The fossil American lion (Panthera atrox) in South America: Palaeobiogeographical implications

By the late 19th and early 20th centuries, authors described several specimens belonging to a very large felid, the size of a lion, from some Late Pleistocene localities at southern Chile and Argentina. These remains were considered as belonging to large and now extinct subspecies of jaguar. In the present contribution based on qualitative and quantitative postcranial and cranial characters, we conclude that that the “Patagonian Panthera” should be considered as representing remains of the American Cave Lion Panthera atrox. Evidence at hand indicates that Panthera atrox was a very large lion-like felid, with reddish skin, that inhabited dry open habitats. Furthermore, in contrast to living jaguars, transported and accumulated its preys in rocky dens. The presence of at least two pantherine lineages (i.e. P. onca and P. atrox) in the Pleistocene of South America indicates that the lineage has a long and complex history in the New World, and that the history of the clade in this landmass is still far from being well understood.     

The geographical and chronological pattern of herpetofaunal Pleistocene extinctions on the Iberian Peninsula

Despite the postulation of the “herpetofaunal stability hypothesis”, which suggests that the herpetofauna underwent relatively few changes during the Pleistocene, the amphibian and squamate faunas of western Europe are known to have progressively diminished during the Pliocene and the beginning of the Pleistocene. Iberian Early Pleistocene sites continue to document the presence of “exotic” herpetofaunal elements that are supposed to have disappeared from the mainland, such as Oriental vipers, agamid lizards, the anguids Pseudopus and Dopasia, and a representative of the green toad group (Bufo viridis sensu lato), as well as possibly gekkonids, a scincid lizard (Lygosominae), the anguid Ragesaurus and blind snake (Scolecophidia). The geographical and temporal pattern of these progressive southward withdrawals and extirpations on the Iberian Peninsula shows that extirpation events occurred in northern Spain at roughly the Olduvai paleomagnetic event and in southern Spain just before the Jaramillo paleomagnetic reversal, thus permitting us to establish a framework that can be used to complement the biochronological zonation of the Iberian Early Pleistocene.     

A new Late Pleistocene non-anthropogenic vertebrate assemblage from the northern Iberian Peninsula: Artazu VII (Arrasate,Basque Country)

Late Pleistocene palaeontological sites without human intervention are limited in the Cantabrian region, and even more so those with a good state of preservation and rich biodiversity. A new vertebrate fossiliferous locality discovered at Kobate Quarry (Arrasate, northern Iberian Peninsula) is presented in this article. This site, in which remains of 40 different vertebrate taxa were accumulated, acted as a natural trap. The preliminary vertebrate faunal list includes five amphibian taxa, four reptiles, seven species of birds and 24 mammalian taxa. While small mammals are represented by 13 small mammal taxa (seven in the Order Rodentia, five in the Order Eulipotyphla, and one in the Order Chiroptera), the large mammal fauna comprises eleven species, including ungulates and carnivores. The palaeoecology inferred from this faunal assemblage suggests the existence of large forested areas with some grassland and a watercourse nearby, within a notably warm and humid climate. These palaeoenvironmental conditions, combined with AMS and AAR results carried out in macrofaunal bone samples, suggest that the deposit from Artazu VII would be located in the first half of the Late Pleistocene, in the Marine Isotope Stage (MIS) 5c.     

A Neandertal mandible from the Cova del Gegant (Sitges, Barcelona, Spain)

A human mandible from the site of Cova del Gegant is described here for the first time and compared with other Middle and Upper Pleistocene representatives of the genus Homo from Europe and Southwest Asia. The specimen was recovered from sediments which also yielded Mousterian stone tools and Pleistocene fauna. The preserved morphology of the mandible, particularly in the region of the mental foramen, clearly aligns it with the Neandertals, making the Cova del Gegant the only known site in Catalonia documenting diagnostic human skeletal remains in association with Middle Paleolithic stone tools. This represents an important new addition to the human fossil record from the Iberian Peninsula and joins the Ba?olas mandible in documenting the course of human evolution in the northern Mediterranean region of Spain.     

The distinction between the extinct Pleistocene European Crane,Grus primigenia,and the extant Asian Sarus Crane,G. antigone

In contradiction with the conclusions of some previousworkers, the extant Sarus Crane, Grus antigone (L.), and the extinct large Crane, Grus primigeniaMilne-Edwards, from the Pleistocene of Europe, cannot be considered as conspecific. In fact, the tibiotarsal epicondyles, particularly the medial epicondyle, the distal end of the humerus, its ventral epicondyle and the adjoining process, are comparatively smaller in Grus primigenia. These findings indicate differences in locomotion.     

The fossil bat assemblage of Sima del Elefante Lower Red Unit (Atapuerca,Spain): First results and contribution to the palaeoenvironmental approach to the site

The fossil bat assemblage from the Lower Red Unit of Sima del Elefante (TELRU) in Atapuerca (Burgos, Spain) has been exhaustively analysed for the first time. Bat fossil assemblages are of particular relevance to palaeoenvironmental approaches to sites. Here we integrate our new data on the chiropteran fauna with the data provided previously by other authors on the basis of the small-vertebrate assemblages and the palynology of the site. Our results are consistent with the earlier results in that they indicate a generally warmer climate than at present in the area and stable environmental conditions throughout the major part of the TELRU sequence. However, fossil bat assemblages sometimes lead to problems when they are used in landscape reconstruction. These problems are discussed here and should be taken into account in future works.     

Biogeography and chronology of the Eurasian extinct rhinoceros Stephanorhinus etruscus (Mammalia,Rhinocerotidae)

Stephanorhinus etruscus is one of the most abundantly recorded and better known Eurasian Early Pleistocene rhinoceroses. Nevertheless, the first and last appearances of this species, as well as its paleogeographic distribution, are controversial and debated in literature. S. etruscus is documented since the latest Pliocene in Spain (Las Higueruelas), Italy (Montopoli and Castelnuovo di Barardenga), France (Perrier–Les Étouaires) and Romania (Iar??–Cariera Veche). During the Early Pleistocene, S. etruscus occurred in several Spanish, French and Italian localities, as well as in The Netherlands (e.g., Tegelen), Germany (e.g., Thiede), Greece (e.g., Aivaliki) and Israel (e.g., Ubeidiya). The last appearance of S. etruscus in Eurasia is debatable. Etruscan rhino populations survived till the Jaramillo subchrone (around 1.1 Ma) in France (Bois-de-Riquet), Romania (Betfia XII) and Hungary (Osztramos 2 and 8), and close to the early–middle Pleistocene transition in Spain (Cueva Victoria, Huéscar 1, Atapuerca TD4, TD6 and TD8), and Italy (Monte delle Piche).     

Palaeoenvironment and palaeoclimate in the western Liguria region (northwestern Italy) during the Last Glacial. The small mammal sequence of Riparo Mochi (Balzi Rossi,Ventimiglia)

Riparo Mochi is considered a key site for the Middle–Upper Palaeolithic chrono-cultural sequence in the Mediterranean area. Climatic and environmental conditions have been reconstructed after the study of the small mammal assemblage by means of linear regression method and the Habitat Weighting method. The climate proxies for the late Mousterian and Proto-Aurignacian (cultural units I to G) indicate an environment with a relative high percentage of forest component during two cold and one relative warm oscillations. Open environments with low temperatures are registered for the Gravettian (cultural Unit D) allowing a possible relation to the Heinrich 3 Event. The final Gravettian (cultural Unit C) shows a relative warm peak, tentatively related to Greenland Interstadial 2. The late Epigravettian (cultural Unit A) is characterized by a high percentage of forest-related environment suggesting that this horizon formed during the Bølling–Allerød Interstadial. Finally, the Riparo Mochi sequence contributes to reconstruct the processes of faunal changes along the northern Tyrrhenian coast in a period of climatic instability, especially during the end of Marine Isotope Stage 3 when foraging groups adopted different mobility strategies between southern France and the Italian peninsula.     

Paleoecological and microenvironmental aspects of the first European hominids inferred from the taphonomy of small mammals (Sima del Elefante,Sierra de Atapuerca,Spain)

Unit TE9 of the Sima del Elefante (Sierra de Atapuerca, Burgos, Spain), where the remains of Homo sp. have been discovered (1.2–1.3 Ma), is also a level rich in small mammals. The taphonomic study of these small vertebrates sheds light on the landscape that provided the setting for the activities of these early hominids and allows us to describe what the cave was like during the formation of the level. Small mammal predators identified in the study indicate that during this period the Sierra de Atapuerca was part of a large biome consisting of semi-open riparian forests with meadows nearby. Postdepositional alterations reveal that the production of fossils took place outside the cave, being transported inside by water currents. During the formation of TE9, the cave presented conditions of high humidity, which made it difficult for hominids to establish occupations inside, although these conditions became somewhat less severe in TE9c, the sublevel where human remains are found.     

On some Pleistocene bird fossils from the south Aegean islandof Karpathos (Greece)

This paper deals with some Pleistocene fossil birds from the south Aegean island of Karpathos. The species which could be identified are surveyed, and the paleogeography of the area is discussed.     

Woody macroremains from the Acheulian site of Torralba: Occurrence and palaeoecology of Pinus cf. sylvestris in the Middle Pleistocene of the Iberian Peninsula

The pieces of wood extracted by the Marquis of Cerralbo at the Acheulian site of Torralba constitute one of the few materials from this famous locality that have not been subjected to detailed study by researchers. Over a century after their extraction, the present paper provides the first anatomical results of these 34 woody remains. The results highlight the presence of the taxon Pinus sylvestris–nigra group (Pinus cf. sylvestris) at the site, thus constituting one of the few and oldest references attributable to the aforementioned taxon for the Iberian Peninsula. The anatomical diagnosis confirms the existence of Middle Pleistocene landscapes with pine woods at elevations at least 200 m lower than at present in the site area. From a taphonomic point of view, the wood samples exhibit the typical morphologies of woody macroremains preserved in continental sedimentary environments, and there was no evidence at all that they had previously been manipulated by man.     

Middle to Late Pleistocene herpetofauna from Scladina and Sous-Saint-Paul caves (Namur,Belgium)

For the first time, the fossil herpetofauna from the Middle and Late Pleistocene of Scladina and Sous-Saint-Paul caves (Sclayn, Belgium) is described. The amphibians and squamate reptiles are represented by one salamander (Salamandra salamandra), three anurans (Pelodytes punctatus, Bufo bufo and Rana temporaria), two lizards (Lacerta cf. agilis and Anguis fragilis) and two snakes (Zamenis longissimus and Vipera cf. berus). The occurrence of the Parsley Frog (Pelodytes punctatus) and the Aesculapian Snake (Zamenis longissimus), which are not currently represented in Belgium, is of particular interest. Scladina also represents one of the northernmost fossil mentions for the Fire Salamander (Salamandra salamandra) although it is within its current distribution in Europe. Finally, the presence of the Adder (Vipera cf. berus) is very probably attested in Scladina whereas today this snake is infrequent and classified as endangered in Belgium.     

Paleoenvironmental reconstruction of the Early Pleistocene site of Quibas (SE Spain) using a rodent assemblage

In this paper we analyze the rodent assemblage from the Early Pleistocene site of Gruta1 (Quibas karstic complex, Murcia, SE Spain), providing taphonomic comments and paleoecological and paleoclimatic data. The studied assemblage includes 209 identified rodent teeth corresponding to at least four taxa. The karstic source of the site and the low presence of digested remains suggest that the accumulation is scatological in origin, with the influence of predators such as owls; the accumulation also shows certain evidence of slight hydrodynamic sorting. For the area around Quibas/Gruta1 the paleoecological study indicates a predominance of rocky areas (31.7%), forested environments (31.7%) and open dry meadows or shrublands (29.7%), which would indicate a mosaic forest environment, and to a lesser extent the presence of open humid land (5.4%) and areas along streams or ponds (1.5%). The distribution of the bioclimatic spectra yields the highest percentage for a Mediterranean climate (37.5%), while the climatic parameters calculated (MAT = 15 °C, MTW = 25.7 °C, MTC = 4.3 °C, MAP = 390 mm) suggest colder and slightly more humid conditions than today at the time of the deposition of the remains. The data also point to a relative decrease in temperature within the phase in which the site was produced, which possibly occurred during a relative cold period between MIS 36 and MIS 40 according to the age.     

A human parietal fragment from the late Early Pleistocene Gran Dolina-TD6 cave site,Sierra de Atapuerca,Spain

The TD6-2 level of the Gran Dolina cave site (Sierra de Atapuerca, Burgos, Spain) has yielded an assemblage of about 170 human fossil remains dated to > 800 ka (probably MIS 21) and assigned to the species Homo antecessor. In this study, we describe for the first time a large portion of a parietal bone (ATD6-100/168). The morphology of the fractures on the bones is compatible with a peri-mortem trauma. The superior parietal areas are flat. There is a large parietal foramen and one smaller accessory parietal foramen. Middle meningeal vessels are not particularly developed, but they are distributed in both anterior and posterior districts, with the parietal vasculature originating from the posterior branch. The meningeal vessels show multiple minor connections with the pericranial and diploic vascular systems. The diploe is not particularly developed, and large diploic channels are not detected. The bone is thin when compared with adult fossil humans, and equivalent to juvenile values. All these characters suggest that the parietal ATD6-100/168 probably belonged to a juvenile individual, with plesiomorphic endocranial traits similar to those described for H. ergaster/erectus. The derived temporal, maxillary and dental traits in Homo antecessor and the primitive parietal morphology further point to distinct (mosaic) patterns of morphological evolution of face and braincase.     

The Middle-to-Upper Palaeolithic transition in Cova Gran (Catalunya, Spain) and the extinction of Neanderthals in the Iberian Peninsula

The excavations carried out in Cova Gran de Santa Linya (Southeastern PrePyrenees, Catalunya, Spain) have unearthed a new archaeological sequence attributable to the Middle Palaeoloithic/Upper Palaeolithic (MP/UP) transition. This article presents data on the stratigraphy, archaeology, and ¹⁴C AMS dates of three Early Upper Palaeolithic and four Late Middle Palaeolithic levels excavated in Cova Gran. All these archaeological levels fall within the 34-32 ka time span, the temporal frame in which major events of Neanderthal extinction took place. The earliest Early Upper Palaeolithic (497D) and the latest Middle Palaeolithic (S1B) levels in Cova Gran are separated by a sterile gap and permit pinpointing the time period in which the Mousterian disappeared from Northeastern Spain. Technological differences between the Early Upper Palaeolithic and Late Middle Palaeolithic industries in Cova Gran support a cultural rupture between the two periods. A series of 12 ¹⁴C AMS dates prompts reflections on the validity of reconstructions based on radiocarbon data. Thus, results from excavations in Cova Gran lead us to discuss the scenarios relating the MP/UP transition in the Iberian Peninsula, a region considered a refuge of late Neanderthal populations.     

Palaeoreas,Majoreas, and Stryfnotherium gen. nov. (Mammalia: Artiodactyla: Bovidae) from the Late Miocene of Greece

A new comparative study of the two crania, LGPUT RZO-03 and LGPUT RZO-68 attributed to Palaeoreas zouavei Bouvrain, 1980 from the Late Miocene of Axios Valley, Greece reveals great morphological and metric differences between them, which require the exclusion of LGPUT RZO-68 from this species and even from Palaeoreas Gaudry, 1861. This decision leaves unsupported many diagnostic features of P. zouavei applied by previous authors and along with recent data from Bulgaria put in doubt the validity of P. zouavei as a whole. As a result, the genus Palaeoreas is re-defined as monotypic and the cranial morphometric changes of P. lindermayeri (Wagner, 1848) are discussed on a temporal basis. An updated taxonomic analysis confirms the presence of P. lindermayeri in the Late Turolian mammal assemblage of Dytiko-1 (Axios Valley, Greece) but rejects its occurrence at Dytiko-2. Instead, a re-examination of old cranial specimens and a study of new ones suggest the presence in both sites of Majoreas Kostopoulos, 2004 a Late Miocene taxon previously mostly known from the eastern banks of the Aegean Sea. A reassessment of the LGPUT RZO-68 cranium, and its comparison with Palaeoreas and Criotherium Forsyth-Major, 1891, suggests that it deserves taxonomic distinction at genus and species levels: Stryfnotherium exophthalmon.     

First record of Entelodontidae (Mammalia,Artiodactyla) from the late Eocene of Southeast Asia

We describe two entelodontid upper premolars that were recovered from the late Eocene of the Krabi coal mine in southern Thailand. The size and morphology of the material suggest that it can be referred to Entelodon aff. E. gobiensis, a species known from the late Eocene to the early Oligocene of northern Asia and southern China. The Thai material documents for the first time the southernmost occurrence of entelodontids in Asia during the Paleogene and also suggests that Eocene Southeast Asian mammal localities might potentially yield further entelodontid remains mostly associated with selenodont ungulates.     

Variability of the upper incisors in the cave bears (Carnivora,Ursidae) from the Caucasus and Urals

Morphometric and morphotypic variability of the cave bear upper incisors from two different geographic regions (Caucasus and Urals), different stratigraphic periods (middle and late Pleistocene), and bearing different mitochondrial haplogroups (kudarensis and ingressus) was studied. The specific diet of the cave bears, i.e. hard vegetables, led to noticeable differences between their incisors and the incisors of the brown bear (Ursus arctos). It was found that the upper incisors of the Caucasian cave bears from different stratigraphic periods demonstrate consistent development of their morphology. The late Pleistocene cave bears from the Urals show a greater similarity to the Caucasian cave bears from earlier periods than with the cave bears from later periods. Our results suggest that the incisor morphology has evolved independently in the Caucasian and Ural cave bears as they belong to different phylogenetic lineages and display different ways of adaptation to local environmental conditions.