceno – Holoceno en el yacimiento de la cueva de Nerja (Ma´laga). Geogaceta 8, 102–104. Jorda´, J.F., Aura, J., Rodrigo, M.J., Pe´rez, M., Badal, E., 2003. El registro paleobiolo´gico ˜ a). cuaternario del yacimiento arqueolo´gico de la Cueva de Nerja (Ma´laga, Espan Boletin de la Real Sociedad Espanola de Historia Natural. Seccion Geologia 98, 73–89. Lario, C.J., Zazo, C., Goy, L.J., Hoyos, M., Hillaire, C., 1998. Episodios marinos del u´ltimo interglacial (estadio isoto´pico 5) del litoral de Ma´laga (SE. Peninsular). In: Elementos de los paisajes de la provincia de Ma´laga, pp. 231–249. Leach, F., 2006. Fishing in Pre-European New Zealand. Archaeofauna 15, 9–266. Lozano, M.C., Vera, J.L., Simo´n, M.D., Corte´s, M., 2003. Taxonomı´a, autoecologı´a y actividad antro´pica de los moluscos e invertebrados marinos de la sala del ˜ a). Plioce´nica 3, 74–87. Vestı´bulo (Cueva de Nerja, Ma´laga, sur de Espan Lozano, M.C., Vera, J.L., Simo´n, M.D., Corte´s, M., 2004. Nuevos datos sobre el registro malacolo´gico del Pleistoceno Superior-Holoceno de la Cueva de Nerja (Ma´laga, ˜ a). Revista de la Sociedad Espan ˜ ola de Paleontologı´a 19, 215–228. Espan Martı´nez, J.M., Peinado, M., 1987. Andalucı´a Oriental. In: Peinado, M., Rivas, S. (Eds.), ˜ a. University of Alcala´ de Henares, Alcala´ de Henares, pp. La vegetacio´n de Espan 231–255. McAndrews, J.H., King, J.E., 1976. Pollen of the North American Quaternary: the top twenty. Geoscience and Man 15, 41–49. Morales, A., 1998. The mobile faunas: reliable seasonal indicators for archaeozoologists? In: Rocek, T.R., Bar-Yosef, O. (Eds.), Seasonality and Sedentism. Archaeological Perspectives from Old and New World Sites, Peabody Museum Bulletin, vol. 6, pp. 25–40. Morales, A., Martı´n, J.M., 1995. Los mamı´feros de la Cueva de Nerja: ana´lisis de las cuadrı´culas NM-80A, NM-80B, y NT-82. In: Trabajos de la Cueva de Nerja, vol. 5. Cave of Nerja Foundation, Ma´laga, pp. 59–159. ˜ as, J.M., 1994. Cueva de Nerja (prov. Ma´laga). A close look Morales, A., Rosello´, E., Can at a twelve thousand years ichthyofaunal sequence from Southern Spain. In: Van Neer, W. (Ed.), Fish Exploitation in the Past. Annales du Muse´e Royale de L’Afrique Centrale, Sciences Zoologiques, vol. 274, pp. 253–262. Morales, A., Rosello´, E., Herna´ndez, F., 1998. Late Upper Palaeolithic subsistence strategies in southern Iberia: Tardiglacial faunas from Cueva de Nerja (Ma´laga, Spain). European Journal of Archaeology 1, 9–50. Pate, J.S., 2001. Carbon isotope discrimination and plant water-use efficiency. In: Unkovich, M., Pate, J.S., McNeill, A., Gibbs, D.J. (Eds.), Stable Isotope Techniques in the Study of Biological Processes and Functioning of Ecosystems. Kluwer, Boston, pp. 19–36. Pe´rez, M., 1986. Avance al estudio de los mamı´feros de la Cueva de Nerja (Ma´laga). In: Trabajos sobre la Cueva de Nerja, vol. 1. Cave of Nerja Foundation, Malaga, pp. 99–106.

M. Corte´s-Sa´nchez et al. / Quaternary Science Reviews 27 (2008) 2176–2193 Pe´rez, M., Raga, J.A., 1998. Los mamı´feros marinos en la vida y en el arte de la prehistoria de la Cueva de Nerja. In: Sanchidria´n, J.L., Simo´n, M.D. (Eds.), Las Culturas del Pleistoceno Superior en Andalucı´a. Cave of Nerja Foundation, Malaga, pp. 251–275. Petit, J.R., Jouzel, J., Raynaud, D., Barkov, N.I., Barnola, J.M., Basile, I., Bender, M., Chappellaz, J., Davis, J., Delaygue, G., Delmotte, M., Kotlyakov, V.M., Legrand, M., Lipenkov, V., Lorius, C., Pe´pin, L., Ritz, C., Saltzman, E., Stievenard, M., 1999. Climate and Atmospheric History of the Past 420,000 years from the Vostok Ice Core, Antarctica. Nature 399, 429–436. Pujol, C., Vergnaud-Grazzini, C., 1989. Palaeoceanography of the last deglaciation in the Alboran Sea (Western Mediterranean). Stable isotopes and planktonic foraminiferal records. Marine Micropaleontology 15, 153–179. Ramos, J., Corte´s, C., Aguilera, A., Lozano, M.C., Vera, J.L., Simo´n, M.D., 2006. El ˜ a, Ma´laga). In: Magdaleniense y Epipaleolı´tico del Complejo del Humo (La Aran Sanchidria´n, J.L., Ma´rquez, A.M., Fullola, J.M. (Eds.), La cuenca mediterra´nea durante el Paleolı´tico Superior 38.000–10.000. IV Simposio de Prehistoria Cueva de Nerja y Reunio´n de la VIII Comisio´n de Paleolı´tico Superior U.I.S.P.P. Cave of Nerja Foundation, Malaga, pp. 326–341. Reitz, E., Wing, E., 1999. Zooarchaeology. Cambridge University Press, Cambridge. Riquelme, J.A., 2004. Cambios faunı´sticos en los registros arqueolo´gicos tardiglaciares e inicios del holoceno en el sur peninsular. In: Sa´nchez de las Heras, C. (Ed.), Sociedades recolectoras y primeros productores. Actas de las Jornadas Tema´ticas Andaluzas de Arqueologı´a. Arqueologı´a Monografı´as. Junta de Andalucı´a, Sevilla, pp. 295–299. Riquelme, J.A., Simo´n, M.D., Corte´s, M., 2005/2006. La fauna de mamı´feros del Solutrense en la Cueva de Nerja. Munibe 57, 255–263. ˜ a. M.A.P.A.– Rivas, S., 1987. Memoria del Mapa de Series de Vegetacio´n de Espan ICONA, Madrid. Rodrigo, M.J., 1994. Remains of Melanogrammus aeglefinus (Linnaeus, 1758) in the Pleistocene–Holocene passage in the cave of Nerja, Ma´laga/Spain. Offa 51, 348–351. Rosello´, E., in preparation. The fishes from the Sala del Vestı´bulo at Cueva de Nerja (Ma´laga, Spain). ˜ as, J.M., 1995. Estudio ictioarqueolo´gico de la Cueva de Rosello´, E., Morales, A., Can Nerja (prov. Ma´laga): Resultados de las campan˜as de 1980 y 1982. In: Pellicer, M., Morales, A. (Eds.), Fauna de la Cueva de Nerja I. Salas de la Mina y de ˜ as 1980–1982. Trabajos sobre la Cueva de Nerja, vol. 5. Cave of la Torca, campan Nerja Foundation, Ma´laga, pp. 163–217. Sa´nchez, M.F., 1993. Criterios de base tafono´mica para la interpretacio´n de ana´lisis palinolo´gicos en cueva: el ejemplo de la regio´n canta´brica. In: Fumanal, M.P.,

2193

Bernabeu, J. (Eds.), Estudios sobre Cuaternario, medios sedimentarios y ha´bitat humano. University of Valencia, Valencia, pp. 117–130. Sa´nchez, A., 1996. Aves fo´siles del Pleistoceno ibe´rico: rasgos clima´ticos, ecolo´gicos y zoogeogra´ficos. Ardeola 43, 207–219. Sa´nchez, A., 2004. Avian zoogeographical patterns during the Quaternary in the Mediterranean region and paleoclimatic interpretation. Ardeola 51, 91–132. Sa´nchez, M.F., in preparation. The birds from the Sala del Vestı´bulo at Cueva de Nerja (Ma´laga, Spain). Sa´nchez, M.F., Cacho, I., Turo´n, J.L., Guiot, J., Sierro, F.J., Peypouquet, J.P., Grimalt, J.O., Shackelton, N.J., 2002. Synchroneity between marine and terrestrial responses to millennial scale climatic variability during the last glacial period in the Mediterranean region. Climate Dynamics 19, 95–105. Serrano, F., Guerra, A., Lozano, M.C., Vera, J.L., 1997. Multivariate analysis of remains of molluscan foods consumed by latest Pleistocene and Holocene humans in Nerja Cave (Ma´laga, Spain). Quaternary Research 48, 215–227. Serrano, F., Lozano, M.C., Vera, J.L., Guerra, A., 1995. Malacofauna en yacimientos prehisto´ricos de la Cueva de Nerja. In: Pellicer, M., Morales, A. (Eds.), Fauna de la Cueva de Nerja I. Salas de la Mina y de la Torca, campan˜as 1980–1982. Trabajos sobre la Cueva de Nerja, vol. 5. Cave of Nerja Foundation, Malaga, pp. 297–373. Shackleton, J.C., Andel, T.H., van, Runnels, C.N., 1984. Coastal Paleogeography of the Central and Western Mediterranean during the last 125,000 years and its Archaeological implications. Journal of Field Archaeology 11, 307–314. Simo´n, M.D., 2003. Una secuencia con mucha prehistoria: la Cueva de Nerja. Mainake 25, 249–274. ˜ o, I., Reyes, E., Linares, J., 1991. Late Quaternary climatic Va´zquez, A., Zamarren changes on the southwestern Balearic slope (Western Mediterranean): isotopic, faunal, and mineralogical relationships. Palaeogeography, Palaeoclimatology, Palaeoecology 81, 215–227. Vera, J.L., Lozano, M.C., Simo´n, M.D., Corte´s, M., 2003. Relevancia del estudio de los moluscos en yacimientos prehisto´ricos, un caso bien documentado: la Cueva de ˜ a). Plioce´nica 3, 88–106. Nerja (Ma´laga, Sur de Espann Vera, J.L., Navarrete, I., Lozano, M.C., 2004. Los escafo´podos fo´siles (Mollusca, Sca˜ a). phopoda) del yacimiento prehisto´rico del Hoyo de la Mina (Ma´laga, Espan Plioce´nica 4, 36–50. ˜ ola. In: I Villaverde, V., 2006. Arte paleolı´tico de la regio´n mediterra´nea espan Jornadas de Patrimonio en la comarca del Guadalteba. Centenario de los descubrimientos del arte prehisto´rico en Ma´laga (1905–2005), Ma´laga, pp. 61–70.