N. García & J.L.Arsuaga Universidad Complutense de

Last Glaciation cold-adapted faunas in the

García, N. & Arsuaga, J.L., 2003 - Last Glaciation cold-adapted faunas in the Iberian Peninsula - in: Reumer, J.W.F., De Vos, J. & Mol, D. (eds.) - ADVANCES IN MAMMOTH RESEARCH (Proceedings of the Second International Mammoth Conference, Rotterdam, May 16-20 1999) - DEINSEA 9: 159-169 [ISSN 0923-9308] Published 24 May 2003

The present paper presents a compilation and update of sites which contain last glaciation faunas in the Iberian peninsula. Some authors consider that, except in the Pyrenees, the Iberian glaciation was restricted to the OIS 2 and they only include this mountain range (sometimes together with the Cantabrian mountain range) in the distribution maps of cold-adapted faunas in Europe. Nevertheless, there were glaciation processes on other mountain systems placed at a more sout- hern latitude than the northern part of the Iberian Peninsula, and cold-adapted species have been found even in the Andalusian territory. The aim of this study is to provide detailed and updated information about Iberian sites with the presence of Mammuthus primigenius, Coelodonta anti- quitatis, Rangifer tarandus, Saiga tatarica, Ovibos moschatus, Gulo gulo and Alopex lagopus. This association represents typical cold-resistent forms of the last glacial in Eurasia and extends also to the Far East and Beringia (Kahlke 1999) so their presence in a number of sites in the Iberian Peninsula suggests cold conditions.

Correspondence: N. García & J.L. Arsuaga, Universidad Complutense de Madrid, ISCIII, C/Sinesio Delgado 4, Pab-14, 28029 Madrid, , e-mail: [email protected] & [email protected]

Keywords: Spain, Late-Pleistocene, fauna

INTRODUCTION In addition to either fossilised remains of For the selection of cold-adapted large mam- arctic fauna or artistic representations, there mals species, we have followed the sugge- is other evidence to indicate that large areas stions of Kahlke (1999: 77), and only consi- of the Iberian Peninsula had very open eco- der those inhabiting the arctic to subarctic or systems with few trees during the OIS 2 cold inner-continental regions: Mammuthus primi- period, and perhaps in earlier cold periods as genius, Coelodonta antiquitatis, Rangifer well. This evidence of periglacial conditions tarandus, Saiga tatarica, Ovibos moschatus, comes from three sources: the pollen record, Gulo gulo and Alopex lagopus. Since the idea glacial landforms and sediments. is to clearly separate the arctic faunas which The pollen spectra of the Cantabrian archae- provide clear data about the glaciation condi- ological (Sánchez Goñi 1993) show tions in the Iberian Peninsula, we have not cold and dry conditions with steppe-like included Ursus spelaeus, Bison priscus, vegetation during the maximum upper pleni- Megaloceros or other species in our study. glacial and lateglacial (Younger Dryas) or Although these species are usually associated OIS 2. These caves are all located below 400 with typical cold-adapted faunas, they are m above sea level. In Carihuela Cave, in also found in woody temperate zones without Southeastern Spain (and at around 1020 m arctic faunas. above sea level), the pollen spectra show

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Figure 1 Art findings from caves or open air sites representing cold adapted species.We have indicated the questionable occur- rences with a question mark in the legends and with a square symbol (instead of the corresponding faunal symbol) on the maps.

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similar forest regressions in the same periods, below 1000 m of altitude and descending to in levels containing thermoclastic scree. Even as low as 340 m in one exceptional case in the Levant caves, where fossil remains of (Asón Gorge). The larger glaciers of the arctic fauna have never been found, the sedi- Iberian Peninsula developed in the Pyrenees, ments assigned to the maximum upper pleni- with some descending to 700-800 m above glacial and late glacial indicate very dry sea level, with lengths of dozens of kilome- conditions with naked soils (Fumanal 1986). ters (52 km in one case, the glacier of Aneu The pollen spectra obtained in lakes or peat in the Noguera Pallaresa river) and with ice bogs seem to follow a similar pattern, such as thicknesses of more than 500 m and in some the continuous sequence from the site of cases more than 900 m. Padul, south of the city of Granada and loca- Although in the past some authors recognized ted at 1000 m above sea level. Grass steppes a Riss and a Würm glaciation, most researchers and open coniferous parklands would have today believe that the Iberian glaciers were been largely dominant during the maximum restricted to the maximum upper pleniglacial upper pleniglacial and lateglacial on the enti- and lateglacial (except, of course, in the re Iberian Peninsula. However, in coastal Pyrenees, where some glacial cirques and refuges of the Atlantic Ocean and the snowy hollows even survive today). Two Cantabrian and Mediterranean seas and levels of moraines corresponding to two lowermost valleys (Costa et al. 1990), deci- major ice advances are usually recognized: duous or Mediterranean woodlands survived the lowest belonging to the maximum upper and later expanded in the Holocene. The pleniglacial and the highest to the late glacial. annual mean temperature was perhaps 10º C However, some authors recognize very degra- colder than that of the present day. ded moraines in the Sierra Nevada, at a lower Before the maximum upper pleniglacial, altitude, that could correspond to an earlier the pollen record of the Iberian Peninsula is ‘Riss’ glaciation (Rubio et al. 1993). much poorer, although in Carihuela Cave a Although there is still not much information maximum lower pleniglacial is also recorded available for , it seems that there and there is a steppe phase recorded in the were no glacial processes nor a cold climate Padul sequence (Dupré 1988) of probably affecting the vegetation until the maximum equivalent age. Glaciers were present in the upper pleniglacial (Raposo 1995). main mountain ranges of the Iberian The chronology of the Last Glacial Peninsula during the maximum upper plenig- Maximum in the Iberian Peninsula has beco- lacial and lateglacial (Gómez & Pérez 1998). me a matter of great interest in relation to the In the Pyrenees, Cantabrian Mountain Range, problem of modern human colonisation of Galaico-Leonés System, Sierra de la Estrela Europe and Neandertal extinction, that took and Sierra de Gredos (the last two in the place during OIS 3 (van Andel & Tzedakis Central System), and Sierra Nevada, valley 1998). In the 1980s, some Spanish scholars glaciers developed. There were also icecaps established that Neandertals were replaced by irradiating valley glaciers in the Galaico- modern humans in the Iberian Peninsula later Leonés System. The Sierra Nevada glaciers than in other areas (Vega-Toscano et al. were the Southernmost ones in Europe. In 1988), and climatically at the beginning of a and the Iberian cold period (‘Würm III’) that would corre- System, the glaciers were restricted to basins, spond to the end of OIS 3. Since then, it has which were the focal point for their nourish- become clearer that in what is nowadays eco- ment, or to deep snowy hollows. Some of the logically the Mediterranean Iberia (i.e. south end moraines descended to a very low altitu- of the Ebro River), the Neandertals lasted for de in the Galaíco-Leonés System and Canta- perhaps more than 10.000 yrs after the first brian Mountain Range, in many instances modern humans arrived in the Atlantic Iberia

161 ADVANCES IN MAMMOTH RESEARCH DEINSEA 9, 2003

Figure 2 Fossil remains of cold adapted species from Iberian sites. Most of the fossil evidence occurs in the northern part of the Iberian Peninsula, but also in southern sites.We have indicated the questionable occurrences with a question mark in the legends and with a square symbol (instead of the corresponding faunal symbol) on the maps.

162 GARCÍA & ARSUAGA: Iberian cold-adapted faunas

(north of the Ebro River). The ‘Ebro frontier’ stone tools. When the fossil remains are out model suggests that the basin of the Ebro of archaeological context it is noted as ‘unde- represented an ecological barrier to the dis- fined cultural level’. The study includes not persal of the first modern humans in Iberia only the fossil remains recovered but also (Duarte et al. 1999). Some authors believe diverse artistic representations (portable and that a climatic deterioration around 30.000 parietal art) depicting arctic animals. However, yBP in the Mediterranean ecosystems caused since in some cases the art interpretations are the dispersal of modern humans south of the open to debate, we have indicated the Ebro river and the subsequent extinction of questionable occurrences with a question the last Neandertals, which would have disap- mark in the tables and with a square symbol, peared together with other Mediterranean instead of the corresponding faunal symbol, endemic fauna and flora (Raposo & Cardoso on the maps. The sites with absolute chrono- 1998). metric dates are also included in the text.

MATERIAL AND METHODS COLD ADAPTED TAXA IN THE The tables include an inventory of cold adap- IBERIAN PENINSULA ted faunal remains from Iberian archaeologi- cal sites and represent a compilation based on Reindeer (Rangifer tarandus) numerous published studies. The maps are elaborated by us to better show the approxi- art findings (Fig. 1) At Guadalajara, two mate geographic location of the sites. No arc- caves show reindeer representations, Cueva tic species have been reported up to now in del Reno, with one engraving (Alcolea et al. Portugal, and for this reason we did not 1997) and Cueva de la Hoz also with one include this territory in the maps. Further- engraving (Balbín & Alcolea 1994; Balbín et more, an approximate chronology of the sites al. 1995). In the open air site of is attempted when possible, in most cases (Salamanca), two engravings have been inter- based on the interpretation of the associated preted as reindeer figures (Balbín & Alcolea

Tabl e 1 Rangifer tarandus remains from Navarra, Guipúzcoa and Vizcaya.

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Table 2 Cantabrian sites with Reindeer (Rangifer tarandus) remains.

1994). In the Cantabrian Mountain range, the Aurignacian level and dated on 37, 340 ± Tito Bustillo cave site () presents 1000 yBP to 35, 480 ± 1000 yBP (Maroto et seven paintings and at Las Monedas cave al. 1996). (Santander) there are four reindeer images (Altuna 1996a). Finally at Altxerri cave Mammoth (Mammuthus primigenius) (Guipúzcoa) six engravings were found (Altuna 1996a). art findings (Fig. 1) In the Cantabrian Mountain Range, there are eight mammoth fossil remains (Fig. 2, Tables 1, 2 and 3) representations in the Pindal cave (Balbín et The first recordings of Rangifer tarandus in al. 1999), and Las Caldas cave presents three the Iberian Peninsula are those from Harlé superimposed plate engravings (Corchón (1908), who only included five sites: 1994, 1997), both sites are in Asturias. Aizbitarte IV, Solutrean- Another site from Asturias is La Lluera I with (Basque Country); Ojebar, Valle and Paloma, one questionable engraving (Gónzalez all Magdalenian (Cantabria); and Serinya, Echegaray & González Sáinz 1994). From Magdalenian (Girona). In Tables 1-3 we Cantabria, one painting from El Castillo cave include more recently excavated sites with and another one from Arco-B are recorded reindeer remains. The five remains of rein- together with two questionable representa- deer antler from Bora Gran d’en Carreres tions from Las Chimeneas cave and La (Girona, Table 3) show signs of extraction Pasiega cave (Ripoll & Ripoll 1992, which indicates that this antler fragments Gónzalez Echegaray & González Sáinz were transported to the sites as a raw material 1994). To the north of Burgos, in the Ojo perhaps over a long distance. Reclau Viver Guareña karst system, one mammoth painting (Girona) is ascribed to the Aurignacian period is found (Ortega & Martín 1986). This is one and dated from 30,190 ± 500 yBP (level A) to of the very few findings with an absolute 40,000 ± 1,400 yBP (level B) (Maroto et al. chronological age ranging between 11,540 1996). L'Arbreda (Girona) includes a very yBP and 10,950 yBP for five samples complete archaeological sequence from the (Corchón et al. 1996). These dates represent Solutrean to the late Mousterian periods and the last occurrence of woolly mammoth at the the Rangifer tarandus remains comes from Iberian Peninsula. In Madrid one engraving level H which is assigned to the early (now lost) at El Reguerillo cave has been

* In Buelna (Asturias, loc. no. 37 in Fig. 2) a proboscidean skeleton was found in a small chamber inside the coastal ‘Cueva de la Gesa’.This cave is only accesible during the low tide. It was attributed to Mammuthus primigenius (Balbin et al. 1999) but Mazo (1998) assigned it to Elephas antiquus.This second opinion was mainly based on the large dimensions of the pelvic bone and femur, but they fit well on the Mammuthus primigenius range (due to the high sexual dimorp- hism). No cranial nor humerus remains (which are the best diagnostic parts of the skeleton) have been recovered yet. The sediments where this elephant remains were found seem to correspond to sandstones of a fossil beach.This evidence is used by A. Mazo as an argument to adscribe the elephant to the last interglacial and so defend the Elephas antiquus interpretation.The discussion about the specific assignment will remain open until the rest of the animal be extracted from this site.

164 GARCÍA & ARSUAGA: Iberian cold-adapted faunas

recorded.There is also one Portugal). The Gruta Figueira Brava ivory sculpture representing a head of a wol- (Sesimbra: Portugal) is dated by 14C. verine carved on a proboscidean tusk at the However, more recent publications (Raposo II (Guadalajara) site. Furthermore, & Cardoso 1998, Raposo 1995) do not inclu- there is one engraving that could represent a de this taxon in the faunal list, so we have mammoth from Los Casares cave interpreted this as a consequence of a revi- (Guadalajara). sion. The lowermost level is 30,050 ± 550 yBP and the uppermost level is 30,930 ± 700 fossil remains (Fig. 2 *, Table 4) The first yBP (Raposo & Cardoso 1998). citation of Mammuthus primigenius in the Iberian Peninsula was that of Obermaier Wooly rhinoceros (Coelodonta (1916), who only included six sites, from antiquitatis) Cantabria (Mina Inadvertida at Pámanes; Minas de las Dolores y Angel at Udías), art findings (Fig. 1) The artistic representa- Oviedo (Cueto de la Mina at Posadas), tions of the wooly rhinoceros in the Iberian Girona (Clot del Lop at Olot; Cau de les Peninsula are very scarce, only two engra- Goyes at San Juliá de Ramis), and in vings are clear, one is the plate from Las Barcelona (Pedralves). According to Aguirre Caldas cave (Asturias) mentioned above (1968) there are ‘E. primigenius (?)’ remains (Corchón 1994, 1997) and the other one is in a fluvial deposit of the Manzanares river found on a cave wall at Los Casares (Madrid). However, in the original publica- (Guadalajara). Finally, at the open-air site of tion by Hernández-Pacheco (1950) only Siega Verde (Salamanca) one engraving is Elephas antiquus remains are recorded. Thus reported by Balbín & Alcolea (1994). no mammoth findings are clearly represented in the Madrid province. The southernmost fossil remains (Fig. 2, Table 5) Harlé Mammuthus primigenius finds in the Iberian (1920) described one skull of Coelodonta Peninsula are those from El Padul peat-bog at antiquitatis from Arenys de Mar (Barcelona). only 20 km from the city of Granada The Coelodonta antiquitatis remains from (Aguirre, 1973). A carpal bone was used for Arroyo Culebro in Madrid province (Arsuaga 14C analyses and an age of 35,790 ± 960 yBP & Aguirre 1979) are represented by two skul- was obtained (Martín pers. comm.). ls, one rostrum and one hemimandible of a Cardoso (1993) reported the presence of young individual (thus 3-4 individuals). The Mammuthus cf. primigenius at Algar Joao chronology is very problematic. Guerin’s Ramos and Gruta Figueira Brava (both in opinion (pers. comm. in Arsuaga & Aguirre

Tabl e 3 Rangifer tarandus remains from sites other than Basques and Cantabrian.

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Table 4 Mammoth (Mammuthus primigenius) fossil remains. See Note.

1979) was that "these remains do not seem is Unquera (Arsuaga & Aguirre 1979) and the very old, in any case, it is not the Rissian other is Udías with one skull, some teeth and form". The fossils were found in a gravel several more fragments. On the Catalonian quarry in a terrace of the small river Arroyo coast, there are several sites that have yielded Culebro, that was assigned ‘geomorphologi- wooly rhinoceros remains: El Toll cally’ to the Early Würm, perhaps equivalent (Barcelona; Moyá in Altuna 1972), to the OIS 4, although in the absence of any L’Arbreda, Bora Gran d’en Carreres (both in geochronological dating or any taxa of bios- Girona). And also at Labeko Koba, tratigraphical relevance, a later age (OIS 2) Urtaigako Leizea, Lezetxiki (at Guipúzcoa) cannot be completely excluded. Among the and Abauntz (Navarra; Altuna 1996a). associated lithic materials also coming from the same Arroyo Culebro gravel exploitation, Saiga (Saiga tatarica) there is a handaxe that would support an Early Würm age, if it was truly associated art findings (Fig. 1) There is only one refe- with the woolly rhinoceros remains, which is rence to two possible saiga engravings at the not totally certain. Arroyo Culebro is the site of Altxerri (Guipúzcoa; Altuna 1996a). southernmost occurrence for this species in the Iberian Peninsula (0º 3' 40'' E and 40º 17' fossil remains (Fig. 2, Table 6) The only 43'' N). There are two Cantabrian sites that Iberian site that has yielded saiga evidence is have yielded wooly rhinoceros remains, one Abauntz (Navarra) (Altuna, 1996b) with only

Table 5 Wooly Rhinoceros (Coelodonta antiquitatis) fossil remains.

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six fossil remains (five phalanges and one CONCLUSIONS tarsal bone). Since all the bones come from Cold-adapted species have been found in a one single foot, they could have been trans- great part of the Iberian Peninsula. The pre- ported as part of a saiga skin from other loca- sence of arctic faunas indicating truly perigla- lity. Although the saiga antelopes were very cial conditions in the Iberian Peninsula before abundant in the Aquitaine large plain (north the last glaciation cycle is not certain. There of the Pyrenees), the hilly landscape of the is an occurrence of Praeovibos sp. in Ata- region surrounding the Abauntz site would puerca, in Gran Dolina upper level 7, of early have been unsuitable for their existence. Middle Pleistocene age (Van der Made 1998). Praeovibos also occurs in the Lower Wolverine (Gulo gulo) Pleistocene of Venta Micena (Moyá-Solá 1987) in Granada (South of Spain). However, art findings (Fig. 1) At Jarama II it seems that Praeovibos is more an indicator (Guadalajara) the Upper Paleolithic ivory sta- of dry climate and steppe conditions (not tue mentioned earlier depicts the head of a necessarily very cold), whereas the Middle wolverine (Adán et al. 1995) and Late Pleistocene musk ox is clearly cold adapted. The same can perhaps be said of fossil remains (Fig. 2, Table 6) From the Vulpes preglacialis (the likely ancestor of the site of Lezetxiki level III (Guipúzcoa) there is Late Pleistocene arctic fox) that also occurs one maxilla. This level is attributed to the in Atapuerca / Gran Dolina Early Pleistocene Gravettian. At the Mairuelegorreta cave site level TD6, as well as in the Early Pleistocene (Alava) a single canine was recovered wit- site of Venta Micena and the Middle Pleisto- hout archaeological context (Altuna 1996a). cene site of Huescar 1 (both in Granada). If these two taxa are not periglacial indicators, Musk Ox (Ovibos moschatus) no arctic taxa are found prior to the OIS 4, Only a single remainder attributed to the thus re-inforcing the hypothesis that very musk ox has been recovered from the cold conditions never existed in the Iberian L’Arbreda (Girona) site. It comes from the Peninsula before the last glaciation (in the Gravettian-Solutrean transitional level OIS 4 and still more exaggerated in the OIS (Maroto et al. 1996). (Fig. 2, Table 6) 2). The exception to this rule could be the reindeer remains assigned by Obermaier Arctic fox (Alopex lagopus) (1916) to level 18 of El Castillo cave. This Amalda level VI (Guipúzcoa) is the only site lower level of the El Castillo sequence was that has yielded arctic fox fossil remains, described as ‘Acheulian’ and could belong to (Fig. 2, Table 6). This level is assigned to the the end of the Middle Pleistocene (OIS 6). Gravettian cultural level by Altuna (1996a).

Tabl e 6 Saiga (Saiga tatarica), musk ox (Ovibos moschatus), wolverine (Gulo gulo) and arctic fox (Alopex lagopus) fossil remains.

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ACKNOWLEDGEMENTS L’Anthropologie 103 (1): 51-92. This study is supported by DCYT: PB96- Balbín Behrman, R. de, Alcolea, J.J., Moreno, F. & Cruz, 1026-CO3-03/96. We are very grateful to L.A., 1995 - Investigaciones arquelógicas en la J.M. Carretero, A. Gracia, I. Martínez, C. Cueva de la Hoz (Santa María del Espino, Lorenzo, R.-D. Kahlke, and D. Mol for their Guadalajara). Una visión del conjunto actualizada - valuable comments and to Rolf Quam for his in: Balbín-Behrmann R. de, Valiente J. & Mussat help with the corrections on the English. M.T. (eds.) - Arqueología en Guadalajara. Patrimonio Histórico-Arqueología, Toledo: 39-53 REFERENCES Balbín Behrman, R. de & Alcolea, J.J., 1994 - Arte Adan Álvarez, G., Arribas, A. Barbadillo, J., Cervera Paleolítico de la meseta Española - in: Brunet T.C. & Estrada, R., García, M.A., Jordá, J.F., Pastor, J., Menéndez M. (eds.) - Arte Paleolítico - Complutum Sánchez, B., Sánchez, A., Sanchiz, B. & Sesé C., 5: 97-138 1995 - Prospecciones y excavaciones arqueológicas Barandiarán, I., 1995 - Arte mueble del Paleolítico en el alto valle del Jarama (Valdesotos, Guadalajara, Cantábrico: una visión de síntesis en 1994 - in: Castilla-La Mancha) - in: Balbín-Behrmann R., Brunet, T.C. & Menéndez Fernández, M. (eds.) - Arte Valiente J. & Mussat M. T. (eds.) - Arqueología en Paleolítico - Complutum 5: 45-79 Guadalajara . Patrimonio Histórico-Arqueología, Canal, J. & Carbonell, E., 1989 - Catalunya Paleolítica - Toledo: 111-124 Patronat Eiximensis, Girona, 443 pp. Aguirre, E., 1968 - Revisión sistemática de los Cardoso, J. 1993 - Contribuiçao para conhecimientos dos Elephantidae por su morfología y morfometría grandes mamíferos do Pleistoceno superior de dentaria - Estudios Geológicos 24: 109-167 Portugal - Unpublished PhD thesis Aguirre, E., Lhenaff, R. & Zazo, C., 1973 - Nuevos Carrión, J.S., Munuera, M. & Navarro, C., 1998 - The fósiles de elefantes en Andalucía - Estudios paleoenvironment of Carihuela Cave (Granada, Geológicos 29: 295-306 Spain): a reconstruction on the basis of palynological Alcolea, J.J., Balbín-Behrman, R. de, García Valero, investigations of cave sediments - Review of M.A. & Jiménez Sanz, P.J., 1997 - Nouvelles décou Palaeobotany and Palynology 99: 317-340 vertes d’art paléolithique dans le centre de la Corchón, S., 1994 - Últimos hallazgos y nuevas intrepre Péninsule Ibérique: la (Valdesotos, taciones del arte mueble paleolítico en el occidente Guadalajara) - L’Anthropologie 101(1): 144-163 asturiano - in: Brunet T.C. &. Menéndez Fernández, Altuna, J., 1972 - Fauna de mamíferos de los yacimien M. (eds.) - Arte Paleolítico- Complutum 5: 234-264 tos prehistóricos de Guipúzcoa - Munibe 24: 238- Corchón, S., 1997 - La corniche Cantabrique entre 315. 15 000 et 13 000 ans BP: la perspective donnée par Altuna, J., 1996 a - Faunas de clima frio en la Peninsula l’art mobilier - L'Anthropologie 101 (1): 114-143 Ibérica durante el Pleistoceno superior - in: Ramil- Corchón, S., Valladas, H., Bécares, J., Arnold, M., Rego P., Fernández Rodríguez, C. & Rodríguez Tisnerat, N. & Cachier, H., 1996 - Datación de las Guitián M. (eds.) - Biogeografía Pleistocena- pinturas y revisión del arte Paleolítico de cueva Holocena de la Península Ibérica - Santiago de Palomera (Ojo Guareña, Burgos, España) - Zephyrus Compostela: Xunta de Galicia: 13-39 49: 37-60 Altuna, J., 1996b - Primer hallazgo de restos óseos de Costa Tenorio, M., García Antón, M., Morla Juaristi, C. antílope Saiga (Saiga tatarica L.) en la Península & Sainz Ollero, H., 1990 - La evolución de los bos Ibérica - Munibe 48: 3-6 ques de la Península Ibérica: Una interpretación Arsuaga, P.M. & Aguirre, E., 1979 - Rinocerontes basada en datos paleobiogeográficos - Ecología. lanudos en la provincia de Madrid (Coelodonta anti- ICONA, Madrid, Fuera de Serie 1: 31-58. quitatis Blumenbach) - Boletin Real Sociedad Duarte Maurício, J., Pettitt, P.B., Souto, P., Trinkaus, E., Española Historia Natural (Geol.) 77: 23-59 Van der Plicht, H. & Zilhão, J., 1999 - The early Balbín Behrman, R. de, Alcolea, J.J. & González Pereda, Upper Palaeolithic human skeleton from the Abrigo M.A., 1999 - Une vision nouvelle de la grotte de El do Lagar Velho (Portugal) and modern human Pindal, Pimiango, Ribadedeva, Asturies - emergence in Iberia - Proceedings National Academy

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