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Meat consumption by early hominins is a hotly debated issue. A key question concerns their access to large mammal carcasses, including megafauna. Currently, the evidence of anthropic cut marks on proboscidean bones older than -or close to- 1.0 Ma are restricted to the archaeological sites of Dmanisi (Georgia), Olduvai (Tanzania), Gona (Ethiopia), Olorgesailie (Kenya) and La Boella (Spain). During an inspection of the almost complete carcass of Mammuthus meridionalis (FN3-5-MPS) from the Oldowan site of Fuente Nueva 3 (Orce, Spain, c. 1.2 Ma), a few traces compatible with human-made cut marks and carnivore tooth marks were found. From this finding and previous interpretations the following questions arise: When and under what conditions was FN3-5-MPS deposited? What is the nature of the marks found on the surface of the bones of this mammoth? To answer, we have conducted a high-resolution analysis of these remains, combining both taphonomic and microstratigraphic data. Our results, using microstratigraphic and micromorphological analyses of sediments based on thin-sections, show that this individual was deposited in a marshy environment. Subsequently, the carcass was exploited by hominins and large felids that left their marks on the surface of some of its bones. For this purpose, the identification and characterisation of both cut marks and tooth marks were performed using high-resolution 3D modelling, geometric morphometrics, and artificially intelligent algorithms. Based on the anatomical position of both the cut and tooth marks, we propose that both the hominins and the saber-toothed cats had early access to the animal. Finally, this paper shows how an interdisciplinary approach can shed detailed light on the particular story regarding the death and processing of the carcass of a female mammoth, deposited at Fuente Nueva 3.

Meat consumption by early hominins is a hotly debated issue. A key question concerns their access to large mammal carcasses, including megafauna. Currently, the evidence of anthropic cut marks on proboscidean bones older than -or close to- 1.0 Ma are restricted to the archaeological sites of Dmanisi (Georgia), Olduvai (Tanzania), Gona (Ethiopia), Olorgesailie (Kenya) and La Boella (Spain). During an inspection of the almost complete carcass of Mammuthus meridionalis (FN3-5-MPS) from the Oldowan site of Fuente Nueva 3 (Orce, Spain, c. 1.2 Ma), a few traces compatible with human-made cut marks and carnivore tooth marks were found. From this finding and previous interpretations the following questions arise: When and under what conditions was FN3-5-MPS deposited? What is the nature of the marks found on the surface of the bones of this mammoth? To answer, we have conducted a high-resolution analysis of these remains, combining both taphonomic and microstratigraphic data. Our results, using microstratigraphic and micromorphological analyses of sediments based on thin-sections, show that this individual was deposited in a marshy environment. Subsequently, the carcass was exploited by hominins and large felids that left their marks on the surface of some of its bones. For this purpose, the identification and characterisation of both cut marks and tooth marks were performed using high-resolution 3D modelling, geometric morphometrics, and artificially intelligent algorithms. Based on the anatomical position of both the cut and tooth marks, we propose that both the hominins and the saber-toothed cats had early access to the animal. Finally, this paper shows how an interdisciplinary approach can shed detailed light on the particular story regarding the death and processing of the carcass of a female mammoth, deposited at Fuente Nueva 3.

This work received financial support from the research projects CGL2015–68333-P (MINECO/FEDER, UE), Research Group BSCHUCM 910607 and Consolidated Research Group EO5 (Aragón Government). Referencias bibliográficas: • Adrover R (1986) Nuevas faunas de roedores en el Mio-Plioceno continental de la región de Teruel (España). Interés bioestratigráfico y paleoecológico. Instituto de Estudios Turolenses, Teruel • Adrover R, Feist M, Hugueney M, Mein P, Moissenet E (1982) L'áge et la mise en relief de la formation détritique culminante de la Sierra Pelarda (Province de Teruel, Espagne). C R Acad Sci Paris 295:231–236 • Aguirre E (1973) Conservación e historia de la Naturaleza. Boletín Estación Central. Ecología 2:89–97 • Aguirre E, Morales J, Soria D (1981) Accumulated bones in a Pliocene cave in Cerro Pelado, Spain. Natl Res Rep 13:69–81 • Agustí J, Santos-Cubero A, Furió M, De Marfa R, Blain H-A, Oms O, Sevilla P (2011) The late Neogene-early quaternary small vertebrate succession from the Almenara-Casablanca karst complex (Castellón, Eastern Spain): chronologic and paleoclimatic context. Quat Int 243:183–191 • Calvo JP, Daams R, Morales J, López-Martínez N, Agustí J, Anadón P, Armenteros I, Cabrera L, Civis J, Corrochano A, Díaz-Molina M, Elizaga E, Hoyos M, Martín-Suárez E, Martínez J, Moissenet E, Muñoz A, Pérez-García A, Pérez-González A, Portero JM, Robles F, Santisteban C, Torres T, van der Meulen J, Vera JA, Mein P (1993) Up-to-date Spanish continental Neogene synthesis and paleoclimatic interpretation. Rev Soc Geol Esp 6:29–40 • Daams R, Meulen A, Van Der Peláez-Campomanes P, Álvarez-Sierra MA (1999) Trends in rodent assemblages from the Aragonian (early-middle Miocene) of the Calatayud-Daroca Basin, Aragón, Spain. In: Agustí J, Rook L, Andrews P (eds) Hominoid evolution and climatic change in Europe (the evolution of Neogene terrestrial ecosystems in Europe), vol 1. Cambridge University Press, Cambridge, pp 390–412 • Hilgen FJ, Lourens LJ, Dam JA Van, with Contr, Corral de Lobato, a karstic site in the area of Molina de Aragón has been studied in a preliminary way. Even though there are not many Neogene karstic sites in the Iberian Chain, they occur in four clusters, with ages ranging from latest middle Miocene (MN7/8) to early Pleistocene (MN17). Correlations between these clusters and the reference stratigraphical units of the Tagus Basin, as well as with local and global events, are proposed. These karstic sites provide a complementary source of fossil vertebrate remains to that of the stratified sites formed lowland. The Heritage significance of such sites arises from the enhanced preservation of rare taxa or associations, and the operation of biotic concentrative processes., Gobierno de Aragón, Depto. de Didáctica de las Ciencias Experimentales , Sociales y Matemáticas, Fac. de Educación, TRUE, pub

The Orce Basin Archaeological Zone (OZAB, Granada, Spain) extends over a surface area of some > 8.5 km2 and constitutes one of the richest Pleistocene vertebrate fossil records in western Europe including one of the oldest hominin presence in this part of Eurasia. Exceptionally rich collections of stone tools have been excavated from both of the Orce Oldowan sites: Barranco León (BL) (1.4 Ma) and Fuente Nueva 3 (FN3) (1.2 Ma), while BL has yielded a hominin deciduous lower molar. We present a multi-proxy approach to determine the palaeoeocological context of these first hominin settlements in Western Europe. By combining results from macrovertebrates, microvertebrates, as well as from pollen and stable isotopes from macrovertebrate tooth enamel, we determine whether the palaeo landscapes were dominated by savanna or open woodland. The results reveal the regional specificities of the OAZB, and also allow us to infer local features within the Orce sites. Overall, our data reveal the dominance of a typically Mediterranean climate and landscape since 1.8 Ma ago. The climatic conditions were generally more humid than at present, with warmer temperatures during the coldest months, indicating a higher net primary productivity (NPP). We find that precipitation and NPP appear to have been limiting factors for hominin presence in the OAZB. Thus, at the older palaeontological site of Venta Micena (1.6 Ma), climatic conditions appear to have been less compatible with hominin presence than during the BL and FN3 sequences, when early hominins inhabiting the OAZB were able to cope with changing climatic and environmental settings. Lastly, the comparison of the isotopic results of the Orce sites with those of the contemporaneous Shungura Formation (Ethiopia) reveals that the habitat in the westernmost part of Eurasia was distinctly unlike a typical African savanna.

Altres ajuts: CERCA Programme/Generalitat de Catalunya, Els Casots is one of the richest fossil vertebrate sites of the Vallès-Penedès Basin (Catalonia, Spain). It was discovered in 1989 and excavated briefly during the 1990s, resulting in the recovery of thousands of remains and the erection of several new mammal species. Excavations were resumed in 2018 and continue to date. Here we provide updated results regarding the age, stratigraphy, biota and palaeoenvironment of the site. The age of the site is well constrained to ~15.9 Ma thanks to recent bio- and magnetostratigraphic data, thus coinciding with the Miocene Climatic Optimum (MCO). The stratigraphic succession at the site area indicates lacustrine to palustrine environments with cyclically oscillating water level. There are several fossiliferous layers that have yielded a vertebrate fauna comprising up to 75 different vertebrate species including amphibians, reptiles, birds and mostly mammals. The finding of several articulated partial skeletons indicate that the site records an autochthonous to parautochthonous assemblage. The abundance and completeness of the vertebrate remains together with a well-constrained age and detailed stratigraphic and palaeoenvironmental data, make els Casots a key site for understanding wetland ecosystems in southern Europe during the MCO.

Small to medium-sized mustelids from the last 18 million years represent a heterogeneous group of carnivorans with a wide-ranging record in the northern hemisphere. They were first referred to the genera Mustela and Martes, but lately ascribed to the latter, and hence considered as the longest-lived genus within Mustelidae. However, a great many of these forms have been based upon fragmentary material and Martes has conformed progressively to a wastebasket nomen for species of uncertain relationships. Here, we describe dentognathic material of a small-sized mustelid from three middle Miocene (MN7 + 8, latest Aragonian) localities of the Iberian Peninsula that constitutes a new genus and species. Aragonictis araid, gen. et sp. nov. represents a distinct taxon if compared with early/middle Miocene forms ascribed to "Martes" spp., especially the similar-sized early Miocene Circamustela? laevidens and the middle Miocene "Martes" caedoti and "Martes" delphinensis. The finding of particular features in A. araid (low p2-3, loss or reduction of the p4 accessory cuspid with its main cuspid centrally located, presence of a sharp, beveled and lingually open m1 talonid, and reduction of M1 lingual platform) indicates affinities with the late Miocene Circamustela in the range of hypercarnivory. Our reassessment of "Martes" indicates possible evidence of cladogenesis for Miocene mustelidae with, at least, two different events being recognized in Europe-the latter during MN7 + 8 to MN9 with presence of Aragonictis and Circamustela. The finding of A. araid further confirms the presence of more densely forested environments than expected in inner Iberia during the latest middle Miocene.

The fossil record of lynxes provides clear evidence of a large range across the North Hemisphere during the Pliocene and Pleistocene. However, their origin, systematics and evolutionary relationships are still fraught with difficulties and controversy. Here we report a complete hemimandible of a medium-sized felid from the Early Pleistocene (MN17, middle Villafranchian, 2.05 Ma) site of La Puebla de Valverde (Teruel, Spain). Based on comparative and multivariate analyses of the lower dentition of 458 individuals of medium-sized Lynx, Caracal and Leptailurus, we confidently ascribe the remains to Lynx aff. issiodorensis. Although the dental proportions are somewhat different from those of the Eurasian L. issiodorensis (smaller canines and more elongated p4/m1), Lynx aff. issiodorensis shows affinities with the Issoire lynx from the contemporaneous site of Saint Vallier (France), sharing a similar morphology of the mandible, reduced canines, and long m1. We further test the hypothesis that examines the presence of the African/Asian Caracal in the European Plio/Pleistocene for C. depereti and C. issiodorensis, and discard the attribution of L. issiodorensis into Caracal. This mandible extends the record of the genus and contributes to update our understanding of the Lynx lineage and its variability within the European fossil record.

This volume honours Professor Jorge Morales – one of the world´s leading experts in fossil carnivorans, a great scholar, and inspiration of many to follow him in the discipline of palaeontology – to mark his recent retirement. In July 2021, Prof. Morales retired after 50 years working and building up a reputation not only as a leading scientist but also as a teacher, facilitator of science, and mentor. This festschrift celebrates his lifelong, outstanding scientific and cultural achievements to mammal palaeontology. It contains contributions by colleagues, former students, and friends that come from far and wide. Articles pay homage to the jubilarian´s research interests, covering the systematics, palaeoecology, biochronology, palaeobiogeography, taphonomy, and geoheritage. On the basis of the written acknowledgements, it is easy to recognise the affection and deep esteem Prof. Morales induces in everyone. If we think about him, our professional and personal lives would not be the same without his support, mentorship, friendship, and good humour. It is a proud privilege to thank him for his career in terms of production and excellence which without doubt placed Spanish palaeontology among the global elite. Prof. Morales is retired, but he remains active in research, and as inspiration to all of us.

Small to medium-sized mustelids from the last 18 million years represent a heterogeneous group of carnivorans with a wide-ranging record in the northern hemisphere. They were first referred to the genera Mustela and Martes, but lately ascribed to the latter, and hence considered as the longest-lived genus within Mustelidae. However, a great many of these forms have been based upon fragmentary material and Martes has conformed progressively to a wastebasket nomen for species of uncertain relationships. Here, we describe dentognathic material of a small-sized mustelid from three middle Miocene (MN7 + 8, latest Aragonian) localities of the Iberian Peninsula that constitutes a new genus and species. Aragonictis araid, gen. et sp. nov. represents a distinct taxon if compared with early/middle Miocene forms ascribed to “Martes” spp., especially the similar-sized early Miocene Circamustela? laevidens and the middle Miocene “Martes” caedoti and “Martes” delphinensis. The finding of particular features in A. araid (low p2–3, loss or reduction of the p4 accessory cuspid with its main cuspid centrally located, presence of a sharp, beveled and lingually open m1 talonid, and reduction of M1 lingual platform) indicates affinities with the late Miocene Circamustela in the range of hypercarnivory. Our reassessment of “Martes” indicates possible evidence of cladogenesis for Miocene mustelidae with, at least, two different events being recognized in Europe—the latter during MN7 + 8 to MN9 with presence of Aragonictis and Circamustela. The finding of A. araid further confirms the presence of more densely forested environments than expected in inner Iberia during the latest middle Miocene.

Els Casots is one of the richest fossil vertebrate sites of the Vallès-Penedès Basin (Catalonia, Spain). It was discovered in 1989 and excavated briefly during the 1990s, resulting in the recovery of thousands of remains and the erection of several new mammal species. Excavations were resumed in 2018 and continue to date. Here we provide updated results regarding the age, stratigraphy, biota and palaeoenvironment of the site. The age of the site is well constrained to ~15.9 Ma thanks to recent bio- and magnetostratigraphic data, thus coinciding with the Miocene Climatic Optimum (MCO). The stratigraphic succession at the site area indicates lacustrine to palustrine environments with cyclically oscillating water level. There are several fossiliferous layers that have yielded a vertebrate fauna comprising up to 75 different vertebrate species including amphibians, reptiles, birds and mostly mammals. The finding of several articulated partial skeletons indicate that the site records an autochthonous to parautochthonous assemblage. The abundance and completeness of the vertebrate remains together with a well-constrained age and detailed stratigraphic and palaeoenvironmental data, make els Casots a key site for understanding wetland ecosystems in southern Europe during the MCO.

Venta Micena is an area containing several palaeontological sites marking the beginning of the Calabrian stage (Early Pleistocene). The richness of the fossil accumulation including species of Asian, African and European origin, makes Venta Micena a key site for the the palaeoecological and palaeoenvironmental study of southern Europe during the Early Pleistocene. Thus, research has been focused on Venta Micena 3, which was originally interpreted as a single palaeosurface associated with a marshy context, in which most of the fauna was accumulated by Pachycrocuta brevirostris. Recent excavations have unearthed a new site, Venta Micena 4, located in the same stratigraphic unit (Unit C) and in close proximity to Venta Micena 3. Here we show the first analyses regarding the taphonomic and spatial nature of this new site, defining two stratigraphic boundaries corresponding to two different depositional events. Furthermore, the taphonomic analyses of fossil remains seem to indicate a different accumulative agent than Pachycrocuta, thus adding more complexity to the palaeobiological interpretation of the Venta Micena area. These results contribute to the discussion of traditional interpretations made from Venta Micena 3

The Guadix-Baza Basin (GBB) in Andalucía, Spain, comprises palaeontological and archaeological sites dating from the Early Pliocene to the Middle Pleistocene, including some of the earliest sites with evidence for the presence of early humans (Homo sp.) in Europe. Thus, the history of climate and environments in this basin contributes significantly to our understanding of the conditions under which early humans spread into Europe during the Early Pleistocene. Here we present estimates of precipitation and primary productivity in the GBB from the Pliocene to the Middle Pleistocene based on dental ecometrics in fossil communities of large herbivorous mammals, and perform an ecometrics-based distribution modelling to analyse the environmental conditions of Early and Middle Pleistocene human sites in Europe. Our results show that Early Pleistocene humans generally occupied on average relatively diverse habitats with ecotones, such as woodlands and savannas, but avoided very open and harsh (cool or dry) environments. During the Middle Pleistocene in Europe, humans occupied a comparatively much broader range of environments than during the Early Pleistocene, but were on average more concentrated in environments where the dental ecometric of mammals indicate wooded palaeoenvironments. In the earliest human occupation sites of the GBB, Barranco Leon and Fuente Nueva 3, the mean annual precipitation and net primary production estimates indicate climatic conditions close to modern Mediterranean sclerophyllous woodland environments, but with slightly higher primary productivity, basin indicating some similarity with East African woodlands. On the other hand, the environments did not resemble African grassland savannas. The browse-dominated diets of ungulates from Barranco León and Fuente Nueva 3 further suggest palaeoenvironments where grasses were a minor component of the vegetation. In the slightly older site of Venta Micena that has no evidence for the presence of hominins

Venta Micena is an area containing several palaeontological sites marking the beginning of the Calabrian stage (Early Pleistocene). The richness of the fossil accumulation including species of Asian, African and European origin, makes Venta Micena a key site for the the palaeoecological and palaeoenvironmental study of southern Europe during the Early Pleistocene. Thus, research has been focused on Venta Micena 3, which was originally interpreted as a single palaeosurface associated with a marshy context, in which most of the fauna was accumulated by Pachycrocuta brevirostris. Recent excavations have unearthed a new site, Venta Micena 4, located in the same stratigraphic unit (Unit C) and in close proximity to Venta Micena 3. Here we show the first analyses regarding the taphonomic and spatial nature of this new site, defining two stratigraphic boundaries corresponding to two different depositional events. Furthermore, the taphonomic analyses of fossil remains seem to indicate a different accumulative agent than Pachycrocuta, thus adding more complexity to the palaeobiological interpretation of the Venta Micena area. These results contribute to the discussion of traditional interpretations made from Venta Micena 3.

The end of the Miocene was an eventful period of changes in climate and geography, and a restructuring of terrestrial plant and mammals. The tendency towards global aridification has attracted much recent interest, making the late Miocene a striking case-study for testing current and near-future scenarios involving global warming. Little is known about the consequences of global changes in temperature and precipitation at regional or local scales. Given its geographical position and extraordinary fossil record, the Iberian Peninsula offers many insights into short- and long-term shifts in climate, and the local response. Here, we explore the diet and ecology of large-mammals through tooth-wear patterns, and examine changes in local climate and habitat conditions in central Spain in a period (9.1–6.3 Ma) for which there exists a dearth of palaeoenvironmental information. Relatively dry climates and open-woodland landscapes evolved locally during the late Vallesian and early Turolian (9.0–7.7 Ma). Unexpectedly, we detect a period of high precipitation and a peak of humidity at the end of the Turolian (7.0 Ma) that prompted the development of wetter, more forested habitats, suggesting that the traditional view of the late Miocene as a steppe landscape is a misconception. We also find a period of relatively drier and warmer conditions from the early Ventian onwards. Overall, our finding that a local episode of increased humidity in central Spain was synchronous with a global warming trend in Europe provides evidence that the greatest climatic changes may have an opposite impact at regional and local scales.