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Re-Evaluating Subsistence Strategies During the Solutrean in the Cantabrian Region (c. 22–19.5 ky cal BP)

Published online by Cambridge University Press:  10 March 2026

Rodrigo Portero Open the ORCID record for Rodrigo Portero [Opens in a new window] ,
María José Fernández Gómez Open the ORCID record for María José Fernández Gómez [Opens in a new window]  and
Esteban Álvarez-Fernández Open the ORCID record for Esteban Álvarez-Fernández [Opens in a new window]
Rodrigo Portero
Affiliation:
Department of Human Sciences, University of La Rioja , Logroño, Spain
María José Fernández Gómez
Affiliation:
Department of Statistics, University of Salamanca , Spain
Esteban Álvarez-Fernández*
Affiliation:
Department of Prehistory, Ancient History and Archaeology, University of Salamanca , Spain
*
Corresponding author: Esteban Álvarez-Fernández; Email: epanik@usal.es
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Abstract

The Solutrean in the Cantabrian region is one of the periods in the Upper Palaeolithic with the highest number of faunal studies conducted in recent decades, which offer valuable insights into how human groups exploited their environment for survival. Here, the authors analyse twenty-four archaeological levels from twelve sites dated between c. 22 and 19.5 ky cal bp, focusing on the exploitation of large mammals through palaeoecological and palaeoeconomic approaches. Their examination of prey acquisition and transport, age profiles, seasonality, nutrition, and energy costs shows that hunting decisions were influenced not only by the economic profitability of prey but also by the environment, topography, climate, animal behaviour, and species abundance. This multifactorial perspective provides an updated view of subsistence strategies at the onset of the Last Glacial Maximum.

Le Solutréen en Cantabrie est une des périodes du Paléolithique supérieur ayant fait l’objet du plus grand nombre d’études archéozoologiques effectuées au cours des dernières décennies. Ces études ont fourni des informations précieuses sur le mode d’exploitation de l’environnement pour subvenir aux besoins des groupes humains de la région. Les auteurs de cet article présentent une analyse paléoécologique et paléoéconomique de vingt-quatre niveaux archéologiques sur douze sites datés entre 22 000 et 19 500 cal bp, centrée sur l’exploitation des grands mammifères. L’examen de leur acquisition, de leur transport, de leur profil d’âge, de la saison de leur chasse et de leur apport en énergie et en nutrition démontre que le bénéfice économique n’était pas seul à influencer les décisions de chasser et que l’environnement, la topographie, le climat, le comportement des animaux et l’abondance des espèces entraient également en jeu. Cette approche multifactorielle permet d’actualiser nos idées sur les stratégies de subsistance au début du dernier maximum glaciaire.

Das kantabrische Solutréen ist eine der Perioden des Jungpaläolithikums, für welche in den letzten Jahrzehnten eine sehr große Anzahl archäozoologischer Studien unternommen wurde. Diese lieferten wertvolle Einblicke in die Art und Weise, wie menschliche Gruppen in der Region ihre Umwelt zur Sicherung ihrer Subsistenz ausbeuteten. In diesem Artikel untersuchen die Verfasser vierundzwanzig Niveaus aus zwölf archäologischen Stätten, die zwischen 22 000 und 19 500 cal bp datiert werden. Sie konzentrieren sich auf die Nutzung von Großsäugern in einer paläo-ökologischen und -wirtschaftlichen Perspektive. Die Untersuchung ihres Erwerbs, Transports, Altersprofils, der Jagdsaison, des Beitrags dieser Tiere zur Ernährung und zum Energiebedarf zeigt, dass die Entscheidung zu jagen nicht nur von der wirtschaftlichen Rentabilität der Beute abhing; die Umgebung, die Topografie, das Klima, das Verhalten der Tiere und die Erhältlichkeit der verschiedenen Tierarten spielten auch eine wichtige Rolle. Dieser multifaktorielle Ansatz ermöglicht es, die Subsistenzstrategien am Anfang des Letzteiszeitlichen Maximums erneut zu bewerten.

Keywords

subsistence patternsarchaeozoologypalaeoeconomypalaeoecologyCantabrian SpainUpper Solutreanmodèles de subsistancearchéozoologiepaléoéconomiepaléoécologieCantabrieSolutréen supérieurSubsistenzmodelleArchäozoologiePaläowirtschaftPaläoökologieKantabrienoberes Solutréen

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Article
Information
European Journal of Archaeology , First View , pp. 1 - 20
Creative Commons
Creative Common License - CCCreative Common License - BY
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0), which permits unrestricted re-use, distribution and reproduction, provided the original article is properly cited.
Copyright
© The Author(s), 2026. Published by Cambridge University Press on behalf of the European Association of Archaeologists

Introduction

The Solutrean in the Cantabrian region is a well-known period that extends from c. 24,000 to 19,500 cal bp, covering periods from the Heinrich Event 2 to Greenland Stadial 2b (Grootes et al., Reference Grootes, Stuiver, White, Johnsen and Jouzel1993). During this time, the region became a refuge for hunter-gatherers from northern Europe, who adapted to technological, geographical, social, and subsistence changes. The Upper Solutrean (c. 22–19.5 ky cal bp) material culture is characterized by a rich assemblage of lithic points, an increase in flat retouch, parallel and covering techniques, bifacial working, and the production of shouldered points and concave base points (Rasilla Vives & Santamaría Álvarez, Reference Rasilla Vives and Santamaría Álvarez2005; Aura et al., Reference Aura, Tiffagom, Jordá Pardo, Duarte, Fernández de la Vega and Santamaria2012; Schmidt, Reference Schmidt2015). For this period, around fifteen sites have been studied in recent decades in terms of the subsistence strategies used in the region (e.g. Yravedra, Reference Yravedra2010; Ríos et al., Reference Ríos, Garate, Gómez-Olivencia, Iriarte, Ríos and Gómez-Olivencia2013; Castaños Ugarte, Reference Castaños Ugarte and Corruchaga2016; Altuna & Mariezkurrena, Reference Altuna, Mariezkurrena and Corchón2017; Álvarez-Fernández et al., Reference Álvarez-Fernández, Bécares, Jordá Pardo, Álvarez-Alonso, Elorza, Garcia-Ibaibarriaga, Schmidt, Cascalheira, Bicho and Weniger2019; Torres-Iglesias et al., Reference Torres-Iglesias, Marín-Arroyo and Rasilla2022).

The Cantabrian region, a well-defined coastal strip over 400 km long between the Cantabrian Sea and the Cantabrian Mountains, hosts an exceptional concentration of Upper Palaeolithic sites. These sites are remarkably homogeneous in their technological, artistic, and symbolic traditions, making the region a coherent unit for studying prehistoric hunter-gatherer societies. Initial studies of subsistence strategies, developed in the 1970s (Altuna, Reference Altuna1972; Freeman, Reference Freeman1973; Straus, Reference Straus1976), proposed a dichotomy between sites specialized in hunting red deer (Cervus elaphus) and those focused on Iberian ibex (Capra pyrenaica), depending on local biotopes. In this regard, the works of Lawrence Straus (Reference Straus1983a, Reference Straus, Clutton-Brock and Grigson1983b) revealed that such hunting behaviours were already emerging during the Solutrean.

In the 1980s, numerous excavations and the publication of data from previously explored caves led to a significant increase in faunal studies. Their findings were compiled in interdisciplinary monographs on sites such as Las Caldas (Corchón, Reference Corchón1981; Soto & Meléndez, Reference Soto, Meléndez and Corchón1981), El Rascaño (Altuna, Reference Altuna, González-Echegaray and Barandiaran1981; González-Echegaray & Barandiarán, Reference González-Echegaray and Barandiaran1981), La Riera (Altuna, Reference Altuna, Straus and Clark1986; Straus & Clark, Reference Straus and Clark1986), and El Juyo (Barandiarán et al., Reference Barandiarán, Freeman, González-Echegaray and Klein1987; Klein & Cruz Uribe, Reference Klein, Cruz-Uribe, Barandiarán, Freeman, González-Echegaray and Klein1987). By the 1990s, the accumulated archaeozoological data made it possible to interpret subsistence changes during the Upper Palaeolithic in the region (Altuna, Reference Altuna1990, Reference Altuna, Moure and Sainz1995; González Sainz, Reference González Sainz and Moure1992; Quesada, Reference Quesada1997a, Reference Quesada1997b). This period also saw the introduction of taphonomic analyses into faunal research (e.g. Pumarejo & Bernaldo de Quirós, Reference Pumarejo and Bernaldo de Quirós1990; Mateos Cachorro, Reference Mateos Cachorro1999).

Since the early 2000s, taphonomic studies in the Cantabrian region have adopted shared methodological frameworks (Yravedra, Reference Yravedra2002; Mateos Cachorro, Reference Mateos Cachorro2003). Recent decades have seen the integration of GIS in subsistence research (e.g. Andrés-Herrero et al., Reference Andrés-Herrero, Becker and Weniger2018; Torres-Iglesias et al., Reference Torres-Iglesias, Marín-Arroyo and Rasilla2022), alongside applications of optimal foraging theory (e.g. Marín-Arroyo, Reference Marín-Arroyo2010), isotopic analyses (e.g. Castaños de la Fuente, Reference Castaños de la Fuente2017), zooarchaeology by mass spectrometry (e.g. Torres-Iglesias et al., Reference Torres-Iglesias, Marín-Arroyo, Welker and Rasilla2024), and genetic studies (e.g. Lira et al., Reference Lira, Tressières, Chauvey, Schiavinato, Tonasso-Calvière and Seguin-Orlando2025).

In this research on subsistence strategies, the Solutrean period did not go unnoticed, as evidenced by the studies conducted in the 1970s by Altuna (Reference Altuna1972), Freeman (Reference Freeman1973), and Straus (Reference Straus1976). Although monographs proliferated in the region throughout the 1980s, only a few sites, such as Las Caldas (Corchón, Reference Corchón1981) and La Riera (Straus & Clark, Reference Straus and Clark1986), included information on Solutrean sequences. By the 1990s and the turn of the millennium, excavations at sites with Solutrean sequences multiplied, and hunting strategy approaches were revised (e.g. Mateos Cachorro, Reference Mateos Cachorro1999; Yravedra, Reference Yravedra2002). High percentages of red deer were also observed in Solutrean sites (e.g. Altuna, Reference Altuna, Moure and Sainz1995; Yravedra, Reference Yravedra2002; Rojo & Menéndez, Reference Rojo and Menéndez2012), and explanations for this phenomenon rest on a progressive evolution towards the specialized hunting of this animal, which for some authors occurred from 22,000 cal bp in the region (Altuna, Reference Altuna1992; Quesada, Reference Quesada1997b). In recent decades, archaeozoology and taphonomy have become the pillars for reconstructing subsistence strategies at sites such as Abrigo de La Viña (Torres-Iglesias et al., Reference Torres-Iglesias, Marín-Arroyo and Rasilla2022) or the cave of El Cierro (Portero et al., Reference Portero, Elorza, Jordá Pardo and Álvarez-Fernández2025).

Despite methodological advances, the biotope-based hunting model proposed in the 1980s remains influential, albeit with refinements (e.g. Yravedra, Reference Yravedra2002; Rojo & Menéndez, Reference Rojo and Menéndez2012; Portero, Reference Portero2022). However, regional studies integrating multiple analytical approaches are still lacking and hence this study addresses the subsistence strategies of Upper Solutrean hunter-gatherers in the Cantabrian region through a combined palaeoecological and palaeoeconomic perspective. Its aim is to correlate diverse factors influencing resource exploitation and thus offer an updated view of large mammal exploitation during this period.

Materials and Methods

To ensure that the comparison of subsistence strategies is founded on the same analytical basis, we selected levels on archaeological sites that met the following three criteria:

  1. 1. Radiocarbon dates: only levels with radiocarbon dates calibrated using OxCal and IntCal20 were included (see Supplementary Material, Table S1).

  2. 2. Faunal remains: data must come from excavations conducted with modern recovery techniques, i.e. from sites excavated post-1970.

  3. 3. Archaeofaunal studies: the levels studied must contain at least thirty faunal remains and be supported by methodologically-sound archaeozoological studies.

With these conditions in place, we compared the subsistence strategies in twenty-four stratigraphic levels at twelve sites, six in the western, four in the central, and two in the eastern Cantabrian region (Figure 1).

Figure 1.

Locations of the Upper Solutrean sites selected for this study. 1) Las Caldas; 2) La Viña; 3) El Cierro; 4) El Buxu; 5) La Riera; 6) Los Canes; 7) El Linar; 8) El Ruso; 9) Cobrante; 10) El Mirón; 11) Arlanpe; 12) Amalda.

Palaeoecological analysis

The palaeoecological analysis employed ecological indices to assess species richness, abundance, and evenness. Biodiversity was evaluated using Simpson’s Diversity Index (D), which accounts for species abundance and is suitable for small samples. The formula used is:

$$ D=\sum \limits_{i=1}^S\frac{n_i\left({n}_i-1\right)}{N\left(N-1\right)} $$

This index will indicate the differences or similarities of faunal comparisons without being greatly affected by the quantity of existing remains, as it is an index that can be used in small samples. To enhance interpretability, we also applied Simpson’s Reciprocal Index (1/D), commonly used in zooarchaeology (e.g. Grayson & Delpech, Reference Grayson and Delpech2002; Portero et al., Reference Portero2022).

Shannon’s Index (H′) was also calculated to assess community heterogeneity, considering both species richness and relative abundance. The formula used is:

$$ H^{\prime }=-\sum pi\ ln\ pi $$

In assemblages dominated by a few taxa, H′ values tend towards zero, indicating low diversity (Grayson & Delpech, Reference Grayson and Delpech2002; Jost, Reference Jost2010; Portero et al., Reference Portero, Fernández-Gómez and Álvarez-Fernández2024b).

To determine the degree of homogeneity of a faunal assemblage, the Shannon homogeneity index or Evenness (E) has been calculated, based on the following equation:

$$ E=-\sum pi\ ln\ pi/\mathit{\ln}\;(S) $$

This index fluctuates between 0 and 1; the closer to 0, the lower the homogeneity (Jost, Reference Jost2010).

The effective number of species (qD) was calculated using the exponential of Shannon’s index:

$$ qD=\mathit{\exp}\left(-\sum pi\ ln\ pi\right) $$

This approach transforms diversity indices into effective species counts, reflecting species richness adjusted for rarity. Diversity results from Simpson’s and Shannon’s indices are visualized through scatter plots, with Simpson’s reciprocal index (1/D) and Shannon’s evenness on the y-axis, and the natural and Napierian logarithms of the minimum number of individuals (MNI) on the x-axis (Grayson & Delpech, Reference Grayson and Delpech2002; Portero et al., Reference Portero2022). To assess the relationship between diversity indices and the number of identified specimens (NISP), Spearman’s rank correlation was applied, revealing the strength, direction, and statistical significance of the associations (p < 0.05).

Prey acquisition and transport

To investigate prey acquisition and transport strategies, we analysed the skeletal profiles of documented taxa to assess the contribution of animal carcasses to human groups (e.g. Yravedra, Reference Yravedra2002; Portero, Reference Portero2022). Taxonomic abundance was used to infer hunting preferences (e.g. Altuna, Reference Altuna1992, Reference Altuna, Moure and Sainz1995; Yravedra, Reference Yravedra2002; Rojo & Menéndez, Reference Rojo and Menéndez2012; Portero et al., Reference Portero, Cueto, Elorza, Marchán-Fernández, Jordá Pardo and Álvarez-Fernández2024a), based on the number of remains and NISP data from Upper Solutrean contexts.

Anatomical representation was examined for evidence of transport decisions. The bones were grouped by anatomical region (cranial, axial, forelimb, hindlimb, and limbs) to identify patterns indicative of selective transport (Yravedra, Reference Yravedra2006).

Age patterns

To study the age ranges, the mortality profiles of mammals at each level were compared, classifying them into age categories: immature, juvenile, adult, and senile. We only considered individuals of species whose presence at a site had an anthropogenic origin. For greater precision in determining hunting strategy, mortality profiles were grouped using ternary plots based on the four zones proposed by Discamps and Costamagno (Reference Discamps and Costamagno2015), adapted to the life cycle of each species:

  • - JPO (Juvenile–Prime–Old) refers to the number of juveniles being greater than that of adults, which is greater than that of senile individuals; typically L-shaped profiles

  • - JOP (Juvenile–Old–Prime) refers to the number of juveniles being greater than that of senile individuals, which is greater than that of adults; characteristic of U-shaped attritional profiles

  • - P (Prime): dominated by adult individuals

  • - (Old): predominance of senile individuals.

Seasonality and catchment areas

The seasonality of site occupations was inferred from age profiles, considering reproductive cycles (mating, gestation, birth, and development) as well as the estimated life expectancy of modern wild species (Krasińska & Krasiński, Reference Krasińska and Krasiński1995; Outram & Rowley-Conwy, Reference Outram and Rowley-Conwy1998; Alados & Escós, Reference Alados, Escós, Salvador and Barja2017; Carranza, Reference Carranza, Salvador and Barja2017; Mateos-Quesada, Reference Mateos-Quesada, Salvador and Barja2017; Pérez-Barbería et al., Reference Pérez-Barbería, García-González, Palacios, Salvador and Barja2017).

The catchment areas for hunting resources were reconstructed by integrating seasonal occupation data derived from the prey’s age-at-death profiles with known seasonal migrations of mammals. Using a digital elevation model (DEM) of the Cantabrian region, we processed provincial maps at a scale of 1:200,000 (MTN 200) of the Spanish National Geographical Institute (IGN) and bathymetric data (GEBCO, 2020) with QGIS v.3.22. This enabled the visualization of potential resource zones, including areas now submerged, based on altitudinal migratory patterns. Seasonal site activity, when available in the literature, was mapped to estimate distances between sites and resource areas throughout the year.

Diet and energy intake

We calculated the calories provided by each taxon consumed at the different sites to establish the nutritional contribution of each prey in the diets of the groups that inhabited the Cantabrian region during the Upper Solutrean. To do this, we adhered to the following procedure (Portero et al. Reference Portero, Fernández-Gómez and Álvarez-Fernández2024b):

  • - The mean weight of each taxon from the proportion of males and females of each species (Clutton-Brock & Iason, Reference Clutton-Brock and Iason1986) was determined.

  • - Meat and fat yields were estimated following Binford’s (Reference Binford1978) observations for caribou and sheep, applicable to cervids and small bovids in the Cantabrian region. For rabbits (Oryctolagus cuniculus), nutritional data were sourced from the Spanish Food Composition Database (BEDCA, 2007). Juvenile and immature individuals were adjusted by reducing adult weights by 33 per cent and 66 per cent, respectively, in line with species life cycle estimates. This allowed calculation of the exploitable biomass per individual.

  • - Caloric values per kilogram of meat and fat were estimated using data from the USDA (2020) and the Spanish Food Composition Database (BEDCA, 2007). These values were multiplied by the estimated edible weight of each specimen (see Supplementary Material, Table S2) to calculate total energy yield. Only species showing clear evidence of anthropogenic manipulation were included, ensuring their contribution reflects human subsistence activities.

  • - Energy contributions were used to identify the most nutritionally significant prey. To assess whether caloric estimates might underestimate smaller species (due to their lower body mass) Spearman’s rank correlation was calculated between the percentage of total calories per taxon and the logarithm of their MNI.

Profit from prey

To determine the economic profitability of each species at Upper Solutrean sites, we considered the search time, handling time, and return rates, respectively. These aspects affect the costs of acquiring prey and, in combination with the energy provided, allow us to rank them in a prey hierarchy (Stiner et al., Reference Stiner, Beaver, Munro, Surovell and Bocquet-Appel2008; Portero et al., Reference Portero, Cueto, Jordá Pardo, Bécares and Álvarez-Fernández2019, Reference Portero, Cueto, Fernández-Gómez and Álvarez-Fernández2022) (Supplementary Material, Table S3).

Results

Palaeoecological analysis

The palaeoecological analysis of twenty-four Upper Solutrean levels (Supplementary Material, Table S4) reveals a significant inverse correlation between NISP and Shannon Evenness (ρ = -0.77; p < 0.001), indicating that higher specimen counts are associated with lower taxonomic homogeneity (Figure 2). El Mirón (levels 125, 122, 121) and Cobrante 4 are the most homogeneous, while Caldas 11 and Riera 16 show the least uniformity. Taxonomic richness is low across all levels, with the effective number of species values ranging from 1.6 to 5 (x̄ = 2.51±0.16). Simpson’s reciprocal index (1/D) also shows a significant inverse correlation with NISP (ρ = -0.47; p < 0.03) (Figure 3). Cierro H1 exhibits the highest dominance of a single taxon, whereas Arlanpe II is the most diverse. Despite most levels showing strong dominance of one taxon, some levels (Cobrante 3, Mirón 122, Arlanpe II, Canes 2A) lack this. Following Quesada’s (Reference Quesada1997a) threshold of 1/D ≥ 2.45 for diversified hunting, Arlanpe II, Mirón 122, Canes 2A, Ruso IVa, and Amalda IV suggest broader subsistence strategies.

Figure 2.

Graph showing the relationship (Spearman ρ) between the taxonomic representation (LN NISP) and the Shannon homogeneity values (Evenness) in Upper Solutrean levels in the Cantabrian region (see Supplementary Material, Table S4).

Figure 3.

Graph showing the relationship (Spearman ρ) between the taxonomic representation (Log. NISP) and Simpson’s reciprocal index (1/D) in Upper Solutrean levels in the Cantabrian region (see Supplementary Material, Table S4).

Red deer (Cervus elaphus) is the most frequent taxon in seventeen out of twenty-four levels and exceeds 60 per cent representation in twelve of these levels (Supplementary Material, Figure S1). Iberian ibex (Capra pyrenaica) is the most abundant in five levels, but in none does it reach 60 per cent of NISP. Only in Amalda IV and Buxu 3 do Pyrenean chamois (Rupicapra pyrenaica) predominate. It is noteworthy that in Buxu 3 Pyrenean chamois is dominant but that in Buxu 1 red deer is the main taxon. A similar situation exists in El Mirón, where Iberian ibex is dominant in levels 121 and 122, while red deer predominates in level 125. Red deer also have a significant presence even in some levels where Iberian ibex and Pyrenean chamois have greater representation, such as in Mirón 121 and 122, Buxu 3, Amalda IV, and Canes 2A. Wild horse (Equus ferus) has some importance in Ruso IVa, Linar 3B, and Caldas 11, always below 27 per cent. Large bovids (Bos/bison sp.) and roe deer (Capreolus capreolus) are present in most levels but have low representation (< 13.5 per cent).

Acquisition and transportation of prey

All skeletal elements of red deer (Cervus elaphus) are represented across the sites. Cranial remains are particularly abundant, exceeding 50 per cent in Caldas 7 and Cierro H1 and H2. Although axial elements are generally less frequent, they are present in all levels and even dominate in La Riera (levels 16, 12, 10) and Ruso IVa, suggesting that complete carcasses were transported. Limb bones are consistently represented, especially autopodia (distal part of the limbs) and zeugopodia (forearm and lower leg), with the highest proportions in Linar 3B (Supplementary Material, Figure S2).

Unlike red deer, Iberian ibex (Capra pyrenaica) shows a high frequency of axial elements across most sites, except at El Cierro, where it is absent. Cranial remains are also abundant, exceeding 50 per cent in some levels, while limb bones are fewer, except in Buxu 3 and Cobrante 3 and 4. The prevalence of axial elements may reflect whole carcass transport due to the species’ smaller size, while its absence in El Cierro could relate to low NISP or distance from the hunting area. For Pyrenean chamois (Rupicapra pyrenaica), skeletal representation is available for Buxu 3 and 1, Caldas 11 and 5–4, and Amalda IV. Cranial elements dominate in Caldas 11 and Buxu 3 and 1, largely due to numerous dental remains. As with Iberian ibex, all skeletal elements of Pyrenean chamois are well represented, suggesting whole carcass transport.

For wild horses (Equus ferus), skeletal data from Caldas 11, 7, 5–4, and Ruso IVa show high proportions of cranial and axial elements, with hindlimbs abundant only in Caldas 11. This pattern supports the hypothesis of whole carcass transport, except in Caldas 11 (Supplementary Material, Figure S2). Other species lack sufficient remains for meaningful comparison.

Age patterns

The age of death of ungulates documented in the Upper Solutrean levels shows a greater number of captures of adult animals than of immature, juvenile, and senile individuals.

For red deer (Cervus elaphus), adults range from 30 to 60 per cent (100 per cent in Mirón 125), while immature specimens vary between 10 and 50 per cent, dominating in Amalda IV. Senile individuals are least represented, peaking at 25 per cent in Buxu 1. Several levels, such as Mirón 121 and 122, show a balanced age distribution. Ternary diagrams reveal a predominance of JPO profiles, with fewer cases showing JOP (Supplementary Material, Figure S3).

For Iberian ibex (Capra pyrenaica), adults dominate across all levels, occasionally sharing representation with juveniles and immature specimens (e.g. Cierro H1, Caldas 9, Mirón 121). Senile individuals are only recorded in Caldas 11 and 5–4. Two instances from Ruso IVa are excluded due to their non-anthropogenic origin (Yravedra et al., Reference Yravedra, Gómez-Castañedo and Muñoz2010). Ternary diagrams show a predominance of P profiles, with few cases in the JPO zone (Supplementary Material, Figure S4).

In seven levels with two or more individuals, Pyrenean chamois (Rupicapra pyrenaica) shows a consistent predominance of adults (≥ 50 per cent). Juvenile and senile individuals are generally ≤ 25 per cent, while immature animals are only noted in Amalda IV and Buxu 3, with a notable presence in the latter. Ternary diagrams indicate mostly P and JPO profiles, with only Caldas 11 showing a JOP profile (Supplementary Material, Figure S5).

For the remaining hunted species, data are too limited for comparative analysis. However, their presence will be considered when assessing seasonal site occupation and their potential dietary contribution to human groups.

Seasonality and catchment areas

Mortality patterns suggest that most ungulate captures took place in spring and summer (e.g. Cierro H1, Riera 16), although hunting in autumn and winter is also evident (e.g. Ruso IVa, Buxu 3). In several cases, broad age ranges prevent precise seasonal interpretations, leaving open the possibility of year-round occupation (e.g. Riera 10, Caldas 11).

Potential catchment areas for key hunting resources can be visualized in a DEM (Figure 4), based on their seasonal movements and cave occupation patterns. Table 1 presents data on seasonality, cave elevation, and distance from the coastline, offering insights into resource accessibility and mobility strategies.

Figure 4.

Digital elevation model of the Cantabrian region with bathymetric data indicating the Upper Solutrean coastline, modelled in QGIS v.3.22. The map includes the seasonal altitudinal distribution of hunted species and the occupation of selected caves. 1) Las Caldas; 2) La Viña; 3) El Cierro; 4) El Buxu; 5) La Riera; 6) Los Canes; 7) El Linar; 8) El Ruso; 9) Cobrante; 10) El Mirón; 11) Arlanpe; 12) Amalda.

Table 1.

Location, altitude, and distance to the current (DCC) and Upper Solutrean (DUSC) coastlines based on marine regression data (Jordá Pardo et al., Reference Jordá Pardo, Carral, Maestro, Álvarez-Alonso, Arias, Bécares, Garcia-Ibaibarriaga, Murelaga, Suárez and Suarez2018), with seasonality estimated for Upper Solutrean ungulate hunting. SP=spring; SU=summer; A=autumn; W=winter; ND=no data.

Spring migration patterns show red deer (Cervus elaphus) moving from low altitudes to areas above 500 m asl, while Iberian ibex (Capra pyrenaica) and Pyrenean chamois (Rupicapra pyrenaica) reach elevations above 1000 m asl. During spring, caves such as Las Caldas, La Viña, El Cierro, La Riera, El Ruso, El Mirón, and Amalda were occupied. Red deer could have been hunted within c. 2 km of these sites, and the presence of juveniles may reflect spring birthing behaviour. In Mirón 121 and 122, despite low MNI, red deer may have been sourced from the valley of the river Asón. Due to the high-altitude location of Iberian ibex and Pyrenean chamois in spring, their hunting appears less frequent, as seen in Buxu 3 and Amalda IV.

During summer, all caves except Viña V are occupied. Red deer migrate to elevations above 500 m asl, coinciding with the birthing and nursing period of hinds, which intensifies hunting. However, their catchment areas are more extensive (c. 5 km), which increases transport costs. In contrast, Iberian ibex and Pyrenean chamois remain at high altitudes but are hunted more frequently than in spring, as seen in Amalda, El Ruso, and Las Caldas, with a focus on young adults. In El Buxu and Amalda, red deer are also hunted, consistent with their summer migration to nearby high-altitude zones.

In autumn, both the number of captures and cave occupation intensity decline. Red deer hunting shifts to lower altitudes (within a radius of c. 2 km), coinciding with the rutting season (Carranza, Reference Carranza, Salvador and Barja2017). Rock-dwelling ungulates (Iberian ibex and Pyrenean chamois) begin descending, increasing their representation in levels where they predominate. Autumn hunting may have been more effective near caves, as mating behaviour leads these species to occupy open areas for harem control (Pérez-Barbería et al., Reference Pérez-Barbería, García-González, Palacios, Salvador and Barja2017).

Winter shows the lowest intensity of site occupation, coinciding with a decline in red deer hunting, despite the species being at its lowest migratory altitudes. In contrast, Pyrenean chamois hunting increases, particularly in Buxu 3 and 1. Additionally, heightened carnivore activity in Amalda IV suggests that the cave was probably not occupied by humans during this season (Yravedra, Reference Yravedra2010).

These results are based on mammal remains and are limited to species identification and chronological markers of death. It is important to note that migratory patterns can vary depending on geography, climate, and species adaptability. Therefore, interpretations of seasonal hunting strategies should be approached with caution.

Diet and energy intake

We estimated the caloric contribution of each taxon to the Upper Solutrean human diets in the Cantabrian region, based on published data from caves where MNI and evidence of consumption were reported. Detailed energy calculations are provided in the Supplementary Material (Table S5).

Considering total caloric estimates, red deer (Cervus elaphus) contributed the largest share to the meat-based diet (40 per cent), followed by wild horse (Equus ferus) (30 per cent) and bovids (Bos/bison sp.) (c. 20 per cent). Iberian ibex (Capra pyrenaica) and Pyrenean chamois (Rupicapra pyrenaica) provided 7 per cent and 3 per cent, respectively. Other taxa, such as roe deer (Capreolus capreolus) and rabbit (Oryctolagus cuniculus), contributed less than 0.3 per cent (Figure 5).

Figure 5.

Percentage of calories and potential meat and fat contributed by the taxa at Upper Solutrean sites in the Cantabrian region.

If we consider the weight that each of the levels represents in the overall picture, we observe that Caldas 11 could have provided the highest caloric energy to Upper Solutrean human groups, with a total of 4,991,944 Kcal (36 per cent) and just over 4368 kg of meat and fat. In contrast, the levels that account for the least energy are Cierro H2, which could have contributed 40,737 Kcal (0.3 per cent) and just under 36.7 kg of meat and fat. A Spearman correlation between %Kcal per taxon and Log. MNI reveals a moderate but non-significant positive relationship, indicating that there was no linear correlation between energy contribution and prey abundance (Supplementary Material, Figure S6).

Species profitability was assessed based on handling time, catchment proximity, and energy yield (Supplementary Material, Table S6). Comparison with regional prey rankings shows that, except for large bovids, species appear in order of economic return.

Discussion

The palaeoecological and palaeoeconomic analysis of twenty-four Upper Solutrean levels in the Cantabrian region gave us a number of insights into the subsistence strategies in the region between c. 22 and 19.5 ky cal bp, summarized here.

Taxonomic data confirm the dominance of red deer (Cervus elaphus). Iberian ibex (Capra pyrenaica) is also significant at sites like Los Canes, El Mirón, and Arlanpe, although never exceeding 60 per cent representation. Pyrenean chamois (Rupicapra pyrenaica) predominates in Buxu 3 and Amalda IV; in the former, taphonomic evidence supports human hunting (Rojo, Reference Rojo2020), whereas in the latter, carnivore activity may account for its presence (Yravedra, Reference Yravedra2010). These findings align with the palaeoecological analyses in this paper, showing low species diversity and dominance by a single taxon; notably, earlier levels (Mirón 125, Cobrante 4) show greater diversity than later levels (Mirón 122 and 121, and Cobrante 3). This may be due to a progressive evolution towards specialized red deer hunting, which, as some authors have indicated, occurs from around 22 ky cal bp, after the peak of glacial expansion (Altuna, Reference Altuna1992; Quesada, Reference Quesada1997b). While climate may have influenced this trend, it is not the sole factor (Yravedra, Reference Yravedra2002). Taphonomic studies confirm the anthropogenic processing of red deer across all sites, while Iberian ibex and Pyrenean chamois were consumed in eleven and nine levels, respectively. The contribution of wild horse is notable in Caldas 11, 9, 7, Linar 3B, and Ruso IV; other species remained below 13.5 per cent NISP.

This taxonomic representation seems to indicate that the meat strategy targeted red deer as the main prey, given their high percentages of NISP and the evidence of anthropic processing of their carcasses. This specialization in red deer hunting during the Upper Solutrean has been observed at other sites in the Cantabrian region (Altuna, Reference Altuna1972, Reference Altuna, Moure and Sainz1995; Freeman, Reference Freeman1973; Quesada, Reference Quesada1997b; Yravedra, Reference Yravedra2002; Rojo & Menéndez, Reference Rojo and Menéndez2012; Portero et al., Reference Portero, Elorza, Jordá Pardo and Álvarez-Fernández2025).

The anatomical distribution of red deer shows complete skeletal representation across all sites, although axial elements are scarce in some levels (e.g. Cierro H1 and H2, Cobrante 3, Linar 3B), suggesting selective transport of high-value parts. In contrast, the abundant axial remains of Iberian ibex and Pyrenean chamois indicate frequent whole-body transport, except in Cierro H1, and Buxu 1 and 3, respectively. Wild horses, present in four levels, shows variable skeletal patterns: cranial in Ruso IVa and Caldas 5–4, axial in Caldas 9, and appendicular in Caldas 11. Although there does not appear to be a consistent pattern in the transport of wild horses, all skeletal elements are represented in these sites.

Mortality profiles vary across levels but generally reflect a hunting strategy focused on immature and young adult red deer, probably targeting hinds and calves (e.g. Caldas 11, 9, 7 and 5–4, Cierro H1, Buxu 3, Amalda IV). The presence of senile individuals, especially in Cierro H1, suggests occasional selective hunting. Although the presence of immature and senile animals may be the result of accumulation by large carnivores such as leopards, lions, or hyenas (e.g. Davis, Reference Davis2002), in all cases the presence of red deer at these sites has been demonstrated to be due to anthropogenic activity.

In the case of the Iberian ibex and Pyrenean chamois, the hunting strategy targeted adult and juvenile individuals in their reproductive stage. Given the scarcity of immature individuals, this may be related to individual hunting of these rock-dwelling animals.

Based on these mortality profiles, we established the seasonality of the sites. In general, a greater number of captures were made during the spring and summer, as seen in Cierro H1, Riera 16, Mirón 121, and Mirón 122. There are, however, also levels that show occupation during the autumn and winter, such as in Viña V, Buxu 3, or Buxu 1. A small group where captures could have been made throughout the year is attested at Caldas 11 and 5–4, Riera 10, Ruso IVa, and Amalda IV, due to the wide range of documented individuals.

Andrés-Herrero et al. (Reference Andrés-Herrero, Becker and Weniger2018) analysed ungulate catchment areas around major Cantabrian river basins, showing that site locations were favourable for hunting red deer, bovids (Bos/bison sp.), and wild boar (Sus scrofa), while Iberian ibex was only profitable in steep, rocky terrain. Our data align with these findings, although wild boar remains lack anthropic marks in the levels we analysed. Seasonal patterns from our catchment map indicate that red deer was intensively hunted during the breeding season within a ≤ 5 km radius of all caves. Although occasionally present in summer, rock-dwelling species (Iberian ibex and Pyrenean chamois) are more abundant in spring and autumn, coinciding with mating and birthing periods. For other taxa, limited sample sizes prevent identification of clear seasonal trends.

Caloric estimates indicate that red deer was the main contributor of meat and fat, accounting for around 39.5 per cent of potential energy. Although less frequent, large prey such as wild horse (c. 29.5 per cent) and bovids (c. 19 per cent) surpassed red deer in energy contribution at Viña V, Caldas 11, and Ruso IVa. Iberian ibex, dominant in five sites, contributed < 7.5 per cent overall and was never the primary energy source. Pyrenean chamois provided < 3.5 per cent of total calories, with notable contributions only at Buxu 3 (c. 49 per cent) and Amalda IV (c. 19 per cent). Other species contributed less than one per cent.

Considering these energy data, a hypothetical group of ten Upper Solutrean hunter-gatherers would need slightly more than eight million calories annually for their subsistence. All this energy would not depend exclusively on the faunal remains we analysed, as an individual would not consume more than 300 g of protein daily (1300 Kcal) without consuming a significant amount of fat, or risking the possibility of diseases such as ‘rabbit starvation’ (Speth, Reference Speth2025). Nonetheless, the dependence on meat resources could account for around half the intake (Binford, Reference Binford2001). Thus, among the analysed levels, only Caldas 11 would provide enough meat and fat resources for a continuous occupation throughout an entire year for the group. This implies that human groups would have to rely on seasonal mobility to sustain themselves. Additionally, other faunal resources would have provided nutritional supplements during the Upper Solutrean, such as marine resources (Álvarez-Fernández, Reference Álvarez-Fernández2008; Álvarez-Fernández & Fernández García, Reference Álvarez Fernández and Fernández-García2012), birds (e.g. Elorza, Reference Elorza1990; Garcia Petit, Reference Garcia Petit and Corruchaga2016; Eastham, Reference Eastham and Corchón2017), and fish (e.g. Adán et al., Reference Adán, Álvarez-Lao, Turrero, Arbizua and García-Vázquez2009; Portero et al., Reference Portero, Elorza, Jordá Pardo and Álvarez-Fernández2025).

Integrating palaeoecological and palaeoeconomic data reveals that red deer, bovids, and wild horse were generally hunted near cave sites, while Iberian ibex often required travel beyond 5 km. From an energy-efficiency perspective, rock-dwelling species (Iberian ibex and Pyrenean chamois) were only advantageous where topography facilitated capture. Considering altitudinal migration, red deer emerges as the most profitable prey due to its high caloric yield, low acquisition cost, frequent encounters, and manageable handling time, especially in coastal valleys (Portero et al., Reference Portero, Cueto, Jordá Pardo, Bécares and Álvarez-Fernández2019). Even wild horses and bovids were more energetically favourable than Iberian ibex and Pyrenean chamois at sites like Buxu 3, Ruso IVa, and Amalda IV.

A hunting strategy focused on red deer may have led to longer recovery times for the species, reducing its availability and requiring human groups to maintain mobility for both survival and ecosystem regeneration (Stiner et al., Reference Stiner, Beaver, Munro, Surovell and Bocquet-Appel2008; Merkle et al., Reference Merkle, Cherry and Fortin2015; Venkataraman et al., Reference Venkataraman, Kraft, Dominy and Endicott2017). Intensive exploitation, especially of females and calves, as observed in some sites, would slow population recovery (Beaver, Reference Beaver2007; Stiner et al., Reference Stiner, Beaver, Munro, Surovell and Bocquet-Appel2008). To avoid resource depletion, groups are likely to have adopted logistical or residential mobility strategies (Binford, Reference Binford1978). In this case, the seasonal occupation of most of the analysed captures indicates that the human groups that inhabited the Cantabrian region during the Solutrean maintained intense mobility.

Conclusions

Based on palaeoecological, archaeozoological, and taphonomic data from faunal remains, our characterization of the subsistence strategies during the Solutrean (c. 22 to 19.5 ky cal bp) has highlighted the significant role that hunted resources played in the diet of human groups in the Cantabrian region. It highlighted the importance of red deer, not only as the main species hunted and consumed but also as the greatest energy resource in the meat diet during this period.

The integration of palaeoecological and palaeoeconomic data demonstrates that hunting decisions among Solutrean groups were shaped by more than just the economic profitability of large mammals. Environment, topography, climate, animal behaviour, and species abundance also played key roles, alongside cost–benefit considerations. These findings refine the subsistence models proposed for the Cantabrian region in the 1970s, offering a more nuanced understanding of human–environment interactions during the Upper Palaeolithic.

The coastal valley model includes sites located near present-day coastlines, typically showing high frequencies of red deer. While this pattern holds during the Upper Solutrean, in some cases, larger and more energetically profitable prey, such as wild horses or bovids, surpassed red deer in meat contribution, indicating that prey selection was guided by caloric return rather than abundance alone.

The mountain model includes sites in steep or mountainous areas, where rock-dwelling species are expected to dominate. While sites like Los Canes, Arlanpe, and Amalda fit this model, only Amalda IV shows rock-dwelling taxa exceeding 60 per cent of NISP during the Upper Solutrean. Moreover, these species never represent the main caloric resource. This suggests that, although topography influences prey accessibility, Solutrean groups prioritized species with higher energy yields, optimizing subsistence strategies beyond environmental constraints.

The interior valley model includes sites located further than thirty km from the present-day coast, near diverse biotopes, and shows greater faunal diversity (e.g. La Viña, Las Caldas, El Cobrante, and El Mirón). These sites feature varying dominance of red deer, Iberian ibex, Pyrenean chamois, and occasionally wild horses and bovids. Stratigraphic shifts in prey dominance, as seen in El Buxu and El Mirón, may reflect ecological changes, species abundance, or seasonal cave use. Despite these fluctuations, red deer and wild horse consistently emerge as the most calorically profitable species, indicating that prey selection was primarily driven by energy return rather than abundance alone.

In sum, our study has shown us that, while the environment and the topography of the surroundings of a given site influence subsistence strategies by determining the proximity and abundance of faunal resources, we must not overlook that human groups tend to prioritize economically more profitable animals, even if their acquisition cost is higher than that of other animals in a given environment, as the energy benefit compensates for the search, capture, and handling costs. This strategy will be modified when the conditions that make these resources profitable change, whether due to changes in the abundance of specific species in the environment, increased difficulty in acquiring them, changes in seasonal behaviours, or to avoid overexploitation of the resource.

Supplementary Material

To view supplementary material for this article, please visit http://doi.org/10.1017/eaa.2025.10028.

Acknowledgements

The research was made possible by a PhD scholarship awarded to Rodrigo Portero from the Junta de Castilla y Leon and European Social Fund, and a Margarita Salas postdoctoral fellowship awarded by the University of Salamanca and the Spanish Ministry of Science (Plan de Recuperación, Transformación y Resiliencia) with EU-Next-Generation funds. We are grateful to L.C. Teira of the University of Cantabria for producing the map illustrated in Figure 1.

Funding

This work was undertaken in the context of the PaleontheMove Project PID2020-114462GB-I00/AEI/10.13039/501100011033 (PI: Esteban Álvarez Fernández), funded by the Programa Estatal de Fomento de Generación de Conocimiento y Fortalecimiento Científico y Tecnológico of the Spanish Ministry of Science and Innovation.

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Figure 0

Figure 1. Locations of the Upper Solutrean sites selected for this study. 1) Las Caldas; 2) La Viña; 3) El Cierro; 4) El Buxu; 5) La Riera; 6) Los Canes; 7) El Linar; 8) El Ruso; 9) Cobrante; 10) El Mirón; 11) Arlanpe; 12) Amalda.

Figure 1

Figure 2. Graph showing the relationship (Spearman ρ) between the taxonomic representation (LN NISP) and the Shannon homogeneity values (Evenness) in Upper Solutrean levels in the Cantabrian region (see Supplementary Material, Table S4).

Figure 2

Figure 3. Graph showing the relationship (Spearman ρ) between the taxonomic representation (Log. NISP) and Simpson’s reciprocal index (1/D) in Upper Solutrean levels in the Cantabrian region (see Supplementary Material, Table S4).

Figure 3

Figure 4. Digital elevation model of the Cantabrian region with bathymetric data indicating the Upper Solutrean coastline, modelled in QGIS v.3.22. The map includes the seasonal altitudinal distribution of hunted species and the occupation of selected caves. 1) Las Caldas; 2) La Viña; 3) El Cierro; 4) El Buxu; 5) La Riera; 6) Los Canes; 7) El Linar; 8) El Ruso; 9) Cobrante; 10) El Mirón; 11) Arlanpe; 12) Amalda.

Figure 4

Table 1. Location, altitude, and distance to the current (DCC) and Upper Solutrean (DUSC) coastlines based on marine regression data (Jordá Pardo et al., 2018), with seasonality estimated for Upper Solutrean ungulate hunting. SP=spring; SU=summer; A=autumn; W=winter; ND=no data.

Figure 5

Figure 5. Percentage of calories and potential meat and fat contributed by the taxa at Upper Solutrean sites in the Cantabrian region.

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