Abstract
The aim of this chapter is to understand, from a multidisciplinary approach, Neanderthal landscapes and environment from layer J records from Abric Romaní rockshelter. The proxy data used are paleontological and paleobotanical records with natural or anthropic orgin. This study includes pollen, charcoal, small vertebrates (micromammals and anphibians), large mammals and malacofauna. The data yielded through these disciplines approached on the basis of different methodologies (anthracology, palynology, paleontology, and dental microwear and mesowear) provide an assemblage of data showing different aspects of the Neanderthal landscapes and environment. These proxies show a diverse landscape with forested and open landscape areas under a cold environment, which locally has yielded taxa reflecting more humidity rates and the presence of Mediterranean taxa reflecting a cooler climate. In this context, Neanderthals exploited a variety of biotopes for the obtaining of resources, pointing out their adaptability capabilities.
Introduction
The Abric Romaní sequence spans a chronological period of ca. 40–70 ka BP with level J dated at ca. 50 ka BP (Bischoff et al. 1988). Different paleoecological studies on the flora and fauna have already provided a vast range of data, with very little variability among their results (Carbonell et al. 1996; Burjachs and Julià 1994; Cáceres et al. 1998; Arteaga et al. 2001; Vaquero et al. 2001; Chacón et al. 2007).
The geological landscape of the Abric Romaní provides different biota for the development of flora and fauna assemblages. The shelter is located at an altitude of 350 m above sea level, faces north-east, and is approximately 200 m from the Anoia River. The geomorphology of the region features different areas, including plains, travertine cliffs and mountain chains (see Vallverdú et al. 2012). According to the geomorphological study, a water source caused dampness in the area above the shelter.
The main objective of this chapter is to provide an overview of the environment surrounding the Abric Romaní in order to contribute further information about the supply and distribution of the biotic resources on which the Neanderthals depended. This approach will also define the natural landscape from a regional and local perspective. This information, based on the distribution of fauna and flora, will allow a greater understanding of the environment and climate during MIS 3. It will also define the Neanderthals’ framework of adaptation to this environment. Finally, we hope to describe the shelter environment as it relates to habitability during the occupations.
This study incorporates different disciplines for the study of macromammals, microvertebrates and malacofauna and the botanical assemblage including pollen and charcoal.
Paleobotany
The paleobotany record from sublevels Ja and Jb includes pollen and charcoal remains. Pollen analyses are intended to provide information about vegetation, identifying different flora categories, which include herbs, trees and shrubs. The charcoal record, however, is subject to human activities and the data is based considerably on local woody plants (trees and shrubs). Both disciplines complement records on the vegetation that jointly contribute to a clearer interpretation of past landscapes and climates.
Methods
The pollen analysis was conducted using seven samples from level J from the stratigraphical section at square P45. The sediments were treated using the method proposed by Goeury and Beaulieu (1979), slightly modified from Girard and Renault-Miskovsky (1969), and according to the protocol developed by Burjachs (1990) and Burjachs et al. (2003). To the pollen diagram, we have added the average histogram of the seven samples.
The charcoal analysis was conducted using 1205 charcoal fragments that were scattered across the surface of the archeological sublevels or hearths. All the charcoal remains have an anthropic origin related to firewood management (see Allué et al. 2012). Charcoal analyses are based on the taxonomic identification of charcoal fragments through examining the three wood anatomy sections under a metallographic light microscope. A reference collection and a European wood anatomy atlas (Schweingruber 1990) were used to support the identifications.
Results
The results of the pollen analyses show a semi-open landscape with 50% arboreal pollen (AP) (Fig. 5.1). The trees dominating this landscape are pines (Pinus) and, to a lesser degree, junipers (cf. Juniperus), accompanied in cold protected and sunny areas by mesophilous taxa (deciduous Quercus, Corylus , Juglans, Rhamnus) and even Mediterranean mesothermophilous taxa (evergreen Quercus, Olea/Phillyrea, Syringa). We also found evidence of riverside vegetation which would include at least Alnus , Salix , Populus, Cyperaceae and Typha or Sparganium.
Pollen diagram from level J. (AP arboreal pollen; NAP non-arboreal pollen)
The shrubs we have identified include Buxus, Ephedra fragilis-type and a liana Vitis. The component of the open landscape would mainly have been composed of grasses (Poaceae), Compositae (Asteraceae) and Artemisia.
Charcoal analyses reveal only two taxa: Pinus sylvestris type (Scots Pine) and Salix (willow). Pinus sylvestris type is the dominant taxa, whereas Salix is represented by a single fragment (Fig. 5.2). There are relatively low percentages of undetermined conifers or undetermined fragments. These categories are due to the size of the fragments or preservation problems which hindered further identification.
Charcoal diagram from sublevels Ja and Jb
Discussion
From these results, we deduced that this region housed semi-open pine groves that would have permitted the proliferation of herbs in their understorey. We suggest that it would have been a rather patchy landscape with forested areas and other open areas dominated by prairies and Mediterranean steppes.
Charcoal analyses suggest that selection occurred concerning the management of resources which were the most abundant in the environment. From this record we can be certain that there was a well developed forested area in the near vicinity of the rockshelter. According to these data, the pine forests were probably mono-specific forests that may have had junipers as the understoried vegetation, which is represented in the pollen record but was not found in our charcoal analyses. Furthermore, the identification of Pinus sylvestris (Scots Pine) in the charcoal record points to the fact that during the period under study mountain range pine forests grew 300 m below their present day location.
In fact, the scarce arboreal vegetation cover would have been directly associated with the semi-cold interstadial (IS 14, Dansgaard et al. 1993) as well as with the scarcity of annual rainfall, poorly interspersed throughout the year. Furthermore, detailed analyses of the diagram show that the site is not located in a very cold region, which is suggested by the scarcity of birch (Betula) and, more importantly, by the presence of Mediterranean taxa (evergreen Quercus, Olea/Phillyrea and Syringa). In the north-eastern Iberian Peninsula, diagrams of Late Pleistocene vegetation found at similar altitudes to that of the Abric Romaní show an increase in Betula in later periods closer to the Last Glacial Maximum (Ros 1987; Burjachs and Renault-Miskovsky 1992), whereas in level J of Abric Romaní there are significant thermo-mesophillous taxa, which point to the existence of refugia.
The MIS 3 climatic conditions that existed during the Dansgaard–Oeschger oscillation corresponding to level J would have been very different from the conditions we find today. The presence of cedar (Cedrus) in the vegetation record suggests that dominant meteorological patterns with easterly blowing winds allowed the pollen to arrive from the North African Atlas mountains, since this tree was extinct at the end of the Pliocene and early Pleistocene in Europe (Beaulieu and Reille 1973; Barrón 1999; Carrión et al. 2000; Ravazzi et al. 2005).
Large Mammal Paleontology
Large mammal remains recovered in level J are fragmentary, especially the postcranial bones (Rosell et al. 2012), and so paleontological identification has mainly been performed using teeth (or a few mandibles or maxillae). Ten species were identified: five Carnivora, two Perissodactyla, and three Artiodactyla.
Systematic Study
Carnivora Bowdich, 1821
Carnivores are not very abundant. Among the carnivore remains, however, there was a relatively high degree of diversity, with a sample made up of five different species:
Bear, Ursus sp. was identified in sublevel Ja with a first phalanx.
Wolf, Canis lupus Linnaeus, 1758 was identified with a cervical vertebra and a deciduous tooth, both in Ja.
Fox, Vulpes vulpes Linnaeus, 1758 with one metapodial in sublevel Ja.
Lynx, Lynx sp. with a second phalanx in Ja, and an undetermined phalanx in Jb.
Hyena, Hyaena sp. was identified with a deciduous tooth in sublevel Jb.
Perissodactyla Owen, 1848
Equidae Gray, 1821
Horse, Equus ferus Boddaert, 1785
The horse remains from Abric Romaní were described by Sánchez (1989). On the upper teeth, the fossettes present a maximum of two foldings, in the mesial and distal parts. The caballine fold is well developed on the premolars and slightly outlined on the molars. The styles are well developed—on the premolars, they present grooves (striations) and on the molars, there is a tendency toward the splitting of the parastyle in two. On the lower teeth, the flexids (pre- and postflexids) are not highly folded. On the premolars, the hypoflexids (vestibular groove) do not penetrate the flexids, and they have a well developed caballine fold. On the molars, the ectoflexid moves toward the double knot but does not reach it; the caballine fold is well outlined only on some specimens. Comparing the metrical data with that from other Spanish sites indicates that the population at Abric Romaní is comparable to the horses of Cueva Horá and Cueva Negra (Sánchez 1989). The Abric Romaní horse would have been an intermediate form between the two Würmian subspecies germanicus and gallicus (Sánchez, 1989).
Rhinocerotidae Gray, 1821
Rhinoceros, Stephanorhinus hemitoechus (Falconer, 1868)
Rhinoceros remains are represented in level J by a deciduous premolar, a permanent third or fourth premolar, a phalanx of an immature individual, as well as a few fragmentary remains. This species was previously identified as Stephanorhinus hemitoechus in other levels at Abric Romaní where more significant remains were discovered.
Cetartiodactyla Montgeland et al., 1997
Cervidae Goldfuss, 1920
Red deer, Cervus elaphus Linnaeus, 1758
For the Cervidae, morphological features were examined on the upper and lower permanent dentition (Lister 1996). All the remains recovered from level J belong to a single species, Cervus elaphus. As described below, no evidence of fallow deer, Dama dama, has been found. All upper and lower teeth are represented in the assemblage from level J; however Table 5.1 only shows measurements of large samples.
The upper molars have columns, buccal cones and well pronounced styles. The central lingual column and its associated cingulum are also strongly marked. Finally, the antero-buccal edge of the teeth is straight toward the base, and not angled as in Dama.
On the lower premolars (Fig. 5.3), the entoconid and the entoconid wing are oriented following the transverse axis of the teeth, and the lingual division between the entoconid and the posterior wing of the hypoconid is not as deep as found in Dama.
Left mandible of red deer (Cervus elaphus) from Abric Romaní (AR’94 Ja P51). Buccal view (a) and lingual view (b). Abbreviations: bc buccal column; lc lingual column; ec entoconid column
On the lower molars (Fig. 5.3), the lingual columns and furrows in between are stronger than in Dama. In addition, unlike in Dama, the entoconid column is straight and does not curve anteriorly. More specifically, the central buccal column is stronger in M2 than in M1. Finally, on M3, there is no clear step between the second and third lobes (entoconid and hypoconid).
The measurements were compared to other Late Pleistocene locations in Spain and southern France (Table 5.1): Cueva Millán, Burgos (Pérez-Legido and Cerdeño 1992); Kiputz IX, Gipuzkoa (Castaños et al. 2006); and Adaouste, Bouches-du-Rhône (Defleur et al. 1994). The population from Abric Romaní fits within the limits of variations of Late Pleistocene populations, with some larger and some smaller dimensions. These differences are certainly related to sample size.
Bovidae Gray, 1821
Aurochs, Bos primigenius Bojanus, 1827
Identifying the dental remains of aurochs (Bos primigenius) and bison (Bison priscus) is not easy, but several criteria have been proposed to help in making the distinction. Molars are particularly useful for identifying to the genus level (Slott-Moller 1990). In all the specimens analyzed, we only found morphological criteria of Bos. Measurements are presented in Table 5.2.
On both the lower and upper molars, the central buccal column is straight, starts on the cervix (cementoenamel junction) and extends up to the occlusal surface. Other criteria for identification are the shape of the pillars on the protocone and hypocone (upper teeth) and the protoconid and hypoconid (lower teeth). In Bos, these pillars are straight, shaped like a column without thickening at the cervix (Moullé 1992).
The upper third and fourth premolars, in mesial view, have sub-parallel lingual and labial edges that do not converge towards the occlusal surface as in Bison. On the upper third molar, the lingual connection between the two lobes forms an open angle. The hypocone is V shaped.
The lower third molar is one of the characteristic remains used for distinguishing between the two genera, and particularly the shape of the third lobe (Slott-Moller, 1990). In Bos, the third lobe forms a continuity with the lingual side of the tooth (paraconule and metaconule), and forms a largely open angle with the labial side (protoconule and hypoconule) (Fig. 5.4).
Right m3 of aurochs (Bos primigenius) from Abric Romaní (AR94 Ja L48 27). Buccal view (a) and lingual view (b). Abbreviations: bc buccal column
Pyrenean chamois, Rupicapra pyrenaica Bonaparte, 1845
Remains identified as Caprinae in level J are very rare. Only four specimens were attributed to the genus Rupicapra: an upper third molar, fragments of an upper second or third molar, a lower third molar (bud), and a fragment of a lower molar. From these specimens, we chose to focus our paleontological analysis on the upper M3 because it is the only complete, well preserved, and measurable specimen.
The third upper molar (AR93-Ja-U43-3) has a reduced metastylar wing (Fig. 5.5) and the lingual side of the second lobe has a rounded section without the typical pinching of Hemitragus. The metastyle forms a small metastylar wing and the distal interstylar surface is smaller than the proximal surface. These features are typical of the species Rupicapra pyrenaica.
Left M3 of Pyrenean chamois (Rupicapra pyrenaica) from Abric Romaní (AR’93 Ja U43 3). Lingual view (a) and buccal view (b). Abbreviations: dis distal insterstylar surface; mw metastylar wing
The metrical analysis (Table 5.3 and Fig. 5.6) shows that the population from Abric Romaní bears a very strong resemblance to the population from Cova de l’Arbreda (Serinyà, Spain), south of the Pyrenees, compared to the reference population from Portel-Ouest (Loubens, France) in the northern Pyrenees. This similarity indicates homogeneity of size and morphology in the populations of chamois in the southern Pyrenees.
Ratio diagram of M3 average measurements for Rupicapra pyrenaica (Standard base: Portel-Ouest cave). See Table 5.3 for description of measurements
Rupicapra pyrenaica is often depicted as a mountain ungulate, but its Pleistocene distribution indicates that it is not only adapted to high altitude habitats. The species preferentially inhabits pastures and rocky areas in diverse mountain regions of Europe. It is found in many Pleistocene locations at low altitudes, such as at Arago Cave (Crégut-Bonnoure 1979) or Les Cèdres (Defleur et al. 1989). The presence of Pyrenean chamois around Abric Romaní is not exceptional for the Mediterranean area. The population may have taken advantage of the travertine cliff.
Dental Microwear and Mesowear Analyses: Reconstruction of Paleodiets and Paleoenvironments
Large mammals, and especially ungulates, are commonly used to create paleoenvironmental reconstructions through various types of proxies (hypsodonty, body mass, biodiversity) and methods of analysis (ecological groups, climatograms, cenograms, multivariate analysis). However, these methods have some limitations, especially when applied to the Pleistocene. First, they are based on parameters that strongly reflect the phylogenetic history of a group, i.e. they are linked to the adaptations of ancestors. Thus, they provide clues, but do not necessarily indicate the immediate behavior or ecology of a population (Solounias et al. 2004). Also, there are many limitations when these methods are applied to archeological sites, because human behavior implies the intentional selection of the animal hunted and brought back to the site (Valensi and Abbassi 1998). We decided to use dental wear analysis methods (meso- and microwear) because they provide insight into the diet of the individuals, i.e. into the immediate behavior of the population. The results of the mesowear analysis reflect the diet of the last weeks or months (Fortelius and Solounias 2000), whereas the microwear analysis yields information on the last few days or week (Solounias and Semprebon 2002). These methods therefore provide a snapshot of the local environment around the site at the time Neanderthals occupied the shelter.
Methods
Mesowear analysis was performed in keeping with that described by Fortelius and Solounias (2000). Mesowear is based on physical properties of ungulate foods as reflected in the relative amounts of attritive (tooth-on-tooth) wear and abrasive (food-on-tooth) wear on the dental enamel of the occlusal surfaces. Low abrasion diets, such as those of browsers, generate an attritional (sharpening) relationship between the upper and lower cusps. In contrast, abrasion-dominated (food-on-tooth) wear, associated with grazing diets, results in more rounded and blunted wear facets and less precise occlusion. Only adults are found to have consistent signs of mesowear (Rivals et al. 2007a) and were therefore selected for study. Mesowear was scored on the paracone of the M2. In the original formulation of the method, mesowear was recorded by characterizing the buccal apices of molar cusps as sharp, rounded, or blunt, and the valleys between the cusps as high or low. The original data reported by Fortelius and Solounias (2000) on extant ungulates were converted into a more simplified univariate score representing a continuum of mesowear stages from the highest and sharpest cusps (0) to cusps that are completely blunted with little or no relief (3) (Rivals et al. 2007b). Intermediate stages of mesowear consisting of more rounded cusp apices with higher and lower levels of cusp relief were assigned scores of 1 and 2, respectively. Teeth with low relief and sharps cusps were assigned a score of 2.5. Individual scores were then averaged for each sample.
Microwear analysis was performed using the method described by Solounias and Semprebon (2002) and Semprebon et al. (2004). Further discussion of this method and its application in dietary reconstruction can also be found in Godfrey et al. (2004) and Palombo et al. (2005). Molar teeth (lower and upper M2) were selected by excluding young and old individuals. High-resolution epoxy casts were made for all available upper and lower second molars in sublevels Ja and Jb. The casts were screened using a stereomicroscope and any specimens exhibiting signs of weathering were excluded. Microwear features were identified and quantified on high resolution epoxy tooth casts at 35× magnification using a stereomicroscope. Most microwear features can be categorized as pits and scratches of various sizes and textures. Pits are circular or sub-circular microwear scars. Small pits are relatively shallow, refract light easily, and appear bright and shiny. Large pits are deeper, wider, and less refractive. Large pits are recorded qualitatively as being present or absent on the wear surface of the tooth. Scratches are elongated microfeatures with straight, parallel sides and can be subcategorized as fine or coarse. Scratch texture is evaluated on the basis of general appearance and light refractive properties (0 = fine scratches only; 1 = mixture of fine and course, 2 = only coarse scratches). Cross scratches are oriented somewhat perpendicularly to the majority of scratches observed on dental enamel (Solounias and Semprebon 2002). To approximate pit and scratch frequency, they are counted in a standard 0.4 × 0.4 mm square area on the lingual (inner) band of enamel on the paracone of the upper second molar.
Results
Mesowear scores recorded for sublevel Ja, where all three main species are present, indicate a high diversity of dietary resources ranging from low abrasion for Cervus elaphus (MWS = 0.5) to high abrasion for Equus ferus (MWS = 2.5) (Table 5.4 and Fig. 5.7). High abrasive resources exploited by Equus ferus indicate a definite grazing behavior for that species (as well as in sublevel Jb). Bos primigenius plots in an area where all dietary categories overlap, which is characteristic of a mixed feeding behavior and indicates that this species fed on both grass and browse. Cervus elaphus have a low mesowear score indicating a diet mainly composed of low abrasive items such as leaves from trees or bushes.
Mesowear scores for the fossil ungulates from Abric Romaní (solid black dots) compared to modern wild ungulates (white dots). Data on modern species from Fortelius and Solounias (2000). Abbreviations: LB leaf browsers, MF mixed feeders, G Grazers
The diversity of dietary behavior observed in sublevel Ja reveals the presence of various habitats in the environment surrounding the Abric Romaní.
For the microwear analysis, our sample consisted of 116 teeth which were deemed suitable for analysis (Table 5.5): 81 from Equus ferus (63 in Ja and 18 in Jb), 22 from Cervus elaphus (20 in Ja and two in Jb), 11 from Bos primigenius in Ja, and two from Rupicapra pyrenaica in Ja. Samples with fewer than eight specimens are generally considered too small to obtain definite results and are usually excluded from the studies. However, because our study includes a low number of different species, we decided to include the results for the two small samples for the purpose of comparison.
The summary statistics of the microwear features found for the three species are presented in Table 5.5. Two species, Cervus elaphus and Equus ferus, were found in both Ja and Jb. The results from only one sample for each of these species are presented in Table 5.5 as the samples overlapped. The microwear parameters recorded for these two species are not significantly different in the two sublevels. This indicates that there are no significant differences between Ja and Jb for Cervus elaphus and Equus ferus: there were no changes in diet for either species during the two occupations (Fig. 5.8).
Bivariate plot of the average numbers of pits and scratches in extant ungulates (data from Solounias and Semprebon 2002) and the fossil ungulates from Abric Romaní. Convex hulls are drawn around extant browsing and grazing taxa for ease of comparison. Abbreviations for the fossil species: BP Bos primigenius CE Cervus elaphus; EF Equus ferus; RP Rupicapra pyrenaica. Abbreviations for extant species: see Fortelius and Solounias (2000)
Microwear data gave definite results for all the species analyzed. Cervus elaphus, with a low number of pits and scratches, plots within the browsing morphospace, confirming the low abrasive diet suggested by mesowear.
Bos primigenius and Equus ferus, with more scratches and fewer pits, plot within or beyond the grazing morphospace. For Equus ferus, this confirms a diet with high grass content. Bos primigenius falls within the grazing morphospace but the mesowear analysis did not reveal very high abrasive levels. This discrepancy between the two methods is characteristic of mixed feeders (Semprebon and Rivals 2007); it reveals changes in diet on a seasonal basis. Bos primigenius was hunted during the season it was preferentially feeding on grasses (Rosell et al. 2012).
Rupicapra pyrenaica falls among the mixed feeders; however, only two teeth were available for analysis. Nevertheless, the pattern observed at Abric Romaní is similar to that found in other samples from other sites in the Mediterranean area (Rivals and Deniaux 2005), indicating a mixed feeding behavior. Compared to other species, the teeth of Rupicapra pyrenaica have a relative high number of pits, which is characteristic of exogenous abrasive particles such as grit, dust, or soil (Semprebon and Rivals 2007) found in animals that either feed close to the ground, or that feed on plants covered by grit or dust.
Discussion
The assemblage of ungulates identified at Abric Romaní is typical of the Late Pleistocene fauna of the Mediterranean area. The species have morphologies and dimensions comparable to other populations from Late Pleistocene locations in Spain and southern France. The presence of horses, red deer, aurochs and Pyrenean chamois in equal numbers illustrate a diverse environment and a variety of habitats in the vicinity of the site.
Our findings suggest that the ungulates hunted by the occupants of level J had very diverse feeding behaviors and enjoyed a large variety of plant resources. Bos primigenius and Equus ferus ingested large amounts of highly abrasive plants, such as grasses. This is especially true of Equus ferus, which was found to subsist on a diet containing a relatively high abrasive content compared to modern ungulates. Because grass is a larger part of the diet of the fossil species when it is abundantly available in the environment, we propose that at the time of their deaths both Bos primigenius and Equus ferus were selectively feeding on grasses. However, the mesowear analysis indicated that Bos primigenius was a mixed feeder, whose diet changed according to the seasons. These two species were probably living in very similar open habitats where grasses were abundant, such as grasslands. On the other hand, Cervus elaphus fed on vegetation with a low abrasive level. It probably inhabited more closed habitats than Bos primigenius and Equus ferus, and had a diet composed of branches, twigs and leaves of woody plant species.
Despite the small sample size, Rupicapra pyrenaica revealed a microwear pattern similar to that found in other Pyrenean chamois populations which inhabited pastures and rocky areas at low altitudes, such as the population from Arago Cave in France (Rivals and Deniaux 2005).
All four species lived in very diverse habitats, thus revealing the diversity of biotopes present around the site during the deposition of level J. This also provides insight into the biotopes exploited by Neanderthals for their subsistence activities at that time. The pattern in level J is quite different from that found in the underlying level M, where Bos primigenius, Equus ferus, and Cervus elaphus were found to live in very similar open habitats (Fernández-Laso et al. 2011).
Small Vertebrates
The small vertebrate assemblage from level J is made up of 201 identified fossils remains. The most representative remains belong to the order of rodents (94.5%), although other taxa such as amphibians (4.5%) and insectivores (1%) are also represented, albeit to a lesser degree (Fig. 5.9).
Distribution of small vertebrates in level J of the Abric Romaní in Number of Identified Specimens (NISP) and Minimum Number of Individuals (MNI)
Systematic Study
We analyzed 29 fossil specimens corresponding to 21 small mammal remains and eight amphibian remains (Fig. 5.9).
Amphibians
Anura Fischer von Waldheim, 1813
Bufonidae Laurenti, 1768
Bufo bufo (Linnaeus, 1758)
One right humerus of a female specimen (Fig. 5.10)
a Bufo bufo, right humerus of female (AR98/Ja/L42), ventral view. b–c Rana temporaria, left and right ilia (AR95/Ja/R51), lateral and posterior views
The genus Bufo is characterized by a thick humeral shaft without a paraventral crest and a slightly laterally displaced condyle. The poor development of the condyle and epicondyles, in ventral view, as well as the size of the fossil (preserved length = 30 mm) are consistent with B. bufo, which is the largest bufonid in western Europe. The absence of a well developed mesial crest and the slightly curved shaft are characteristic of the female.
The common European toad (B. bufo) has an extensive Eurasian range. It lives in nearly all environments, even in dry areas. Its only requirement seems to be, during its breeding season, the presence of calm or low energy water, preferably permanent and with vegetation (Lizana 2004). In Catalonia, the distribution of this species is relatively continuous and very common, although it seems to be rarer on the Lleida plain and in the highest mountain areas of the Pyrenees (Llorente et al. 1995).
Ranidae Rafinesque, 1814
Rana temporaria Linnaeus, 1758
Two ilia, one vertebra, one radioulna, two tibiofibulae and one phalanx (Fig. 5.10).
Among the representatives of the family Ranidae, the ilium permits a relatively certain attribution at the species level (Böhme 1977; Esteban and Sanchiz 1985, 1991; Bailón 1999; Gleed-Owen 2000; Blain 2005). In the genera Rana (brown frogs) and Pelophylax (green frogs), the ilium has a more or less high dorsal crest on the ilial shaft, thee pars ascendens is relatively short and the postero-medial side is smooth and without any sulcus interiliacus. The two fossil ilia are incomplete, as they lack the anterior part. The juncture ilioischiatica has a fairly straight medial outline, and the ventral acetabular wall is thin, a particular characteristic of brown frogs. The acetabular diameter vs. thickness (d/t ratio, sensu Gleed-Owen 2000) is 3.71 and 3.57, which is well within the range of 2.75–4.00 of brown frogs, and well outside the range of 2.12–2.88 measured for green frogs (Gleed-Owen 2000). The tuber superior is unangled and the pars descendens is postero-ventrally directed, whereas R. arvalis has an angular tuber superior and R. dalmatina and R. iberica a ventrally-directed pars descendens (Gleed-Owen 2000 and personal observation). The dorsal crest is rather low although it lacks the typical inflection beyond the tuber superior. According to measurements made by Esteban and Sanchiz (1985, 1991), the height of the dorsal crest is consistent with R. temporaria. Other bone fragments fit well into the genus Rana.
The common brown frog (R. temporaria) is a broad-range Eurasian species. In Catalonia, the species only lives in a narrow strip in the north consisting of pre-Pyrenean and Pyrenean areas with high levels of rainfall (average annual precipitation over 800 mm) and cool temperatures (average annual temperature <12°C and average January temperature <5°C) (Llorente et al. 1995, Balcells 1975). At present, its southernmost representatives correspond to a quite isolated population in the Massif of Montseny (Llorente et al. 1995). This population can be found at altitudes ranging from 300 to 2800 m above sea level with larger numbers between 1000 and 1800 m, and lives in various non-Mediterranean environments, such as peat bogs, alpine meadows and wooded river biotopes.
Mammals
Rodentia Bowditch, 1821
Muridae Iliger, 1811
Arvicolinae Gray, 1821
Iberomys cabrerae Thomas, 1906
Three first lower molars (M1) (Fig. 5.11)
a First lower right molar Iberomys cabrerae (AR95/Ja-b/L45), b–c first lower left molar Iberomys cabrerae (AR95/Ja-b/L45—AR95/Jb/K49). Occlusal view
The Iberomys lineage has Iberomys cabrerae, the Iberian vole, as its sole extant representative. It is descended from the Iberomys brecciensis species, which appears in the Middle Pleistocene. The Iberian vole is characterized by its relatively long and wide lower molar (M1), the reduction of the triangles of the anteroconid complex (ACC), a long and narrow fifth lingual salient angle (LSA5), which is a measure of its latero-medial asymmetry, also known as labio-lingual asymmetry (Cuenca-Bescós et al. 1995), a fourth buccal salient angle (BSA4) with a quadrangular shape, and enamel completely covering the labial wall of the ACC (Ayarzagüena and López Martínez 1976). These characteristics are present in the specimens from level J (Fig. 5.11). An analysis of the variability in the size of M1 of the Abric Romaní specimens compared to other specimens from the Middle Pleistocene (Galería in Cuenca-Bescós et al. 1999), the Late Pleistocene (Cova del Gegant in López-García et al. 2008) and the Holocene (Cova Foradada; Bronze Age) establishes that the size falls within the range of the extant Iberomys cabrerae (Fig. 5.12).
Comparison of the first lower molars (m1) length (A) and width (B) of Iberomys cabrerae from level J of the Abric Romaní (n = 3) with Iberomys cabrerae from the Cova del Gegant (n = 7) and Cova Foradada (n = 5), and with Iberomys brecciensis from the Middle Pleistocene of Atapuerca-Galeria (n = 9; Cuenca-Bescós et al. 1999). Measurements are shown in mm
The Iberian vole (I. cabrerae) is an endemic species of the Iberian Peninsula, which today has a restricted and discontinuous geographic range (around the Central System, southern Iberian System, sub-bethics Mountains and several points in the Aragon Pyrenees). The Iberian vole is not currently present in Catalonia. This species is characteristic of a Mediterranean climate, with a preferential habitat featuring perennial vegetation and water streams (Blanco 1998a).
Microtus arvalis (Pallas, 1779)
Two first lower molars (M1)
The genus Microtus is characterized by the presence of closed triangles (T4–T5) in the lower first molars (M1). Furthermore, the symmetric and parallel disposition of the triangles T4/T5 and especially T6/T7, which display a round morphology of the anteroconid complex (ACC), are specific characteristics of the species Microtus arvalis.
The common vole (M. arvalis) is found in temperate regions of southern Europe. It is a rodent which mainly lives in open and not overly wet meadows. It is normally found at altitudes ranging from 900 to 2000 m above sea level with an average annual precipitation exceeding 800 mm. In Catalonia, this species lives in the upper Pyrenees and at several locations in the northern pre-Pyrenees (Blanco 1998a; Gosàlbez 1987).
Arvicola sapidus Miller, 1908
12 first lower molars (M1)
The genus Arvicola is characterized by the presence of a posterior fold in the first lower molar (M1) followed by three closed triangles. The disposition of an acute angle in the lingual side, towards the internal edge of the anterior cusp (AC) in the first lower molar and, the “Arvicola” enamel pattern, thicker on the distal side than the proximal side, are morphological characteristics of the species Arvicola sapidus.
The southern water vole (A. sapidus) has a restricted range in Europe, and can be found throughout most of France and the Iberian Peninsula. In Iberia, it has a homogenous territorial distribution which includes Catalonia. It is a rodent species that prefers permanent water streams, such as rivers or irrigation ditches with slow-moving, deep water at constant levels (Blanco 1998a).
Murinae Illiger, 1811
Apodemus sylvaticus (Linnaeus, 1758)
One mandible (M1–M2) and maxilla (M1–M3)
The genus Apodemus is characterized by the presence of a low occlusal surface with six main cusps in the first lower molar (M1). The anterolingual and anterolabial cusps of the M1 are arranged in an X shape. In some specimens, these cusps are separated by a deep, narrow furrow. The posterior cusp (PC) of the M1 is low, rounded and well developed. On the labial side of the M1, there are two or three accessory cusps (C) and one proximal tubercle (TMA) (Cuenca-Bescós et al. 1997). Moreover, the conjunction between the tubercles T4 and T7 on the first upper molar (M1) and the development of the tubercle T9 on the second upper molar (M2) (Pasquier 1974) are the morphologic characteristics that ascribe our material to the Apodemus sylvaticus species.
The wood mouse (A. sylvaticus) has a large range in eastern and western Europe. It is distributed continuously throughout the Iberian Peninsula, and in Catalonia, it is found from the Pyrenees to the Mediterranean coast. A. sylvaticus is a generalist species which lives in a variety of habitat types, although it prefers the marginal zones of deciduous forests with the presence of water streams. It is found at altitudes of up to 2,000 m (Gosàlbez 1987).
Erinaceomorpha Gregory, 1910
Talpidae Fischer, 1817
Talpinae Fischer, 1817
Talpa Linnaeus, 1758
One scapula and one radius
Two species of moles are present on the Iberian Peninsula today: Talpa europea (common mole) and Talpa occidentalis (Iberian mole). According to Cleef-Roders and Hoek-Ostende (2001) the diagnostic characteristics for distinguishing between the two species are the morphology and measurements of the dentition and mandible and the measurements of the humerus. Due to the scarcity of diagnostic elements, we have considered our specimen to be Talpa sp.
The mole is a eurithermic genus, found equally in cold and warm environments, but it prefers deep soils with a high degree of humidity (Blanco 1998b).
Microvertebrate Taphonomy
The accumulation of small vertebrates is usually a consequence of the predation of nocturnal or diurnal birds of prey and small carnivorous mammals (Mellett 1974; Mayhew 1977; Korth 1979). The small vertebrate fossil assemblage cannot therefore be considered an exact reflection of life in the paleocommunity because the specimens found here could have been selected by predator diet criteria. A taphonomic (biostratinomic) study based on a descriptive-systematic method developed by Andrews (1990) allows us to identify the predator which left the accumulation, and these data in turn permit us to make paleoecologic interpretations of the fossil assemblages (Andrews 1983; Andrews and Nesbit-Evans 1983; Andrews 1990; Kowalski 1990; Fernández-Jalvo and Andrews 1992; Denys et al. 1995; Fernández-Jalvo 1995; Cuenca-Bescós 2003).
The rodent assemblage is made up of 13 individuals and 122 remains. The mean anatomic representation of elements is low (29.6%) (Table 5.6A). The Pc/C index (Table 5.6B) indicates that postcranial remains are better represented than cranial elements and we found a very high degree of fracture in the cranial and postcranial elements analyzed. This is related to post-depositional incidents, which break and reduce the number of elements left at the site (Cáceres et al. 2012).
Most of the elements we analyzed did not show signs of digestion (Fig. 5.13). Light digestion is basically only observed on femur and humerus bones. Moderate digestion is only represented on an isolated lower first molar (M1) of Microtus arvalis (Fig. 5.14). According to Andrews (1990) this degree of alteration is included in modification category 1, which comprises several bird predators. However, only two of these predators could possibly be responsible for the small vertebrate accumulations of level J: Asio otus (long-eared owl) or Tyto alba (barn owl).
Percentage of digestion degree of the small vertebrates
Habitat interpretation through the small vertebrate association. The data represent the percentage of the association taxa by habitat
Both species can be found in semi-open forests with large clearings nearby, at altitudes below 1300–1500 m. Generally, Tyto alba prefers to hunt in clearings or semi-clearings and nest in hollow trees and rocky outcrops. Asio otus is associated with deciduous, coniferous (especially pine) or mixed forests near watercourses (König et al. 1999).
Discussion
Iberomys cabrerae is a species endemic to the Iberian Peninsula. Although it can no longer be found here, it is well represented at numerous sites on the Iberian Peninsula dating to the second half of the Late Pleistocene and the beginning of the Holocene, such as Cova de les Cendres (Alicante) (Guillem-Calatayud 1999), Cova Bolumini (Alicante) (Guillem-Clatayud 1999), Baños de Mula (Murcia) (Agustí et al. 1990), El Portalón (Atapuerca, Burgos) (López-García et al. 2010a) and Cova Colomera (Lleida) (Oms et al. 2008; López-García et al. 2010b). Their movement from the Iberian Peninsula may be linked to two factors. First, there may have been competition with other species that shared the same ecological habitat, such as Microtus agrestis (common vole)—a species that today is very abundant in areas that are not overrun by the Iberian vole (Iberomys cabrerae). The second factor may have been environmental changes that occurred after the Bronze Age. The Iberian vole is commonly associated with wet meadows, and its move from the peninsula was quite likely brought about by increased dryness, such as during the Medieval Warm Period between 800 and 1300 AD.
The common frog (Rana temporaria) is a Euro-Siberian species with a habitat ranging from eastern Europe to the Urals, reaching as far as Norway in the north. In Spain, it has a continuous distribution along the north, where it finds a high degree of humidity and a cool climate (Esteban and García-París 2004). In the fossil record from the late Lower Pleistocene, Gran Dolina (TD5, Burgos) and Cal Guardiola (Barcelona) have produced the oldest evidence of these frogs on the Iberian Peninsula to date (Blain et al. 2008; Blain et al. 2009). The presence of R. temporaria at the Abric Romaní, as well as at Gran Dolina and Cal Guardiola, at the moment represents the only record of this species located slightly south of its present distribution, and is probably related to colder and wetter periods.
The small vertebrate assemblage of level J is dominated by taxa linked to woodland edges (50%) such as Apodemus sylvaticus and Rana temporaria, and taxa related to open wet meadows (23%) and water streams (22%) such as Iberomys cabrerae, Talpa sp., Arvicola sapidus and Rana temporaria. This association indicates the predominance of wetter conditions than are currently found in the region.
The intersection of the present distributions of all the species occurring in one location may indicate possible climatic conditions (Blain et al. 2008; Martínez and Sanchiz 2005). In order to evaluate paleoclimatic parameters we use the principle of mutual climatic range, which consists of defining the climatic conditions of the area currently inhabited by the extant fauna from the site. This method yields a total of two 10 × 10 km UTM squares situated in the external ranges of the Pyrenees in Huesca. Such an intersection for level J suggests mean annual temperatures (MAT) (recent data are from Font Tullot 2000) equal to 8 ± 1.4°C, (minimum = 7°C, maximum = 9°C), and mean annual precipitation (MAP) equal to 875 ± 35 mm (min. = 850 mm, max. = 900 mm). The mean temperature of the coldest month (MTC) for level J is 2.75 ± 0.35°C (max. = 3°C, min. = 2.5°C), and for the warmest month (MTW), 20.25 ± 0.35°C (max. = 20.5°C, min. = 20°C) (López-García and Cuenca-Bescós 2010).
Malacofauna
The malacofauna studied comes from sublevel Ja of the Abric Romaní. All the specimens identified belong to continental species, both terrestrial and aquatic, which at present are common in the north-east of the Iberian Peninsula. The information available on their present habitat allows us to understand the environmental conditions in place during their lifetimes and contributes to our understanding of the habitability of the shelter.
Materials and Methods
The studied assemblage includes 746 mollusk remains collected in 90 squares, involving five of the six zones into which the site was divided (Fig. 5.15). The identification of the continental malacofauna is based on Locard (1893, 1894); Haas (1929, 1991), Bech (1990), Martínez Ortí and Robles (2003) and Ruiz et al. (2006).
Spatial distribution of malacofauna according to their habitat
The conchological study included the following primary aspects: size and description of the general shape; number of whorls in adult specimens; surface sculpture; shape of the aperture; presence and morphology of umbilicus.
To study the distribution of species within the grid established on the site, Diversity (Margalef 1978) (H) and Equitability (Beerbower and Jordan 1969) (E) indices were applied. Diversity provides information about the homogeneity of populations in the assemblage, whereas equitability establishes the relation between the diversity index and its maximum value for that number of species.
Systematic Study
Class Gastropoda
Subclass Prosobranchia
Order Archeogastropoda
Family Hydrobiidae
Mercuria confusa (Frauenfeld, 1863)
Mercuria confusa is found throughout the Mediterranean region and in some northern European countries (England, The Netherlands and Belgium) (Adam 1960). In Catalonia it is a common species, except in the mountains (Bech 1990). It lives in clean, calm, stagnant or little moving waters.
Nehoratia ateni (Boeters, 1969)
Nehoratia ateni inhabits the province of Lleida (Spain) (Bech 1990). This species lives on rocks in warm non-sulfurous moving waters.
Subclass Pulmomata
Order Basommatophora
Family Lymnaidae
Lymnaea palustri s (Müller, 1774)
Lymnaea palustris inhabits Europe, Asia, North Africa and probably North America. In Catalonia, it appears along the coast in the provinces of Girona and Barcelona, as well as in the Llobregat river basin (Bech 1990). It lives in marshes and in areas featuring stagnant water.
Lymnaea peregra (Müller, 1774)
Lymnaea peregra is found in Europe, Iraq, the Canary Islands and north-western Africa. In Catalonia it is found in the provinces of Barcelona and Girona (Bech 1990). L. peregra inhabits stagnant waters.
Lymnaea (Galba) trunculata (Müller, 1774)
Lymnaea (Galba) trunculata is a holartic species (Hudendock 1951). It is a very common species in Catalonia (Bech 1990) and lives in small masses of stagnant water, and in pits. L. trunculata is usually adhered to plants and also lives outside of water.
Order Stylommatophora
Family Succineidae
Oxyloma elegans (Risso, 1826)
Oxyloma elegans is distributed in Europe, North Africa, and western and northern Asia. In Catalonia, it appears in the Llobregat basin, as well as in the delta and in the Montmell range (Baix Penedès) (Bech 1990). Absorbent, it is typical of permanently wet places such as irrigation ditches, pools, spring and creek banks, in herbaceous vegetation, aquatic plants, stones and wet land, as well as in irrigation cultures. Its size seems to diminish with altitude.
Succinea putris (Linné, 1758)
This is a Euro-Siberian species (Kerney and Cameron 1979). In Catalonia it appears in the Llobregat river basin and in the Pyrenean tributaries of the Ebro River (Bech 1990). It lives in very wet locations, stagnant or slow waters, and on aquatic plants.
Family Cochlicopidae
Cochlicopa lubrica (Müller, 1774)
Cochlicopa lubrica is a holartic species (Kerney and Cameron 1979). It is a very common species in Catalonia (Bech 1990), living in wet, shady places. It is found under stones or within dead leaves in wet areas.
Family Zonitidae
Subfamily Gastrodontinae
Zonitoides (Zonitoides) nitidus (Müller, 1774)
This is a Holartic species (Kerney and Cameron 1979). At present, Z. nitidus inhabits the entire Iberian Peninsula and is very common in the Catalan region (Bech 1990). It lives near irrigation ditches, in gutters alongside roads, on river banks, in flooded meadows and wooded areas and within riverside vegetation. It is found under stones, within dead leaves or in decomposed vegetation.
Family Helicidae
Subfamily Helicellinae
Cernuella (Cernuella) virgata (Da Costa, 1778)
This species is found in the Mediterranean region and in western Europe (Kerney and Cameron 1979). In general, it lives in two types of habitat, not usually mountainous. It is commonly found in dry locations on the coast and in the steppes of the interior of Catalonia (Bech 1990), generally adhered to plants.
Helicella bolenensis (Locard, 1882)
The distribution and habitat of this species is the western Mediterranean zone (Kerney and Cameron 1979). They live in dry, sandy areas and during dry periods they affix themselves to rocks, whereas in rainy periods they live on grass or in shrubs.
Helicella cistorum (Morelet, 1845)
This species can be found throughout the Iberian Peninsula (Ruiz et al. 2006). It is associated with bush areas (rockroses, heathers, etc.) with or without leafy trees (oaks) and is often found under stones.
Xerotricha aff. apicina (Lamarck, 1822)
This species is distributed in the Mediterranean area, mainly in areas close to the coastline under stones and dead leaves.
Xerotricha conspurcata (Draparnaud, 1801)
This species is distributed along the western Mediterranean (Bech 1990). It is a very widespread species and inhabits a large variety of habitats.
Xerotricha huidobroi (Azpeitia, 1925)
X. huidobroi is distributed along the Spanish Levante (Almeria, Alicante, Granada and Murcia) and Catalonia (Bech 1990). It lives preferably in dry and steppe zones.
Xeroplexa barcinensis (Bourguignat, 1868)
This very polymorphic species lives on the Catalan coast, and in Valencia and France (Bech 1990). It inhabits steppic areas near paths. The largest specimens live in low areas, and their size diminishes with altitude. The smallest specimens live at altitudes of 1000–1200 m (Ruiz et al. 2006).
Xeroplexa montserratensis (Hidalgo, 1870)
X. montserratensis is found in the province of Barcelona (Bech 1990) in dry, calcareous areas with poor vegetation.
Xeroplexa murcica (Guirao, 1859)
X. murcica inhabits the Mediterranean area (Bech 1990) in Mediterranean bush regions with pastures and a limestone substratum. It can be found under stones and in vegetation.
Candidula camporroblensis Siro de Fez, 1944
This species lives in the Spanish Levante (Altimira 1959). In Catalonia, it is located in the Baix Camp and Tarragonés areas (Tarragona) (Bech 1990). C. camporroblensis inhabits arid and dry zones adhering to plants or on the soil among grasses.
Monacha sp. Fitzinger, 1833
This genus is found throughout the Mediterranean area (Kerney and Cameron, 1979). In Catalonia it has been recorded in the Baix Camp and Alt Camp regions (Bech 1990). The Monacha species inhabits submountainous regions, and is also found in wetter zones. It is commonly found in grass.
Fructicum sp. Haas, 1929
The genus Fructicum inhabits the Mediterranean area (Kerney and Cameron 1979). It is found in wet areas.
Subfamily Helicinae
Cepaea nemoralis (Linné, 1758)
This western European species (Kerney and Cameron, 1979) is very common. It lives in the cold and wet areas of the Pyrenees, as well as in the dry and warm lands of Tarragona (Bech 1990), especially near water courses.
Theba pisana (Müller, 1774)
T. pisana inhabits the entire Mediterranean area (Kerney and Cameron 1979). It lives in warm, dry, arid areas, especially with a marine influence, and adheres itself to plants.
Results
All the identified remains correspond to terrestrial and aquatic continental gastropods. We have identified 28 different taxa: 22 terrestrial and six aquatic species (Table 5.7).
Although the skeletal preservation in the specimens is very good, their fragility has caused many of them to break. In some cases, this taphonomic characteristic impedes identification at the species level. In our case, up to 200 specimens from the total assemblage have been identified only to the genus level (eight genera). The largest number of specimens was concentrated in zone 3 at the site, and included both terrestrial and aquatic gastropods.
The most abundant taxa are Cepaea nemoralis and Cernuella virgata, both terrestrial and characteristic of dry environments, although Cepaea nemoralis can also inhabit cold, wet regions. The next most abundant taxon is Helicella bolenensis, which lives in sandy and dry areas. The path distribution of these three taxa in sublevel Ja is very similar, with a concentration in zone 3, zone 4 and zone 6 (Fig. 5.15). The remaining species are dispersed in the same areas of the site. The distribution of the aquatic species is concentrated in zone 3, with some single specimens in zone 4 and one in zone 6. At present, all these taxa inhabit stagnant waters with very little current. The most abundant aquatic species is Lymnaea peregra which is distributed in two of the three areas.
The diversity analyses show quite stable environments. The aquatic malacofauna was found to be highly diverse (H = 1.76; E = 0.91) (Table 5.8).
Discussion
Gastropods at the site can be classified into three groups according to habitat: wet terrestrial, dry terrestrial, and aquatic. 97% of the specimens are terrestrial malacofauna, and only the remaining 2.3% are aquatic. The analysis of the bulk of terrestrial gastropods indicates that most of the species are associated with dry environments (97.2%), whereas only 2.75% were found to be associated with wet environments.
The diversity and equitability indices for the aquatic malacofauna suggest a very stable environment, although the low number of specimens in the sample warrants a cautious interpretation. When these results are compared to those previously obtained for level I (Carbonell et al. 2002), various remarkable differences become apparent (Table 5.8). The percentages of aquatic gastropods in level I are higher (71%) than in sublevel Ja, as is the total number of specimens. The relationship between species that prefer wet environments (10.6%) and those that tend to live in dry environments (89.4%) shows an increase in wet-condition species in level I compared to sublevel Ja, although the percentage of dry-environment species is still significantly higher. The diversity analyses applied to the terrestrial mollusks found in level I found a more unstable environment than that found in sublevel Ja. The same occurs with the aquatic gastropod values in level I, which have lower diversity and equitability indices, which would indicate a more unstable environment than that in sublevel Ja.
These data suggest that the malacofauna in sublevel Ja basically corresponds to a dry and stable terrestrial environment, whereas the malacofauna in level I corresponds to a wetter, more unstable environment. Moreover, whereas aquatic malacofauna is dominant in level I, it is clearly less common in sublevel Ja.
Discussion
The results of these multidisciplinary studies paint a picture of a diverse regional landscape in which open woodlands exists side by side with clearings (Table 5.9). Forests were not very dense (48.3% AP in pollen analysis, 50% in small vertebrates), and were probably rather monospecific (99–100% pine in charcoal analyses).
We also know that the open areas had different features depending on their geological subtract, soil type, sun exposure, geomorphology, the dominant direction of wind, and other factors including seasonal variations which affected their appearance. Therefore, we suggest that the area may have functioned as a prairie (13.2% in pollen analysis, 23% in small vertebrates) during the rainy season and as Mediterranean steppe (29.2% in pollen analysis, 5% in small vertebrates) in drier seasons.
In addition, there were wet local areas above the cliff related to the travertine formations of the Capelló and at the Anoia River below the slope of the shelter. Pollen analyses yielded 18.4% hygrophilous taxa and 5.6% riverside trees; small vertebrates show 22% wet prairies; and malacology analyses indicate 28.4% wet-area and 4.7% aquatic taxa.
As for the climate during the period studied, we interpret a cold environment, moderated by the coolness and the proximity of the Mediterranean Sea. This interstadial environment permits the presence of refugia areas of Mediterranean taxa (7.5% Mediterranean taxa in palynology, 2.7% in malacology). The marked seasonality of the Mediterranean climate and the physiographical heterogeneity would have permitted the duality between dry and mesic taxa. Furthermore, the presence of riverside and aquatic taxa points to permanent local water sources throughout the year, whereas the malacological record shows a primarily dry environment inside the shelter, making its conditions for habitability quite favorable.
In short, the environments surrounding the Abric Romaní shelter during the occupation of level J were quite diverse. There were local hygro-hydrophilous components related to water (pools, marshes and rivers), which were responsible for the travertine formations and, on a regional scale, open pine forests, as well as meso-thermophilous taxa forming small forests sharing the territory with scrubland and open areas (plains and cliffs), prairies or Mediterranean steppes.
The study of the environment and landscapes that surrounded the Abric Romaní site during the occupations of sublevels Ja and Jb suggests that biotic resources were available and abundant. Data provided by anthropic and natural records verify that Neanderthals selected their resources from a wide range of possibilities and different biotopes (Finlayson and Carrión 2007; Stringer et al. 2008; Allué et al. 2012; Rosell et al. 2012; Vaquero et al. 2012). Hunting, collecting wood for fuel, and finding wood for manufacturing artifacts depend on specific strategies which include technology, energy waste, organization, and occupation length. Furthermore, biotic resources describe a similar pattern throughout the sequence, which suggests an ongoing behavior. Such behavior is very closely related to the surrounding landscape and the climatic conditions which ensure the supply of essential raw materials.
Plant resources were distributed over a large area, among which the most abundant records are related to wood. According to pollen and charcoal records, woody plants were dominant on nearby slopes and the riverbanks. Pine groves must have covered a vast area near the shelter due to the ubiquitous character of Scots Pine. The abundance of cf. Juniperus in the pollen record suggests that it too was important, and we think these were probably small common junipers. Other taxa present in the pollen record might have grown in remoter and more protected areas, suggesting the presence of refugia. The woody vegetation along the river was well established, although the record includes only a single fragment of willow in the charcoal assemblage.
The large mammals hunted by Neanderthals were distributed throughout a large diversity of biotopes. These biotopes were identified through dental wear analyses which indicate the immediate dietary adaptation of the populations and thus the habitat in which the game was preferentially hunted. Horses and aurochs fed mainly on highly abrasive items such as grass, so we can assume that they inhabited and were hunted in open landscapes, probably on the plain below the shelter. On the other hand, the microwear signs found on the dentition of the red deer are characteristic of a diet composed of low abrasive items, such as ligneous plants, indicating that red deer were present in forested habitats. Finally, the chamois probably inhabited and was hunted on or around rocky and abrupt areas which are abundant near the site.
These results indicate that all the biotopes around the shelter were known and exploited by Neanderthal groups during their hunting activities. The woody environment was preferably exploited for wood supply whereas a larger area was explored for hunting or for the exploitation of lithic raw materials.
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Acknowledgements
We would like to stress our gratitude to the students, researchers and others that have contributed to the field work at the Abric Romaní during the past years. This research has been funded by the Spanish Ministerio de Ciencia e Innovación (MICINN) with the projects HAR2008-01984/HIST and HAR2010-19957/HIST and Generalitat de Catalunya SGR813. J.M.L.-G. research has been supported by a postdoctoral grant from Juan de la Cierva Subprogram (JCI-2009-04026), with the financial sponsorship of the Spanish Ministry of Science and Innovation. M.L.B research has been supported by a predoctoral grant from UAU-CSIC. We thank the anonymous reviewers whose comments helped to improve our manuscript.
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Allué, E. et al. (2012). Neanderthal Landscapes and Their Home Environment: Flora and Fauna Records from Level J. In: Carbonell i Roura, E. (eds) High Resolution Archaeology and Neanderthal Behavior. Vertebrate Paleobiology and Paleoanthropology. Springer, Dordrecht. https://doi.org/10.1007/978-94-007-3922-2_5
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