The field vole, an abundant and widespread microtine rodent, is a complex comprised of three cryptic species: the short-tailed field vole (Microtus agrestis) which is present over much of Eurasia, the Mediterranean field vole (Microtus lavernedii) in southern Europe, and the Portuguese field vole (Microtus rozianus) in western Spain and Portugal. Previous research has shown high genomic differentiation of these three lineages. However, the details of the process underlying their divergence remain unknown. We analyse 70 mitogenomes and 16 nuclear genomes of modern specimens, and 83 mitogenomes and 12 nuclear genomes of ancient specimens spanning the last 75 thousand years (ka). We estimate the divergence of Portuguese from short-tailed and Mediterranean field voles to be ca. 220 ka ago and of the latter two species to be ca. 110 ka ago, earlier than previous estimates involving only modern sequences. The divergence times we obtain match those between major mitochondrial lineages of cold-adapted and steppe rodents in Europe. We find signatures of gene flow within and between field vole lineages, with some analyses suggesting a hybrid origin of the Mediterranean lineage. Ancient specimens from the Italian Peninsula reveal a previously unrecognised lineage that show evidence of genetic exchange with other populations. The pattern of genetic variation in the field vole species complex demonstrates the impact of stadial-interstadial cycles in generating recurrent episodes of allopatry and connectivity of populations, a situation which could only be revealed by our dense genomic sampling over time.

The turtle Marocokatognathus jimenezi n. g., n. sp., known by a dentary, unique by its long, narrow and upward curved beak, is defined as Chelonioidea, not Dermochelyoidea. The morphology of the dentary indicates its skull had a secondary palate. By comparison with Chelonioid turtles, the variations of that structure are classified here in morphotypes, of practical taxonomical use and partly of phyletic incidence, notably as a Chelonioidea member. The new fossil is attributed to Cheloniidae and related to extant forms by:- its general dentary morphology including a Carettine shape of the bony triturating surface and the same relative proportions of distances between structural elements (posterior border of the symphysis, coronoid process, foramen mentale, in relation to corresponding skull structures);- the assignation to a same morphotype 1 of secondary palate, by an elongated snout for an elongated lower jaw, the choanal border anterior to palatine fossae, palate not developed beyond ca. two thirds of the snout length, correlatively preserving long dentary branches anteriorly and posteriorly to the coronoid process;- and overall by the posterior implantation of the mandibular ramphotheca on the mandibular symphysis with a medial symphyseal facet, specific of extant Cheloniidae. The possible diet and prey capture processes are examined, proposing a seizure of various prey of shape adapted to the narrow and long beak, followed immediately by their aspiration and swallowing. By its novelty, the new form emphasizes the great diversity of fossil turtles from Phosphates of Morocco, which are inventoried, as of the whole world. (c) 2025 Elsevier Masson SAS. All rights are reserved, including those for text and data mining, AI training, and similar technologies.

Abundant crocodylian fossil remains from the Early Miocene locality of Monte la Pila, which have not been studied previously, are described here. This fossil site is located in the municipality of Lardero, a few kilometers from Logroño (La Rioja, Spain), in the Ebro Basin (late Aquitanian–early Burdigalian). The recovered material includes one hundredand twenty-one disarticulated and isolated specimens. Despite the incompleteness of the remains, certain diagnostic features within the genus Diplocynodon are unambiguouslyrecognized in the studied material. Based on the identification of some of these characters, the material has been assigned to Diplocynodon sp. This assignment confirms the presence of this taxon in the late Aquitanian–early Burdigalian of the Iberian Peninsula.It also provides valuable data on the anatomical characteristics and distribution of the crocodile fauna on the Iberian Peninsula during the Lower Miocene.

The Tortonian Pena Adrian Formation represents the youngest depositional unit of the Miranda-Trebino basin (Basque-Cantabrian Pyrenees), which developed on the Southern Basque-Cantabrian Pyrenees from late Eocene to Late Miocene times. The formation is a 50-160 m thick succession of alluvial detrital grading to lacustrine carbonates that contain rich and varied calcareous fossil biota (gastropods, ostracods and charophytes). The fossil association characterizes warm temperate, shallow lakes with vegetated bottoms and well-oxygenated and alkaline fresh waters. Integration of stratigraphic, sedimentological, paleontological and C-O stable isotopic data allows the differentiation of a wide range of sedimentary facies, the construction of a depositional model and the definition of distinct evolutionary phases and relation to allogenic processes. Up to 3 metre-thick facies sequences record repetitive water-level changes, likely reflecting short-term climate changes. Overall, the succession outlines an asymmetric cycle of gradual expansion and faster contraction of a shallow ramp-like lake system evolving under oscillating climatic conditions. C and O stable isotopes are consistent with decreasing salinity and increasing precipitation/evaporation balance trough time. Excellent preservation of aragonitic and bimineralic gastropods characterizes the open lacustrine deposits, whereas shell dissolution and neomorphism are distinct in the palustrine ones. This contrasting degree of preservation of calcareous biota clearly reflects changes in the physico-chemical conditions that prevailed during sedimentation and early burial. The findings add to the knowledge of carbonate lake basins, help discern the factors that controlled their evolution and highlight specific depositional and preservation conditions for gastropod-rich carbonate records.

This study examines the factors that controlled the formation of Quaternary calcareous tufa deposits along the valley of the Inglares River (Álava, north Spain), currently fed by a karst-carbonate aquifer. It is based on stratigraphic, chronological (Amino Acid Racemisation, AAR), sedimentological, and δ13C and δ18O analyses, complemented by palaeontology (molluscs, ostracods and charophytes). The examined deposits occur as isolated bodies in the uppermost and downmost stretches of the valley, reaching 45 and 25m in thickness, respectively. AAR dating classified them into two groups: Middle Late Pleistocene (MIS 5e) and Middle and Late Holocene (MIS 1). The former only occurred in the downstream stretch. Up to eleven carbonate facies and minor allochthonous coarse detrital facies have been characterised and arranged into four distinct facies associations. Their features and bedding geometries suggested two main depositional settings: a low- to moderate-slope stepped stretch with small barrage cascades, dammed areas, and abundant palustrine zones (downstream system) and a high-slope stretch with steep stepped cascades and pools (upstream system). These settings respond to bedrock lithology and structural changes throughout the valley, which appear to be the principal factors controlling the tufa depositional architecture. Based on δ13C, an increase in aridity was inferred from the Middle–Late Pleistocene to Holocene. The isotopic differences between the upstream and downstream Holocene tufa might reflect the spatial evolution of δ13C and δ18O in stream water through the approximately 8km long surveyed transect. Erosion due to sudden changes in water discharge may have caused the stratigraphic gap between the two tufa groups.

The study of the small mammals from the Mesolithic site of El Toral III (Llanes, Asturias) is reported in this paper. A total of 9 species have been identified, six belonging to the Order Rodentia and three to Order Eulipotyphla. All units are very poor in terms of the number of micromammal remains recovered, a figure that is in line with the high human occupation estimated for the site. Paleoecological data from all levels indicate a clear predominance of forest, with levels 20 and 21 showing a slight decrease in the masses in relation to the rest of the samples. Therefore, it was probably at that time when the less humid conditions and mild temperatures, recorded in this site, took place.

The stable isotopes (C and O), bulk mineralogy and sedimentary facies types of the Miocene lacustrine palustrine carbonates of the Cucho Section (Miranda-Trebiño basin, N Iberia) are studied. The results indicate deposition in a freshwater carbonate lake recording significant biogenic CO2 input, changing Precipitation/Evaporation rates and variations in vegetation cover and pedogenesis on palustrine areas. Short-lived stages of higher salinity and evaporative conditions are recorded by the relative abundance of fine-grained dolomite. The highest δ13C values suggest 12C sequestration in anoxic bottom waters due to water column stratification. These new data support the general deepening trend of the lake system established from previous sedimentological analyses.

The chronology of lower Miocene Iberian small mammal faunas is still poorly constrained given the scarcity of well dated sedimentary successions including small mammal fossil localities. Such scarcity has prevented also an accurate understanding of the response of European terrestrial ecosystem to global changes across the Miocene climate optimum (MCO), one of the best analogues of present-day global warming. Here we present an updated fossil small mammal record of the Bardenas Reales de Navarra (western Ebro basin, Spain), where an expanded lower to middle Miocene continental succession is superbly exposed. Previous and new magnetostratigraphic results from this succession have enabled us to propose, along with additional magnetostratigraphically-dated Iberian faunas, a new chronology for local zones Y to D (Mammal Neogene zones MN2 to MN5). In addition to that, the studied small mammal faunas point to a gradual increase in temperature and humidity conditions in SW Europe between 20 and 15.5 Ma, which appears to be coupled with the progressive shift towards warmer regional (Atlantic) and global conditions across the MCO, thereby pointing to gradual changes in oceanic circulation as the main driver of this period of global warmth. The evolution of sedimentary facies appears to indicate a threshold response of the Ebro basin hydrological balance to the MCO, whereas pedogenic formation of magnetic minerals seems to be linked to periods of enhanced climate variability. These results highlight the need of combining different paleoenvironmental indicators in order to obtain a reliable view of the response of continental ecosystems to global warming.

The Loza-Portilla Formation consists of 100-300 m of alluvial siliciclastics and palustrine-lacustrine carbonates and minor evaporites, infilling a tectonically active syncline basin (Miranda-Trebi & ntilde;o Basin, Basque-Cantabrian Pyrenees), which developed over the south Pyrenean Thrust Front during middle to late Eocene times. Detailed facies characterisation and correlation allowed the reconstruction of the stratigraphic architecture of the formation, which consists of three unconformity-bounded depositional sequences, representative of three successive phases of carbonate lake development. A model of low-energy shallow lake with ramp margins and wide palustrine fringes fits well with the architecture of the three stacked carbonate-dominated sequences, with stratal architectures characteristic of shallow balanced-fill lake basins. A range of limestones representative of different palustrine and lacustrine environments is distinguished, commonly arranged in metre-thick facies sequences and cycles reflecting water-level changes and marked shoreline shifts, likely associated with prevailing tectonic and climate conditions. Compressional tectonism and associated halokinetic movements exerted major control on depositional trends and distribution of the main depocentres. The Loza-Portilla Formation is of major palaeogeographical significance in the tecto-sedimentary evolution of the Pyrenean domain, labelling the first phase of uplift and emersion that affected large areas of the Basque-Cantabrian (western) Pyrenees, prior to end-Eocene generalised continentalisation. The Loza-Portilla Formation (middle-upper Eocene) from the Miranda-Trebi & ntilde;o Basin (Western Pyrenees, N Spain) consisits of alluvial to palustrine-lacustrine facies associations arranged in three depositional sequences. The lacustrine deposition was controlled by a semiarid climate, compressional tectonism and diapiric influences.image

This work is focused on a Lower–Middle Miocene section exposed on the northern flank of Miranda Trebiño piggy-back basin (Cucho section). The sedimentological and stratigraphic study of the section, composed of up to 160 m of terrigenous, mixed (marls), coal and carbonate deposits, allowed characterizing three depositional environments: distal alluvial, lacustrine and palustrine. The stratigraphic arrangement of different facies reflects five 28-43 m thick sequences which are related to progradation and retrogradation of alluvial systems and consequent retraction and expansion of the lacustrine and palustrine wetlands. The alluvial deposits have been associated with the alluvial fans of the northern basin margin, which was activated by the compressive tectonics prevailing during the time of deposition.

The common vole is a temperate rodent widespread across Europe. Phylogeographic studies of its extant populations suggested the Last Glacial Maximum (LGM) as one of the main drivers of the species’ population history. However, analyses based solely on extant genetic diversity may not recover the full complexity of Late Pleistocene population dynamics. To reconstruct the population history of the common vole through the Last Glacial Period, we analysed a 4.2 kb-long fragment of mitochondrial DNA of 148 ancient and 51 modern specimens, sampled from across Europe, and covering the last 60 thousand years (ka). We estimate the time to the most recent common ancestor of Last Glacial common vole lineages to 90 ka ago and the diversification of the main extant lineages to between 55 and 40 ka ago, substantially earlier than previously estimated. Our data suggests multiple lineage turnovers in Europe at the end of Marine Isotope Stage (MIS) 3 and around the Pleistocene/Holocene transition. Conversely, data from the Western Carpathians suggest continuity throughout the LGM. This further suggests that climate amelioration during MIS 2 had little impact on common voles and that the main driver of population dynamics was the reduction of open habitats during the interstadial periods.

This work presents the sedimentological study of the Upper Miocene-Pliocene Oko Limestones, exposed in the Antzin-Murieta Basin, in Navarre. Two stratigraphic sections of the carbonate unit have been studied: the Oko cemetery and Ega road sections. The former exposes facies that characterize a fluvial carbonate environment, with a distinct development of microbial oncolitic deposits. The coeval deposits exposed in the Ega road section, instead, largely represent a palustrine-lacustrine setting that evolved under alternating conditions of ponding and desiccation. Data integration from these sections and the intermediate areas, allows the interpretation of a fluvial to palustrine-lacustrine wetland with a marked east-southeast trend, which developed coevally to the uplift of the Piedramillera thrust sheet.

This work presents the results of the paleontological study of the Pena Adrian section (Miranda de Ebro, Burgos), formed by alluvial and palustrine-lacustrine facies from the Upper Miocene. Two levels of marls have been sampled, PAF15 and PAF24 (A and B) where different genera and species of gastropods, charophytes and ostracods have been identified. From the stratigraphic and paleontological study, it has been possible to recognize the environmental conditions of the lacustrine system and their variation throughout the succession, showing phases of higher and lower energy until the final retraction of the lake system.

Este trabajo presenta los resultados del estudio paleontológico de la sección de Peña Adrián (Miranda de Ebro, Burgos), formada por facies aluviales y palustre-lacustres del Mioceno Superior. Se han muestreado dos niveles margosos, PAF15 y PAF24 (A y B) donde se han identificado diferentes géneros y especies de gasterópodos, carofitas, y ostrácodos. A partir del estudio estratigráfico y paleontológico ha sido posible conocer las condiciones ambientales del sistema lacustre y su variación a lo largo de la sucesión, mostrando fases de mayor y menor energía hasta la colmatación y retracción final del sistema.

A systematic study of vertebrate remains from Mesolithic to Chalcolithic/Bronze Age levels in Lumentxa Cave has examined a total of 12,390 remains belonging to large mammals, small mammals, amphibians and reptiles, which have been identified at genus/species level. They are distributed among 37 taxa, corresponding to a minimum number of 328 individuals. The analysis of the fauna has allowed the reconstruction of the lifestyle of the human groups that inhabited Lumentxa, and which originated part of the large mammal accumulation from the Neolithic to Bronze Age periods. Changes in food production, in natural resource management and in settlement patterns originated a new way in which humans and the environment interacted.

Se describen por primera vez restos fósiles de cocodrilo del Mioceno Inferior (Formación Tudela, Rambliense, Biozonas MN2b-3) de las Bardenas Reales de Navarra, región situada en el sector noroccidental de la Depresión del Ebro. El material consiste en fragmentos craneales y mandibulares, dientes, huesos del esqueleto axial y apendicular y del escudo dérmico. La mayor parte dl los restos están desarticulados y pertenecen a varios individuos, habiéndose reconocido tanto formas juveniles como adultas. La escasa variabilidad morfológica observada en la asociación fósil sugiere que todos los restos pertenecen al mismo taxón. El material se atribuye a Diplocynodon sp. (Eusuchia, Alligatoroidea). Dicha atribución se basa en caracteres mandibulares, como la presencia de alveolos dentarios tercero y cuarto del mismo tamaño y confluentes, esplenial excluido de la sínfisis mandibular, y en rasgos de los osteodermos ventrales, que están formados por osificaciones pares. La principal característica de los dentarios es una sínfisis mandibular muy corta, que no sobrepasa en longitud el borde posterior del tercer alveolo. Los yacimientos de las Bardenas Reales de Navarra han proporcionado el material más abundante y diverso de Diplocynodon conocido en el Mioceno de la Península Ibérica.

ABSTRACT Abundant turtle fossils from two lower Miocene localities in which this lineage had not previously been studied are described herein. The localities are Monte Pila and Fuenmayor, which are 10 km apart, both located in the Autonomous Community of La Rioja (northern Spain), in the Ebro Basin. Information about the turtles from the lower Miocene levels in this basin was until now restricted to the finds made in the MN3 biozone of the Bardenas Reales of Navarre. Several turtle taxa had been recognized there. However, the material attributed to each of them was scarce. Monte Pila and Fuenmayor are located in biozone MN2. The remains of turtles are abundant in both sites. A terrestrial lineage, recognized at Monte Pila, is not represented in the Bardenas Reales fauna of Navarre. It represents the first record of Titanochelon Pérez-García & Vlachos, 2014 in La Rioja and is one of the oldest member of this genus, being assigned to Titanochelon cf. bolivari (Hernández-Pacheco, 1917). The only lineage of turtles identified in Fuenmayor, and the most abundant in Monte Pila, is Ptychogasterinae. All material of this clade of freshwater turtles identified there is compatible with a single form, for which a high range of intraspecific variability is recognized. It is attributed to Ptychogaster (Temnoclemmys) cf. bardenensis Murelaga, Lapparent de Broin, Pereda Suberbiola & Astibia, 1999, this species having been recognized, until now, as exclusive to the Bardenas Reales of Navarre.

Abstract. Sites corresponding to Marine Isotope Stage MIS 5c and MIS 4 like Artazu VIII, with a continental record but no hominin imprints or carnivore activities, are very scarce in the Iberian Peninsula. The Artazu VIII cave fill (Arrasate, northern Spain) was discovered in 2013 and is presented here for the first time, including detailed stratigraphic and sedimentological descriptions of the differentiated 12 levels, as well as an ordered chronology of most levels, covering at least 36.000 years. In addition, the small mammal assemblage yielded a total of 8,129 identifiable skeletal remains corresponding to 14 taxa. The conditions inferred from the ecological preferences of the small mammal associations have been correlated with four different stages and substages from MIS 5c to MIS 4 in which the woodland mass expanded and retreated, depending on the relative temperature and humidity in each period. Finally, considering the species variability and estimated palaeoenvironment, a comparison with the NGRIP δ18O curve and other sites with the same chronology has been performed, showing that Artazu VIII is one of the few deposits attributable to the Marine Isotopic Stages previously mentioned and the only one that correlates to the Greenland Interstadial 18 from the Iberian Peninsula.

Bothremydidae is the most abundant clade of turtles in the Campanian and Maastrichtian (Upper Cretaceous) fossil record of southwestern Europe. Several members of Foxemydina Gaffney, Tong & Meylan, 2006 are known in an area that includes Southern France and the North-Eastern half of Spain. The problematic ‘Polysternon’ atlanticum is the worst characterized, lacking a diagnosis that allows its specific validity to be confirmed, and whose generic attribution has been recognized as doubtful. Its presence was exclusively proposed in its type locality, the upper Campanian quarry of Laño, in Treviño County (Burgos Province, North of Spain). Despite the fact that knowledge about Bothremydidae has markedly increased after the description of ‘Polysternon’ atlanticum Lapparent de Broin & Murelaga, 1996, no new information about this species has been published since the 1990s. The analysis of abundant unpublished material of the bothremydid from Laño allows us to confirm the validity of this species. As a consequence of this study, it is not only identified in its type locality, but also in other Spanish regions and in the south of France. The diversity of Bothremydidae Baur, 1891 in the Upper Cretaceous of Europe is lower than previously considered. Thus, the species ‘Iberoccitanemys convenarum’ (Laurent, Tong & Claude, 2002), originally defined for the French record, and subsequently also identified in Spain, is identified here as a synonym of the species described in Laño. An emended diagnosis for the upper Campanian to upper Maastrichtian, Iberoccitanemys atlanticum (Lapparent de Broin & Murelaga, 1996) n. comb., is proposed.