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The Sobrarbe-Pirineos UNESCO Global Geopark shows an extremely well-developed underground karst relief as a result of the great abundance and thickness of its limestone formations. The most important Pleistocene vertebrate site within the Geopark is Coro Tracito Cave at Tella. The fossil association is made up exclusively of bones belonging to Ursus spelaeus from the upper Pleistocene, accumulated over several thousand years. Based on scientific analysis of the fossil bones, an interesting public outreach project has been organized, involving the refurbishment of the site within the cave and the creation of a permanent exhibition called the Tella Cave Bear Museum. These two infrastructures are visited by thousands of tourists each year and constitute the main geoscientific tourist attraction of the Sobrarbe-Pirineos Geopark.

The red-toothed shrews (Soricinae) are the most widespread subfamily of shrews, distributed from northern South America to North America and Eurasia. Within this subfamily, the tribe Nectogalini includes the fossil species Nesiotites hidalgo recorded from the Late Pleistocene to Holocene of the Balearic Islands (Western Mediterranean). Although there is a consensus about the close relationship between the extinct red-toothed shrew genera Nesiotites and Asoriculus based on morphology, molecular data are necessary to further evaluate the phylogenetic relationships of the Balearic fossils. We obtained a near complete mitochondrial genome of N. hidalgo, allowing the first molecular phylogenetic analysis of this species. Analyses based on 15, 167 bp of the mitochondrial genome placed N. hidalgo as close relative to the extant Himalayan shrew (Soriculus nigrescens), and a combined analysis using molecular and morphological data confirm that N. hidalgo and Asoriculus gibberodon are sister-taxa with S. nigrescens as the immediate outgroup. Molecular clock and divergence estimates suggest that the split between N. hidalgo and its closest living relative occurred around 6.44 Ma, which is in agreement with the previously proposed colonisation of the Balearic Islands from mainland Europe by nectogaline shrews during the Messinian Salinity Crisis (5.97–5.33 My ago). Our results highlight that it is possible to retrieve genetic data from extinct small mammals from marginal environments for DNA preservation. Additional finds from the fossil record of Soricinae from the Eurasian Late Miocene/Early Pliocene are needed to shed further light on the still confusing taxonomy and paleobiogeography of this clade.

Current knowledge of the evolution of the postcranial skeleton in the genus Homo is hampered by a geographically and chronologically scattered fossil record. Here we present a complete characterization of the postcranium of the middle Pleistocene paleodeme from the Sima de los Huesos (SH) and its paleobiological implications. The SH hominins show the following: (i) wide bodies, a plesiomorphic char- acter in the genus Homo inherited from their early hominin ancestors; (ii) statures that can be found in modern human middle-latitude pop- ulations that first appeared 1.6–1.5 Mya; and (iii) large femoral heads in some individuals, a trait that first appeared during the middle Pleistocene in Africa and Europe. The intrapopulational size variation in SH shows that the level of dimorphism was similar to modern humans (MH), but the SH hominins were less encephalized than Ne- andertals. SH shares many postcranial anatomical features with Ne- andertals. Although most of these features appear to be either plesiomorphic retentions or are of uncertain phylogenetic polarity, a few represent Neandertal apomorphies. Nevertheless, the full suite of Neandertal-derived features is not yet present in the SH popula- tion. The postcranial evidence is consistent with the hypothesis based on the cranial morphology that the SH hominins are a sister group to the later Neandertals. Comparison of the SH postcranial skeleton to other hominins suggests that the evolution of the postcranium oc- curred in a mosaic mode, both at a general and at a detailed level.

Galve (Teruel, Spain) is not a single fossil site but an ensemble of more than 50 localities with Mesozoic terrestrial vertebrate remains of Tithonian-Lower Barremian age. Galve fossil sites belong to the Maestrazgo basin (Central Iberian Range), Gaive Subbasin, and appear in four geological formations: Higueruelas Formation (Tithonian), Villar del Arzobispo Formation (upper Tithonian-middle Berriasian), El Castellar Formation (uppermost Hauterivianlowermost Barremian) and Camarillas Formation (lower Barremian). Most sites contain bony remains, but there are also paleoichnological and paleoological sites, with dinosaur and other reptilian tracks and eggshells. Mammals and dinosaurs are the best-known Gaive vertebrates, but some studies on sharks, bony fishes, amphibians, squamates and crocodiles have been published. Some groups, like turtles and pterosaurs, have not practically been studied yet. Galve is the type locality of severa1 taxa: the shark Lonchidion microselachos, the amphibian Galverpeton ibericum, the dinosaur Aragosaurus ischiaticus, the mammals Galveodon nannothus, Lavocatia alfambrensis, Eobaatar hispanicus, Parendotherium herreroi, Spalacotherium henkeli and Pocamus pepelui, and the dinosaur eggshell Macroolithus turolensis. In this paper we revise the knowledge on Galve vertebrates, and update the faunal list of its fossil sites, taking into account, for the first time, the stratigraphic distribution of the taxa.En Galve (Teruel) hay más de 50 localidades con restos de vertebrados continentales en las Formaciones Higueruelas (Titónico), Villar del Arzobispo (Titánico superior-Berriasiense medio), El Castellar (Hauteriviense terminal-Barremiense basal) y Camarillas (Barremiense inferior). Por tanto, «Galve» no es un único yacimiento de vertebrados mesozoicos, sino una localidad con numerosos yacimientos de vertebrados del intervalo Titónico-Barremiense, que geológicamente pertenecen a la Cuenca cretácica inferior del Maestrazgo (Cordillera Ibérica Central), Subcuenca de Galve. La mayor parte de los yacimientos contienen restos óseos, pero también son abundantes los yacimientos paleoicnológicos y paleoológicos. Los vertebrados mejor conocidos son los mamíferos y dinosaurios, aunque también hay estudios sobre los tiburones, peces óseos, anfibios, escamosos y cocodrilos. Algunos grupos, como tortugas y pterosaurios, permanecen prácticamente sin estudiar. Galve es la localidad tipo de varios taxones de vertebrados: el tiburón Lonchidion microselachos, el anfibio Galverpeton ibericum, el dinosaurio Aragosaurus ischiaticus, los mamíferos Galveodon nannothus, Lavocatia alfambrensis, Eobaatar hispanicus, Parendotherium herreroi, Spalacotherium henkeli y Pocamus pepelui, y la cáscara de huevo de dinosaurio Macroolithus turolensis. Es este trabajo se revisa el estado de conocimientos sobre los vertebrados de Galve, y se actualiza la lista faunística de sus yacimientos, siendo la primera vez que se realiza teniendo en cuenta la distribución estratigráfica de los taxones.

Galve (Teruel, Spain) is not a single fossil site but an ensemble of more than 50 localities with Mesozoic terrestrial vertebrate remains of Tithonian-Lower Barremian age. Galve fossil sites belong to the Maestrazgo basin (Central Iberian Range), Gaive Subbasin, and appear in four geological formations: Higueruelas Formation (Tithonian), Villar del Arzobispo Formation (upper Tithonian-middle Berriasian), El Castellar Formation (uppermost Hauterivianlowermost Barremian) and Camarillas Formation (lower Barremian). Most sites contain bony remains, but there are also paleoichnological and paleoological sites, with dinosaur and other reptilian tracks and eggshells. Mammals and dinosaurs are the best-known Gaive vertebrates, but some studies on sharks, bony fishes, amphibians, squamates and crocodiles have been published. Some groups, like turtles and pterosaurs, have not practically been studied yet. Galve is the type locality of severa1 taxa: the shark Lonchidion microselachos, the amphibian Galverpeton ibericum, the dinosaur Aragosaurus ischiaticus, the mammals Galveodon nannothus, Lavocatia alfambrensis, Eobaatar hispanicus, Parendotherium herreroi, Spalacotherium henkeli and Pocamus pepelui, and the dinosaur eggshell Macroolithus turolensis. In this paper we revise the knowledge on Galve vertebrates, and update the faunal list of its fossil sites, taking into account, for the first time, the stratigraphic distribution of the taxa. En Galve (Teruel) hay más de 50 localidades con restos de vertebrados continentales en las Formaciones Higueruelas (Titónico), Villar del Arzobispo (Titánico superior-Berriasiense medio), El Castellar (Hauteriviense terminal-Barremiense basal) y Camarillas (Barremiense inferior). Por tanto, «Galve» no es un único yacimiento de vertebrados mesozoicos, sino una localidad con numerosos yacimientos de vertebrados del intervalo Titónico-Barremiense, que geológicamente pertenecen a la Cuenca cretácica inferior del Maestrazgo (Cordillera Ibérica Central), Subcuenca de Galve. La mayor parte de los yacimientos contienen restos óseos, pero también son abundantes los yacimientos paleoicnológicos y paleoológicos. Los vertebrados mejor conocidos son los mamíferos y dinosaurios, aunque también hay estudios sobre los tiburones, peces óseos, anfibios, escamosos y cocodrilos. Algunos grupos, como tortugas y pterosaurios, permanecen prácticamente sin estudiar. Galve es la localidad tipo de varios taxones de vertebrados: el tiburón Lonchidion microselachos, el anfibio Galverpeton ibericum, el dinosaurio Aragosaurus ischiaticus, los mamíferos Galveodon nannothus, Lavocatia alfambrensis, Eobaatar hispanicus, Parendotherium herreroi, Spalacotherium henkeli y Pocamus pepelui, y la cáscara de huevo de dinosaurio Macroolithus turolensis. Es este trabajo se revisa el estado de conocimientos sobre los vertebrados de Galve, y se actualiza la lista faunística de sus yacimientos, siendo la primera vez que se realiza teniendo en cuenta la distribución estratigráfica de los taxones.

10.1016/j.quascirev.2014.05.031 The first migration out of Africa undertaken by the genus Homo is documented in Georgia at 1.8 Myr (Dmanisi) and some 0.4 Myr afterwards in the Middle East (¿Ubeidiya). However, the debate on when the European continent was populated for the first time remains open. The first human presence in Europe prior to the Jaramillo subchron (1.07¿0.99 Myr) is evidenced at Fuente Nueva 3 and Barranco León D (Orce) and at Sima del Elefante (Atapuerca), an occupation that seems to have continued through the Jaramillo at Gran Dolina TD3¿4 and TD5 (Atapuerca), at Vallparadís (Barcelona), and up to the Matuyama¿Brunhes boundary at Gran Dolina TD6. Even so, those who still defend a `short chronology¿ espouse an intermittent early population limited to the Mediterranean area, delaying the first occupation until after the Jaramillo. These hypotheses fail to explain what factors were behind the absence of population in Europe prior to this period, bearing in mind that there were populations of hominins at the gates of Europe between 1 and 0.5 Myr before the first archaeological record documented in Western Europe. Paleomagnetic analyses of the archaeological sites are rarely able to detect the Jaramillo subchron due to its short duration, while the radiometric dating methods (U-series/ESR) usually applied are limited in the accuracy they can achieve for the chronologies in question. These limitations make it necessary to depend on the biostratigraphy of small and large mammals to ascertain with precision the time of the first colonization of the continent. Accordingly, in the present article we discuss the chronological data from the older Iberian archaeological sites using biostratigraphic data to establish an archaeological sequence that demonstrates the

The Middle Pleistocene coincides with an important climate change, from this time ahead, the 100ky glacial cycles become a dominant feature. The opening of the landscapes, and a series of dispersal events of prairie taxa as the microtines, cricetids, ungulates and carnivores as well as Homo heidelbergensis mark this event. The rareness of fossil faunas of this age makes the Cueva de los Huesos de Obon one of the most important localities of this age in Aragon. Here we provide the first account on the Middle Pleistocene mammals from the Cueva de los Huesos, Obon, Teruel. The fossil mammalian species recovered in Obon are: Canis lupus, Vulpes vulpes, Felis sylvestris, Meles meles Hyaenidae and Carnivora indet., Stephanorhinus cf. hemitoechus, Equus sp. Gr. E. mosbachensis, Cervus elaphus priscus, Hemitragus sp. Erinaceus europaeus, Rhinolophus sp., Allocricetus bursae correzensis, Apodemus sp. This association is somewhat similar to that in the Trinchera Galería, Trinchera Dolina 8b-11 and Sima de los Huesos localities of Atapuerca

The Middle Pleistocene coincides with an important climate change, from this time ahead, the 100ky glacial cycles become a dominant feature. The opening of the landscapes, and a series of dispersal events of prairie taxa as the microtines, cricetids, ungulates and carnivores as well as Homo heidelbergensis mark this event. The rareness of fossil faunas of this age makes the Cueva de los Huesos de Obon one of the most important localities of this age in Aragon. Here we provide the first account on the Middle Pleistocene mammals from the Cueva de los Huesos, Obon, Teruel. The fossil mammalian species recovered in Obon are: Canis lupus, Vulpes vulpes, Felis sylvestris, Meles meles Hyaenidae and Carnivora indet., Stephanorhinus cf. hemitoechus, Equus sp. Gr. E. mosbachensis, Cervus elaphus priscus, Hemitragus sp. Erinaceus europaeus, Rhinolophus sp., Allocricetus bursae correzensis, Apodemus sp. This association is somewhat similar to that in the Trinchera Galería, Trinchera Dolina 8b-11 and Sima de los Huesos localities of Atapuerca

The association of dinosaurs of the Villar del Arzobispo Formation in Galve (Teruel, Aragon, Spain) represents a «jurassic association» by the abundance of primitive clades like basal neosauropods and the scarcity of ornithopods. This fact could indicate the persistence of the Jurassic dinosaur faunas in this part of Laurasia, at least up to the Berriasian. Anyway the absence of stegosaurians could be interpreted as the first evidence of the turnover that would come in the Cretaceous faunas. Our data indicate that the climatic change did not suddenly affected to the dinosaurs of the Tethyan province. On the contrary, the turnover to the modern faunas took place gradually at least during the Berriasian

The association of dinosaurs of the Villar del Arzobispo Formation in Galve (Teruel, Aragon, Spain) represents a «jurassic association» by the abundance of primitive clades like basal neosauropods and the scarcity of ornithopods. This fact could indicate the persistence of the Jurassic dinosaur faunas in this part of Laurasia, at least up to the Berriasian. Anyway the absence of stegosaurians could be interpreted as the first evidence of the turnover that would come in the Cretaceous faunas. Our data indicate that the climatic change did not suddenly affected to the dinosaurs of the Tethyan province. On the contrary, the turnover to the modern faunas took place gradually at least during the Berriasian

An elephant tusk has been found in a fluvial terrace of a small creek affluent of the Martin river. The tooth is slightly curved, almost round in section and becomes increasingly smaller towards the tip. The specimens measurements and morphology lead us to propose it as a task of the "wood elephant" of the Pleistocene of Europe: Paleoloxodon antiquus, probably of an adult male

An elephant tusk has been found in a fluvial terrace of a small creek affluent of the Martin river. The tooth is slightly curved, almost round in section and becomes increasingly smaller towards the tip. The specimens measurements and morphology lead us to propose it as a task of the "wood elephant" of the Pleistocene of Europe: Paleoloxodon antiquus, probably of an adult male

In this paper we present the first results of metrical analysis of premolar and molar lengths and metacarpal length and width of Ursus spelaeus Ros.-Hein. from Coro Tracito cave, the first cave bear high mountain site in Spain. According to their dental dentition size the Coro Tracito bears can be grouped into de the ?normal sized? cave bears of Spain. Their slender metacarpal bones place them between into the Ismail paw? cave bear group. The lower dentine aspartic add razemization ratios allowd us to date the Coro Tracito cave bear population as a tardy population that probably inhabited the area at the 3 Oxigen Stage Episode

Discoveries of isolated teeth of dromeosaurid (theropod) dinosaurs from the Upper Barremian of Spain, provided the first evidence that the range of dromeosaurid extended to southern Europe in the Lower Cretaceous. We report here the dromeosaurid teeth from Vallipon (Artoles Formation) in the Iberian Chain (Teruel). The teeth of Vallipon are distinguished from other curved theropod teeth by their strong lateral compression, by the shape of their denticles and by small size. Generally denticles appear only the distal carinae of the teeth. Nevertheless some teeth have denticles on both anterior and posterior margins, although the mesial denticles are much smaller than the distal denticles

Discoveries of isolated teeth of dromeosaurid (theropod) dinosaurs from the Upper Barremian of Spain, provided the first evidence that the range of dromeosaurid extended to southern Europe in the Lower Cretaceous. We report here the dromeosaurid teeth from Vallipon (Artoles Formation) in the Iberian Chain (Teruel). The teeth of Vallipon are distinguished from other curved theropod teeth by their strong lateral compression, by the shape of their denticles and by small size. Generally denticles appear only the distal carinae of the teeth. Nevertheless some teeth have denticles on both anterior and posterior margins, although the mesial denticles are much smaller than the distal denticles

In this paper we present the first results of metrical analysis of premolar and molar lengths and metacarpal length and width of Ursus spelaeus Ros.-Hein. from Coro Tracito cave, the first cave bear high mountain site in Spain. According to their dental dentition size the Coro Tracito bears can be grouped into de the ?normal sized? cave bears of Spain. Their slender metacarpal bones place them between into the Ismail paw? cave bear group. The lower dentine aspartic add razemization ratios allowd us to date the Coro Tracito cave bear population as a tardy population that probably inhabited the area at the 3 Oxigen Stage Episode

The Iberomys lineage is represented by the species I. huescarensis (Ruiz Bustos, 1988), at the lower levels (TD3 to lower TD8) of Trinchera Dolina (Sierra de Atapuerca, Burgos), and not by I. brecciensis Ciebel, 1847 as stated previously. Both species mainly differ in the communication of triangles T4 and T5 of the first lower molar dental pattern, broad in I. huescarensis and narrow in I. brecciensis. I. huescarensis is the ancestral species of the I. brecciensis-cabrerae lineage, and characterizes the late Lower Pleistocene and the transition Lower-Middle Pleistocene in the Iberian Peninsula

Up to the moment, two species of Iguanodon had been cited in the Lower Cretaceous of Spain: I. cf. atherfieldensis and I. bernissartensis. In general, the size has been the diagnostic character to separate I. atherfieldensis from I. bernissartensis, and many of the spanish Iguanodon remains would be regarded as Iguanodontidae indet or Iguanodon sp. A third species (1. cf. fittoni) is situated in the Jurassic-Cretaceous transition

Up to the moment, two species of Iguanodon had been cited in the Lower Cretaceous of Spain: I. cf. atherfieldensis and I. bernissartensis. In general, the size has been the diagnostic character to separate I. atherfieldensis from I. bernissartensis, and many of the spanish Iguanodon remains would be regarded as Iguanodontidae indet or Iguanodon sp. A third species (1. cf. fittoni) is situated in the Jurassic-Cretaceous transition

The Iberomys lineage is represented by the species I. huescarensis (Ruiz Bustos, 1988), at the lower levels (TD3 to lower TD8) of Trinchera Dolina (Sierra de Atapuerca, Burgos), and not by I. brecciensis Ciebel, 1847 as stated previously. Both species mainly differ in the communication of triangles T4 and T5 of the first lower molar dental pattern, broad in I. huescarensis and narrow in I. brecciensis. I. huescarensis is the ancestral species of the I. brecciensis-cabrerae lineage, and characterizes the late Lower Pleistocene and the transition Lower-Middle Pleistocene in the Iberian Peninsula

One new locality in the continental deposits of Almazán Basin is studied. The Rodentia fauna: Eucricetodon aff. aquitanicus, Pseudodryomys simplicidens, Peridyromys murinus y Heteroxerus rubricati is characteristc of the Z local Zone of the Ramblian (Lower Miocene). Also several Lagomorpha, Lipotyphla and Artiodactyla indeterminados have been found, Estudio de una nueva localidad en los depósitos de la Cuenca De Almazán. La fauna Rodentia: Eucricetodon aff. aquitanicus, Pseudodryomys simplicidens, Peridyromys murinus y Heteroxerus rubricati es característic de la Zona local Z del Ramblar (Mioceno inferior). También se han encontrado varios Lagomorpha, Lipotyphla y Artiodactyla indeterminados.

One new locality in the continental deposits of Almazán Basin is studied. The Rodentia fauna: Eucricetodon aff. aquitanicus, Pseudodryomys simplicidens, Peridyromys murinus y Heteroxerus rubricati is characteristc of the Z local Zone of the Ramblian (Lower Miocene). Also several Lagomorpha, Lipotyphla and Artiodactyla indeterminados have been found

The Sciuridae are the family of rodents that comprises all extant and fossil squirrels. In the Ramblian and especially in the Aragonian of the Tertiary basin of Calatayud-Montalbán (Aragón, Spain) the remains of Sciuridae are relatively abundant. A major problem solved in this paper is the distinction of Ml and M2, both in the upper and the lower dentition. Traditionally, in the literature these elements are treated as a unit Ml,2. We managed to separate them, on the basis of the study of material of two extant species, Atlantoxerus getulus and Sciurus vulgaris. In our study of the fossil Sciuridae from the Calatayud-Montalbán Basin 9 species were recognized, belonging to 6 genera: Heteroxerus rubricati, H. grivensis, Atlantoxerus idubedensis sp. nov., A. blacki, Aragoxerus ignis, Spermophilinus besana sp. nov., S. bredai, Freudenthalia daamsi sp. nov. y Palaeosciurus feignouxi. Aragoxerus and Freudenthalia are new genera. Atlantoxerus idubedensis sp. nov. is a small Atlantoxerus, similar to primitive Heteroxerus, and characterized by a reduced or absent anterior cingulum in M1, M2 and M3. Spermophilinus besana sp. nov. is the smallest Spermophilinus species known so far. In the lower teeth the anterolophid is separated from the protoconid by a deep furrow, and the entoconid is reduced or absent. Aragoxerus gen. nov. (tribe Xerini) contains only one species, A. ignis, formerly assigned to Heteroxerus. Although Aragoxerus is quite similar to Heteroxerus, its smaller size, simple dental pattern, absence of metaconule and anterior cingulum, and the lack of accessory ornamentations permit us to distinguish it as a separate genus. Freudenthalia gen. nov. is a monospecific genus of uncertain systematic position, tentatively placed in the tribe Sciurini. Freudenthalia daamsi sp. nov. is the only species of this genus. It is characterized by a metaloph completely fused to the protocone, a loop-shaped metaconule, and the absence of the entolophid. Within the genus Heteroxerus two pairs of names are synonimized: H. vireti is a synonym of H. ru