Introduction

The Mesolithic in the Mediterranean region of the Iberian Peninsula was a period of significant cultural and environmental changes. Temperatures and humidity were gradually increasing and sea level was rising from the end of the Pleistocene and into the Holocene (Cacho et al. 2010). Landscape changes included the spread of oak forests (Carrión 2005) that was accompanied by increased populations of forest adapted animal species like roe deer (Capreolus capreolus) and wild boar (Sus scrofa), and also red deer (Cervus elaphus) although this species could also live in other biotopes (e.g. open grasslands, meadows, river valleys). As a result, Mesolithic environments differed dramatically from those in the Upper Palaeolithic. Generally observed patterns point to logistic mobility considering the uneven and patchy distribution of the main prey consumed around inland and wetland/coastal territories, favoring the incorporation of a wide range of small species in the diet. This type of mobility seems to respond efficiently to subsistence strategies adapted to the new environmental conditions and also involved logistic movements for particular resource procurement and social practices and interactions aimed at maintaining social networks locally and regionally (Martí et al. 2009; Whallon 2006). In contrast to the late Pleistocene period, hunter-gatherers in this region shifted their subsistence practices, and new forms of economic and cultural activity emerged, including visible changes in stone tool technologies and the production of portable art (García-Puchol 2005; López-Montalvo et al. 2024). In eastern Iberia, from a general technological overview, one of the most remarkable aspects of technological shifts was the rise of a notched and denticulated lithic technocomplex at the end of the XI millennium cal BP (Notches and denticulates Mesolithic -NDM-), followed by another major shift linked with the emergence of the blades and trapezes technocomplex along the western Mediterranean in the middle of the IX millennium cal BP (Geometric Mesolithic -GM) (Marchand and Perrin 2017; Martí et al. 2009).

One of the key points for understanding these changes is the composition of the human diet. In order to comprehend the economic behaviour of these last hunter-gatherers, it is essential to have faunal bone assemblages and their corresponding zooarchaeological and taphonomic studies. Their results provide relevant data on hunting patterns (diversified or specialised), the exploitation of different biotopes and prey (for food and also for obtaining raw materials), as well as on cooking and meat preparation practices. Archaeological evidence of these types of activities is present throughout the Iberian Mediterranean, and has been gradually increasing. However, there are fewer sites with complete zooarchaeological studies from Mesolithic contexts than from Late Upper Palaeolithic (e.g. Magdalenian) ones. Almost twenty sites with faunal remains are currently published in the Mediterranean area of the Iberian Peninsula, although their distribution is not uniform (Fig. 1). In the northeast and southeast regions there are very few sites, but most of them are concentrated in the central area. On the other hand, not all the assemblages have provided relevant data, mainly because in many of them the number of faunal remains is very low, there is a low percentage of identified bones, or because studies are still in progress.

Fig. 1
figure 1

Location of Cocina cave and other Mesolithic sites cited in the text. Map made with QGIS 3.16 (QGIS 2022) from (SCUAM 2013) map

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Based on the analysed assemblages, the general hypothesis is that Mesolithic hunter-gatherers focused their subsistence activities mainly on the nearest biotopes, but there is also striking evidence for contact with more remote areas between inland and coastal areas, as marine species (fish and shellfish) could be found at inland sites (Aura 2019; Martí et al. 2009; Pascual Benito and García-Puchol 2015; Román et al. 2020). In Mediterranean Iberia, Mesolithic hunting was much more diversified than in the Magdalenian and Epipalaeolithic, which was largely focused on one medium-sized ungulate (either red deer or Iberian ibex (Capra pyrenaica)) and rabbits (Oryctolagus cuniculus) (e.g. Aura et al. 2002; Real 2020b; Villaverde et al. 2012). In contrast, Mesolithic hunting included a wider range of ungulate species, along with other types of resources such as carnivores, shellfish, fish and even snails and turtles (e.g. Fernández-López de Pablo et al. 2013a, 2015; Lloveras et al. 2021; Pascual Benito and García-Puchol 2015; Pérez Ripoll and García-Puchol 2017). Nevertheless, our knowledge of these subsistence patterns is still incomplete and we need new assemblages or a thorough and comprehensive study of bone assemblages from known studied sites.

One of the key sites in the Mesolithic archaeological tradition of the Western Mediterranean is Cocina cave. This is an archaeological site with a long research history and a significant site for the Mesolithic of the Iberian Peninsula. Several excavations were conducted between 1941 and 2018 that were directed by different research teams, and thus had different objectives and used varying methodologies. The most recent project (2015–18) focused on understanding the chronology and stratigraphy of the site, resulting in the documentation of two chronological phases of occupation (Cocina A and Cocina B) during the Late Mesolithic (García-Puchol et al. 2023). In addition, earlier excavations resulted in a rich assemblage of archaeological materials to analyse the technological, subsistence and cultural features of the human groups from the Late Mesolithic. Its long research trajectory, including recent excavations, make Cocina cave a prominent site in the study of the subsistence and social actions of Mesolithic groups in Eastern Iberia.

For all of these reasons, we present a complete overview of the Cocina cave faunal assemblage summarizing the zooarchaeological and taphonomic analyses of the faunal remains from the different excavation phases. We compiled the raw data from previous research conducted by M. Pérez Ripoll and analysed the bone remains from recent excavations (2015–18) to assess subsistence activities and the occupation patterns of the hunter-gatherers from Cocina cave. We present the taxonomic and body part identifications to assess diet, hunting, and transport decisions, along with skeletal modifications to characterize processing and consumption. Secondly, we describe chronological variation to assess potential changes in seasonality and site functionality. Finally, we compare the results from Cocina cave with those from other Mesolithic sites in the Mediterranean Iberia, from the Catalan coastal area to the south, to assess the degree to which hunting decisions at Cocina cave were unique or followed broader cultural patterns. We also discuss the main characteristics that define the Mesolithic subsistence activities more generally, and to what extent there is a change in landscape exploitation compared to previous phases of the Final Upper Palaeolithic (e.g. Aura et al. 2009; Morales Pérez 2015; Real 2021).

Regional setting

The Mediterranean Iberia basin is a wide and narrow area with bioclimatic and orographic variations, but a constant human occupation during the Late Pleistocene and Early Holocene (e.g. Aura et al. 2006; García-Puchol et al. 2015b; Barton et al. 2018; Fernández-López de Pablo et al. 2019; Fyfe et al. 2019). New environmental conditions in the Early Holocene are reflected in the progressive increase of temperature and humidity resulting in the expansion of the Mediterranean forest reflected by charcoal analysis (including deciduous oak and Pinacea) (Carrión 2005). Mesolithic sites are mostly located in mid-mountain zones (Fig. 1), between 1000–300 m.a.s.l., but openair sites have also been recognised in lowland areas such as Cativera (Tarragona), El Collao (also known as El Collado) (Oliva) or Benàmer (Muro de Alcoi) (Fernández-López de Pablo et al. 2015; Martí et al. 2009; Tormo Cuñat 2011; Vaquero 2006). The set of mid-mountain sites –with faunal studies– along the Mediterranean basin includes a few sites such as Cueva de Nerja (level NV-IIIc) (Málaga) in the south and Abric Agut (Anoia) and Molí del Salt (Tarragona) in the north, (Ibáñez López 2006; Morales Pérez 2015; Rufà et al. 2017), these last related to the NDM. The greatest number of sites, and the most complete sequences, have been described in the central region, encompassing eastern Mediterranean territories administratively included in the Valencian region and Lower Aragon. From this wide area, we know several assemblages that in some cases reveal both NDM (10.000 to 8500 cal BP) and GM occupations (from 8500 to 7500 cal BP). These techno-typological contexts have been described in a detailed manner including the discussion about the break between the different knapped techniques and toolkits from an expeditive knap** for removing flakes (NDM) and an elaborate regular blade production focused on obtaining geometric projectiles tools (GM). In regards to subsistence strategies, some sites in this central Mediterranean area have provided small assemblages of animal bones such Mas Gelat (Alcoi), Abric de la Falguera (Alcoi), Casa Corona (Villena) or El Lagrimal (Villena-Yecla) (Fernández-López de Pablo et al. 2015; Miret i Estruch et al. 2006; Pérez Ripoll 1991, 2006). Others, however, offer a larger assemblage and, thus, more information about the Mesolithic diet: Cingle del Mas Cremat (Portell de Morella), Cingle del Mas Nou (Ares del Maestrat), Balma de la Fontanella (Vilafranca), Benàmer (Muro d’Alcoi) Coves de Santa Maira (Castell de Castells), Tossal de la Roca (Vall d’Alcalà) and Cocina cave (Dos Aguas) (Lloveras et al. 2021; Morales Pérez 2013; Morales and Llorente 2019; Pérez Ripoll and Martínez Valle 1995; Tormo Cuñat 2011; Vicente Gabarda et al. 2009; Vizcaino León 2010).

Cocina cave (Dos Aguas, Valencia)

Cocina cave is an archaeological site in eastern Spain, located in Dos Aguas, Valencia. The site is a cave at 408 m.a.s.l. with a wide entrance. It is situated in the heart of a rugged mid-mountain landscape, but from which there is a visual dominance of the plain area close to the coast, currently 38 km away. The cave was the focus of several archaeological excavations. The first excavations were carried out in the 1940s (1941, 1942, 1943, 1945) under the direction of Luis Pericot (Pericot 1945). This work positioned the site as a key archaeological site for studying the Mesolithic period in the region. Beginning in the 1970s, Javier Fortea excavated for several seasons (1974–1981) and focused on the central area of the site using new methodologies for the time allowing for a more accurate recording of materials and stratigraphy (Fortea 1971, 1973; Fortea et al. 1987). Despite early recognition of the potential significance of the site, very few of the materials were analysed or published until O. García-Puchol (together with co-directors Joaquim Juan-Cabanilles and Sarah B. McClure) conducted new excavations and analyses of materials from all previous studies (Gallello et al. 2021; García-Puchol et al. 2014, 2015a, 2016, 2018a, 2018b; Pardo-Gordó et al. 2017, 2018; Pascual Benito and García-Puchol 2015). Excavation focused on several strategic test pits to understand the stratigraphy of the cave, while a large suite of AMS radiocarbon dates and material analyses were conducted to clarify the chronology, taphonomy, palaeoenvironmental and cultural sequence attribution throughout the cave (García-Puchol et al. 2018a; Pardo-Gordó et al. 2017). Recent multistage Bayesian modeling of new radiocarbon dates and statistical analysis based on cultural variability and spatial density record resulted in a new general correlation sequence between the different excavation areas and a new understanding of site occupation (García-Puchol et al. 2023) (Suppl. Table 1). The prehistoric sequence of the cavity mainly includes the last Mesolithic levels (GM), and posterior occupations dating to the Neolithic and Bronze Age periods (Fig. 2). The GM levels have been subdivided into phases (A and B) and, where possible, a series of subphases correlated the different-grained information resulting from varied fieldwork methodologies (García-Puchol et al. 2023). The excavation campaigns vary considerably in the extent and volume excavated, and the depth of the test pits, reaching 4 m in Pericot’s campaigns. The collection of faunal remains also varied considerably. The assemblages from the 1940s were highly selected and consist of mostly larger taxa and elements that were readily identifiable, likely as a result of hand collection during excavation. Despite much more accurate recording of materials in situ in the Fortea excavations, the faunal remains that remained in the collection post-excavation also appear to have been selected as small fragments are largely missing. In contrast, recent excavations water-sieved all dirt in 2 mm screens and all bones were collected.

Fig. 2
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Site map of the cavity modified from García-Puchol et al. (2023) with indication of archaeological fieldwork, (red = Pericot, purple = Fortea, green = 2015–18). The isolines have been spaced by 25 cm. The warm colors refer to the highest surfaces (increasing from yellow to red), and the cold colors refer to the lowest surfaces (increasing from green to blue) (maps made by Global Geomàtica, València). The Fortea’s grid has been translated to the entire cavity (squares of 1 m) although the currently named grid refers to cardinal references (N, S, E, O) from 0 XY point added to the letters and numbers (example KE2S)

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The faunal remains recovered in the old excavations (1940s and 1970s) were studied by Professor Manuel Pérez Ripoll and partially published in Fortea et al. (1987) and Pérez Ripoll and García-Puchol (2017, 2019). Beginning in 2013, new research on the previously excavated artifacts and excavations at the site itself revised the stratigraphy and chronology of the older excavations and M. Pérez Ripoll resumed the study of the archaeofauna assemblages from the excavations of both Pericot and Fortea until his untimely death in April 2020. M. Pérez Ripoll completed the analysis of the archaeofaunal remains from the old excavations but was unable to publish the final results in full. In addition, the assemblages from the 2015–18 excavations remained unstudied. This paper presents the results of his last research project in conjunction with reporting the archaeofaunal results from the most recent excavation units.

Materials and methods

Materials

We present the zooarchaeological and taphonomic analysis of faunal remains of the Mesolithic levels from different excavations conducted at Cocina cave (Table 1), as the following:

  • The faunal remains recovered in the 1941 field season came from the sector "Cata Sur" (13 layers) and the sector "Ampliación" (13 layers).

  • The materials from the 1942 field season were recovered from the subsectors “Surest” (layer 1), “Sur” (layer 1), A (layer 2), B (layers 2 to 9), BC (layer 11), C (layers 2 to 9) and D (layers 2 to 6).

  • The faunal assemblage from 1943 came from sectors A (layers 1–6), B (layers 1–7), and C-D-E (layers 1–9).

  • The faunal assemblage from 1945 came from sectors E1 (layers 1–17) and E2a (layers 1–13).

  • The fauna from the 1974–81 field seasons came from level H (living floors H1, H2, H3 and H4) of sector E. Fortea described these living floors based on several hypotheses around the place occupied by a possible heart, although recent detailed spatial analysis has concluded about the problems for confirming their consistency (Pardo-Gordó et al. 2018). Nevertheless, we consider theirs as proper archaeological layers.

  • The materials from the 2015–18 excavations came from two test pits. From test pit 3 (TP3) we analysed the stratigraphic unit 4 (UE4). From test pit 4 (TP4) we included units S45a and S45b.

Table 1 Summary of excavation campaigns, analysis, and publication history of the faunal assemblages from Cocina cave. We include all available data from M. Pérez Ripoll’s analyses along with new data generated from the 2015–18 excavation campaigns
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We analysed all assemblages recovered in the different campaigns, including the faunal remains that were not clearly assigned to subunits. This is a relevant aspect in that proceeding in this manner allows us to obtain a more complete view for the broad occupation sequence recognized at the cavity. In this sense, Pericot’s fieldwork resulted in the greatest excavation volume and the most complete sequence, although with a minor spatial and temporal resolution. García-Puchol et al. (2023) present the discussion on the integrity of the sequence and the proposal for building the smaller temporal units. Based on the high degree of selection of faunal bones in Pericot’s and Fortea’s campaigns, we decided to group the assemblages by the more general phases described for the Mesolithic (phase A and B) (Suppl. Table 1). The most detailed data comes from the 2015–18 campaigns. The new chronological framework indicates a primary occupation horizon during the Late Mesolithic (from the first centuries of the IXth millennium and including the first half of the VIIIth millennium cal BP) as documented both by a distinctive lithic tradition with trapezes (phase A) and triangles (phase B), and a series of AMS radiocarbon dates (García-Puchol et al. 2018a, 2023; Pardo-Gordó et al. 2018). The upper layers of some sectors show mixtures of material from Mesolithic episodes and from more recent Neolithic, Chalcolithic, and possibly Bronze Age occupations (García-Puchol et al. 2023; Pericot 1945). Here we classify these layers as mixed and do not include them in our discussions, although the data are presented in the supplementary materials (Suppl. Table 2; Suppl. Text 1).

Methods

As mentioned above, M. Pérez Ripoll studied the assemblages from 1941, 1942, 1943, 1945 and 1974–81. He published part of the results in various venues, including the study of the macromammals from the 1941 and 1942 excavations (Pérez Ripoll and García-Puchol 2017), the study of the leporids from the 1943, 1945, and 1974–81 excavations (Pérez Ripoll and García-Puchol 2019), and a short, general overview of the faunal remains from the 1974–81 excavations (Pardo-Gordó et al. 2018). We compiled the raw data of his database from all these studies, which included taxonomic and anatomical data (only by NISP), and the presence of modifications on bone remains (only type of mark and element affected). Age at death and seasonality data are limited to the assemblages from 1941 and 1942. In the process of compiling these data, some quantifications were updated, which explains some slight variations from his previously published data. The faunal remains from the 2015–18 excavations (Test Pit 3 -TP3- and Test Pit 4 -TP4-) were studied by C. Real and A. Sanchis using a more complete methodology outlined below. Therefore, regarding the taphonomic analysis, the discussion is based mostly on our study (2015–18 assemblage), because the modifications are described in more detail in terms of morphology, distribution, intensity, etc., although the assemblages studied by M. Pérez Ripoll were also used to support our assessments.

Taxonomic and anatomical identifications were made using the Laboratorio Gil-Mascarell (University of Valencia) and Gabinet de Fauna Quaternària Innocenci Sarrión (Museu de Prehistòria de València) reference bone collection. Indeterminate remains were classified by weight (very small, small, medium and large) and type of bone (long bones, flat bones, articular bones) (Blasco 2011; Cáceres 2002). The number of remains (NR), the number of identified specimens (NISP), the minimum number of elements (MNE) and the minimum number of individuals (MNI) (Brain 1981; Lyman 2008) were used where possible to quantify the remains. For the analysis of the Iberian ibex assemblage of 2015–18 excavations, the standardised minimal animal units (%MAU) were also used and correlated with the bone density to identify attrition processes (Binford 1984; Lyman 1994).

Age at death was established by bone epiphyseal fusion, tooth eruption and dental wear (Silver 1963). In the case of Cervus elaphus, Mariezkurrena (1983) and Azorit (2011) were followed for tooth eruption/wear and Mariezkurrena (1983) for epiphyseal fusion. For Capra pyrenaica, age was established by tooth eruption/wear (Pérez Ripoll 1988) and epiphyseal fusion (Serrano et al. 2004) and for Sus scrofa Bull and Payne (1982) was used. Regarding leporids, the age is based on epiphysis fusion (see references compiled in Sanchis 2012).

The bone surface was observed with a stereomicroscope (Nikon SMZ-10A) to identify fractures and anthropogenic and non-anthropogenic modifications. The work of Villa and Mahieu (1991) was followed for the analysis of breakage. In the case of 2015–18 materials, we also applied our morphotypes of fracture (Real et al. 2022). Fractures were classified as recent (excavation origin), fresh, dry, mixed (fresh-dry) and indeterminate. Cut marks, percussion marks and burnt bones were recorded as anthropogenic modifications (e.g. Binford 1981; Galán et al. 2009; Nicholson 1993; Pérez Ripoll 1992; Potts and Shipman 1981; Shipman and Rose 1983; Soulier and Costamagno 2017; Stiner et al. 1995; Vettese et al. 2020). Non-anthropogenic modifications followed classic criteria (e.g. Andrews 1990; Binford 1981; Domínguez-Rodrigo and Piqueras 2003; Fernández‐Jalvo and Andrews 1992; Sala et al. 2012; Yravedra 2006) and include tooth marks and digestion. Specific references were followed to identify and describe modifications on leporid bones (Hockett and Haws 2002; Lloveras and Nadal 2015; Lloveras et al. 2009; Pérez Ripoll 1993, 2004, 2005; Rosado-Méndez et al. 2016; Sanchis 2000, 2012; Sanchis et al. 2011).

Data were recorded in Excel spreadsheets and combined into a Filemaker Pro v18 database. Data analysis and visualization were conducted in Excel pivot tables. All data used in this study are presented in the Supplementary tables: Suppl. Tables 3, 4 and 5 summarize the data from Pericot’s excavations; Suppl. Table 6 from Forteas’ excavations; and, Suppl. Table 7 for 2015–18 excavations. Moreover, we provide the data from the analysis of the Pericot’s assemblage separately, 1941 plus 1942 and 1943 plus 1945 in Supplementary Text 1.

Results

Pericot Excavations

Taxonomic and anatomical composition

A total of 5274 bones and teeth were collected from all of the areas that Pericot excavated 1941–1945. Of these, 1423 came from mixed contexts, either where we were unable to distinguish between Late Mesolithic phases, or where additional evidence for more recent time periods, including the Neolithic, Chalcolithic, and even Bronze Age, were interspersed. Detailed descriptions of the data for each sector are available in the supplementary material (Suppl. Table 3, 4, 5). Here we summarize the evidence for the Mesolithic assemblages that were identifiable to early (Cocina A) and late (Cocina B) phases of the Late Mesolithic.

Secure assemblages from the early phase of the Late Mesolithic (Table 2; Fig. 3) component of the site (Cocina A; n = 1889) consisted of largely Iberian ibex (Capra pyrenaica; 71.9%), followed by red deer (Cervus elaphus; 13.9%), and rabbit (Oryctolagus cuniculus; 12.7%). Other species are represented by few remains, in many cases only one bone, but point to the diversity of the assemblage: equid (Equus sp.; 0.1%), aurochs (Bos primigenius; 0.1%), wild boar (Sus scrofa; 0.6%), fox (Vulpes vulpes; 0.4%), lynx (Lynx pardinus; 0.1%), wild cat (Felis silvestris; 0.1%), and partridge (Alectoris rufa; 0.2%).

Table 2 Number of identified specimens (NISP) of taxonomic categories and their relative proportions from Pericot’s excavations by chronological phase
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Fig. 3
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Taxonomic composition of faunal assemblage (%NISP) by excavation sector and chronological phase

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The late phase Late Mesolithic faunal assemblage (Cocina B; n = 1962) of Pericot’s excavations is largely similar (Table 2; Fig. 3), with a clear dominance of Iberian ibex (72.7%), followed by rabbit (13.1%) and red deer (10.6%). However, some slight variation is visible in the other species represented. First, wild boar (1.5%), equid (0.7%), and aurochs (0.5%) are more common than in the earlier phase. In addition, other species are present that were not found in the earlier phase: roe deer (Capreolus capreolus; 0.3%), canid (0.1%), and badger (Meles meles; 0.1%). Other animals were found in roughly the same proportion as earlier in time: fox (0.3%), lynx (0.1%), wild cat (0.1%), and partridge (0.1%).

The anatomical profiles of the three main species (Iberian ibex, red deer and rabbit) are presented by phase and according to NISP as the MNE is not available (Table 3). In both phases of the Late Mesolithic represented in the assemblage, there is a predominance of isolated teeth and mandibles of Iberian ibex, followed by the remains of the axial skeleton (Fig. 4). The bones of the limbs are less represented, and even the greater density elements such as astragali and phalanges are limited in number in comparison to skull and axial elements. This pattern is slightly different with red deer (Fig. 5), largely due to fewer isolated teeth; in this case, the Cocina A phase assemblage consists of 41.2% limb elements, while the Cocina B phase has a greater proportion of skull and axial fragments (68.1%). Regarding rabbits (Fig. 6), mandibles and tibiae dominate, while the rest of the skeleton is under-represented.

Table 3 Anatomical composition of the main species (Iberian ibex, red deer and rabbit) by phase from Pericot’s excavations
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Fig. 4
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Anatomical composition of the Iberian ibex remains (%NISP) by excavation sector and chronological phase

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Fig. 5
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Anatomical composition of the red deer remains (%NISP) by excavation sector and chronological phase

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Fig. 6
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Anatomical composition of the rabbit remains (%NISP) by excavation sector and chronological phase

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Furthermore, age at death data for Iberian ibex is available for part of the 1942 assemblage, specifically layer 1 of the SE sector. Age at death of Iberian ibex helps characterize hunting patterns by the human groups from Cocina cave during the Late Mesolithic. Unfortunately, and unbeknownst to Pérez Ripoll at the time of analysis, this particular level is considered a mixed context with both Mesolithic and some evidence for more recent time periods. Although most Iberian Ibex remains could be Mesolithic in date, we cannot confirm this assumption. With caution, we report the findings here. This analysis is based on a set of 5 maxillae, 23 mandibles and 319 isolated teeth, corresponding to 45 individuals (see detailed methodology in Pérez Ripoll and García-Puchol 2017). The results of the study indicate that young and subadult Iberian ibexes (phase II: 13 to 25 months; and mainly phase III: 25 to 60 months) are the predominant age groups in the assemblage and clearly outnumber adults and senile individuals. Given the typical birth season of Iberian ibex during late spring and early summer (Alados and Escos 1988), animals of this age in the assemblage suggest that hunting may have been conducted throughout the year.

Fragmentation and modifications

Modifications and patterns of fragmentation (n = 1526) found on various species in the Late Mesolithic assemblages are related mainly to human processing and consumption (Table 4; Fig. 7). These are cut marks (n = 171 Cocina A; n = 105 Cocina B), fractures for marrow extraction (n = 407 Cocina A; n = 338 Cocina B) and human bite marks in the case of rabbit (n = 1 Cocina A), and burnt bones (n = 161 Cocina A; n = 328 Cocina B) (Table 4). There are other tooth marks (n = 2 Cocina A; n = 13 Cocina B), but without a clear origin.

Table 4 Modifications present (NISP) on the Pericot faunal assemblage by type and species
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Fig. 7
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Proportions (%NISP) of modification type found on all taxa by excavation sector and chronological phase. Numbers inside circles designate the total number of bone fragments (NISP) with recorded modifications

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All these modifications mainly affect the principal species (Iberian ibex, red deer, rabbit, and wild boar) (Fig. 8, 9, 10), although they were identified in smaller quantities on bones of equids, aurochs, roe deer and badgers. There is some variation in types of modifications depending on the phase of the Late Mesolithic in question. During the earlier phase (Cocina A), 39.2% of the assemblage had some sort of modification, and fractures were the most commonly identified mark (55%), followed by cut marks (23.1%) and evidence of burning (21.7%). On the other hand, 40.1% of the Cocina B assemblage had some sort of modification. While fractures were also the most numerous of these (43.1%), evidence for burning was much greater (41.8%) and only 13.4% of the assemblage had evidence of cut marks.

Fig. 8
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Examples of fresh fractures on Iberian ibex bones (a, c-g, i) and anthropogenic modifications: cut marks (a, b, c, d, h), percussion mark (e), and burnt bone (b). Bones: a. mandible, b. astragalus, c. humerus, d. scapula, e. tibia, f. humerus, g. vertebra, h-i. skull

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Fig. 9
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Examples of fresh fractures on red deer bones (a-e) and anthropogenic modifications: cut marks (f, g), percussion marks (b, c, h), and burnt bone (a, d). Bones: a. phalanx, b. coxal, c. ulna, d. humerus, e. tibia/astragalus, f. coxal, g. rib, h. metatarsal

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Fig. 10
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Examples of fresh fractures on rabbit bones (a-g) and anthropogenic modifications: tooth marks (c), burnt bones (e, f, g), and cut marks (f, g). Bones: a. humerus, radius, ulna, b. femur, c. coxal, d-g. tibia

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When we examine the available data by taxon, we see a difference between Iberian ibex, red deer, and rabbits. In the earlier Late Mesolithic phase, modifications of Iberian ibex remains included many fractures (53.9%), followed by evidence of burning (23%) and cut marks (22.8%). Burning was much more common in the later phase of the Late Mesolithic levels (47.3%), while cut marks significantly declined (11.2%) and fractures also declined somewhat (39.4%). This shift is not visible in red deer, where there are only a few changes in documentation rates between Cocina A and B assemblages, with fractures dominating both periods (49.6% Cocina A; 45.1% Cocina B), followed by cut marks (25.2% Cocina A; 24.5% Cocina B), and evidence of burning (24.4% Cocina A; 28.4% Cocina B). Finally, modifications on rabbit bone are largely the same between the two chronological phases with the rates of fractures (61.7% Cocina A; 58.9% Cocina B) and cut marks (22.7% Cocina A; 12.8% Cocina B) remaining similar, but with a clear increase of burning (15.6% Cocina A; 27% Cocina B) (Fig. 10).

Fresh fracture evidence by percussion is numerous and is present on long bones, skulls, axial remains and phalanges (Fig. 9). Moreover, there is some evidence of fresh fracture by bite and tooth marks on rabbit bones that seem to have an anthropic origin. However, the tooth marks present on ungulate bones, although M. Pérez Ripoll identified some of them (mainly those on ribs) as anthropic (Pérez Ripoll and García Puchol 2017), we do not have enough evidence to support this affirmation (Table 4).

Fractures also show variable values depending on species. Among Iberian ibex long bones, fragments of diaphysis are the best represented part, but also distal parts of humerus, radius, tibia and metapodia, and proximal part of femur and metatarsal. In the case of red deer, the pattern is center around diaphysis too, and distal of metapodia, and both proximal and distal of radius.

Long bones of rabbits are also fractured by human bites (Fig. 10 a, b, d), and, some mandibles and above all the coxal of rabbits show signs of human teeth chewing during the removal of the meat adhering to the bones. These are small, fine, parallel grooves and other more pronounced grooves ending in small indentations in the case of the ilium (Fig. 10 c). Non-anthropic modifications, in the form of tooth marks (pits, furrowing), are present on ungulate bones from Cocina A and Cocina B (Fig. 11 b, d).

Fig. 11
figure 11

Examples of carnivore modifications on red deer bones: scores on an astragalus (a) and humerus (b); and Iberian ibex bones: pits on a radius (c) and puncture on a hyoid (d)

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Assemblages from 1974 to 1981

Taxonomic and anatomical composition

The Mesolithic faunal remains (n = 1173) from this portion of the site were excavated in Sector E and data are available from four contexts, originally interpreted by Fortea as living floors: H1 (n = 325); H2 (n = 342); H3 (n = 299); and H4 (n = 207). Data is presented here by chronological phase (Cocina A: n = 506; Cocina B: n = 667) and by level in the Supplementary Documentation (Suppl. Table 6).

The Mesolithic faunal remains (n = 1173) from this portion of the site were excavated in Sector E and data are available from four contexts, originally interpreted by Fortea as living floors: H1 (n = 325); H2 (n = 342); H3 (n = 299); and H4 (n = 207). Data is presented here by chronological phase (Cocina A: n = 506; Cocina B: n = 667) and by level in the Supplementary Documentation (Suppl. Table 6).

As is the case in the Pericot section of the site, Iberian ibex dominate the assemblage in both phases of the Late Mesolithic (Fig. 3). In the early phase, Iberian ibex represents 70.6% of the assemblage, followed by red deer (17.4%), and rabbit (9.9%) (Table 5). Other species include equid (0.2%) and wild boar (1.6%), and a single bone each of a chamois (Rupicapra rupicapra) and a canid were also identified. In the Cocina B phase, we find a similar distribution of the primary species with Iberian ibex dominating the assemblage (67.5%), followed by red deer (16.5%) and an increase in rabbit bone (12.7%). In addition, aurochs (0.3%), equid (0.5%, and badger (0.1%) were present, along with one confirmed (0.2%) and two possible domestic Caprinae bones. It should be noted that the presence of domesticated sheep or goat in this upper level has been shown to be the result of mixing due to subsequent Neolithic activities in the cave, and is not a result of Mesolithic forager activities (Pardo-Gordó et al. 2018).

Table 5 Number of identified specimens (NISP) of taxonomic categories and their relative proportions from Fortea’s excavations by chronological phase
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Anatomical profiles are available for Iberian ibex, red deer, and rabbit, but only from H1, the uppermost stratum (Cocina B; Table 6). In the case of Iberian ibex, 61.9% of the identified elements pertain to limb bones, with a particularly large number of hind limbs (tibiae; femora), while 21.4% come from the skull and 16.7% from the axial skeleton (Fig. 4).

Table 6 Anatomical composition of the main species (Iberian ibex, red deer and rabbit) by phase from Fortea’s excavations. Note that anatomical data was only available for the faunal assemblage from H1 of Fortea’s excavation area that dates to the Cocina B phase of the Late Mesolithic
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In the case of red deer, the majority of the elements identified are also limb bones (68.8%), while 27.1% of the assemblage comes from the axial skeleton and only 4.1% are skull fragments, consisting of a mandible and a loose tooth (Fig. 5). Although the assemblage is much smaller than Iberian ibex, there is a predominance of forelimb elements. Similarly, the small assemblage of rabbit bone consists of mostly limb elements (68.8%) with only a very few axial and cranial remains (Fig. 6).

Although these data represent only a small portion of the assemblage from this part of the cave, there are striking differences in the anatomical profiles of the same taxa from the Pericot sectors. In particular, relatively more limb bones of Iberian ibex and red deer were recovered than was the case in the Pericot sections, and we suggest that this likely reflects variations in recovery methods rather than differences in past human behavior.

Fragmentation and modifications

Bone modifications (n = 265) in the form of cut marks, fractures, and burnt bones were only identified in the three main species: Iberian ibex, red deer, and rabbit, and represent 21.6% of the assemblage (Table 7). Although fractures are most common in both periods, cut marks were only recorded on 15.8% of the Cocina A modified bone, but on 25.3% of the Cocina B modified bone (Fig. 7). When we look at taxonomic variation, cut marks decrease through time for red deer and rabbits, but increase with Iberian ibex from 5.2% in the early Mesolithic to 19.6% in the late Mesolithic phase. Burnt bones are only present in H1 of Cocina B (7.1%).

Table 7 Modifications present (NISP) on the Fortea faunal assemblage by type and species
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Assemblages from 2015 to 2018

Taxonomic and anatomical composition

We analysed a total of 1309 remains from the excavation of 2015–18: 19 remains from TP3 and 1290 remains from TP4 (Table 8; Suppl. Table 7). In general, 17.9% were taxonomically and anatomically identified.

Table 8 Number of identified specimens (NISP) of taxonomic categories and their relative proportions from the 2015–18 excavations by test pit
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TP3 is composed only of three remains of Iberian ibex (mandible, ulna, tibia), one bone of red deer (calcaneus) and two of rabbit (tibia, metapodial) (Table 9). There are also six fragments (mainly long bones) of indeterminate medium-sized animals and seven indeterminate remains. Only one individual of each species was calculated.

Table 9 Anatomical composition of the main species (Iberian ibex, red deer and rabbit) by phase from 2015–18 excavations
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The TP4 assemblage includes different species of ungulates, carnivores, rodents and one bird bone (Table 8). The most abundant taxa in both units are Iberian ibex (34.2%), Caprinae (38.2%) and red deer (17.1%), but wild boar is also present (1.8%). Carnivores are represented by one long bone of Carnivora (UE5a) and one second phalange of wild cat and one second metatarsal of lynx (UE5b). Among small mammals, a few bones of leporids (4.9%) were identified: tibia, mandible, forelimb and metapodial (Table 9; Fig. 6).

All anatomical groups of Iberian Ibex (including Caprinae remains) are present in both units of TP4 (Table 9; Fig. 4). Based on the relative proportion of %MAU, mandible, humerus, radius, and hindlimb (except for the tarsal) show a percentage of over 50% (Table 10; Fig. 12). The absence or lower quantities of some elements could be due to conservation problems because the correlation between %MAU and the bone density is positive (r = 0.7522, p < 0.001), suggesting that elements with less dense bone may not have survived taphonomically. In the case of red deer, although most of the elements are present, isolated teeth, antler and metapodials are more abundant based on the %NISP (Table 9; Fig. 5). However, the relative proportion of %MAU shows dominance of limbs in comparison with the cranial and axial skeleton (Table 10; Fig. 12). Moreover, phalanges and compact bones from the carpal and tarsal are practically absent too. Wild boar is represented by a mandible, a second metatarsal, and a first and third phalanx.

Table 10 Anatomical composition of Iberian ibex and red deer by Minimum Number of Elements (MNE) and Minimum Anatomical Units (%MAU) from TP4 of the 2015–18 excavations
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Fig. 12
figure 12

Anatomical composition of Iberian ibex and red deer remains (%NISP) of unit TP4 from the 2015–18 excavation, based on MNE and %MAU

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Regarding the MNI and their age at death, the TP4 assemblage is composed of one adult wild boar, two red deer and two adult leporids. Eight Iberian ibexes were identified: a neonate represented by a very young humerus; one juvenile indicated by several unfused elements; two subadults (around 3 years of age), based on unfused epiphyses and the age of a mandible and a maxilla using tooth eruption and wear (phase III); three adults (around 5 years of age) based on tooth eruption and wear of a mandible and two maxillae (phase IV), and the MNE of mandibles and astragali; and a very old (senile) individual indicated by a mandible with tooth eruption and wear patterns in phase V.

The indeterminate remains of TP4 are abundant (82.3%) (Table 8). Most of the remains are small fragments without taxonomic or anatomical identifications. Bones of medium-sized animals are also abundant, primarily fragments of long bones, but also some articular ends, vertebrae and ribs.

Fragmentation and modifications

All remains from TP3 are fragmented and 2.2% of TP4 are complete remains (Table 11). These bones are mainly teeth of Iberian ibex and red deer, phalanges, carpal/tarsal elements and unfused articular ends of metapodial of Iberian ibex; but also, a calcaneus of red deer and a metatarsal and a phalange of wild boar.

Table 11 Complete and fragmented bones from TP4 of 2015–18 excavations
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A total of 35.4% of the remains present fresh fractures, but there are also some indeterminate (38.4%), dry (18.3%) and mixed (7.9%) fractures (Table 11). Among the types of fractures present on Iberian ibex and red deer (Table 12), fragmented diaphyses of long bones with incomplete circumference (humerus, radius, femur and tibia) stand out. There are no cylinders in the assemblage. Metapodials were better conserved as fragments of proximal or distal parts, while phalanges were mainly found as fragments of proximal and distal parts (Fig. 9 a). In the case of leporids, despite the few remains, all long bones are present as diaphysis cylinders.

Table 12 Portions of long bones, metapodials and phalanges of Iberian ibex, red deer, and rabbit. Abbreviations: complete proximal part (px), fragmented proximal part (fg. px), complete diaphysis (di), fragmented diaphysis (fg. di), complete distal part (dt), fragmented distal part (fg. dt)
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Modifications (n = 200) are only documented on TP4 bones (Table 13). Anthropogenic modifications are present mainly as burnt bones (91%), but also as cut marks (5.5%) and percussion marks (1.5%). In the case of Iberian ibex, there are two slicing marks on a burnt astragalus done during the disarticulation (Fig. 8 b), multiple slicing marks on two femur diaphyses caused during defleshing. A simple and longitudinal scraper mark on a diaphysis of a humerus that resulted from the removal of the periosteum. Multiple, transversal and intense slices on the ramus of two mandibles are the result of skinning (Fig. 8 a); and a simple slice on the diaphysis of a metatarsal maybe also have been caused by skinning. Other indeterminate long bones from medium-sized animals present slicing marks from defleshing. There are also cut marks on an indeterminate long bone of Carnivora.

Table 13 Modifications present (NISP) on the 2015–18 faunal assemblage from TP4 by type and species
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Regarding the percussion marks, notches were identified on two metapodials of Iberian ibex, and a percussion flake on a metatarsal of red deer (Fig. 9 h). Burning was identified mostly on indeterminate remains in both units, and charring (84%) is more abundant than calcination (16%). There is only one metapodial of Iberian ibex partially affected.

Non-anthropogenic modifications are few. Multiple tooth marks (Table 13) on an antler of red deer and a coxal and astragalus of Iberian ibex, as well as pits on a radius of Iberian ibex and a long bone of medium-sized animal (Fig. 11 a, c), suggest carnivore gnawing of these specimens.

Discussion

Animal exploitation from Mesolithic hunter-gatherers of Cocina cave

The Cocina cave faunal assemblage provides relevant data from two different phases of the Late Mesolithic: early phase (Cocina A) and late phase (Cocina B). The data from Cocina A come only from the Pericot excavations (partial data is also presented for Fortea excavation), but data from Cocina B are available from all excavation phases.

Ungulates are the main prey, although carnivores, leporids and birds are present in lower quantities. Comparing both phases, there is a slight difference regarding the taxonomic spectrum: between Cocina A and B the quantity of Iberian ibex increases a bit, and red deer decreases; and, other ungulates like equids, auroch and wild boar are more common in the earlier phase. Nevertheless, Iberian ibex is the best-represented taxon in both phases (more than 70% of the NISP), which is feasible considering the location of the cave in a steep environment. However, the appearance of other taxa, even with lower values, typical of open spaces such as aurochs and equids, or of woodland areas such as red deer, wild boar and roe deer, indicates the existence of altitudinal and/or seasonal movements of the groups.

Although elsewhere body part distributions have provided insights into hunting and food preparation processes (e.g., Lupo 2006), the known biases in the Pericot collection of faunal remains limits the anatomical profiles of the three main species (Iberian ibex, red deer and rabbit) and our ability to meaningfully interpret the patterns. However, we recognize that elements of the entire skeleton are present, which would indicate that hunters brought complete animals to the site and not only some portions of a carcass. On the other hand, less information and interpretation are not possible due to only having NISP counts (Pericot and Fortea data). In the study of the 2015–18 collection, the MNE and the %MAU show a different pattern in comparison with the %NISP representation (see Fig. 4–5 and 12). In this case, limbs and cranial elements of Iberian ibex are well-represented, except for some small bones, but the axial skeleton is poor. This bias is likely due to attritional processes, and could also affect the red deer assemblage. Therefore, the prey transport could be complete.

The case of rabbits is a bit complicated. The very small number of remains in the Fortea assemblage and the recent excavations (2015–18) do not allow for an interpretation of the transport and differential preservation of these prey; especially if we take into account the limited space of the last excavations. Nonetheless, rabbits are small prey that could be easily transported completely to the cave. On the other hand, there is a large assemblage of rabbit remains from the Pericot excavations, in which the anatomical composition in both phases is composed mainly of mandibles and tibiae. In this case, the absence of other elements could be due to sampling methods of Pericot's excavations.

Modifications and fractures of faunal assemblage from both Late Mesolithic phases are related mainly to anthropic activities. Based on the data from the 2015–18 excavation, the fragmentation rate is high, only a few teeth and small elements remain complete, and fresh fractures are the most common type. Among ungulates (Iberian ibex and red deer), fracture impacts are located on the ends of the long bone diaphysis, near the joints, and are evidence of actions aimed at accessing the marrow. Some bones show particularities in their fracturing, such as the Iberian ibex skulls which are fractured transversally behind the start of the horns. This morphotype of fracture has been observed in other anthropic assemblages dominated by Iberian ibex, such as those from the Early Upper Palaeolithic levels of the Cova de les Malladetes (Sanchis et al. 2023). Vertebrae are fractured in half and evidence of fracturing related to percussion. Small bones are also fractured, such as the first and second phalanges of both species, which appear with different fracture morphotypes: sagittal, proximal, distal, and lateral. The long bones of rabbits with fresh fractures seem to have been produced by human bites aimed at facilitating the disarticulation of some limbs (mandibular ramus, olecranon of the ulna, distal joint of the radius and tibia) and fracture the articular parts to suck out the marrow (mainly in the tibia and to a lesser extent in the humerus and femur). This action has created cylinders of diaphysis and epiphysis as has already been pointed out in other assemblages (Hockett and Haws 2002; Pérez Ripoll 2005; Sanchis et al. 2016).

Cut marks show variable values depending on the sector and taxon, and Cocina A fauna had a larger proportion of cut marks than the later phase Cocina B Mesolithic (based on Pericot's collection). Among the Iberian ibex and red deer, most of the cut marks are located on joints and are related to disarticulation, with fewer located on long bone diaphysis. Collection biases may have contributed to this pattern since very few long bone shafts were collected. However, in the 2015–18 collection there is also evidence of skinning, defleshing and removal of the periosteum. In the case of rabbits, there are slicing marks on the diaphysis of the tibia, which seems to be linked to defleshing, although the consumption seems to be immediately because there is not enough evidence to suggest that carcass processing is linked to meat preservation (Pérez Ripoll and García-Puchol 2019).

Carnivore modifications are present on a few Iberian ibex and red deer bones, mostly from Cocina B. It is difficult, however, to make any kind of assessment of the possible agents causing these modifications, both because of the small number of remains with modifications and their measures. Pits on cortical bone (n = 3) have dimensions (3.75 × 1.90 mm; 2.50 × 1.90 mm; 3.50 × 2.50 mm) in the range of those of various references of current predators (for example, see Fig. 8 in Sauqué and Sanchis 2017). In spite of the impossible to determine a specific carnivore, their activities seem to be occasional and we cannot relate to the use of the cave as a den. These may be sporadic visits by carnivores in search of food leftovers for scavenging.

The Mesolithic from the Mediterranean Iberia: an economic overview

The Mesolithic archaeological record from Mediterranean Iberia includes multiple sites, open and cave sites, that have provided insight into the subsistence activities of the last hunter-gatherers. The foragers at Cocina cave hunted a diversity of taxa, rabbits appear to have had only a limited importance in their diets and, there was a clear connection to coastal areas visible in the material record, such as fish and marine shellfish (Pascual Benito and García-Puchol 2015), as we describe later. These characteristics of the subsistence behavior seen in Cocina cave are common among most of the sites in this area such as Balma de la Fontanella, Benàmer, Falguera, Coves de Santa Maira, Tossal de la Roca or Cueva de Nerja among others (Aura et al. 2006; Cacho et al. 1995; LLoveras et al. 2021; Morales Pérez 2013; 2015; Pérez Ripoll 2006; Román et al. 2023; Tormo Cuñat 2011), sites located in similar mid-mountain environments where Iberian ibex also predominates. Although this subsistence pattern began during the Epipaleolithic, it was a drastic change from the Upper Palaeolithic.

The taxonomic spectrum of the Mesolithic human diet includes mostly the same species already seen during the Final Upper Palaeolithic (Aura et al. 2002; Real 2020b; Villaverde et al. 2012). Faunal remains from Upper Palaeolithic and Epipalaeolithic sites were grouped into three analytical categories: leporids (rabbits and hares); red deer or Iberian ibex as the “main” taxon; and then a variety of “minor” taxa found in assemblages that may vary in size (Aura et al. 2009; Morales Pérez 2015; Real 2021). We have also applied this classification to the Mesolithic sites, and the proportion between species, however, is different (Fig. 13). In the Upper Palaeolithic assemblages, leporids typically represent 80–90% of faunal assemblages, while red deer or Iberian ibex constitute only 6–20% of faunal remains. In contrast, Mesolithic assemblages witness a clear decrease in leporids with less than 27%, while red deer or Iberian ibex increase to 40–70% of faunal remains. Similarly, the proportion of ‘minor’ taxa increased in the Mesolithic as well, perhaps signaling a diversification of hunting practices as shown in the results of the Simpson diversity index in cases like Cocina (Cocina A 1-D = 0.32, Cocina B 1-D = 0.36) or Falguera (1-D = 0.34) (Suppl. Table 8). This change indicates a different exploitation pattern of a landscape where the Mediterranean forest and the species (red deer, roe deer, wild boar) that prefer this biotope spread from the beginning of the Holocene (Carrión 2005). In this more wooded landscape, people seem to have practiced a less specialised hunting that some have described as a wide broad-spectrum economy and related to high mobility (LLoveras et al. 2021).

Fig. 13
figure 13

Proportion of leporids, main taxon (red deer or Iberian ibex) and minority taxa from Magdalenian, Epipalaeolithic, and Mesolithic sites: Cova del Matutano (Mat.) (Olária et al. 1981), Cova dels Blaus (Bla.) (Martínez Valle 1996), Tossal de la Roca (T.R.) (Pérez Ripoll and Martínez Valle 1995), Coves de Santa Maira (S.M.) (Morales Pérez 2015), Cova de les Cendres (C.C.) (Real 2021), Balma de la Fontanella (Font.) (Lloveres et al. 2021), Cingle del Mas Nou (M.N.) (Morales and Llorente 2019)

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Carnivores are also included in this taxonomic diversity. Species of canids, felids and mustelids are present in the assemblage of Cocina cave, as well as in the other Mesolithic sites (Fernández-López de Pablo et al. 2015; Llorente 2015; Lloveras et al. 2021; Morales Pérez 2013; Morales and Llorente 2019; Pérez Ripoll 2006; Pérez Ripoll and Martínez Valle 1995). Fox is especially relevant among the other species, and humans exploited all their resources including meat, fur and bones (Pascual Benito 2017). During this phase, human consumption of terrestrial gastropods (Sphincterochila candidissima and Iberus alonensis) were documented in open-air sites such as Arenal de la Virgen and Casa Corona (Fernández-López de Pablo et al. 2013a) and also in Cocina cave (Pascual Benito, pers. com.). These gastropods are common in other Valencian Mesolithic sites, but their human consumption has not been confirmed.

Particularly noteworthy is the drop in rabbit consumption in the Mesolithic. The percentage of this small prey decreased to 45% in sites such as Cueva del Lagrimal or Cueva de Nerja; around 25% in Balma de la Fontanella and Tossal de la Roca; and even less than 15% in Coves de Santa Maira, Cingle del Mas Nou and Benàmer (Morales Pérez 2013, 2015; Pérez Ripoll 1991; Pérez Ripoll and Martínez Valle 1995; Román et al. 2023; Tormo Cuñat 2011). Therefore, the number of rabbit remains is closer to what is typically found in Middle Palaeolithic assemblages rather than the Final Upper Palaeolithic regional assemblages (e.g. Morin et al. 2019; Pérez Ripoll and Villaverde 2015; Real 2020a; Sanchis and Fernández Peris 2008). This is not a progressive decrease, but an abrupt change in the Mesolithic at least in central-eastern Iberia (Fa et al. 2013; Pérez Ripoll and García-Puchol 2019). Some researchers have suggested this change was the result of the spread of the forest and a related shift in economic behaviour (Aura et al. 2009). However, we suggest that the causes are still not clear and seem to have been more related to different mobility and occupation patterns than the availability of this prey in the environment, since forestation began earlier (during the Epipaleolithic) but the consumption dropped later in time. Other areas, however, such as the north-eastern of the Mediterranean basin, continued to consume higher quantities of rabbits (Rosado-Méndez et al. 2018; Rufà et al. 2017; Vaquero 2006). This is the case of the Abric Agut or Molí del Salt site, where in the Mesolithic level (Sup), as well as in the previous Magdalenian levels, leporids are the main prey and represent 73% of the assemblage.

Lastly, the third characteristic is the inclusion of marine resources in human activities. Shellfish or fish remains are present in some inland Mesolithic sites such as Fontanella, Cova Fosca, Coves de Santa Maira, Cocina cave, as well as the coastal site of Cativera and El Collao, and were not used only for ornament production (in the case of the shellfish) but also as part of their diet (Aparicio 2008; Aura et al. 2009; Morales et al. 2017; Pascual Benito and García-Puchol 2015; Román et al. 2020). The presence of these resources proves the existence of a strong relationship between the coastal areas and the inland sites and is in line with the high mobility of these last hunter-gatherer societies. Moreover, it offers a picture in which the species linked to marine biotopes (shellfish, fish, birds and mammals) played a more relevant role in the human diet than in previous phases (Aura 2019; Román et al. 2020). Indeed, isotopic analyses of human remains from Mesolithic sites support this consumption (Fernández-López de Pablo et al. 2013b; Fernández et al. 2020; García-Guixé et al. 2006; McClure et al. 2023; Salazar-García et al. 2014), although it is still unclear to what extent marine and lagoon products were important in the forager diet. The results of the δ13C and δ15N analyses of human bone at some sites were interpreted as aquatic products were dominant in some individuals' diets (Fontanals Coll et al. 2023). However, at Cocina cave, the δ13C and δ15N stable isotopic data from several faunal and human remains are consistent with consumers living in a terrestrial C3 environment (McClure et al. 2023), and there are variations among humans and even among herbivores. The enrichment of the δ13C value seen in humans is mirrored in herbivores and could be due to other factors in the terrestrial environment rather than marine or aquatic animal consumption. Therefore, it is possible that marine resources were part of the Mesolithic diet, however, to what extent the individuals received a high or low percentage of marine protein is still an open discussion.

On the other hand, another type of resource found in Mesolithic sites in this area (e.g. in El Collao, Mas Gelat or Tossal de la Roca) are freshwater turtles or terrapins. These are mainly remains of bone plates and to a lesser extent appendicular bones belonging to the species Mauremys leprosa (Mediterranean turtle) and Emys orbicularis (European pond turtle). In the case of El Collao, some remains of these species show evidence of human consumption, in the form of thermal alterations during the roasting of the specimens and fractures for the separation of the back and plastron (Fernández-López de Pablo et al. 2015; Moya 2024).

Some optimal foraging models regarding resource availability and economic behavior focus on a logistical subsistence organization including smaller residential mobile groups (collectors) (Bettinger et al. 2015; Binford 1980; Kelly 2013). This scenario is supported by the data from the last Mesolithic episodes in eastern Iberia: 1) a similar species spectrum among different sites; 2) a diversity in the lithic record in terms of an expedient vs curated character (Cortell-Nicolau et al. 2019); and 3) the evidence of prevalent structures like necropolises (Gibaja et al. 2015) and contexts revealing special social activities (Cortell-Nicolau et al. 2023; García-Puchol et al. 2023). Some proposals have pointed to a small band organization with a mobility radius of 25–30 km and including coastal/inland movements in eastern Iberia (Martí et al. 2009). Although our knowledge of open-air sites vs caves and rock shelters is partial and biased, the current archaeological record allows us to infer that both explain a complementary spatial locational pattern including the exploitation of aquatic resources from marine or marsh biotopes. The consumption of marine shells (mainly Cardium edule) and fishes detected in Cocina demonstrates this pattern. A recent paper analyzed the potential of lithics for defining mobility patterns this area (Cortell-Nicolau et al. 2019), concluding that the variability in correlation between lithic volumetric density (LVD) and retouch frequency emphasizes differences in the optimization of raw materials and lithic variability, consequently pointing to a logistical mobility structure (including long-term camps complemented with short-term camps for specific functions). The diversity index results are more significant and infer two groups of sites, pointing to greater diversity and long-term diversified camps (e.g., Mas Cremat, Los Baños, Secans, Botiqueria), and less diversity as is expected for hunting spots (e.g., Cocina, Falguera among others) (Cortell-Nicolau et al. 2019). Interestingly, the different lithic subunits of Cocina are coincident with a similar pattern revealed by the faunal record.

On the other hand, García-Puchol et al. (2018a, b) emphasize the variable spatial distribution patterns of the last hunter-gatherer groups in eastern Iberia with a changing settlement distribution for Mesolithic occupations in the first half of the VIII millennium (Cocina B) and involving differential persistence between areas. Some have argued that the effects of 8200 cal BP climate event had something to do with this settlement visibility issue (Fernández-López de Pablo et al. 2015), however, the continuity in Cocina cave is noteworthy (Bernabeu et al. 2014; García-Puchol et al. 2017). As, in other areas, the explanation for this pattern was linked with other factors that affected social network connectivity as a prelude to the consequences triggered by the neolithic expansion. The reliability of ungulate species such as Iberian ibex and red deer in this region offset some of the variation in the more coastal areas.

Conclusions

In general, there is great stability in the faunal record throughout the Mesolithic, not only at Cocina cave but also in other sites (e.g. Lloveras et al. 2021). Human groups that occupied Cocina cave had the same subsistence pattern in both phases (Cocina A and Cocina B). Iberian ibex was the main hunted prey, transported completely to the site. These last hunter-gatherers exploited ungulates for meat and marrow, but also for raw material to elaborate the bone industry (see Pascual Benito 2017). Other prey was occasionally exploited, even small animals like rabbits, but with less intensity than in previous periods (e.g. Pérez Ripoll 2004; Real 2020a). Occupations were seasonal, human groups used the cave at least in Spring, Summer, and to a lesser extent in Autumn (Pérez Ripoll and García-Puchol 2017).

Nevertheless, while the faunal record at Cocina cave shows continuity throughout the Mesolithic phases, other archaeological materials (e.g. lithic, plaquettes) were changing (Cortell-Nicolau et al. 2023; García-Puchol et al. 2023; López-Montalvo et al. 2024). Perhaps it was precisely the presence of stable and predictable game that allowed people to return to Cocina cave through time and that supported the broadscale cultural shifts visible in other aspects of the archaeological record.

Research on Mesolithic subsistence in eastern Iberia indicates that small, mobile groups had a logistical organisation. Mobility included coastal and inland movements, with camps of varying durations. The distribution of settlement patterns in this area is variable for Late Mesolithic groups. While in some cases it has been explained on the basis of climatic changes, the continuity in Cocina cave could be related to the connectivity of social networks before the arrival of the Neolithic groups.

These questions, as well as the unknown causes for abrupt change in the subsistence pattern in comparison with the Final Upper Palaeolithic, should remain the subject of future research. The number of Pleistocene-Holocene transition sites needs to be expanded, as well as the analyses of faunal remains from different perspectives to assess the evolution of the economic behavior of the last hunter-gatherers.