Lagomorph exploitation during the Upper Palaeolithic in the Northern Iberian Peninsula. New evidence from Coímbre Cave (Asturias, Spain)

Lagomorph exploitation during the Upper Palaeolithic in the Northern Iberian Peninsula. New evidence from Coímbre Cave (Asturias, Spain)

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Accepted Manuscript Lagomorph exploitation during the Upper Palaeolithic in the Northern Iberian Peninsula. New evidence from Coímbre Cave (Asturias, Spain) J. Yravedra, D. Herranz, C. Sesé, P. López-Cisneros, G.J. Linares-Matás, M. PernasHernández, A. Arrizabalaga, J.F. Jordá-aPardo, D. Álvarez-Alonso PII:

S1040-6182(17)31622-1

DOI:

10.1016/j.quaint.2018.06.016

Reference:

JQI 7476

To appear in:

Quaternary International

Received Date: 27 December 2017 Revised Date:

27 May 2018

Accepted Date: 17 June 2018

Please cite this article as: Yravedra, J., Herranz, D., Sesé, C., López-Cisneros, P., Linares-Matás, G.J., Pernas-Hernández, M., Arrizabalaga, A., Jordá-aPardo, J.F., Álvarez-Alonso, D., Lagomorph exploitation during the Upper Palaeolithic in the Northern Iberian Peninsula. New evidence from Coímbre Cave (Asturias, Spain), Quaternary International (2018), doi: 10.1016/j.quaint.2018.06.016. This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain.

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Lagomorph exploitation during the Upper Palaeolithic in the Northern Iberian Peninsula. New evidence from Coímbre Cave (Asturias, Spain) Yravedra, J.1; Herranz, D.1; Sesé, C.2; López-Cisneros, P.3; Linares-Matás, G. J.4; Pernas-

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Hernández, M1; Arrizabalaga, A.5; Jordá-Pardo, J. F.3; Álvarez-Alonso, D. 3

1. Department of Prehistory, Complutense University, Prof. Aranguren s/n, 28040 Madrid, Spain.

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2. Spanish National Museum of Natural Sciences, CSIC, Dpt. of Paleobiology, C/ José Gutiérrez Abascal 2, 28006 Madrid, Spain, c/e. 3. Department of Prehistory, Faculty of History and Geography. UNED. C/ Senda del Rey SN 28040 Madrid, Spain.

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4. St. Hugh’s College, University of Oxford. St. Margaret’s Road, OX2 6LE, Oxford, United Kingdom. 5. Department of Geography, History and Art History, University of the Basque Country, UPV, EHU.

Abstract

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Traditionally, the Iberian Peninsula has been considered to be a “land of rabbits”, a notion reinforced through the frequent appearance of these animals throughout the Palaeolithic on Mediterranean sites. However, the Cantabrian coast has shown a different pattern, with rabbits being scarce or exceptional at most Northern peninsular sites, with only a few evidences of

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exploitation. Nevertheless, lagomorphs represent around 10% of the bone assemblage at the Upper Magdalenian levels of Coímbre Cave (Peñamellera Alta, Asturias). In this paper, we conduct a taphonomical analysis of the rabbit assemblage from Coímbre Cave. We note that

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bones have been exposed to several taphonomical processes, including carnivore tooth marks, chemical alterations on teeth caused by raptor digestion, and the presence of cut-marks on some bone surfaces. Therefore, we argue that the rabbit assemblage at Coímbre is the result of a complex taphonomical history, with evidence for both anthropogenic activity and the actions of other biological agents. This new evidence retrieved from Coímbre Cave further highlights its exceptionality within other Cantabrian sites. Keywords: Magdalenian, Lagomorphs, Taphonomy, Cut-Marks, Upper Palaeolithic, Coímbre

ACCEPTED MANUSCRIPT 1. Introduction The etymological origin of terms such as ‘Iberia’ and ‘Spain’ is related to the Phoenicians, who designated this region, the Iberian Peninsula, as “land of rabbits” (Linch et al, 2007). Indeed, since Palaeolithic times, some areas within the Iberian Peninsula contained plenty of these animals. In the Middle Pleistocene, some sites present abundant rabbit remains,

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such as Áridos, with more than 6,000 specimens, in addition to other assemblages from the Manzanares Valley (López 1980; Sesé et al., 2011), or Bolomor cave, with evidence of early human exploitation of these resources (Sanchis and Fernández Peris, 2008; Blasco and Fernández-Peris, 2012; Sanchis, 2012). Nevertheless, it is not until the Upper Palaeolithic when the abundance of these animal becomes overwhelming, with several sites showing very high

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percentages of the total faunal assemblage; rabbits reach over 90% NISP at many sites located in the areas of Valencia and Alicante (Villaverde and Martínez Valle, 1992; Villaverde et al.,

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1998, 2017; Olaria, 1999), as well as Catalonia (Ibañez and Saladie, 2004; Vaquero, 2005; Rufà et al, 2017), Andalusia (Aura et al., 2002; Yravedra, 2008), Portugal (Hocket and Haws, 2002; Bicho et al., 2006) and inner territories of Iberia, such as Estebanvela (Yravedra and Andrés, 2013; Yravedra et al., 2017).

In contrast, when we explore the faunal record of Cantabrian sites, the absence of rabbits from lists of identified species is noteworthy. Even though this area has yielded an

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enormous quantity of faunal remains, the assemblages dating from the Middle Palaeolithic and the Upper Palaeolithic comprise mainly herbivores, with a significant scarcity of rabbits throughout northern Iberian sites, from Galicia to the Pyrenees (Altuna, 1972; Straus, 1992; Yravedra, 2001; Sesé, 2005).

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In this paper, we provide evidence of an important rabbit accumulation discovered during recent excavations at Coímbre cave. This data is in stark contrast with other Cantabrian

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sites, being so far a unique exception. In order to understand this pattern, we have carried out a zooarchaeological and taphonomical analysis of the lagomorphs found at Coímbre Cave. Our evidence prompts a reinterpretation of the nature of the lack of leporid representation at Cantabrian sites, and sheds further light into the subsistence strategies of late Upper Palaeolithic populations of this region.

2. Coímbre cave: location and the faunal record Coímbre cave is located in Peñamellera Alta (Asturias, Northern Spain) on the southern margin of the Sierra del Cuera mountains; near the Picos de Europa mountain range (-UTM: Zone 30 T, X. 363.165, Y. 4.798.482-). The site is 145m above sea level and 42m above the left

ACCEPTED MANUSCRIPT meander of the Besnes River, a tributary of the Cares River (Figure 1). The site is surrounded by steep hill slopes but shallow valleys do exist nearby. Open habitats are found only 7-8 km away from the site, by the confluence of the Cares and Deva rivers, an area with several Upper Palaeolithic archaeological sites (Figure 1). Figure 1

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The cave of Coímbre has been known for its prehistoric art since its discovery in 1971 (Moure and Gil 1972). The first archaeological surveys were carried out between 2008 and 2012 (Álvarez-Alonso et al. 2009, 2011, 2013, 2016). The cave presents four different zones: A, B, C, and D. Coímbre Zones A and B show two different Palaeolithic occupations, but here we

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present only the evidence from Zone B.

Zone B comprises a total surface of 100 m2, of which 4 m2 have been excavated

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(Álvarez-Alonso et al. 2013; 2016). We have identified Upper Magdalenian (Level Co.B.1 – including the divisions: 1a, 1b, 1c1, 1c2, and 1c3-), Middle Magdalenian (Level Co.B.2), Lower Magdalenian (Levels Co.B.4 and Co.B.5.1), and Gravettian (Level Co.B.6) occupations levels (Figure 2). Both Magdalenian and Gravettian layers show associated faunal and lithic associations. The Gravettian layer is dominated by quartzite as raw material, but flint plays still a significant role in the assemblage. Bone tools and personal ornaments, mostly made of shells

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and red deer teeth, have also been identified (Álvarez-Alonso et al. 2011, 2013). Coímbre cave is dated to the final part of the Last Glacial period, corresponding to the end of MIS 3 and MIS 2—the Last Glacial Maximum and the Late Glacial Period. The absence of pollen and phytoliths in the deposits has prevented a palynological reconstruction of the site,

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but the charcoal record (Fabaceae, Cytisus, Ulex, Rhamus, Arbutus, Juniperus and Sorbus) as well as the avian and micromammal fossil remains (Arvicola amphibius; Microtus gr. arvalisagrestis; Microtus lusitanicus; Chionomys nivalis, Alectoris/perdix sp; Pyrrhocorax

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pyrrhocorax; Pyrrhocorax graculus and Corvidae indet) infer an open and cold environment (Elorza, 2017; Uzquiano, 2017). Figure 2

The first Magdalenian occupations at Coímbre Cave date between 20,730 and 20,270

cal BP. The start of the Magdalenian sequence (levels Co.B.5.1 and Co. B.4), were deposited during a short warm phase of the Last Glacial Maximum. After these levels, there is a hiatus of 1500 years, represented by the barren level Co.B.3; and Co.B.2 (Middle Magdalenian) formed during Heinrich H1(GS 2a), the coldest moment in the whole sequence. Co. B.4-Co.B.2 show similar paleoenvironmental features to Co.B.6, but Co.B.2 presents colder conditions, as characterised by species such as Juniperus, Artemisia and Chenopodiaceae in the pollen record,

ACCEPTED MANUSCRIPT and Mergus merganser in the avian fossil record. Co.B.1 (Upper Magdalenian) spans the last cold phase of GS 2a (Oldest Dryas) in the Late Glacial Maximum, and the temperate GI 1e (Bølling), marking the start of the Late Glacial Interstadial (GI 1), when temperatures rose outstandingly. This represents the warmest period in the entire sequence at Coímbre Cave (Álvarez-Alonso et al., 2016).

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Following Yravedra et al (2017), the faunal record is characterised by two different patterns. During the Gravettian, ungulates such as aurochs, followed by iberian ibex and red deer, played a major role, whereas iberian ibex predominates throughout the Magdalenian period, followed by red deer in the NISP (Figure 3) and MNI (Figure 4). In relation to taxonomic profiles, we highlight that red deer reaches 10% of the NISP and 21% of the MNI in

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the Magdalenian sequence, a rather unique phenomenon among Palaeolithic sites of the Cantabrian region, as this species is normally represented by only a few remains. In relation to

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the taphonomical analysis of the macrovertebrates (mammals), we highlight the anthropic origin of the faunal assemblage, given the fact that the fauna was selected, transported and consumed by humans (Yravedra et al., 2017). Even in the Gravettian level, a possible exploitation of bones as fuel was noted, according to Yravedra et al. (2016). The evidence for carnivory or the intervention of other biological agents in the formation of the macrovertebrate assemblage is restricted to limited scavenging (Yravedra et al., 2017). We now focus our

site. Figure 3 and 4

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attention on the archaeozoological and taphonomical analysis of the lagomorphs found at the

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3. Sample and Methods

For this study, we analysed a sample of 574 postcranial remains of rabbits according to Callou (1997), corresponding to at least 48 individuals on the basis of the MNI of this

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assemblage, as well as 168 cranial rabbit remains, according to Sesé (2017), following the methodology outlined by Callou (1997). However, we shall focus our analysis on the remains and individuals of Level 1, which contains 561 remains and a MNI of 44 individuals. The sample of the levels 2, 4, and 5 is not very representative and does not allow drawing a definitive conclusion.

When we compare this sample with the large lagomorph assemblages of the Mediterranean region, the rabbit accumulation at Coímbre cave could be considered minor. However, this sample involves many more remains than the number of rabbit bones found at all other Palaeolithic sites in Northern Spain together, on the basis of both NISP and MNI. This particular condition makes this concentration of remarkable significance, and worth of further

ACCEPTED MANUSCRIPT study. We have used the number of identified specimens (NISP) as described by Lyman (1994), and the minimum number of individuals (MNI) based on Brain’s model (1969) to quantify lagomorph remains. The skeletal profiles for limb bones take into account both shafts and epyphyses. In the analysis of the patterns of mortality profiles, we have not divided the sample into cohorts, since nearly all remains belong to adult individuals.

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For the analysis of the lagomorphs and the identification of the species we have taken into account the P3, because it is considered to be most diagnostic element according to López Martínez (1989), as well as all dental elements, mandibles and maxilla, following Chaline et al (1974). The lagomorph fossil remains analysed in this paper were collected in the field, by wet sieving all the soil retrieved, using 0.2 mm mesh sieves. The quantities of processed soil

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retrieved are approximately: Level 1 = 1,6 m3; Level 2 = 0,2 m3; Level 4 = 0,23 m3, and Level 6 = 0,24 m3

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For the systematic taphonomical study of bone surface modifications, we used 20X pocket-hand lenses and 40-60X binocular lenses. In the identification of cut-marks and carnivore tooth marks, we recorded the diagnostic criteria and features as defined by Bunn (1982) and Blumenschine (1988, 1995), respectively, and Landt (2007) with regards to the difference between human and carnivore tooth mark. Bone surface modifications also include the comparative assessment of epiphyseal and shaft areas (Blumenschine, 1988, 1995). Tooth

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mark and cut-mark frequencies have been documented for the whole sample, but we base our estimated percentages only on the basis of bones with well-preserved surfaces. Burning indexes were analysis in the identification of charred, burnt, and calcined remains. Among the traces left by mammalian carnivore activities, we noted furrowing and alterations due to digestion

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processes. From these processes, and following Andrews (1990), digested bones were studied within 4 different categories-stages: 1) Digested bones show a slightly polished bone surface affecting to <25% of the entire surface; 2) Digestion modifications affects up to a surface of

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75%, and show polished, rounding, pitting holes; 3) The whole bone surface is affected by alterations of the digestion and the same modifications of the stage 2, together with fractures in some cases, and 4) Digestion has destroyed the bone surface with high-intensity levels of corrosion, complicating the identification of the bone element. The presence of peak-bird marks was noted, following the descriptive guidelines of Yravedra (2002, 2006). Lastly, in the assessment of other alterations, we have followed the guidelines proposed by Behrensmeyer (1978) for weathering and those of Parson and Brett (1996) for water abrasion. Bones were divided into different categories on the basis of its maximum length, in order to generate bone fragmentation indexes: fragments smaller than 2 cm, fragments between 2.1-3 cm, fragments between 3.1-4cm, fragments between 4.1-5 cm, and fragments larger than 5

ACCEPTED MANUSCRIPT cm. The analysis of the degree of circumference follows Bunn’s (1982) 3 categories: O, >75% of circumference of large bones; C, between 25 and 75% of circumference and I, with less than 25%. These three categories allow us to estimate the maximum length of a fragment bone in relation to a complete bone specimen.

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4. Results

Taxonomical identification shows that all lagomorphs found at Coímbre Cave belong to the species Oryctolagus cuniculus. The mortality profiles show that the entire assemblage comprises adult specimens, with the exception of a few sub-adult teeth and postcranial bones.

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Rabbit features prominently in the faunal assemblage of Level 1, being the second best represented macro-vertebrate species in terms of their MNI (Figs. 2-3); however, its proportion

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decreases from Level 2 onwards, where they only show a more limited presence, with low MNI counts (Table 1). Table 1 shows that this pattern is also mirrored when we consider rabbit numbers in relation to macrofaunal remains.

Table 1.

When looking at rabbit skeletal profiles from Level 1, we note that all anatomical sections are represented, with a relative abundance of axial elements and lower appendicular

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elements (Figure 5). In terms of the proportion of long limb bones, there are no significant different between anterior or posterior appendicular elements. Even though posterior bones are slightly more abundant, the numbers of pelvises in relation to scapulae, femurs in relation to

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humerii, and tibiae in relation to radii are more or less equivalent (Figure 5). Figure 5.

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The pattern observed in the aforementioned skeletal profiles are in agreement with the analysis of the lagomorph bone surface alterations (Table 2), as well as the taphonomical analysis of the microfauna (Sesé, 2017). 99% of the bones present good preservation surfaces, with low incidence of alterations produced by processes such as weathering, trampling, or water abrasion. This means that the bone assemblage has not experienced major spatial displacements, and the position of initial deposition of the remains ought to be in close proximity to the position of discovery. Moreover, all these data is consistent with the taphonomical study of the macrofaunal remains (Yravedra 2017; Yravedra et al., 2017). Table 2.

ACCEPTED MANUSCRIPT When considering the range of taphonomical alterations caused by other biological, non-anthropogenic agents, it can be noted that some bones show tooth marks and traces derived from digestion processes, which means that some of these bones may have been altered by carnivores and/or raptors. The presence of tooth marks on 53 rabbit remains (9,4% of the remains), does suggest the intervention of a carnivore in their accumulation, at least in part. The traces are distributed across all anatomical elements, particularly long limb bones (Figure 6).

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Furrowing is documented on less than 1% of the NISP, and tooth-marked bones with at least than 5 tooth marks represent less than 5% of the sample. Digestion-induced alterations are another indicator of the activity of mammalian carnivores or raptors; nonetheless, the corrosion index at Coímbre is relatively low, with 83% of all the altered bones showing a degree of

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corrosion of only between I and II. We should also take into account the fact that some of the assemblage could have been brought in by raptors, which may be considered responsible for the accumulation of most of the other microfaunal remains (Sesé, 2005). On the basis of the tooth

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mark evidence, we could infer the activity of mammalian carnivores on the rabbit assemblage; on the other hand, the low degree of digestion-derived alterations might be more linked with the actions of nocturnal raptors.

Table 3.

Figure 6

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We have also documented taphonomical alterations by humans at the site. For example, 5,7% of rabbit bone remains show cut-marks and 87 (15%) of the bones have some degree of thermic alterations. Cut-marks appear to be distributed across all anatomical sections, including mandibles, as well as axial and appendicular elements (Figure 7). Nonetheless, cut-marks are

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particularly located on the surfaces of long limb bones; all long limb bones within the rabbit assemblage show cut-marks. We argue that most of these cut-marks are derived from butchery

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processes, on the basis of their orientation and location on the bone, although some isolated marks on a mandible are linked to skinning (Fig. 7), and some are more readily associated with the process of disarticulation. Bones with varying degrees of thermal alterations were also recorded, mainly burnt bones, with only a few charred remains; besides, no calcined bones have been identified yet. It is worth noting that some macrofaunal remains were also affected by thermal alteration (Yravedra et al., 2017). On the basis of the range of thermal alterations experienced by the rabbit bones, we argue that there is no conclusive evidence of intentional roasting processes. Figures 7, 8

ACCEPTED MANUSCRIPT The rabbit assemblage is highly fragmented (Figure 9a), with 82% of rabbit remains being smaller than 3 cm. This high breakage rate is further illustrated through the analysis of the degree of circumference exhibited by long limb bones; no long limb bone within the sample presents more than 25% of the original bone circumference (Figure 9b). Moreover, this clear breakage pattern is also attested in the analysis of the total percentage of bone representation, with around 90% of long limb bones maintaining less than 25% of their original length. Notches

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were not observed in the rabbit bone assemblage, although some long bones such as tibiae do show bending breakage and green fractures. Only some examples of specific elements, such as humerii, femurs, ulnae or tibiae are found as complete specimens. We also note that there are certain patterns of bone breakage within the assemblage. In tibiae, diaphyses appear normally

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with a breakage on the middle, keeping one of the epiphyses, or only as cylinders (Fig. 10). The presence of disjointed epiphyses is also a frequent pattern for tibia remains at the site.

5.

Discussion and Conclusion

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Figures 9, 10

5.1. The role of the rabbit at Coímbre cave and the regional context Coímbre Cave, an inland site by the Picos de Europa mountain range, shows an Upper Pleistocene faunal assemblage characterized the predominance of auroch (Bos primigenius) as

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the main exploited species during the Gravettian and by Iberian ibex during the Magdalenian period. However, what makes Coímbre Cave particularly significant is the considerable lagomorph accumulation at Level 1. Some rabbit remains have been found at other Palaeolithic sites of the northern Iberian Peninsula, such as Bolinkoba, Lezetxiki, Amalda, Abauntz, Ekain,

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Urtiaga; Erralla, Ermitia, Zatoya; Morín, Castillo, el Horno, Rascaño, Lastrilla, Pendo, Ruso, La Garma, Tito Bustillo, la Riera; Buxu, La Lluera, Las Caldas y Balmori (Altuna, 1972; Straus, 1992; Yravedra, 2001); however, the number of remains at each of these sites is remarkably

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lower than at Coímbre.

Independently of the site formation processes responsible for the accumulation of the lagomorph assemblage at Coímbre Cave (humans, carnivores, birds, other natural processes, etc.), this important accumulation suggests that the environment surrounding Coímbre Cave was particularly conducive to the spread of this species.

Blanco (1998) y Palomo and Gisbert

(2002) suggest that rabbits prefer to live in warm environments, with less than 500mm of annual rainfall, and in locations no higher than 1,500 meters above sea level. Northern Spain is a humid-climate region, with over 1000mm of annual rainfall, as well as ubiquitous limestone formations, which hamper the establishment of burrows. Given these circumstances, the low frequencies of rabbit remains noted at Palaeolithic sites of this region are to be expected.

ACCEPTED MANUSCRIPT The surrounding landscape of Coímbre Cave is characterised by a predominance of limestone formations, and today the climate would seem too wet for the ecological niche of rabbits. In fact, the number and frequency of rabbit remains at Levels 2 and 4 of Coímbre is considerably lower than at Level 1, even after taking into account that less soil from these levels was sieved; the frequencies noted for Levels 2 and 4 resemble the patterns noted at other Palaeolithic sites within the region. New data from sites such as Arangas, with a similar chronology to Coímbre

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Level 4, also shows a similar pattern of low rabbit representation (Cueto et al., 2015).

. It is relevant to explore what conditions may have changed this ecological scenario. The paleoenvironmental study of the microfauna from Level 1 shows increasingly drier conditions, with progressive aridification (Sesé, 2017). Moreover, the existence of nearby quartzite

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formations would have allowed rabbits to establish their burrows. These two particular circumstances appear to have contributed to a regional demographic expansion of this animal

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across parts of northern Iberia towards the end of the Pleistocene, evidenced by the successful adaptation of rabbits to the Coímbre environment. Future archaozoological analysis on other nearby archaeological sites, such as Llonín cave, might shed further light on the importance of the rabbit within the Coímbre ecosystem and its surroundings. At the Holocene site of Los Canes, with humid-warm conditions, rabbit is not well represented either (Arias, 2013). Thus, we encourage future regional studies to assess whether the relative abundance of rabbit was a local phenomenon of the Coímbre ecosystem during the late Upper Pleistocene, or if this pattern

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is better seen as a wider regional phenomenon during the Pleistocene-Holocene transition.

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5.2. Who brought the rabbit to Coímbre cave? Our taphonomical analysis confirms that several agents were involved in the accumulation process of the rabbit bone assemblage at Coímbre cave. Direct indicators such as

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cut-marks and tooth marks, and other indirect evidence, such as breakage patterns or partially digested bones, suggest that mammalian carnivores, raptors and humans all played a relevant role in the accumulation and alteration of the lagomorph remains. In order to determine which agent or agents were the most significant actors in the

accumulation process, we refer to a series of reference frameworks (Table 4), compiled from the work of many scholars (Hocket, 1993; Schmitt and Juel, 1994; Schmitt, 1995; Sanchís, 2000; Yravedra, 2002, 2006; Hocket and Haws, 2002; Bournery et al., 2004; Cochard, 2004a, 2004b; Lloveras et al., 2008a, b, 2009, 2012, 2018; Rodríguez-Hidalgo et al., 2015). We review the evidence for the defining features of rabbit bone assemblage on the basis of different depositional processes.

ACCEPTED MANUSCRIPT First, bones in naturally-accumulated lagomorph assemblages tend to show a certain degree of anatomical connection, a predominance of juvenile and/or infant individuals, complete skeletal profiles, unaltered bone surfaces, and low fragmentation rates. On the other hand, at nocturnal raptor-generated assemblages, there is also a high frequency of juvenile individuals, with a predominance of appendicular and cranial remains; sometimes the forelimbs show some degree of anatomical connection (Yravedra, 2006; see Fig 2 of Lloveras et al 2009).

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Moreover, in terms of taphonomic alterations in nocturnal raptor-generated assemblages, there are moderate breakage patterns, presence of puncture marks, holes, polished bones, and moderate digestion-derived bone surface corrosion. The diurnal raptors produce a slightly different pattern to nocturnal raptors: there are no partial anatomical connections, the skeletal

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profiles are characterized by an abundance of cranial and lower limb bones such as the metacarpal, metatarsal, tarsal, carpal, etc. Furthermore, the assemblage shows a larger degree of bone fragmentation, with lower frequencies of bone modifications such as holes, pits or other

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marks on bones.

With regards to canid or felid carnivores assemblages, a similar pattern can be observed: the age profiles show a predominance of adults, with some presence of young individuals, bones do not tend to appear in anatomical connection, and appendicular remains are the better represented in skeletal part profiles, with subtle differences among canids with respect to felids: Bone assemblages accumulated by agents such as fox or coyote show a predominance of

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appendicular bones with virtually no representation of vertebrae and ribs, while the Iberian lynx or the wild cat produce bone accumulations with a predominance of cranial and lower limb bones and a moderate representation of ribs and vertebrae, usually below 20% of the NISP, according to the studies of Lloveras et al., 2008b, 2018; Rodríguez Hidalgo et al., 2015). Both

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sets of bone assemblages generated by mammalian carnivores show a high degree of fragmentation, with canid assemblages bearing more intense digestion-derived corrosion, and

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wildcats showing more corrosive activity than the Iberian lynx. With regards to the presence of tooth marks, foxes produce more tooth marks than felids. The presence of cylinders is exceptional in fox assemblages, and they have not been observed in felid assemblages. In contrast to these three potential case-scenarios mentioned above, adult individuals

feature prominently in mortality profiles when humans are the main taphonomical agent. The assemblage may show some relatively similar epiphyses to shafts percentages, with few bones in anatomical connection, other than phalanges and metacarpals. The number of fore limbs and hind limb bones is usually similar, there are some cylinders in the assemblage, and most importantly, bone surfaces may show cut-marks, a taphonomical alteration which is exclusively produced by human agents.

ACCEPTED MANUSCRIPT Table 4 Given the lack of anatomical connections, and the highly fragmented nature of the bone remains, we argue that it is very unlikely that the Coímbre rabbit assemblage was deposited without the intervention of a taphonomical agent. The absence of evidence for young individuals, peak-bird marks, or anatomical connection, together with the high fragmentation

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frequencies (figure 9) and the predominance of axial elements (figure 5) suggests that raptors were not primarily responsible for the accumulation of the rabbit assemblage at Coímbre. Similarly, the different skeletal profiles found at Coímbre in comparison to typical felid or canid bones assemblages, particularly given the large representation of axial bones (figure 5), the presence of epiphyses, and the relatively low degree of digestion-induced surface alterations

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(table 3), are factors that suggest that mammalian carnivores were not the main taphonomical agent responsible for the accumulation of the rabbit bone assemblage at Coímbre Cave, even

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though mammalian carnivores undoubtedly contributed to its formation and alteration as secondary agents, on the basis of the presence of some tooth mark on bones. Similarly, the low impact of digestion-derived corrosion on bone surfaces might indicate that noctural raptors contributed to a certain extent to the bone accumulation; as shown by Lloveras et al (2008a, b: 2009, 2012, 2018) nocturnal raptors produce less intense digestive corrosion on bones than mammalian carnivores or diurnal raptors.

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Therefore, we argue that the nature of the Coímbre assemblage aligns most closely with the patterns that we would expect from an anthropogenic assemblage (Table 4). For example, all skeletal parts are represented (Figure 5), with the presence of epiphyses in a similar proportion to shafts, some cylinders, and a similar quantity of fore and hind limb bones (Figure 5).

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Furthermore, adult individuals dominate the mortality profiles, and there are taphonomical alterations associated to human activities, such as cut-marks (Figure 7) and burnt bones, with

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high fragmentation frequencies (Figure 9), and systematic bone breakage patterns (Figure 10). In conclusion, the evidence of rabbit exploitation agrees with typical Magdalenian

subsistence strategies. The species Iberian ibex was still dominant, but there was a progressive and general exploitation of alternative resources, with the inclusion into the diet of small mammals, birds, marine resources and freshwater fish (Álvarez-Fernández, 2011). Rabbit hunting and consumption at Coímbre is thus consistent with diversification dynamics that characterise the Pleistocene-Holocene transition at the end of the Upper Palaeolithic in the Iberian Peninsula.

ACCEPTED MANUSCRIPT Acknowledgements This research was part of Project: "Paleoecología y Poblamiento en la Cuenca Media del Río Cares durante el Pleistoceno Superior: La cueva de Coímbre (Alles, Peñamellera Alta)", autorized by the Dirección General de Patrimonio Cultural de la Consejería de Cultura del Principado de Asturias with founds from the Research Group: Investigación de Alto

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Rendimiento de Prehistoria de la Universidad del País Vasco (IT-288-07) and the project HAR2008-3976/HIST del Ministerio de Educación for the season of 2008, the Sociedad de Ciencias de Aranzadi in 2009 and the Mª. Cristina Masaveu Peterson Foundation since 2010. We would like to thank to Peñamellera Alta’s Council for their help during the archaeological fieldworks, and also to the Speleological Team “l’Espertuyu Cavernícola” for its logistical

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support. We are grateful for the collaboration of M. de Andrés Herrero, A. Calvo, J. Rojo, O. Fuente, M. A. Valles, N. García, R. Obeso, P. Obeso, D. Rueda and J. Santa Eugenia and the

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different students of the UNED and University Complutense laboratories.

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ACCEPTED MANUSCRIPT Villaverde, V., Real, C., Roman, D., Albert, R.M., Badal, E., Bel, M.A., Bergadà, M.M., de Oliveira, P., Eixea, A., Esteban, I., Martínez-Alfaro, A., Martínez-Varea, C. M., Pérez-Ripoll, M. (2017), The early Upper Palaeolithic of Cova de les Cendres (Alicante, Spain), Quaternary International, in press, https://doi.org/10.1016/j.quaint.2017.11.051. Yravedra, J. 2001. Zooarqueológía de la Península Ibérica. Implicaciones Tafonómicas y

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Anthropological Sciences. DOI 10.1007/s12520-016-0317Yravedra, ,J., Álvarez-Alonso, D., Estaca, V., Lopez-Cisneros, P., Andrés-Cueto, M., Arrizabalaga, A., Jordá Pardo, J. Elorza, M., Iriarte-Chiapusso, M.J., Sesé, C., Uzquiano, P. 2017. Selection and exploitation of macro-vertebrate resources during the Upper Paleolithic in Northern Spain. New evidence from Coímbre cave (Peñamellera Alta, Asturias). Oxford Archaeology. 36 (4), 331–354.

ACCEPTED MANUSCRIPT Captions Figures and Tables Figure 1. Coímbre Cave: geographical location (Asturias), regional context and nearby Upper Palaeolithic sites. Figure 2. Chronostratigraphic sequence at Coímbre cave.

and Gravettian levels of Coímbre cave (Yravedra et al., 2017)

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Figure 3. Animal frequencies in %NISP (Number of Identified Specimens) for the Magdalenian

Figure 4. Animal frequencies in %MNI (Minimum Number of Individuals) for the Magdalenian and Gravettian levels of Coímbre cave (Yravedra et al., 2017)

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Figura 5. Skeletal profiles of postcranial elements at Coímbre Cave. Craneal elements are excluded in this figure.

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Figure 6. Anatomical distribution of tooth marks identified at Coímbre

Figure 7. Anatomical distribution of cut-marks identified at Coímbre, with an emphasis on lagomorph mandible, ribs, and tibiae

Figure 8. Cut-mark frequencies on rabbit long limb bones

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Figure 9. Bone Breakage patterns on rabbit bones from Coímbre Cave. 9a. Breakage patterns according to bone size. 9b. Long limb bone breakage patterns according to the degree of circumference. 9c. Breakage patterns according to extant length in relation to the whole bone

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Figure 10. Systematic breakage patterns on tibia Table 1. MNI y %MNI of the mammals identified at the sequence of Coímbre cave. Data from the sublevels 1a, 1b, 1c1, 1c2 and 1c3 are quantified together as “Level 1”. See Supplementary

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File for NISP.

Table 2. Taphonomical alterations on rabbit bones at Coímbre Cave Table 3. Carnivore alterations at Coímbre cave Table 4. Main characteristics of rabbit bone assemblages depending on the agent responsible for the accumulation: naturally-deposited assemblages, and accumulations generated by birds, carnivores, and humans. These models are compared to the evidence found at Coímbre cave.

ACCEPTED MANUSCRIPT Lower Magdalenian

Gravettian

Level 1

Level 2

Level 4

Level 6

15680-14230 cal BP

17160-14230 cal BP

19970-18720 cal BP 29660-28560 cal BP

%MNI

MNI

%MNI

MNI

%MNI

Erinaceus europaeus

0

0

1

3

1

1

Sorex gr. araneus-coronatus

1

1

1

1

Talpa gr. europaea-occidentalis

5

3

1

3

Galemys pyrenaicus

0

0

1

3

Arvicola amphibious

38

22

7

19

Microtus gr. arvalis-agrestis

52

30

16

44

Microtus lusitanicus

4

2

2

Microtus oeconomus

10

6

5

Chionomys nivalis

20

11

2

Apodemus gr. sylvaticus-flavicollis

4

2

Oryctolagus cuniculus

43

24

TOTAL

177

100

MNI

Bos / Bison Equus ferus

MNI

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MNI

%MNI

1

1

1

1

12

17

1

20

40

56

1

20

1

20

1

20

6

2

3

14

5

7

6

6

8

1

1

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MNI / %MNI

Middle Magdalenian

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Cal BP

Upper Magdalenian

3

2

3

1

20

36

100

72

100

5

100

%MNI

MNI

%MNI

MNI

%MNI

MNI

%MNI

6

3

1

6

2

11

4

40

6

3

1

6

1

5

1

10

17

9

1

6

2

11

1

10

73

41

9

56

9

47

3

30

4

2

1

6

22

12

2

13

3

16

2

1

1

1

3

2

2

1

1

1

Oryctolagus cuniculus

43

24

1

6

2

11

1

10

Total

180

16

100

19

100

10

100

Cervus elaphus Capra pyrenaica Capreolus capreolus Rupicapra pyrenaica

Ursus arctos Vulpes vulpes Canis lupus Meles meles

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Sus scrofa

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MNI / %MNI

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1

Table 1.

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% NISP 1

% Weathering

1

% Water alteration

1

% Sedimentary Corrosion

6

% Trampling

3

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% Bad preservation surfaces

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Level 1: NISP = 561

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Table 2.

NISP

Sample

561

Tooth Marks More than 5 tooth marks per specimen

Digestion III Digestion IV

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Furrowing

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Digestion II

Table 3.

53

9

4

2

248

44

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Digestion Digestion I

%NISP

133

54

72

29

32

13

11

4

7

1

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Presence of young individuals

Yes

Yes

Yes (only juveniles)

Anatomical connection

Yes

Sometimes, mostly the hind limbs

No

Complete skeleton

Predominantly cranial and limb bones

Predominantly Cranial and lower limb bones

All

Similar proportion

Similar proportion

Absent

Absent

Absent

Similar

(Variability among studies)

Skeletal profiles

Proportion Diaphyses to Epiphyses Diaphysiary Cylinders

No (only Adults) No (only phalanges & metapodials)

Complete skeleton

Complete skeleton

More distal epiphyses

Similar proportion

Both but more diaphyses

Absent

Yes

Some

More hind limbs

Similar proportion

Similar

Very intense

Very intense

Moderate

Very intense

No

2%

0.5%

1-10%

<1%

Cut and tooth marks

Cut and tooth marks

None

Low

Low

Low

Low

Few, mostly on the edges

68%, low intensity

98%, low intensity

99%, high intensity

99%, high intensity

No

95% < 5 tm/bone 50%, small degree

No

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Moderate

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References

More fore limbs

Low frequencies Variable proportions among studies

Predominantly adults (90%) No (only phalanges & metapodials)

No

Appendicular Cranial and bones and lower lower axial appendicular bones More hind limb bones Absence of epiphysis

Coimbre

No

Digested Bone Charred bone

but mainly adults (80%)

Humans

Very intense over 80% bones under 3cm

Bone Fragmentation Traces both carnivoreraptor alterations Number of marks per bone

No

Felid Carnivores Yes,

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More hind limbs Fore/hind limbs

Canid Carnivores Yes, but mostly adults (5080%)

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Diurnal raptor

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Nocturnal raptor

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Natural deposition

No

No

Yravedra, 2008

Hocket 1993; Hocket and Haws, 2002; Sanchis 2000; Yravedra 2002, 2006; Bourneay et al., 2004; Cochard, 2004a, b; Lloveras et al., 2009

Table 4.

No

Schmitt, 1995; Hocket and Haws, 2002; Lloveras et al 2008

Sanchis 2000; Lloveras et al., 2012; Schmitt & Juel, 1994

Lloveras et al 2008b, 2018, Rodríguez Hidalgo et al 2015

Yes

15%

Pérez Ripoll 1992, 2004, 2006

Present study

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