Mammalian turnover as an indicator of climatic and anthropogenic landscape modification: A new Meghalayan record (Late Holocene) in northern Iberia

Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 Available online 1 March 2023 0031-0182/© 2023 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/). Mammalian turnover as an indicator of climatic and anthropogenic landscape modification: A new Meghalayan record (Late Holocene) in northern Iberia Adri´ an ´ Alvarez-Vena a , * , Ana B. Marín-Arroyo b , Diego J. ´ Alvarez-Lao a , C´ esar Laplana c , Martín Arriolabengoa d , Daniel Ballesteros e , Arantza Aranburu d , Peru Bilbao d , ´ Angel Astorqui f , Yolanda Díaz-Casado f a Department of Geology, University of Oviedo, C/ Jesús Arias de Velasco s/n, 33005 Oviedo, Spain b EvoAdapta Group, Universidad de Cantabria, 39005 Santander, Spain c Museo Arqueol´ ogico y Paleontol´ ogico de la Comunidad de Madrid, Pza. Bernardas s/n, 28801, Alcal´ a de Henares, Madrid, Spain d Department of Mineralogy and Petrology, University of the Basque Country, UPV/EHU, Leioa, Spain e Department of Geodynamics, University of Granada, Campus de Fuentenueva s/n, Granada, Spain f Tanea Documentaci´ on y Conservaci´ on SL, C/ Juan Jos´ e P´ erez del Molino 16, 39006 Santander, Spain ARTICLE INFO Editor: Dr. Howard Falcon-Lang Keywords: Climatic change Chalcolithic Bronze Age Iron Age Micromys minutus Mus musculus ABSTRACT The Punta Lucero III cave is a natural trap where abundant vertebrate remains were accumulated during the Meghalayan (Late Holocene). To better understand the paleoenvironmental conditions in which this record was accumulated, the micromammal assemblage, comprising a minimum number of 1396 individuals belonging to 19 taxa, was studied using the Mutual Ecogeographic Range and the Habitat Weighting Method. Throughout ~2600 years, the micromammal community's quick turnover reflected a shift from patchy forests and humid meadows to open, shrubbier grasslands. The Late Holocene Thermal Maximum's humid and mild climatic conditions underwent a cooling and aridification phase, coeval with the Iron Age Cold Epoch. These concluded in a slight temperature rising, coeval with the Roman Warm Period. Macromammals experienced a shift from wild populations to domestic herds. Therefore, this work discusses a broader context for this mammalian turnover from a human cultural perspective. 1. Introduction The Holocene was considered a relatively warm and stable epoch compared to the last glacial period (Dansgaard et al., 1993). Nevertheless, abrupt climatic changes and cooling events identified throughout the Holocene (Alley et al., 1997; Bond et al., 1997; Mayewski et al., 2004; Walker et al., 2018), even if weaker than those of the Last Glacial Cycle, have been intense enough to cause the rise and collapse of significant cultures and civilisations worldwide (Van Geel et al., 1996; deMenocal, 2001; Büntgen et al., 2011). Principal warming or cooling trends at millennial timescales are caused by orbital forcing (deMenocal et al., 2000; Mayewski et al., 2004; Wanner et al., 2008) in addition to fluctuations in total solar irradiance and peaks of volcanic activity, which are also related to abrupt climate shifts (Bray, 1971; Van Geel et al., 1996; Bond et al., 2001; Mayewski et al., 2004; Wanner et al., 2008; Steinhilber et al., 2009; Miller et al., 2012; Sigl et al., 2015; Borzenkova et al., 2015). However, since the end of the Pleistocene and, especially during the Holocene, human activity has become an increasing force shaping the terrestrial and marine ecosystems (Vitousek et al., 1997; Crutzen, 2002; Doughty et al., 2010; Ellis, 2011; Barnosky, 2008, 2013; Boivin et al., 2016), causing a chemical fingerprint, both on a regional and global scale (Ruddiman and Thomson, 2001). From Mid-Holocene onwards, human activities such as animal husbandry, agriculture, and metallurgy, have had a significant impact on the landscapes of the regions inhabited by humans, including the Iberian Peninsula, making anthropogenic and climatic signals often indistinguishable in pollen records (Carri´ on et al., 2010). Due to this climatic and anthropogenic influence, mammals have experienced significant range shifts, extinction events, extirpations, and domestication processes (Rosengren et al., 2021). In this sense, the Holocene warming * Corresponding author. E-mail address: [email protected] (A. ´ Alvarez-Vena). Contents lists available at ScienceDirect Palaeogeography, Palaeoclimatology, Palaeoecology journal homepage: www.elsevier.com/locate/palaeo https://doi.org/10.1016/j.palaeo.2023.111476 Received 15 January 2023; Received in revised form 23 February 2023; Accepted 24 February 2023 Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 2 and the development of human cultures also led to the turnover of the Iberian mammal communities. While domestic herds progressively replaced wild macromammals, micromammal assemblages, whose geographical distribution was affected by habitat changes (climatic or human-driven), also reflected the arrival of allochthonous commensal species (Altuna, 1980; Cuenca-Besc´ os et al., 2009; L´ opez-García et al., 2013; Ba˜ nuls-Cardona et al., 2017a; Domínguez García et al., 2019; Domínguez García et al., 2020; Domínguez-García et al., 2022; Mocl´ an et al., 2023). For this reason, macroand, particularly, micromammals are a valuable source of paleoenvironmental information that has been successfully analysed from diverse perspectives at key sites of the Cantabrian Region (e.g., Altuna, 1980; L´ opez-García, 2008; Cuenca-Besc´ os et al., 2009, 2012; ´ Alvarez-Lao, 2014; Ballesteros et al., 2020; ´ Alvarez-Vena et al., 2021; Jones et al., 2019, 2021). Iberian macromammal assemblages from Holocene sites are mainly associated with archaeological contexts (Altuna, 1980; Mariezkurrena, 1990; Casta˜ nos, 1997; Altuna and Mariezkurrena, 2012; Vega-Maeso et al., 2016). Consequently, they are biased by anthropogenic activity. On the other hand, micromammal accumulations usually result from the predatory activity of raptors or mammalian carnivores (Andrews, 1990; García-Morato et al., 2023). Although these remains can be found in assemblages that do not necessarily involve predation (as in the case of natural traps, e.g., ´ Alvarez-Lao et al., 2020), these kinds of sites are scarce in the literature. In our study case, accidental falling and nocturnal birds of prey are the most likely accumulating agents of the micromammals' assemblage. Still, according to the generalist habits of owls, they accurately represent the environment in which they hunt (Andrews, 1990). In this context, the site presented here is of great relevance as it yielded a Holocene mammal association mainly formed in a natural pitfall trap (at least in the case of macromammals), i.e., without bias by humans or specialised predators, offering an exceptional opportunity to understand the climatic change and human-environment interactions during the Meghalayan (Late Holocene) at the Cantabrian Region. 2. The Punta Lucero III site 2.1. Geographic context The Punta Lucero III site (PL-III) is located at the eastern end of the Cantabrian Region, a strip of land (30–50 km wide) with rugged relief that extends along the north of the Iberian Peninsula. It is a karst shaft (vertical cave) situated on the northern slope of the Punta Lucero mount (43◦21′34.44”N, 3◦6′10.31”W, 240 m asl; Fig. 1) in the municipality of Zierbena/Ciervana (Bizkaia, Basque Country, northern Spain). This mountain flanks the west bank of the Nervi´ on River mouth, where the Bilbao Port has been built (Fig. 1C-D). The site formed within the Lower Cretaceous limestone (calcarenite), sandstone and shale (Garrote et al., 1993) and forms part of a karst coastal area developed in a vadose environment during the Quaternary (Aranburu et al., 2015). Currently, the cave is located at the highest point of a quarry, whose exploitation began during the construction of the Bilbao seaport. However, before these works, it was located halfway up the slope, between the top of the mount and the Cantabrian Sea (Fig. 1E). Nowadays, this territory is part of the Euro-Siberian biogeographic region. The site is in the Atlantic climatic domain, characterised by temperate summers, mild winters, and no dry season (type Cfb according to the K¨ oppen-Geiger classification; Beck et al., 2018). The mean annual temperature (MAT) at Zierbena is 13.9 ◦C, with maximum and minimum month-mean temperatures of 19.8 and 8.7 ◦C, respectively. Precipitation is distributed throughout the year, reaching an annual mean of ~1150 mm (Couto et al., 2011). However, the plant communities in this area have 30% of Mediterranean species (Patino et al., 2002). Some of these taxa expanded due to human activities such as the repeated use of fire to obtain pasture, as is the case of the holm oak (Quercus ilex) and the Kermes oak (Quercus coccifera), adapted to live in more extreme conditions than the Atlantic species (Patino et al., 2002). Direct evidence of the human presence in Punta Lucero dates to times even before the formation of the studied sequence, as evidenced by the human remains (Homo sapiens bones) found at mount ridge by G´ omezOlivencia et al. (2015) in the sites of Punta Lucero II (5566–5318 cal yr BP) and Covach´ on III (4796–4424 cal yr BP), at ~200 and ~ 500 m from the PL-III shaft respectively. In the nearby site of Pico Ramos, it has also been found a sepulchral context of Chalcolithic age (5862–4295 cal yr BP) containing human skeletal remains of a minimum of 104 individuals (Baraybar and de la Rúa, 1995; Zapata, 1995). The palynological study of Pico Ramos Chalcolithic burials depicted an open landscape with signals of anthropisation, in which a mixed-oak grove formation of temperate and humid character was developed (Iriarte, 1994). 2.2. Stratigraphy Before the quarry exploitation, the studied sequence consisted of a cave infill (Fig. 2) whose entrance was above the current level. The homogeneous sedimentary fill, mainly consisting of large limestone clasts and loamy sediment, was sampled at different depth intervals from its top towards its base, sometimes coinciding with stratigraphic levels of contrasting characteristics and, on others, within more thick and homogeneous sets. Due to the narrow excavation area, it was impossible to preserve a stratigraphic profile, so we divided the sequence into different stratigraphic sections each time we extracted a group of large limestone blocks so that bone samples could be grouped. Sections or levels were labelled F to A, from bottom to top of the preserved sequence. The depth was measured from the top of the remnant deposit, partially exposed after the last blasting works in the quarry. To identify differences in the sedimentary sequence, petrographic and mineralogical analyses were made in stratigraphic levels F, D, and C. Overall, they disclose matrix-supported microfacies, with fine to medium sub-angular quartz grains, mainly of sedimentary rocks but also few chert and metamorphic quartz (Appendix 1). Level F is at the bottom of the sequence, lying over the unexcavated deposit. This level was established by its high microcharcoal content in greyish sediment. Level F was under a big limestone block; sediment containing this block was named Level E. Level D was determined after finding the first cervid remains. Level C was labelled after the excavation of Level B, which was an unstable infill. Samples collected during the surface cleaning and the removal of isolated bones before the excavation of Level B were grouped in Level A since most of them fell from the lateral or above. Therefore, Level A must be considered with caution as a reworked sample. 3. Materials and methods The mammal remains studied in this work come from the rescue excavations in the Punta Lucero quarry (Fig. 1D; Fig. 2) in September 2019. The stabilisation blasting at the quarry was supposed to destroy the cave. However, this did not happen, so the remaining stratigraphic sequence is still preserved at the site. 3.1. Stratigraphic study The stratigraphic section of the PL-III site was divided into levels based on the nature, colour, granulometry, cohesivity, sedimentary structures, and relative abundance of paleontological remains (e.g., Arriolabengoa et al., 2015). The top of the PL-III site was destroyed by quarrying; thus, level depth was measured considering the excavation surface in September 2019. The stratigraphic section was complemented by microscopical observations of unconsolidated sediments and X-Ray Diffraction (XRD) analysis to characterise mineralogy and microfacies and to infer the sediment provenance. See Appendix 1 for further details. A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 3 Fig. 1. Geographical context for the Punta Lucero III site. (A) Location of the Cantabrian Region (B) in the Iberian Peninsula. (B) Other sites mentioned in this work: 1 Valdavara-1; 2 Monte Areo mire; 3 El Olivo; 4 La Güelga; 5 Torca del Le´ on; 6 El Espinoso; 7 El Sertal; 8 Cueto la Avellanosa; 9 Cueva del Cobre; 10 La Molina peatbog; 11 La Garma; 12 Kaite; 13 El Mir´ on; 14 Cueva Mayor; 15 Mirador; 16 Zalama peatbog; 17 Pico Ramos; 18 Arenaza; 19 Punta Lucero-II and Covach´ on-III; 20 Kobaederra; 21 Amalda; 22 Herriko Barra. (C) Hypsometric background with 100 m altitude intervals of the Punta Lucero surroundings. (D–E) Orthophotography of the Punta Lucero surroundings: (D) after the stabilisation works in the quarry that allowed the discovery of the Punta Lucero shaft, and (E) before the construction of the Bilbao Port. Cartography and orthophotography from the geoEuskadi online GIS-viewer (Eusko Jaurlaritza/Gobierno Vasco). A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 4 3.2. Dating Three bone samples and two charcoals were selected for AMS radiocarbon dating and sent to the Beta Analytic laboratory (Miami, USA). Obtained ages and published dates discussed in this work were calibrated through the IntCal20 database (Reimer et al., 2020) in OxCal v.4.4.4 (Bronk Ramsey, 2009), considering the 2 σ standard deviation (95.4% probability). Fig. 2. Characterisation of the Punta Lucero III excavation. (A) Orthophoto of the excavation area. The projection of the sections in b and c is indicated. (B) A–B cross-section of the shaft. (C) C–D cross-section of the shaft. (D) Stratigraphic profile of the shaft infill. Small mammal NISPi omits the Lepus remains. Photogrammetric reconstructions of backgrounds from sections of B–D provided by GIM-Geomatics. A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 5 3.3. Collecting and sorting 3.3.1. Micromammals A total of 24 samples of sediment (~10 kg each) were collected and water-screened using two superimposed sieves of 0.5 and 2 mm. The materials in the fine fraction (0.5–2 mm) were sorted by checking the samples with a binocular microscope under 10×magnification. The obtained bone remains were then classified and studied at the Geology Department of the University of Oviedo (Spain). Taxonomic identifications were carried out employing a stereomicroscope Nikon SMZ800N, equipped with a 16 Mpx digital camera. Measurements were taken on the microscope photographs using Adobe Photoshop CC software. Due to their diagnostic value and potential of preservation, the anatomical elements selected for taxonomic identifications were the following: p3 and postcranial skeleton for lagomorphs; isolated molars for murids; m1 for arvicolines (also considering the M3 for the Terricola subgenus, and the M1 and M2 for the species belonging to the Microtus (Agricola) agrestis group); mandibles and maxillae for soricids and erinaceids; and upper molars and humeri for talpids. A comparative collection of micromammal skeletal remains obtained from recent barn owl pellets was also used. Hares were identified following Palacios and L´ opez Martínez (1980), Llorente (2010), and Pelletier (2018). The general identification of rodents and shrews followed Rom´ an (2019) and Nores (1989), respectively. Detailed taxonomical and morphological analyses were based on: Krapp and Niethammer (1982), Nadachowski (1984), and Luzi and L´ opez-García (2019) for Microtus (Agricola) ex gr. agrestis (which includes the species Mi. (Ag.) lavernedii, Mi. (Ag.) rozianus and Mi. (Ag.) agrestis) and Microtus (Microtus) arvalis lower dentition; Nores et al. (1982), Pem´ an (1983) and Barti (2006) for Neomys mandibles; Pasquier (1974), Nores (1988) and Knitlov´ a and Hor´ aˇ cek (2017) for Apodemus (Sylvaemus) dentition; Darviche et al. (2006) for Mus; and Niethammer (1990) and Guti´ errez et al. (2019) for Talpa humeri and upper molars respectively. The taxonomic classification followed the systematics proposed by Wilson et al. (2016, 2017) and Wilson and Mittermeier (2018), also considering the work of Kryˇ stufek et al. (2020) for Clethrionomys and Kryˇ stufek and Shenbrot (2022) for Microtus (Terricola) pyrenaicus. The number of specimens was obtained by counting the most frequent taxonomically identifiable element per species (NISPi, see Lyman, 1984). The relative abundance of each micromammal species was based on the minimum number of individuals (MNI), which was calculated considering the laterality of the most frequent diagnostic element. 3.3.2. Macromammals The main macromammal remains from each level were recovered directly from the excavation surface and coordinated individually. In addition, sediment samples for mechanical sieving of the smaller bones and fragments were collected and labelled according to depth. Anatomical and taxonomical identifications followed the works of Pales and Lambert (1971), Schmid (1972), Barone (1976), and Hillson (2005). The Osteological Collection of the EvoAdapta group at the University of Cantabria was also used for comparison purposes. The age at death has been estimated based on the molar development and the epiphyseal fusion degree of the limb bones. Age estimations followed the criteria published by Noddle (1974), Silver (1980), Mariezkurrena (1983), Azorit et al. (2002), and Tom´ e and Vigne (2003). Relative abundance is based on the number of identified specimens (NISP) with previous reassembling of the fragmented bones. 3.4. Preliminary identification of predatory alterations To evaluate if the likely cause of the micromammal accumulation was other than the accidental falling to the cave, the enamel of arvicoline molars was analysed in search of alterations caused by digestion (and, consequently, predation), following Andrews (1990) and Fern´ andez-Jalvo et al. (2016). The observed digestion signals were counted and classified according to the following categories: absent, light, moderate, heavy, and extreme (Andrews, 1990; Fern´ andez-Jalvo et al., 2016). Macromammal bones were also observed using a Leica stereomicroscope at 10–80×magnification in search of predatory or natural alterations following Binford (1981) and Marín-Arroyo (2010). 3.5. Paleoenvironment The landscape surrounding the PL-III shaft was inferred using the Habitat Weighting Method (HWM; Evans et al., 1981; Andrews, 2006; Blain et al., 2008; Cuenca-Besc´ os et al., 2009; L´ opez-García et al., 2014), which is based on the habitat-type preferences of each micromammal taxon. The obtained habitat types are the following: open dry (OD) comprises meadows under seasonal climate change, dry grasslands, and scrublands; open humid (OH) corresponds to evergreen meadows with dense pastures and suitable topsoil; open woodland (OW) represents woodland margins and forest patches with moderate ground cover; woodland (Wo) indicates mature forest; and water (Wa) corresponds to areas along freshwater streams, lakes, and ponds. The environmental preferences of each species were obtained from Nores (1989), Wilson et al. (2016, 2017), and Wilson and Mittermeier (2018). Hare (Lepus europaeus) remains were obtained simultaneously by excavation (throughout the complete sequence) and sieving. This makes them overrepresented in the sampling compared to the other micromammal species, which were only sampled in specific sections, so they have been excluded from landscape reconstructions. The house mouse (Mus musculus) is a commensal species of humans that commonly inhabits anthropic environments; therefore, it has not been included in the HWM calculations. 3.6. Paleoclimate The climatic conditions at the time of each level's accumulation were estimated using the Mutual Ecogeographic Range (MER) method, based on microvertebrate associations (e.g., Martínez-Solano and Sanchiz, 2005; Blain et al., 2009, 2016; Fagoaga et al., 2019). This method is frequently used to calculate the mean annual temperature (MAT), the mean temperature of the warmest month (MTW), the mean temperature of the coldest month (MTC), and the mean annual precipitation (MAP). In this work, we have also used this method to obtain monthly rainfall and temperature values to observe seasonal patterns. These parameters were calculated by getting the common geographical areas (represented in a net of 10 ×10 km UTM squares) of each micromammal species association (including hares) from the PL-III sequence. The climatic data of the shared UTM squares for each assemblage and the current climatic data from the PL-III site were obtained using the online application agroclimap.aemet.es of the Iberian Climate Atlas (Couto et al., 2011). The current distribution of each species was obtained from Palomo et al. (2007). 4. Results 4.1. Radiocarbon dating Obtained dates ranged from 4402 to 836 cal yr BP (Table 1). Four samples yielded an age according to their stratigraphical position. In contrast, sample Beta-541,219, a charcoal collected at the bottom of the excavated deposit, was likely an outlier as it provided the youngest date of the set (1055–836 cal yr BP). 4.2. Predatory activity Teeth with signs of digestion are scarce in the micromammals' sample (<1%), but in the few cases of digested molars, they reached heavy to extreme corrosion. According to the classification proposed by A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 6 Andrews (1990), this suggests that category-5 predators, mainly mammalian carnivores, ingested those few individuals. However, most of the specimens observed (99%) do not show predation features. The macromammal sample does not show butchery or burning (cooking) features. The only predator signals in the PL-III bone accumulation are those produced by carnivores (Table 5), affecting 6.2% of the specimens. Carnivore tooth marks are more common at levels C (12.3%) and B (9%), being scarcer at levels E (5.9%), F (5.3%), and D (4.6%). 4.3. Micromammal record The Micromammal assemblage of the PL-III sequence yielded a minimum (MNI) of 1396 individuals belonging to 19 taxa (Table 2; Figs. 3–6; Appendix 3). Among rodents, arvicolines (Fig. 3; Table S3.1) are represented by genera Arvicola (Ar. sapidus), Microtus (Mi. (Agricola) lavernedii, Mi. (Microtus) arvalis, Mi. (Terricola) sp., Mi. (Terricola) pyrenaicus, and Mi. (Terricola) ex gr. lusitanicus–duodecimcostatus), and Clethrionomys (Cl. glareolus); additionally, murids (Fig. 4a–c) include three species belonging to three genera: Micromys minutus (Fig. 5A; Table S3.2), Mus musculus, and Apodemus (Sylvaemus) sylvaticus (Fig. S3.1; Table S3.3). In the eulipotyphlans order, the soricids family (Fig. 4d–i’) yielded six taxa belonging to genera Crocidura (Cr. russula and Cr. gueldenstaedtii: Fig. S3.2; Table S3.4), Sorex (Socoronatus and So. minutus) and Neomys (N. anomalus and N. fodiens niethammeri: Fig. 5B; Table S3.5); talpids and erinaceids are represented by one species each: Talpa aquitania (Fig. S3.3; Fig. S3.4; Table S3.6) and Erinaceus europaeus respectively. Lastly, the morphology and size of the postcranial skeleton (Fig. 5C; Fig. S3.5a–d’; Table S3.7) and the lower third premolars (Fig. S3.5b) allowed us to assign the lagomorph remains to the species Lepus europaeus. Arvicolines yielded the most significant number of individuals from levels F, D, and B (Table 2; Fig. 6), being Mi. (Ag.) lavernedii the best-represented species at these levels (35.6–42.9%). Conversely, the lowest proportion of this arvicoline, reached at Level C (19.4–7.8%), matches a considerable increase in the abundance of Cr. russula, a soricid that is scarcely represented at the bottom of the sequence (levels F and D: 0.8% and 0% respectively). We identified two types of micromammal associations in the PL-III sequence according to their shared distribution range (Appendix 4; Fig. 7). The first one, coming from Level F, cohabits nowadays in areas at the northern slope of the Cantabrian mountains (Fig. 7A), where oceanic influence is higher. Conversely, species associations from levels C and B, at present, mainly coexist at the southern slope of the Cantabrian mountains (Fig. 7B–C) in areas of a more continental climate. 4.4. Macromammal record The studied sample comprises 3220 remains (NSP), of which 1569 (48.7%) could be taxonomically classified (Table 3). Among domestic ungulates, bovids dominate the assemblage (Fig. 8a–f), mainly sheep and goats (Ovis orientalis aries: Fig. 8a–a’; and Capra aegagrus hircus: Fig. 8b–d) at the top section of the stratigraphic sequence (levels C–B: Fig. 9; Table 3), followed by cattle (Bos primigenius taurus; Fig. 8e–f). Conversely, equids (Equus sp.) are barely represented, although these have yielded remarkably well-preserved remains (Fig. 8i). All the suid remains correspond to infantile (mostly) or young individuals (Fig. 8j–k): dentitions still retain the deciduous teeth in wearing (M3/m3 are not yet erupted at any of the individuals), and limb bones are not yet fully grown (epiphyses were not fused at the time of death). Consequently, it was not possible to ascribe these remains to the domestic pig or the wild boar, so they are classified as Sus scrofa ssp. Wild ungulates comprise two cervid species: the red deer (Cervus elaphus: Fig. 8g) and the roe deer (Capreolus capreolus: Fig. 8h–h’); both, especially the red deer, are well represented at the lower section of the sequence (levels F–D: Fig. 9; Table 3). Finally, most carnivore remains belong to the European badger (Meles meles). However, remains of canids have also been found, which, due to their small size, are ascribed to dogs (Canis lupus familiaris). The age profile of the assemblage (Table 4) is dominated by infantile (55.8%) and juvenile (24.4%) individuals, while adults (19.8%) are the less abundant group. In the case of Equus sp., Cervus elaphus and Sus scrofa ssp. samples, these reach 80% of infantile individuals. 4.5. Paleoenvironment The different micromammal assemblages found at each of the studied layers of the PL-III sequence describe an open landscape in an evolving context (Table 6, Fig. 10D). Highest proportion of moisture indicators (OH: 47.9%) is at the bottom of the sequence (Level F), where lowest values of aridity were also obtained (OD: 0.6%). The mature forest signal is weak in most of the levels (Wo: <1%), reaching the highest values also at Level F (Wo: 2.4%). However, aridity indicators increase towards de top of the deposit, especially at Level C (OD: 42.5–52.9%). Regarding the micromammal sample from Level D, this is not quantitatively enough (MNI: 14) to make reliable interpretations, though, except for the higher proportion of aquatic indicators (ΔWa: +20%) and the slightly increasing aridity in this level (ΔOD: +4.7%), a certain similarity among levels D and F samples can be observed. The open landscape inferred from micromammal assemblage is consistent with the stratigraphic study of the PL-III site (Appendix 1). This study suggests sediments resulting from soil erosion and run-off processes in the surroundings of the paleontological site. Both processes are common in an open landscape with little vegetation to protect the soil from erosion and improve water infiltration rather than runoff processes (García-Ruiz, 2010). 4.6. Paleoclimate According to MER results (Appendix 4; Fig. 11; Fig. 12A), mean temperatures during the formation of Level F (MAT: 12.5 ±0.7 ◦C), which are slightly under the present-day data (MAT: 13.9 ±0.2 ◦C), are decreasing towards levels C (MAT: 9.9 ±1.7 ◦C) and B (MAT: 10.3 ± 1.3 ◦C), especially the winter temperatures (Figs. 13–14a). The mean rainfall of Level F (MAP: 1359 ±298 mm) is slightly over current mean values (MAP: 1156 ±17 mm), though it also decreases in levels C (MAP: 968 ±319 mm) and B (MAP: 893 ±339 mm). Monthly temperature and rainfall estimations (Appendix 4; Fig. 11) obtained for Level F display the typical Atlantic pattern of the Cantabrian coast. However, the climographs of levels C and B show an increasing continental trend, with more extreme differences between Table 1 Radiocarbon dates from Punta Lucero. Level Depth (cm) Material Method Lab Ref 14 C Deviation cal yr BP a B 77 Bone AMS Beta-629,438 1980 30 1992–1830 C 242 Bone AMS Beta-541,215 2630 30 2837–2723 D 296 Charcoal AMS Beta-541,218 3070 30 3366–3185 F 430 Bone AMS Beta-541,217 3830 30 4402–4097 F 445 Charcoal AMS Beta-541,219 1040 30 1055–836 a Calibrated using the software OxCal v.4.4.4 (Bronk Ramsey, 2009) against the IntCal20 curve (Reimer et al., 2020). Further data is provided in Appendix 2. A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 7 winter and summer temperatures and a considerable decrease in rainfall throughout the year. 5. Discussion 5.1. Origin of the mammal assemblage Predation signals observed in the micromammal sample only affect a few isolated molars (<1%), which reach a high–extreme degree of enamel corrosion produced by digestion. These specimens could get the deposit in the stomach of the trapped carnivores (Canis lupus familiaris or Meles meles) since no other features exclusively produced by predators were observed. The taphonomic signals caused by the intense rockfall or the excavation procedures could be masking those produced by nocturnal birds of prey or other predators, which cannot definitively be ruled out of acting as accumulating agents. In any case, no bias is observed regarding the prey typeor size range. Therefore, the most likely causes of the micromammal sample accumulation are accidental falling in the cave, pellet regurgitation by nocturnal birds of prey, or a combination of both. We do not exclude that run-off processes deposited some micromammal remains after small transport (<5–10 m) from the vicinity of the PL-III site (Appendix 1). Otherwise, the low transport was insufficient to round off the skeletal/dental remains. Regarding the macromammal sample, the proportion of carnivore tooth marks (6.2%: Table 5) is significantly lower than that observed in badger dens by Arilla et al. (2020) and Mallye et al. (2008), whose samples showed this type of alteration in the 40% and the 20–30% of the bones, respectively. The mortality profile of the macromammal sample (Table 4) also displays clear differences with the obtained at badger burrows by Arilla et al. (2020), where the 64% of the individuals are adults, 28% are juvenile, and 8% are infantile. Conversely, this pattern inverses at PL-III (adult 15.9%; juvenile 23.8%; infantile 60.3%). Natural pitfall traps, such as the one studied by ´ Alvarez-Lao (2014), also provided samples dominated by young individuals since those are more likely to get involved in potentially fatal situations. In this case, carcasses of dead animals trapped in the cave would have been scavenged by badgers (Meles meles), whose tooth marks are compatible with those observed in PL-III. Therefore, badgers, the best-represented carnivore in the assemblage, could have fallen into the cave while chasing the smell of the cadavers. 5.2. Paleoenvironment and paleoclimate The micromammal accumulation of the PL-III sequence stands out by its great abundance, species richness, and state of preservation. Most of the identified species inhabit the region nowadays; however, when compared to a sampling on regurgitation pellets performed by Gonz´ alez Oreja et al. (1993), some differences can be highlighted: modern assemblages in the cave surroundings (Fig. 6) include the brown rat (Rattus norvegicus) while lacking other taxa such as the bank vole (Clethrionomys glareolus) and the Güldenst¨ adt's shrew (Crocidura gueldenstaedtii), which are displaced to less-anthropized inland areas (Palomo et al., 2007), or the common vole (Mi. (Mi.) arvalis), whose presence at north-western Iberia is today limited to the southern slope of the Cantabrian mountains. Concerning the evolution of the PL-III micromammal association (Fig. 6), some taxonomic and biogeographical remarks of paleoenvironmental value can be addressed. Among rodents, species of the field vole group (Microtus (Agricola) ex gr. agrestis) are some of the most frequent micromammals in modern and fossil assemblages of the region. The phylogeny of this group has been under continuous revisions, leading to the recognition of three different species under similar morphological features (Paup´ erio et al., 2012; Wilson et al., 2017). At the time of the formation of this paleontological site, the northern and southern lineages of the Mi. (Ag.) agrestis group (Mi. (Ag.) lavernedii, Mi. (Ag.) rozianus and Mi. (Ag.) agrestis) had already split since, according to Table 2 Micromammal species from Punta Lucero and their weighted habitat-preferences. Level F Level D Level C-bottom Level C-middle Level C-upper Level B Habitat type (Sample z: 445–420) (Sample z: 296–288) (Sample z: 280–269) (Sample z: 222–208) (Sample z: 208–177) (Sample z: 0–93) NISP i MNI %MNI NISP i MNI %MNI NISP i MNI %MNI NISP i MNI %MNI NISP i MNI %MNI NISP i MNI %MNI Wa OD OH OW Wo Arvicola sapidus 6 4 0.4 4 3 21.4 2 1 1.7 1 Microtus (Agricola) lavernedii 746 389 35.6 8 6 42.9 41 21 18.6 6 4 7.8 11 7 19.4 47 24 40.0 0.5 0.5 Microtus (Microtus) arvalis 1 1 7.1 11 6 5.3 2 1 2.0 1 1 2.8 13 7 11.7 0.75 0.25 Microtus (Terricola) sp. 266 21 1.9 1 1 7.1 4 2 1.8 1 1 2.0 0.5 0.5 Mi. (Te.) ex gr. lusitanicus-duodecimcostatus 176 88 8.1 1 1 1.7 0.5 0.5 Mi. (Te.) pyrenaicus 57 32 2.9 0.5 0.5 Clethrionomys glareolus 16 11 1.0 2 1 0.9 1 Mus musculus 2 1 – 2 1 – – – – – – Apodemus (Sylvaemus) sylvaticus 315 161 14.7 2 2 14.3 17 10 8.8 7 4 7.8 1 1 2.8 5 4 6.7 1 Micromys minutus 10 7 0.6 1 Sorex minutus 35 20 1.8 0.25 0.75 Sorex coronatus 532 274 25.1 7 5 4.4 7 4 7.8 1 1 2.8 11 5 8.3 0.75 0.25 Neomys fodiens niethammeri 32 16 1.5 0.75 0.25 Neomys anomalus 2 2 1.8 1 1 2.8 0.25 0.75 Crocidura russula 15 9 0.8 112 58 51.3 62 35 68.6 45 24 66.7 24 13 21.7 0.75 0.25 Crocidura gueldenstaedtii 96 49 4.5 0.5 0.5 Talpa aquitania 20 10 0.9 1 1 7.1 10 8 7.1 3 2 3.9 1 1 2.8 5 4 6.7 0.75 0.25 Erinaceus europaeus 1 1 0.1 1 1 1.7 0.25 0.25 0.5 Total 2325 1093 100 17 14 206 113 90 52 61 36 103 60 A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 8 Paup´ erio et al. (2012), they did at around the Last Glacial Maximum (LGM). The frequency of M1 with a postero-lingual triangle, like the one observed in the M2, is a discriminant feature among Mi. (Ag.) ex gr. agrestis species (Krapp and Niethammer, 1982; Wilson et al., 2017). The low proportion of M1 presenting this triangle (1.5%: Table S3.1) has allowed us to ascribe the field-vole specimens from PL-III to the Mediterranean field vole (Mi. (Ag.) lavernedii), which is the current representative of this group in the region. Nowadays, Mi. (Ag.) lavernedii spreads throughout the Cantabrian Region (Palomo et al., 2007). Conversely, Microtus (Microtus) arvalis (common vole), a species of more Fig. 3. Selected arvicoline specimens from Punta Lucero III. Arvicola sapidus: left m1–m3 from Level F in occlusal (a) view. Microtus (Agricola) lavernedii: left m1–m3 from Level F in occlusal (b) view and left M1–M3 from Level C in occlusal (c) view. Microtus (Microtus) arvalis: left m1–m3 from Level C in occlusal (d) view, and left M1–M3 from Level C in occlusal (e) view. Microtus (Terricola) sp.: left m1 from Level F in occlusal (f) view, and left m1–m3 from Level F in occlusal (g) view. Microtus (Terricola) ex gr. lusitanicus–duodecimcostatus: left M3 from Level F in occlusal (h) view and left M3 from Level F in occlusal (i) view. Microtus (Terricola) pyrenaicus: left M3 from Level F in occlusal (j) view and left M3 from Level F in occlusal (k) view. Clethrionomys glareolus: left m1–m3 from Level F in occlusal (l) view. AL: anterior loop. T: triangle. Scale bar 1 mm. A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 9 continental requirements (Nores, 1989), lives south of the watershed, in the northern half of the Iberian Plateau (Nores, 1989; Palomo et al., 2007). Concerning proportion among these species, in Level C, a decrease in the abundance of Mi. (Ag.) lavernedii is observed along with the appearance of Mi. (Mi.) arvalis. However, Mi. (Ag.) lavernedii prevails at Level B, coinciding with the highest abundance ratio of Mi. (Mi.) arvalis in the sequence. It can also be seen that when Mi. (Ag.) lavernedii increases, also does Arvicola sapidus, a temperate-affinity water vole from southwestern Europe. This pattern, in which the initial absence of Mi. (Mi.) arvalis is followed by its increasing abundance, is also observed at La Güelga (Asturias, ~48–37 ka, ´ Alvarez-Vena et al., 2021), an MIS 3 site in which an alternation of climatic changes from temperate to cold stadials was reported in a scenario of increasing aridity. Regarding the remnant vole taxa from PL-III is also remarkable the occurrence of at least two species of the Terricola subgenus: Mi. (Te.) pyrenaicus (the Pyrenean pine vole) and Mi. (Te.) ex gr. lusitanicus–duodecimcostatus. Observed morphotypes of the M1 point to the presence of both Mediterranean (Mi. (Te.) duodecimcostatus) and Lusitanian (Mi. (Te.) lusitanicus) pine voles but overlapping morphological variability did not make us possible to discriminate between these species, which was even more difficult due to the co-occurrence of the Pyrenean pine vole. For this reason, the total MNI of the Terricola subgenus has been calculated based on the m1. However, the proportion of individuals of each species belonging to this subgenus has been Fig. 4. Selected murid, soricid, and talpid specimens from Punta Lucero III. Micromys minutus: left mandible from Level F with m1 in lingual (a) and occlusal (a’) views. Mus musculus: Left maxilla fragment from Level F with M1–M2 in occlusal (b) view. Apodemus (Sylvaemus) sylvaticus: left M1–M3 series from Level F in occlusal (c) view. Neomys fodiens niethammeri: right mandible from Level F in posterior (d) and labial (d’) views. Neomys anomalus: right mandible from Level C in posterior (e) and labial (e’) views. Sorex coronatus: right mandible from Level F in posterior (f) and labial (f’) views. Sorex minutus: right mandible from Level F in posterior (g) and labial (g’) views. Crocidura gueldenstaedtii: right mandible from Level F in posterior (h) and labial (h’) views. Crocidura russula: right mandible from Level C in posterior (i) and labial (i’) views. Talpa aquitania: left humerus from Level C in posterior (j) view. Analogous anatomical elements have been flipped horizontally to show them from the same side for comparison purposes. Scale bars 1 mm. A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 16 From ~5 ka, steppe pastoralists of the Yamnaya culture expanded eastand westward, linking Asia and Europe (Allentoft et al., 2015; Lazaridis et al., 2022). This created a geographic corridor which, aided by the development of horse-riding and chariotry (Anthony, 2007; Anthony and Brown, 2011; Wilkin et al., 2021), led to the dispersal of crops, herds, and commensal species from one continent to another. Regarding the Iberian Peninsula, an increasing gene inflow from Northern and Central European human populations with steppe ancestry has been detected (Olalde et al., 2019; Villalba-Mouco et al., 2021; Patterson et al., 2022), dating the earliest evidence from ~4.5–4 ka, from individuals who coexisted with locals without steppe ancestry (Olalde et al., 2019). This incoming pastoralist peopling to the region agrees with the increasing anthropic pressure linked to animal husbandry observed at PL-III and the coeval records mentioned. However, the increase in domestic herds and deforestation is not the only way those migrations are perceived in the PL-III deposit. The harvest mouse (Micromys minutus), identified at Level F, and the house mouse (Mus musculus), found at levels F and C, are reported from PL-III at a significantly early chronology. According to Hor´ aˇ cek et al. (2013), Pliocene and Pleistocene European records previously ascribed to the Micromys genus belong to Parapodemus coronensis Schaub, 1930, which did not have continuity in the Late Pleistocene. This agrees with the work of Fig. 11. Temperature and rainfall estimations based on the Mutual Ecogeographic Range method. (A) Box and whisker plot of the monthly values obtained through the Mutual Ecogeographic Range method (Appendix 4) for each micromammal association of the Punta Lucero III sequence (levels F, C, and B). (B) Monthly mean values of estimated temperature and rainfall from levels F, C, and B compared with present-day data from the Punta Lucero III location. (C) Mutual Ecogeographic Range method estimations: mean annual temperature (MAT), mean temperature of the warmest month (MTW), mean temperature of the coldest month (MTC), mean annual precipitation (MAP). A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 17 Yasuda et al. (2005), who, based on the phylogeographic pattern of mtDNA, propose that the extant species originated in East Asia. The fossil record suggests that Micromys minutus reached Europe during the Holocene (e.g., Mistrot, 2011, 2013a, 2013b; Serjeantson, 2011; Krajcarz et al., 2020; Royer et al., 2021), but the scarcity of reliable records to date does not allow us to reconstruct a precise timing. Concerning the Iberian Peninsula, the species has been identified at two different sites: Level 2 of Valdavara-1 (Lugo, Galicia; 5305–4978 cal yr BP; L´ opez-García et al., 2011) and Level 1 of Amalda (Gipuzkoa, Basque Country; Pem´ an, 1990), which yielded three dates ranging from 3875 to 1179 cal kyr BP (the oldest considered aberrant by Altuna, 1990). Considering the measurements published in L´ opez-García et al. (2011), the material ascribed to Micromys minutus in Valdavara-1 falls in size with small specimens of Apodemus sylvaticus (Fig. 5A), which could indicate that it is this species and not the previous one, in the absence of reviewing the identified material. This means that the reported in this work could be the oldest record of this species in the Iberian Peninsula. The spread of the harvest mouse across Eurasia was probably boosted by the development of agriculture, especially by millet crops, which, conversely to other cereals, do not belong to the first set of Neolithic crops from the Middle East. Instead, this cereal was initially domesticated in north-eastern China at ~8 ka (Liu et al., 2004; Crawford et al., 2013) and later spread to the rest of Central Eurasia and Eastern Europe (Miller et al., 2016; Filipovi´ c et al., 2020). Foxtail millet (Setaria italica), among other remains of cereals, has been found at the nearby sites of Kobaederra (Level 1) and Arenaza (Level 9) at 5280–4855 cal yr BP and 4085–3693 cal yr BP respectively (Zapata Pe˜ na, 1999, 2002). Therefore, the harvest mouse occurrence at PL-III (Level F: 4402–4097 cal yr BP) and the arrival of millet crops to the region share similar chronological ranges. The earliest evidence of millet consumption by humans in the Cantabrian Region (3163–3051 cal yr BP) has been documented at El Espinoso (Asturias; Gonz´ alez-Rabanal et al., 2022) at ~100 km west of PL-III by individuals with a significant proportion of steppe ancestry (30%, Patterson et al., 2022). Therefore, steppe migrations could be related to harvest mouse arrival in the Cantabrian Region. About the house mouse (Mus musculus), its record in southwestern Europe has been reviewed by Cucchi et al. (2005) and Domínguez García et al. (2019), who found that most of the reports of the species lack published reliable identifications or come from unclear contexts. Even new data from the Cantabrian Region published by Martínez-Villa et al. (2022) and Cernadas-Garrido et al. (2023) may correspond to intrusive specimens and materials of mixed–chronology, respectively. The earliest evidence of Mus musculus domesticus in Iberia comes from Iron Age at the Alorda Parc site (Valenzuela-Lamas et al., 2011). Domínguez García et al. (2019) also place the earliest reliable presence of the house mouse at the Iberian Peninsula within the Iron Age, at ~3 ka, supporting the hypothesis of a dispersal linked to the increase in the sea trade between eastern and western Mediterranean (Cucchi et al., 2005), driven by the commercial expansion of the Phoenicians and Greeks (Cucchi et al., 2012). The oldest records of the house mouse (Mus musculus musculus) from inland Europe come from Eastern Europe and date from the end of the Neolithic (~6.5 ka; Cucchi et al., 2011, 2020). Based on this unexpected finding, Cucchi et al. (2012) proposed a centre of synanthropization for this subspecies in eastern Europe north of the Black Sea. Therefore, a route from the steppes through inland Europe could be an alternative pathway for the arrival of Mus musculus to the Cantabrian Region. This is consistent with the reliable finding of Mus musculus ssp. in northern Italy (Tosina, Milan) dated at ~6 ka (Bona, 2020). However, further research and direct radiocarbon dating of the PL-III specimens need to be done to avoid misinterpretation of this record. Fig. 12. Compared paleoclimatic and paleoenvironmental reconstructions of the Punta Lucero III sequence. (A) Mutual Ecogeographic Range estimations of levels F, C, and B: mean annual precipitation (MAP), mean temperature of the warmest month (MTW), mean annual temperature (MAT), mean temperature of the coldest month (MTC). Dash lines and asterisk columns indicate the presentday values obtained from the Iberian Climate Atlas (Couto et al., 2011). (B) Type of landscape evolution at the surroundings of the Punta Lucero III shaft inferred through the Habitat Weighting method. Levels and samples are plotted according to depth. The column with an asterisk corresponds to a modern micromammal sample collected in the Punta Lucero area from barn owl regurgitation pellets by Gonz´ alez Oreja et al. (1993). Table 5 Proportion (%) of skeletal remains showing carnivore tooth marks. Level A Level B Level C Level D Level E Level F Total NSP* 46 188 414 989 707 876 3220 Tooth-marked specimens 0 17 51 45 42 46 201 Frequency (%) 0 9 12.3 4.6 5.9 5.3 6.2 * Number of specimens (includes specimens that cannot be identified to taxon). A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 18 6. Conclusions The mammal assemblage studied in this work was mainly produced by accidental falling in the PL-III cave, which acted as a natural trap through time. Nocturnal birds of prey could also be implied in the accumulation of the micromammal assemblage since our analyses did not allow us to rule out these predators. The studied samples provided a large number of remains corresponding to 18 micromammal and nine macromammal taxa. Micromammal community underwent a quick turnover which reflects a shift of the environmental conditions that took place during the formation of the deposit: the moist mosaic of forest and meadows observed in Level F (4,402–4097 cal yr BP) turned into an open and shrubbier grassland during the accumulation of levels C (2,837–2723 cal yr BP) and B (1,992–1830 cal yr BP). The earliest Iberian records of Mus musculus and, most likely, of Micromys minutus here reported could be related to the dispersal of the steppe pastoralist. Still, direct radiocarbon dating needs to be done to obtain a more precise date for these remains. Simultaneously, macromammals experienced a shift from wild populations to domestic herds, which, along with the abundant charcoal fragments recovered in the sediment, suggest that anthropogenic fires for the development of livestock pastures caused forest disappearance and did not allow them to recover. Temperature and rainfall estimations reveal that the humid and mild climatic conditions observed at the bottom of the sequence, which occurred during the Late Holocene Thermal Maximum, went through a cooling and aridification phase, coeval with the Iron Age Cold Epoch, and concluded in a slight warming, coeval with the Roman Warm Period. Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Data availability No data was used for the research described in the article. Acknowledgements AB.M-A. developed part of this research as part of the ERC Consolidator Grant (SUBSILIENCE ref. 818299). We thank J.A. Delgado for his technical work on studying macromammal assemblage. Financial support was provided by the Bilbao Port Authority (Autoridad Portuaria de Bilbao) within the project “Estabilizaci´ on del sector occidental de la Cantera de Punta Lucero en el Puerto de Bilbao”. We are also grateful to Juan Manuel L´ opez-García and the anonymous reviewer for their suggestions and comments that strongly improved the manuscript. Appendix A. Supplementary data Supplementary data to this article can be found online at https://doi. org/10.1016/j.palaeo.2023.111476. References Allentoft, M., Sikora, M., Sj¨ ogren, K.G., et al., 2015. Population genomics of Bronze Age Eurasia. Nature 522, 167–172. Alley, R.B., Mayewski, P.A., Sowers, T., Stuiver, M., Taylor, K.C., Clark, P.U., 1997. Holocene climatic instability: a prominent, widespread event 8200 yr ago. Geology 25, 483–486. Altuna, J., 1980. Historia de la domesticaci´ on animal, en el País Vasco, desde sus orígenes hasta la romanizaci´ on. Munibe 30, 1–163. Altuna, J., 1990. Situaci´ on y descripci´ on de la cueva de Amalda. Historia de las excavaciones. Descripci´ on del relleno. Estructuras en el yacimiento. Dataciones de radiocarbono. Otros yacimientos del valle. In: La cueva de Amalda (Zestoa, País Vasco): ocupaciones paleolíticas y postpaleolíticas, pp. 9–31. Colecci´ on Barandiaran (4). Eusko Ikaskuntza. Altuna, J., Arias Cabal, P., 1999. Nuevas dataciones absolutas Para el Neolítico de la Cueva de Arenaza (Bizkaia). Munibe (Antropologia-Arkeologia) 51, 161–171. Altuna, J., Mariezkurrena, K., 2012. Macromammalian remains from the Holocene levels of El Mir´ on Cave. In: Straus, L.G., Gonz´ alez Morales, M. (Eds.), El Mir´ on Cave, Cantabrian Spain. University of New Mexico Press, Albuquerque, pp. 288–318. ´ Alvarez-Alonso, D., ´ Alvarez-Fern´ Andez, E., de Andr´ es-Herrero, M., Ballesteros, D., García-Ibaibarriaga, N., Jim´ enez-S´ Anchez, M., Jord´ A-Pardo, J.F., Yravedra, J., 2018. Excavaciones en la Cueva del Olivo (Pruvia, Llanera). Campa˜ nas 2013–2016. Excavaciones Arqueol´ ogicas en Asturias 2013–2016, 121–132. ´ Alvarez-Lao, D.J., 2014. The Jou Puerta cave (Asturias, NW Spain): a MIS 3 large mammal assemblage with mixture of cold and temperate elements. Palaeogeogr. Palaeoclimatol. Palaeoecol. 393, 1–19. ´ Alvarez-Lao, D.J., ´ Alvarez-Vena, A., Ballesteros, D., García, N., Laplana, C., 2020. A cave lion (Panthera spelaea) skeleton from Torca del Le´ on (NW Iberia): micromammals indicate a temperate and forest environment corresponding to GI-11 (MIS 3). Quat. Sci. Rev. 229, 106123. ´ Alvarez-Vena, A., ´ Alvarez-Lao, D.J., Laplana, C., Quesada, J.M., Rojo, J., GarcíaS´ anchez, E., Men´ endez, M., 2021. Environmental context for the late Pleistocene Table 6 Weighted (%) habitat-preferences a with 95% confidence intervals b of the small mammal assemblage from each level of Punta Lucero. MNI Wa OD OH OW Wo Zierbena modern (Gonz´ alez-Oreja et al., 1993) 773 0.00 36.58 13.71 49.61 0.10 95% CI 0.00–0.48 33.21–40.12 11.37–16.34 45.97–53.13 0.00–0.72 Level B (Sample z: 0–93) 60 1.67 25.00 32.50 40.00 0.83 95% CI 0.04–8.94 14.72–37.86 21.69–46.69 27.56–53.46 0.00–5.96 Level C-upper (Sample z: 208–177) 36 0.69 52.08 15.97 31.25 0.00 95% CI 0.00–9.74 35.49–69.59 6.37–32.81 16.35–48.11 0.00–9.74 Level C-middle (Sample z: 222–208) 51 0.00 52.94 13.73 33.33 0.00 95% CI 0.00–6.98 38.46–67.07 5.70–26.26 20.76–47.92 0.00–6.98 Level C-bottom (Sample z: 280–269) 113 0.44 42.48 20.13 36.06 0.88 95% CI 0.00–3.21 33.23–52.13 13.36–28.96 27.45–45.86 0.02–4.83 Level D (Sample z: 296–288) 14 21.43 5.36 30.36 42.86 0.00 95% CI 4.66–50.80 0.18–33.87 8.39–58.10 17.66–71.14 0.00–23.16 Level F (Sample z: 445–420) 1092 1.47 0.62 47.50 47.99 2.43 95% CI 0.84–2.37 0.26–1.32 44.53–50.54 44.99–51.00 1.64–3.58 a Wa, water; OH, open humid; OD, open dry; OW, open woodland; Wo, woodland. b 95% CI: multiple proportions 95% confidence intervals calculated using the Clopper-Pearson method (Clopper and Pearson, 1934) using the PAST 4.0 software (Hammer et al., 2001). A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 19 (MIS 3) transition from neanderthals to early modern humans: analysis of small mammals from La Güelga Cave, Asturias, northern Spain. Palaeogeogr. Palaeoclimatol. Palaeoecol. 562, 110096. Andersen, K.K., Azuma, N., Barnola, J.M., et al., 2004. High resolution climate record of the Northern Hemisphere reaching into the last glacial interglacial period. Nature 431, 147–151. Andrews, P., 1990. Owls, Caves and Fossils. Predation, Preservation and Accumulation of Small Mammal Bones in Caves, With an Analysis of the Pleistocene Cave Faunas from Westbury-sub-Mendip. The University of Chicago Press, Somerset, UK, London. Andrews, P., 2006. Taphonomic effects of faunal impoverishment and faunal mixing. Palaeogeogr. Palaeoclimatol. Palaeoecol. 241, 572–589. Anthony, D.W., 2007. The Horse, the Wheel, and Language: How Bronze-Age Riders from the Eurasian Steppes Shaped the Modern World. Princeton University Press. Anthony, D.W., Brown, D.R., 2011. The secondary products revolution, horse-riding, and mounted warfare. J. World Prehist. 24, 131. Aranburu, A., Arriolabengoa, M., Iriarte, E., Giralt, S., Yusta, I., Martínez-Pillado, V., del Val, M., Moreno, J., Jim´ enez-S´ anchez, M., 2015. Karst landscape evolution in the littoral area of the Bay of Biscay (north Iberian Peninsula). Quat. Int. 364, 217–230. Arriolabengoa, M., Iriarte, E., Aranburu, A., Yusta, I., Arrizabalaga, A., 2015. Provenance study of endokarst fine sediments through mineralogical and geochemical data (Lezetxiki II cave, northern Iberia). Quat. Int. 364, 231–243. Arilla, M., Ruf` a, A., Rosell, J., Blasco, R., 2020. Small carnivores’ cave-dwelling: neotaphonomic study of a badger (Meles meles) sett and its archaeological implications. Hist. Biol. 32 (7), 951–965. Azorit, E., Analla, M., Carrasco, R., Calvo, J.A., Mu˜ noz-Cobo, J., 2002. Teeth eruption pattern in red deer (Cervus elaphus hispanicus) in southern Spain. Anales de Biología 24, 107–114. Baldini, L.M., Baldini, J.U.L., McDermott, F., Arias, P., Cueto, M., Fairchild, I.J., Hoffmann, D.L., Mattey, D.P., Müller, W., Nita, D.C., Onta˜ n´ on, R., García-Monc´ o, C., Richards, D.A., 2019. North Iberian temperature and rainfall seasonality over the Younger Dryas and Holocene. Quat. Sci. Rev. 226, 105998. Ballesteros, D., ´ Alvarez-Vena, A., Monod-Del Dago, M., Rodríguez-Rodríguez, L., Sanjurjo-S´ anchez, J., ´ Alvarez-Lao, D., P´ erez-Mejías, C., Valenzuela, P., DeFelipe, I., Laplana, C., Cheng, H., Jim´ enez-S´ anchez, M., 2020. Paleoenvironmental evolution of Picos de Europa (Spain) during marine isotopic stages 5c to 3 combining glacial reconstruction, cave sedimentology and paleontological findings. Quat. Sci. Rev. 248, 106581. Ba˜ nuls-Cardona, S., L´ opez-García, J.M., 2016. Climatic and environmental conditions from the Neolithic to the Bronze Age (7000–3000 BP) in the Iberian Peninsula assessed using small-mammal assemblages. Comptes Rendus Palevol 15, 958–967. Ba˜ nuls-Cardona, S., L´ opez-García, J.M., Verg` es, J.M., 2013. Palaeoenvironmental and palaeoclimatic approach of the middle bronze age (level MIR 4) from El Mirador Cave (Sierra de Atapuerca, Burgos, Spain). Quaternaire 24 (2), 217–223. Ba˜ nuls-Cardona, S., L´ opez-García, J.M., Morales Hidalgo, J.I., Cuenca-Besc´ os, G., Verg` es, J.M., 2017a. Lateglacial to late Holocene palaeoclimatic and palaeoenvironmental reconstruction of El Mirador cave (Sierra de Atapuerca, Burgos, Spain) using the small-mammal assemblages. Palaeogeogr. Palaeoclimatol. Palaeoecol. 471, 71–81. Ba˜ nuls-Cardona, S., Martín Rodríguez, P., L´ opez-García, J.M., Morales, J.I., CuencaBesc´ os, G., Verg` es, J.M., 2017b. Human impact on small-mammal diversity during the middleto late-Holocene in Iberia: the case of El Mirador cave (Sierra de Atapuerca, Burgos, Spain). The Holocene 27 (8), 1067–1077. Baraybar, J.P., de la Rúa, C., 1995. Anthropological study of the population of Pico Ramos (Muskiz, Biscay). Some considerations regarding demography, health and subsistence. In: Zapata, L. (Ed.), The Chalcolithic burial deposit of the cave Pico Ramos (Muskiz, Biskay). Munibe (Antropologia-Arkeologia) 47, pp. 151–175. Barnosky, A.D., 2008. Megafauna biomass tradeoff as a driver of quaternary and future extinctions. Proceedings of the National Academy of Sciences 105 (1), 11543–11548. Barnosky, A.D., 2013. Palaeontological evidence for defining the Anthropocene. Geol. Soc. Lond. Spec. Publ. 395 (1), 149–165. Barone, R., 1976. Anatomie compar´ ee des mammif` eres domestiques. Tome I Ost´ eologie, Vigot Fr´ eres Editeurs, París. Barti, L., 2006. Az ´ allkapcsi lyuk (Foramen mentale) helyzete mint kieg´ eszít´ o hat´ aroz´ ob´ elyeg a Neomys fajok (Mammalia, Insectivora Soricidae) biztosabb elkül¨ onít´ es´ ere. Acta Siculica 1, 191–199. Beck, H.E., Zimmermann, N.E., McVicar, T.R., Vergopolan, N., Berg, A., Wood, E.F., 2018. Present and future K¨ oppen-Geiger climate classification maps at 1-km resolution. Sci. Data 5, 180214. Biedma, L., Rom´ an, J., Calzada, J., Friis, G., Godoy, J.A., 2018. Phylogeography of Crocidura suaveolens (Mammalia: soricidae) in Iberia has been shaped by competitive exclusion by C. Russula. Biol. J. Linn. Soc. 123 (1), 81–95. Biedma, L., Calzada, J., Godoy, J.A., Rom´ an, J., 2020. Local habitat specialization as an evolutionary response to interspecific competition between two sympatric shrews. J. Mammal. 101 (1), 80–91. Binford, L.R., 1981. Bones. Ancient Men and Modern Myths, Studies in Archaeology. Academic Press, New York. Blain, H.-A., Bailon, S., Cuenca-Besc´ os, G., 2008. The Early-Middle Pleistocene palaeoenvironmental change based on the squamate reptile and amphibian proxies at the Gran Dolina site, Atapuerca, Spain. Palaeogeogr. Palaeoclimatol. Palaeoecol. 261, 177–192. Blain, H.-A., Bailon, S., Cuenca-Besc´ os, G., Arsuaga, J.L., Bermúdez de Castro, J.M., Carbonell, E., 2009. Long-term climate record inferred from early-middle Pleistocene amphibian and squamate reptile assemblages at the Gran Dolina Cave, Atapuerca, Spain. J. Hum. Evol. 56 (1), 55–65. Blain, H.-A., Lozano-Fern´ andez, I., Agustí, J., Bailon, S., Men´ endez, L.G., Espígares, P.O. M., Ros-Montoya, S., Jim´ enez, J.M.A., Toro-Moyano, I., Martínez-Navarro, B., Sala, R., 2016. Refining upon the climatic background of the Early Pleistocene hominid settlement in western Europe: Barranco Le´ on and Fuente Nueva-3 (GuadixBaza Basin, SE Spain). Quat. Sci. Rev. 144, 132–144. Boivin, N.L., Zeder, M.A., Fuller, D.Q., Crowther, A., Larson, G., Erlandson, J.M., Denham, T., Petraglia, M.D., 2016. Ecological consequences of human niche construction: Examining long-term anthropogenic shaping of global species distributions. Proc. Nat. Acad. Sci. 113 (23), 6388–6396. Bona, F., 2020. . Earliest evidence of Mus musculus ssp. in Western Europe during the Late Neolithic (Tosina, Mantova, Northern Italy): new insights on the house mice migratory waves. Hystrix Ital. J. Mammal. 31 (2), 111–116. Bond, G., Showers, W., Cheseby, M., Lotti, R., Almasi, P., deMenocal, P., Priore, P., Cullen, H., Hajdas, I., Bonani, G., 1997. A pervasive millennial-scale cycle in North Atlantic Holocene and glacial climates. Science 278 (5341), 1257–1266. Bond, G., Kromer, B., Beer, J., Muscheler, R., Evans, M., Showers, W., Hoffmann, S., Lotti-Bond, R., Hajdas, I., Bonani, G., 2001. Persistent solar influence on North Atlantic climate during the Holocene. Science 294, 2130–2136. Borzenkova, I., Zorita, E., Borisova, O., Kalnin¸a, L., Kisielien˙ e, D., Koff, T., Kuznetsov, D., Lemdahl, G., Sapelko, T., Stanˇ cikait˙ e, M., Subetto, D., 2015. Climate change during the Holocene (Past 12,000 Years). In: The BACC II Author Team, (Eds) Second Assessment of Climate Change for the Baltic Sea Basin. Regional Climate Studies. Springer, Cham, pp. 25–49. Bray, J.R., 1971. Solar–climate relationships in the post-Pleistocene. Science 171, 1242–1243. Bronk Ramsey, C., 2009. Bayesian analysis of radiocarbon dates. Radiocarbon 51 (1), 337–360. Büntgen, U., Tegel, W., Nicolussi, K., McCormick, M., Frank, D., Trouet, V., Kaplan, J.O., Herzig, F., Heussner, K.-U., Wanner, H., Luterbacher, J., Esper, J., 2011. 2500 years of European climate variability and human susceptibility. Science 331 (6017), 578–582. Carracedo, V., Cunill, R., García-Codron, J.C., P` elachs, A., P´ erez-Obiol, R., Soriano, J.M., 2018. History of fires and vegetation since the Neolithic in the Cantabrian Mountains (Spain). Land Degrad. Dev. 29 (7), 2060–2072. Carri´ on, J.S., Fern´ andez, S., Gonz´ alez-Samp´ eriz, P., Gil-Romera, G., Badal, E., Carri´ onMarco, Y., L´ opez-Merino, L., L´ opez-S´ aez, J.A., Fierro, E., Burjachs, F., 2010. Expected trends and surprises in the Lateglacial and Holocene vegetation history of the Iberian Peninsula and Balearic Islands. Rev. Palaeobot. Palynol. 162, 458–475. Casta˜ nos, P., 1997. Estudio arqueozool´ ogico de la fauna de Pe˜ na Larga (Crip´ an, ´ Alava). In: Fern´ andez Eraso, J. (Ed.), Pe˜ na Larga: memoria de las excavaciones arqueol´ ogicas 1985–1989. Memorias de yacimientos alaveses. n◦4. Diputaci´ on Foral de ´ Alava, Vitoria-Gasteiz, pp. 127–134. Cernadas-Garrido, A., ´ Alvarez-Vena, A., ´ Alvarez-Lao, D.J., 2023. Palaeobiological study of the Holocene micromammal assemblage from the Cueva del Hueso (Castrill´ on, Asturias, NW Spain). Spanish J. Palaeontol. 38 (1) https://doi.org/10.7203/ sjp.25962 in press. Clopper, C.J., Pearson, E.S., 1934. The use of confidence or fiducial limits illustrated in the case of the binomial. Biometrika 26 (4), 404–413. Couto, M.A., S´ anchez, G., Tavares, C.D., Barcel´ o, A.M., Nunes, L.F., Herr´ aez, C.F., Pires, V., Marques, J., Mendes, L., Chazarra, A., Cunha, S., Mendes, M., Neto, J., Silva, A., 2011. Atlas Clim´ atico Ib´ erico-Iberian Climate Atlas. Instituto de Meteorologia de Portugal and Agencia Estatal de Meteorología, Ministerio de Medio Ambiente y Medio Rural y Marino (Eds.). Crawford, G.W., Chen, X., Luan, F., 2013. A preliminary analysis of plant remains from the Yuezhuang site, Changqing district, Jinan, Shandong province. Jianghan Kaogu (2), 107–113. Crutzen, P.J., 2002. Geology of mankind. Nature 415, 23. Cucchi, T., Vigne, J.-D., Auffray, J.-C., 2005. First occurrence of the house mouse (Mus musculus domesticus Schwarz & Schwarz, 1943) in the Western Mediterranean: a zooarchaeological revision of subfossil occurrences. Biol. J. Linn. Soc. 84, 429–445. Cucchi, T., B˘ al˘ as¸escu, A., Bem, C., Radu, V., Vigne, J.-D., Tresset, A., 2011. New insights into the invasive process of the eastern house mouse (Mus musculus musculus): evidence from the burnt houses of Chalcolithic Romania. The Holocene 21 (8), 1195–1202. Cucchi, T., Auffray, J.-C., Vigne, J.-D., 2012. On the origin of the house mouse synanthropy and dispersal in the Near East and Europe: zooarchaeological review and perspectives. In: Machol´ an, M., Baird, S.J.E., Munclinger, P., Pi´ alek, J. (Eds.), Evolution of the House Mouse. Cambridge Studies in Morphology and Molecules: New Paradigms in Evolutionary Biology. Cambridge University Press, Cambridge, pp. 65–93. Cucchi, T., Papayianni, K., Cersoy, S., et al., 2020. Tracking the near Eastern origins and European dispersal of the western house mouse. Sci. Rep. 10, 8276. Cuenca-Besc´ os, G., Straus, L.G., Gonz´ alez Morales, M., García Pimineta, J.C., 2009. The reconstruction of past environments through small mammals: from the Mousterian to Bronze Age in El Mir´ on cave (Cantabria, Spain). J. Archaeol. Sci. 36, 947–955. Cuenca-Besc´ os, G., Marín-Arroyo, A.B., Martínez, I., Gonz´ alez Morales, M.R., Straus, L. G., 2012. Relationship between Magdalenian subsistence and environmental change: the mammalian evidence from El Mir´ on (Spain). Quat. Int. 272–273, 125–137. Dansgaard, W., Johnsen, S.J., Clausen, H.B., Dahl-Jensen, D., Gundestrup, N.S., Hammer, C.U., Hvidberg, C.S., Steffensen, J.P., Sveinbj¨ ornsdottir, A.E., Jouzel, J., Bond, G., 1993. Evidence for general instability of past climate from a 250-kyr icecore record. Nature 364, 218–220. Darviche, D., Orth, A., Michaux, J., 2006. Mus spretus et M. Musculus (Rodentia, Mammalia) en zone m´ editerran´ eenne: diff´ erenciation biom´ etrique et morphologique: application ` a des fossiles marocains pl´ eistoc` enes. Mammalia 70 (1–2), 90–97. A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 20 deMenocal, P., Ortiz, J., Guilderson, T., Adkins, J., Sarnthein, M., Baker, L., Yarusinsky, M., 2000. Abrupt onset and termination of the African humid period: rapid climate responses to gradual insolation forcing. Quat. Sci. Rev. 19, 347–361. deMenocal, P.B., 2001. Cultural responses to climate Change during the Late Holocene. Science 292 (5517), 667–673. Domínguez García, ´ A.C., Laplana, C., Sevilla, P., Blain, H.-A., Palomares Zumajo, N., de Lugo, Benítez, Enrich, L., 2019. New data on the introduction and dispersal process of small mammals in southwestern Europe during the Holocene: Castillejo del Bonete site (southeastern Spain). Quat. Sci. Rev. 225, 106008. Domínguez García, ´ A.C., Laplana, C., Sevilla, P., 2020. Early reliable evidence of the Etruscan shrew (Suncus etruscus) in southwestern Europe during ancient times. Reconstructing its dispersal process along the Mediterranean Basin. Quat. Sci. Rev. 250, 106690. Domínguez-García, ´ A.C., Laplana, C., Sevilla, P., ´ Alvarez-Vena, A., Collado Giraldo, H., 2022. Small mammals of the Holocene sequence of Postes Cave (SW Spain): biogeographic and palaeoenvironmental implications for southwestern Iberia. Hist. Biol. https://doi.org/10.1080/08912963.2022.2045981. Doughty, C.E., Wolf, A., Field, C.B., 2010. Biophysical feedbacks between the Pleistocene megafauna extinction and climate: the first human-induced global warming? Geophys. Res. Lett. 37, L15703. Ellis, E.C., 2011. Anthropogenic transformation of the terrestrial biosphere. Phil. Trans. R. Soc. A 369, 1010–1035. Evans, E.M.N., Van Couvering, J.A.H., Andrews, P., 1981. Paleoecology of Miocene sites in Western Kenya. J. Hum. Evol. 10, 99–116. Fagoaga, A., Blain, H.-A., Marquina-Blasco, R., Laplana, C., Sillero, N., Hern´ andez, C.M., Mallol, C., Galv´ an, B., Ruiz-S´ anchez, F.J., 2019. Improving the accuracy of small vertebrate-based palaeoclimatic reconstructions derived from the Mutual Ecogeographic Range. A case study using geographic information systems and UDAODA discrimination methodology. Quat. Sci. Rev. 223, 105969. Fern´ andez-Jalvo, Y., Andrews, P., Denys, C., Ses´ e, C., Stoetzel, E., Marin-Monfort, D., Pesquero, D., 2016. Taphonomy for taxonomists: implications of predation in small mammal studies. Quat. Sci. Rev. 139, 138–157. Filipovi´ c, D., Meadows, J., Dal Corso, M., et al., 2020. New AMS 14C dates track the arrival and spread of broomcorn millet cultivation and agricultural change in prehistoric Europe. Sci. Rep. 10, 13698. García-Morato, S., Marin-Monfort, D., Ba˜ nuls-Cardona, S., Cuenca-Besc´ os, G., Verg` es, J. M., Fern´ andez-Jalvo, Y., 2023. Solving a ‘puzzle’. The global 4.2 ka Bond Event at El Mirador cave (Sierra de Atapuerca, Burgos, Spain) and the importance of small mammal taphonomy to the interpretation of past environments and their climatic controls. The Holocene 33 (3), 296–309. García-Ruiz, J.M., 2010. The effects of land uses on soil erosion in Spain: a review. Catena 81, 1–11. Garrote, A., García, J., Mu˜ noz, L., Arriola, G., Eguiguren, E., García, I., 1993. Mapa geol´ ogico del País Vasco 1:25.000 Hoja 37-III Zierbena. Ente Vasco de la Energía. G´ omez-Olivencia, A., Cubas, M., Sala, N., Pantoja, A., Garcia-Ibaibarriaga, N., RiosGaraizar, J., Regalado, E., Libano, I., Solar, G., Arlegi, M., Moreno, J., 2015. Restos humanos calcolíticos de dos nuevos yacimientos de Punta Lucero (Zierbena, Bizkaia). Kobie Serie Paleoantropología 34, 5–18. Gonz´ alez Oreja, J.A., Lorenzo Rodolfo, J.C., P´ erez de Ana, J.M., 1993. Nota sobre la alimentaci´ on de la lechuza común en dos zonas de Vizcaya. Estudios del Museo de Ciencias Naturales de ´ Alava 8, 227–230. Gonz´ alez-Rabanal, B., Marín-Arroyo, A.B., Cristiani, E., Zupancich, A., Gonz´ alezMorales, M.R., 2022. The arrival of millets to the Atlantic coast of northern Iberia. Sci. Rep. 12, 18589. Guti´ errez, J., Aleix-Mata, G., Lamelas, L., Arroyo, M., Marchal, J.A., S´ anchez, A., 2019. Karyotype analysis of the new Talpa species Talpa Aquitania (Talpidae; Insectivora) from Northern Spain. Cytogenet. Genome Res. 159 (1), 26–31. Hammer, Ø., Harper, D.A.T., Ryan, P.D., 2001. PAST: Paleontological statistics software package for education and data analysis. Palaeontol. Electron. 4 (1), 1–9. Hillson, S., 2005. Teeth. In: Manuals in Archaeology. Cambridge University Press. Hor´ aˇ cek, I., Knitlov´ a, M., Wagner, J., Kordos, L., Nadachowski, A., 2013. Late Cenozoic history of the Genus Micromys (Mammalia, Rodentia) in Central Europe. PLoS ONE 8 (5), e62498. Iriarte, M.J., 1994. Estudio palinol´ ogico del nivel sepulcral del yacimiento arqueol´ ogico de Pico Ramos (Muskiz, Bizkaia). Cuadernos de Secci´ on. Prehistoria-Arqueología 5, 161–179. Iriarte, M.J., Mujika, J., Tarri˜ no, A., 2005. Herriko Barra (Zarautz-Gipuzkoa): caract´ erisation industrielle et ´ economique des premiers groupes de producteurs sur le littoral basque. In: Marchand, G., Tresset, A. (Eds.), Actas del Colloque Unit´ e et diversit´ e des processus de n´ eolithisation sur la façade atlantique de l'Europe. Bulletin de la Soci´ et´ e Pr´ ehistorique Française XXXVI, pp. 127–136. Jalut, G., Amat, A.E., Bonnet, L., Gauquelin, T., Fontugne, M., 2000. Holocene climatic changes in the Western Mediterranean, from south-East France to south-East Spain. Palaeogeogr. Palaeoclimatol. Palaeoecol. 160 (3–4), 255–290. Jones, J.R., Richards, M.P., Reade, H., Bernaldo de Quir´ os, F., Marín-Arroyo, A.B., 2019. Multi-Isotope investigations of ungulate bones and teeth from El Castillo and Covalejos caves (Cantabria, Spain): Implications for paleoenvironment reconstructions across the Middle-Upper Palaeolithic transition. J. Archaeol. Sci. Rep. 23, 1029–1042. Jones, J.R., Marín-Arroyo, A.B., Corch´ on Rodríguez, M.S., Richards, M.P., 2021. After the Last Glacial Maximum in the refugium of northern Iberia: environmental shifts, demographic pressure and changing economic strategies at Las Caldas Cave (Asturias, Spain). Quat. Sci. Rev. 262, 106931. Knitlov´ a, M., Hor´ aˇ cek, I., 2017. Late Pleistocene-Holocene paleobiogeography of the genus Apodemus in Central Europe. PLoS One 12 (3), e0173668. Krajcarz, M.T., Szymanek, M., Krajcarz, M., Pereswiet-Soltan, A., Alexandrowicz, W.P., Sudoł-Procyk, M., 2020. Shelter in Smollen III - a unique example of stratified Holocene clastic cave sediments in Central Europe, a lithostratigraphic stratotype and a record of regional paleoecology. PLOS ONE 15 (2), e0228546. Krapp, F., Niethammer, J., 1982. Microtus agrestis (Linnaeus, 1761) – Erdmaus. In: Niethammer, J., Krapp, F. (Eds.), Handbuch der S¨ augetiere Europas. Band 2/1. Nagetiere II. Akademische Verlagsgesellschaft, Wiesbaden, pp. 349–373. Kryˇ stufek, B., Shenbrot, G.I., 2022. Voles and Lemmings (Arvicolinae) of the Palaearctic Region. University Press, Maribor. https://doi.org/10.18690/um.fnm.2.2022. Kryˇ stufek, B., Tesakov, A.S., Lebedev, V.S., Bannikova, A.A., Abramson, N.I., Shenbrot, G., 2020. Back to the future: the proper name for red-backed voles is Clethrionomys Tilesius and not Myodes Pallas. Mammalia 84 (2), 214–217. Laskar, J., Robutel, P., Joutel, F., Gastineau, M., Correia, A.C.M., Levrard, B., 2004. A long-term numerical solution for the insolation quantities of the Earth. Astronomy Astrophys. 428 (1), 261–285. Lazaridis, I., Alpaslan-Roodenberg, S., Acar, A., et al., 2022. The genetic history of the Southern Arc: a bridge between West Asia and Europe. Science 377 (6609). Liu, C., Kong, Z., Lang, S.D., 2004. Dadiwan yizhi nongye zhiwu yicun yu renlei shengcun huanjing de tantao (A discussion on agricultural and botanical remains and the human ecology of Dadiwan site). Zhongyuan Wenwu (Cultural Relics Central China) 4, 25–29. Llorente, L., 2010. The Hares from Cova Fosca (Castell´ on, Spain). Archaeofauna 19, 59–97. L´ opez-Fuster, M.J., Ventura, J., Miralles, M., Casti´ en, E., 1990. Craniometrical characteristics of Neomys fodiens (Pennant, 1771) (Mammalia, Insectívora) from the northeastern Iberian Peninsula. Acta Theriol. 35 (3–4), 269–276. L´ opez-García, J.M., 2008. Evoluci´ on de la diversidad taxon´ omica de los micromamíferos en la Península Ib´ erica y cambios Paleoambientales durante el Pleistoceno Superior. Unpublished Ph.D. Universitat Rovira i Virgili, Tarragona. L´ opez-García, J.M., Blain, H.-A., Cuenca-Besc´ os, G., Alonso, C., Alonso, S., Vaquero, M., 2011. Small vertebrates (Amphibia, Squamata, Mammalia) from the late PleistoceneHolocene of the Valdavara-1 cave (Galicia, northwestern Spain). Geobios 44, 253–269. L´ opez-García, J.M., Blain, H.-A., Morales, J.I., Lorenzo, C., Ba˜ nuls-Cardona, S., CuencaBesc´ os, G., 2013. Small-mammal diversity in Spain during the late Pleistocene to early Holocene: climate, landscape, and human impact. Geology 41 (2), 267–270. L´ opez-García, J.M., Berto, C., Colamussi, V., Dalla Valle, C., Lo Vetro, D., Luzi, E., Malavasi, G., Martini, F., Sala, B., 2014. Palaeoenvironmental and palaeoclimatic reconstruction of the lastest Pleistocene-Holocene sequence from Grotta del Romito (Calabria, southern Italy) using the small-mammal assemblages. Palaeogeogr. Palaeoclimatol. Palaeoecol. 409, 169–179. L´ opez Merino, L., 2009. Paleoambiente y antropizaci´ on en Asturias durante el Holoceno. PhD Thesis. Universidad Aut´ onoma de Madrid. Departamento de Ecología, Madrid. L´ opez-Merino, L., Martínez-Cortizas, A., L´ opez-S´ aez, J.A., 2010. Early agriculture and palaeoenvironmental history in the North of the Iberian Peninsula: a multi-proxy analysis of the Monte Areo mire (Asturias, Spain). J. Archaeol. Sci. 37, 1978–1988. Luzi, E., L´ opez-García, J.M., 2019. Patterns of variation in Microtus arvalis and Microtus agrestis populations from Middle to late Pleistocene in southwestern Europe. Hist. Biol. 31 (5), 535–543. Lyman, R.L., 1984. Bone density and differential survivorship of fossil classes. J. Anthropol. Archaeol. 3, 259–299. Mallye, J.-B., Cochard, D., Laroulandie, V., 2008. Accumulations osseuses en p´ eriph´ erie de terriers de petits carnivores : les stigmates de pr´ edation et de fr´ equentation. Annales Pal´ eontol. 94 (3), 187–208. Mariezkurrena, K., 1983. Contribuci´ on al conocimiento del desarrollo de la dentici´ on y el esqueleto postcraneal de Cervus elaphus. Munibe (Antropologia-Arkeologia) 35, 149–202. Mariezkurrena, K., 1990. Caza y Domesticaci´ on durante el Neolítico y Edad de los Metales en el País Vasco. Munibe (Antropologia-Arkeologia) 42, 241–252. Marín-Arroyo, A.B., 2010. Arqueozoología en el cant´ abrico oriental durante la transici´ on Pleistoceno/Holoceno. La cueva del Mir´ on, PubliCan, Ediciones de la Universidad de Cantabria, Santander. Martín-Chivelet, J., Mu˜ noz-García, M.B., Edwards, R.L., Turrero, M.J., Ortega, A.I., 2011. Land surface temperature changes in Northern Iberia since 4000 yr BP, based on δ13C of speleothems. Glob. Planet. Change 77 (1–2), 1–12. Martínez-Solano, I., Sanchiz, B., 2005. Anfibios y Reptiles del Pleistoceno medio de Ambrona. Zona Arqueol´ ogica 5, 232–239. Martínez-Villa, A., ´ Alvarez-Fern´ andez, E., ´ Alvarez-Vena, A., Arrojo, L., Ballesteros, D., Cubas, M., Drak, L., Llorente Rodríguez, L., Martín-Jarque, S., Gil, M., 2022. New insights into upper Palaeolithic and Mesolithic occupations in Les Pedroses cave (Asturies, North Spain). J. Archaeol. Sci. Rep. 45, 103592. Mayewski, P.A., Rohling, E.E., Stager, J.C., Karl´ en, W., Maasch, K.A., Meeker, L.D., Meyerson, E.A., Gasse, F., van Kreveld, S., Holmgren, K., Lee-Thorp, J., Rosqvist, G., Rack, F., Staubwasser, M., Schneider, R.R., Steig, E.J., 2004. Holocene climate variability. Quat. Res. 62 (3), 243–255. Miller, G.H., Geirsdottir, A., Zhong, Y., Larsen, D., Otto-Bliesner, B.L., Holland, M.M., Bailey, D.A., Refsnider, K.A., Lehman, S.J., Southon, J.R., Anderson, C., Bj¨ ornsson, H., Thordarson, T., 2012. Abrupt onset of the Little Ice Age triggered by volcanism and sustained by sea-ice/ocean feedbacks. Geophys. Res. Lett. 39, L02708. Miller, N.F., Spengler, R.N., Frachetti, M., 2016. Millet cultivation across Eurasia: origins, spread, and the influence of seasonal climate. The Holocene 26 (10), 1566–1575. Mistrot, V., 2011. Les micromammif` eres du site gallo-romain de Melun-Grüber (Seine-etMarne). Bulletin de l'Association des Naturalistes de la Vall´ ee du Loing 87 (3), 132–141. A. ´ Alvarez-Vena et al. Palaeogeography, Palaeoclimatology, Palaeoecology 616 (2023) 111476 21 Mistrot, V., 2013. Les micromammif` eres des couches 37 ` a 33 de la grotte du Gardon: une emprise agricole naissante?. In: Perrin T., Voruz J.-L. (dir.) La grotte du Gardon (Ain) Volume II. Du N´ eolithique moyen II au Bronze ancien (couches 46 ` a 33), pp. 353–356. Mistrot, V., 2013. Les micromammif` eres des couches 47 ` a 39 de la grotte du Gardon: une anthropisation du milieu tr` es limit´ ee.. In: Perrin T., Voruz J.-L. (dir.) La grotte du Gardon (Ain) Volume II. Du N´ eolithique moyen II au Bronze ancien (couches 46 ` a 33), pp. 229–232. Mocl´ an, A., Domínguez-García, A.C., Stoetzel, E., Cucchi, T., Sevilla, P., Laplana, C., 2023. Machine Learning interspecific identification of mouse first lower molars (genus Mus Linnaeus, 1758) and application to fossil remains from the Estrecho Cave (Spain). Quat. Sci. Rev. 299, 107877. Mu˜ noz Sobrino, C., Ramil-Rego, P., G´ omez-Orellana, L., Díaz Varela, R.A., 2005. Palynological data on major Holocene climatic events in NW Iberia. Boreas 34, 381–400. Nadachowski, A., 1984. Taxonomic value of anteroconid measurements of M1 in common and field voles. Acta Theriol. 29 (10), 123–143. Nicolas, V., Martínez-Vargas, J., Hugot, J.-P., 2015. Talpa aquitania nov. sp. (Talpidae, Soricomorpha) a new mole species from Southwest France and North Spain. Bulletin de l’Acad´ emie V´ et´ erinaire de France 168 (4), 329–334. Nicolas, V., Martínez-Vargas, J., Hugot, J.-P., 2017. Talpa aquitania sp. nov. (Talpidae, Soricomorpha), a new mole species from SW France and N Spain. Mammalia 81 (6), 641–642. Niethammer, J., 1990. Talpa. In: Niethammer, J., Krapp, F. (Eds.), Handbuch der S¨ augetiere Europas. Band 3/1. Insektenfresser-Insectivora, Herrentiere-Primates. Aula-Verlag, Wiesbaden, pp. 93–161. Noddle, B., 1974. Ages of epiphyseal closure in feral and domestic goats and ages of dental eruption. J. Archaeol. Sci. 1 (2), 195–204. Nores, C., 1988. Diferenciaci´ on biom´ etrica de Apodemus sylvaticus y Apodemus flavicollis en la Cordillera Cant´ abrica. Primeros resultados. Revista de Biología de la Universidad de Oviedo 6, 109–116. Nores, C., 1989. Variaci´ on temporal y espacial de micromamíferos: determinaci´ on mediante an´ alisis de egagr´ opilas de Tyto alba. Ph.D. Thesis. Departamento de Biología de Organismos y Sistemas, Facultad de Biología, Universidad de Oviedo. Nores, C., S´ anchez-Canals, J.L., Castro, A., Gonz´ alez, G.R., 1982. Variation du genre Neomys Kaup, 1829 (Mammalia, Insectívora) dans le secteur cantabro-galicien de la p´ eninsule Ib´ erique. Mammalia 46, 361–373. Olalde, I., Mallick, S., Patterson, N., et al., 2019. The genomic history of the Iberian Peninsula over the past 8000 years. Science 363 (6432), 1230–1234. Ordiales, A., Su´ arez-Bilbao, A., Garcia-Ibaibarriaga, N., Ibarra, J.L., Murelaga, X., 2015. Estudio de los micromamíferos de los lechos de la Edad del Bronce de la Cueva de Arenaza I (Galdames, Bizkaia). Geogaceta 58, 51–54. Palacios, F., L´ opez Martínez, N., 1980. Morfología dentaria de las liebres europeas (Lagomorpha, Leporidae). Do˜ nana Acta Vertebrata 7 (1), 61–81. Pales, L., Lambert, C., 1971. Atlas Osteologique Pour Servir ` a l’identific´ ation Des Mammif´ eres du Quaternaire. Editions du Centre National de la Recherche Scientifique, Paris. Palomo, L.J., Gisbert, J., Blanco, J.C., 2007. Atlas Y Libro Rojo de los Mamíferos Terrestres de Espa˜ na. Direcci´ on General para la Biodiversidad-SECEM-SECEMU, Madrid, Spain. Pasquier, L., 1974. Dynamique ´ evolutive d’un sous-genre de Muridae, Apodemus (Sylvaemus). Etude biometrique des caract` eres dentaires des populations fossiles et actuelles d’Europe Occidentale. Th` ese. Universit´ e des Sciences et Techniques du Lanquedoc, Montpellier. Patino, S., Valencia, J., Elorza, J., Prieto, A., 2002. La flora del monte Serantes. Bizkaiko Gaiak / Temas Vizcaínos 331–332, 183 pp. Patterson, N., Isakov, M., Booth, T., et al., 2022. Large-scale migration into Britain during the Middle to Late Bronze Age. Nature 601, 588–594. Paup´ erio, J., Herman, J.S., Melo-Ferreira, J., Jaarola, M., Alves, P.C., Searle, J.B., 2012. Cryptic speciation in the field vole: a multilocus approach confirms three highly divergent lineages in Eurasia. Mol. Ecol. 21, 6015–6032. Pelletier, M., 2018. ´ Evolution morphom´ etrique et Biog´ eographie des L´ eporid´ es dans les environnements m´ editerran´ eens au Pl´ eistoc` ene. Implications socio-´ economiques pour les soci´ et´ es humaines. Th´ ese. Aix-Marseille Universit´ e. Pem´ an, E., 1983. Biometría y sistem´ atica del g´ enero Neomys Kamp 1771 (Mammalia, Insectivora) en el País Vasco. Munibe 35 (1–2), 115–148. Pem´ an, E., 1990. Los micromamíferos de la cueva de Amalda y su significado. Comentarios sobre Pliomys lenki (Heller, 1930) (Rodentia, Mammalia). In: La cueva de Amalda (Zestoa, País Vasco): ocupaciones paleolíticas y postpaleolíticas. Colecci´ on Barandiaran (4). Eusko Ikaskuntza, pp. 225–238. Pe˜ na-Chocarro, L., Zapata, L., Iriarte, M.J., Gonz´ alez-Morales, M.R., Straus, L.G., 2005. The oldest agriculture in northern Atlantic Spain: new evidence from El Mir´ on Cave (Ramales de la Victoria, Cantabria). J. Archaeol. Sci. 32 (4), 579–587. P´ erez-Díaz, S., L´ opez-S´ aez, J.A., Pontevedra-Pombal, X., Souto-Souto, M., Galop, D., 2016. 8000 years of vegetation history in the northern Iberian Peninsula inferred from the palaeoenvironmental study of the Zalama ombrotrophic bog (BasqueCantabrian Mountains, Spain). Boreas 45 (4), 658–672. P´ erez-Díaz, S., L´ opez-S´ aez, J.A., Nú˜ nez de la Fuente, S., Ruiz-Alonso, M., 2018. Early farmers, megalithic builders and the shaping of the cultural landscapes during the Holocene in Northern Iberian mountains. A palaeoenvironmental perspective. J. Archaeol. Sci. Rep. 18, 463–474. P´ erez-Obiol, R., García-Codron, J.C., P` elachs, A., P´ erez-Haase, A., Soriano, J.M., 2016. Landscape dynamics and fire activity since 6740 cal yr BP in the Cantabrian region (La Molina peat bog, Puente Viesgo, Spain). Quat. Sci. Rev. 135, 65–78. Ramil-Rego, P., Rodríguez-Guiti´ an, M., Mu˜ noz-Sobrino, C., 1998. Sclerophyllous vegetation dynamics in the north of the Iberian peninsula during the last 16,000 years. Glob. Ecol. Biogeogr. Lett. 7, 335–351. Reimer, P.J., Austin, W.E.N., Bard, E., et al., 2020. The IntCal20 Northern Hemisphere Radiocarbon Age Calibration Curve (0–55 cal kBP). Radiocarbon 62 (4), 725–757. Rom´ an, J., 2019. Manual para la identificaci´ on de los cr´ aneos de los roedores de la península ib´ erica, islas baleares y canarias. Manuales de Mastozoología de la SECEM. Rosengren, E., Acatrinei, A., Cruceru, N., Dehasque, M., Haliuc, A., Lord, E., Mircea, C.I., Rusu, I., M´ armol-S´ anchez, E., Kelemen, B.S., Meleg, I.N., 2021. Ancient. Diversity 13 (8), 370. Royer, A., Laroulandie, V., Bailon, S., Boudadi-Maligne, M., Costamagno, S., Danger, M., Mallye, J.-B., Rofes, J., 2021. Des restes de faune aux pal´ eoenvironnements de Peyzaret. Estudios 43, 35–48. Ruddiman, W.F., Thomson, J.S., 2001. The case for human causes of increased atmospheric CH4 over the last 5000 years. Quat. Sci. Rev. 20, 1769–1777. Ruprecht, A.J., 1971. Taxonomic value of mandible measurements in Soricidae (Insectívora). Acta Theriol. 16 (21), 341–357. S´ anchez, A., 1983. Estudio comparativo de las faunas pleistoc´ enicas y actuales de micromamíferos (Insectívoros y Roedores) en Puenteviesgo (Santander). Unpublished bachelor’s thesis. Facultad de Biología, UCM in press. Schmid, E., 1972. Atlas of Animal Bones. Elsevier Publishing Company, London. Serjeantson, D., 2011. In: Review of animal remains from the Neolithic and early Bronze age of southern Britain (4000 BC–1500 BC).Research Department Report Series, pp. 1–170. Sigl, M., Winstrup, M., McConnell, J., et al., 2015. Timing and climate forcing of volcanic eruptions for the past 2,500 years. Nature 523, 543–549. Silver, I.A., 1980. La determinaci´ on de la edad en los animales dom´ esticos. In: Brothwell, D.R., Higgs, E. (Eds.), Ciencia en Arqueología, Madrid, Fondo de Cultura Econ´ omica, pp. 290–307. Steinhilber, F., Abreu, J.A., Beer, J., Brunner, I., Christl, M., Fischer, H., Heikkil¨ a, U., Kubik, P.W., Mann, M.E., McCracken, K.G., Miller, H., Miyahara, H., Oerter, H., Wilhelms, F., 2012. 9,400 years of cosmic radiation and solar activity from ice cores and tree rings. Proc. Nat. Acad. Sci. 109 (16), 5967–5971. Steinhilber, F., Beer, J., Fr¨ ohlich, C., 2009. Total solar irradiance during the Holocene. Geophys. Res. Lett. 36 (19), L19704. Tom´ e, C., Vigne, J.D., 2003. Roe deer (Capreolus capreolus) age at death estimates: new methods and modern reference data for tooth eruption and wear, and for epiphyseal fusion. Archaeofauna 12, 157–173. Valenzuela-Lamas, S., Baylac, M., Cucchi, T., Vigne, J.-D., 2011. House mouse dispersal in Iron Age Spain: a geometric morphometrics appraisal. Biol. J. Linn. Soc. 102 (3), 483–497. Van Geel, B., Buurman, J., Waterbolk, H.T., 1996. Archaeological and palaeoecological indications of an abrupt climate change in the Netherlands, and evidence for climatological teleconnections around 2650 BP. J. Quat. Sci. 11 (6), 451–460. Vega-Maeso, C., Carmona-Ballestero, E., Sierra Sainz-Aja, A., Marín-Arroyo, A.B., 2016. El Abrigo de la Casta˜ nera (Cantabria, Spain): a Chalcolithic cattle stable? Quat. Int. 414, 226–235. Villalba-Mouco, V., Oliart, C., Rihuete-Herrada, C., et al., 2021. Genomic transformation and social organization during the Copper Age-Bronze Age transition in southern Iberia. Sci.Adv. 7 (47), eabi7038. Vitousek, P.M., Mooney, H.A., Lubchenco, J., Melillo, J.M., 1997. Human domination of earth’s ecosystems. Science 277, 494–499. Walker, M., Head, M.J., Berkelhammer, M., Bj¨ orck, S., Cheng, H., Cwynar, L., Fisher, D., Gkinis, V., Long, A., Lowe, J., Newnham, R., Rasmussen, S.O., Weiss, H., 2018. Formal ratification of the subdivision of the Holocene Series/Epoch (Quaternary System/Period): two new Global Boundary Stratotype Sections and Points (GSSPs) and three new stages/subseries. Episodes 41, 213–223. Wanner, H., Beer, J., Bütikofer, J., Crowley, T.J., Cubasch, U., Flückiger, J., Goosse, H., Grosjean, M., Joos, F., Kaplan, J.O., Küttel, M., Müller, S.A., Prentice, I.C., Solomina, O., Stocker, T.F., Tarasov, P., Wagner, M., Widmann, M., 2008. Midto late Holocene climate change: an overview. Quat. Sci. Rev. 27 (19–20), 1791–1828. Wilkin, S., Ventresca Miller, A., Fernandes, R., et al., 2021. Dairying enabled early Bronze Age Yamnaya steppe expansions. Nature 598, 629–633. Wilson, D.E., Lacher, T.E., Mittermeier, R.A., 2016. Handbook of the Mammals of the World: 6. Lagomorphs and Rodents I. Lynx Edicions, Barcelona, Spain. Wilson, D.E., Lacher, T.E., Mittermeier, R.A., 2017. Handbook of the Mammals of the World: 7. Rodents II. Lynx Edicions, Barcelona, Spain. Wilson, D.E., Mittermeier, R.A., 2018. Handbook of the Mammals of the World: 8. Insectivores, Sloths and Colugos. Lynx Edicions, Barcelona, Spain. Yasuda, S.P., Vogel, P., Tsuchiya, K., Han, S.-H., Lin, L.-K., Suzuki, H., 2005. Phylogeographic patterning of mtDNA in the widely distributed harvest mouse (Micromys minutus) suggests dramatic cycles of range contraction and expansion during the midto late Pleistocene. Can. J. Zool. 83 (11), 1411–1420. Zapata, L., 1995. The excavation of the burial cave Pico Ramos (Muskiz, Biscay). The ornamental and bone industry. In: Zapata, L. (Ed.), The Chalcolithic burial deposit of the cave Pico Ramos (Muskiz, Biskay). Munibe (Antropologia-Arkeologia) 47, pp. 35–90. Zapata Pe˜ na, L., 1999. El combustible y la agricultura prehist´ orica. Estudio arqueobot´ anico de los yacimientos de Arenaza, Kanpanoste Goikoa y Kobaederra (Fuelwood and prehistoric agriculture. Archaeobotanical analyses from the cave sites Arenaza, Kanpanoste Goikoa and Kobaederra). Isturitz 10, 305–337. Zapata Pe˜ na, L., 2002. Origen de la agricultura en el País Vasco y transformaciones en el paisaje: an´ alisis de restos vegetales arqueol´ ogicos. Kobie 4, 223 p. Zapata Pe˜ na, L., Pe˜ na-Chocarro, L., P´ erez Jord´ a, G., Stika, H.-P., 2005. Difusi´ on de la agricultura en la Península Ib´ erica. In: Arias, P., Onta˜ n´ on, R., García-Monc´ o, C. (Eds.), III Congreso del Neolítico en la Península Ib´ erica, pp. 103–113. A. ´ Alvarez-Vena et al.