Early Pleistocene stratigraphy, sedimentary environments, and formation contexts at Dmanisi in the Georgian Caucasus.

The Early Pleistocene site of Dmanisi is now well known for its large number of fossils of early Homo erectus as well as associated artifacts and faunal remains, recovered mainly in pipe-related geologic features. Testing in the M5 unit 100 m to the west of the main excavations revealed a thick stratigraphy with no evidence of pipes or gullies, indicating that the geologic record at Dmanisi included spatially distinct sedimentary environments that needed further investigation. Here we report the results of a geoarchaeological program to collect data bearing on contexts and formation processes over a large area of the promontory. That work has defined over 40,000 m2 of in situ deposits with artifacts and faunas. Stratum A ashes bury the uppermost Mashavera Basalt, which we have dated to 1.8 Ma in the M5 block. The Stratum A deposits contain stratified occupations that accumulated quickly and offer good potential for recovery of in situ materials. Stratum B1 deposits above the A/B unconformity include all of the pipe and gully facies at Dmanisi, reflecting a brief but very intense phase of geomorphic change. Those deposits contain the majority of faunas and all of the hominin fossils. B1 slope facies offer excellent formation contexts away from the piped area, and all B1 deposits are sealed by Stratum B2 over the whole promontory. Strata B2 to B5 register a return to slope facies, with no further evidence of pipes or gullies. Those deposits also present excellent contexts for recovery of in situ occupations. Overall, Dmanisi's geologic history preserves an exceptional record of the activities and environmental context of occupations during the first colonization of Eurasia.

Chronological and environmental context of the first hominin dispersal into Western Europe: The case of Barranco León (Guadix-Baza Basin, SE Spain).

The early Pleistocene site of Barranco León (Guadix-Baza Basin, SE Spain), dated to 1.4 Ma (millions of years ago), records the oldest hominin occurrence in Western Europe, as evidenced by the discovery of one tooth and thousands of Mode 1 stone tools. In this paper a detailed analysis of the microvertebrate content of the D1 and D2 units from this site is presented. The early Pleistocene in the Guadix-Baza Basin is characterized by a sharp climatic deterioration, which possibly impeded the settlement of this region by the early hominin population from the southern Caucasus. Shortly afterwards, when the climatic conditions were again favorable, a hominin presence is suddenly evidenced at the units D1 and D2 of Barranco León. According to the microvertebrate analysis of these units, the mean annual temperature at the time of deposition was significantly higher than 13 °C, with prevalent humid conditions. However, although most of the species were inhabitants of water edges, an open landscape was present in the vicinity of the lake. The data reported here clearly support the idea that the early hominin occupation of Europe was strongly constrained by climatic and environmental conditions, rather than by physiography or cultural factors.

Earliest human occupations at Dmanisi (Georgian Caucasus) dated to 1.85-1.78 Ma.

The early Pleistocene colonization of temperate Eurasia by Homo erectus was not only a significant biogeographic event but also a major evolutionary threshold. Dmanisi's rich collection of hominin fossils, revealing a population that was small-brained with both primitive and derived skeletal traits, has been dated to the earliest Upper Matuyama chron (ca. 1.77 Ma). Here we present archaeological and geologic evidence that push back Dmanisi's first occupations to shortly after 1.85 Ma and document repeated use of the site over the last half of the Olduvai subchron, 1.85-1.78 Ma. These discoveries show that the southern Caucasus was occupied repeatedly before Dmanisi's hominin fossil assemblage accumulated, strengthening the probability that this was part of a core area for the colonization of Eurasia. The secure age for Dmanisi's first occupations reveals that Eurasia was probably occupied before Homo erectus appears in the East African fossil record.

The oldest human fossil in Europe, from Orce (Spain).

The Orce region has one of the best late Pliocene and early Pleistocene continental paleobiological records of Europe. It is situated in the northeastern sector of the intramontane Guadix-Baza Basin (Granada, Andalusia, southern Spain). Here we describe a new fossil hominin tooth from the site of Barranco León, dated between 1.02 and 1.73 Ma (millions of years ago) by Electron Spin Resonance (ESR), which, in combination with paleomagnetic and biochronologic data, is estimated to be close to 1.4 Ma. While the range of dates obtained from these various methods overlaps with those published for the Sima del Elefante hominin locality (1.2 Ma), the overwhelming majority of evidence points to an older age. Thus, at the moment, the Barranco León hominin is the oldest from Western Europe.

Postcranial evidence from early Homo from Dmanisi, Georgia.

The Plio-Pleistocene site of Dmanisi, Georgia, has yielded a rich fossil and archaeological record documenting an early presence of the genus Homo outside Africa. Although the craniomandibular morphology of early Homo is well known as a result of finds from Dmanisi and African localities, data about its postcranial morphology are still relatively scarce. Here we describe newly excavated postcranial material from Dmanisi comprising a partial skeleton of an adolescent individual, associated with skull D2700/D2735, and the remains from three adult individuals. This material shows that the postcranial anatomy of the Dmanisi hominins has a surprising mosaic of primitive and derived features. The primitive features include a small body size, a low encephalization quotient and absence of humeral torsion; the derived features include modern-human-like body proportions and lower limb morphology indicative of the capability for long-distance travel. Thus, the earliest known hominins to have lived outside of Africa in the temperate zones of Eurasia did not yet display the full set of derived skeletal features.

Carbon isotopic record of terrestrial ecosystems spanning the Late Miocene extinction of Oreopithecus bambolii, Baccinello Basin (Tuscany, Italy).

Oreopithecus bambolii is a Late Miocene hominoid with an extensive fossil record in the Baccinello Basin (Tuscany, Italy), and was the only western European hominoid to survive a major extinction event ca. 9.6 Ma (millions of years ago). Oreopithecus lived in the insular Tusco-Sardinian paleobioprovince, where it evolved many unique anatomical specializations that make it important for understanding the mechanisms and history of Late Miocene hominoid evolution. The eventual extinction of Oreopithecus and its associated fauna ca. 6.5 Ma has generally been attributed to interaction with species that arrived from continental Europe following tectonic collision of the Tusco-Sardinian province with mainland Italy, but palynological, paleontological, and sedimentological records indicate an environmental shift toward more variable climate across the extinction event. To explore the possibility of environmental change as a contributing factor in the extinction of Oreopithecus, we developed a stable carbon and oxygen isotope record from organic matter in paleosols from the Baccinello Basin. These data show very low temporal and spatial variability (indicating plant ecosystem stability through time and space) and provide no evidence for ecologically significant changes in floral composition spanning the extinction event, suggesting that environmental change was not an underlying cause for the extinction of Oreopithecus and its associated fauna. The carbon isotope values fall entirely within the range of isotopic variability for modern plants following the C(3) photosynthetic pathway (trees, shrubs, cool-season grasses), indicating that C(4) vegetation (warm-season grasses) was not an important component of biomass. When corrected for temporal variation in the carbon isotopic composition of atmospheric carbon dioxide, the paleosol carbon isotope values are consistent with predicted values based on modern plants and the Baccinello palynoflora, supporting the reliability of paleosol isotopic records as paleoecological proxies.

Geochemical and sedimentary constraints on the formation of the Venta Micena early Pleistocene site (Guadix-Baza Basin, Spain).

Despite the paleontological relevance of the terrestrial Early Pleistocene Venta Micena bonebed (Baza Basin, Spain), it lacks a comprehensive geochemical/sedimentological study. Here, we demonstrate that the 1.5-m-thick Venta Micena limestone formed in a relatively small freshwater wetland/pond located at the periphery of the large saline Baza paleolake. Two microfacies are observed, with high and low contents of invertebrate fossils, and which originated in the centre and margin of the wetland, respectively. X-ray diffraction (XRD) mineralogy and paleohydrological characterization based on ostracod and bulk-rock geochemistry (δ13C and δ18O) indicate that the limestone reflects a general lowstand of the Baza lake, permitting the differentiation of freshwater wetlands that were fed by adjacent sources. Conversely, during highstands, the Baza lake flooded the Venta Micena area and the freshwater fauna was replaced by a saline one. Bulk-rock isotopic data indicate that the lower interval C1 of the limestone (bone-rich in marginal settings) displays general negative values, while the upper interval C2 (bone free) displays less negative values. The bones of predated mammals accumulated in the marginal areas, which were flooded and buried by recurring water-table fluctuations. Lake dynamics played a critical role in bone accumulation, which was previously considered as representing a hyena den.

Taphonomic and spatial analyses from the Early Pleistocene site of Venta Micena 4 (Orce, Guadix-Baza Basin, southern Spain).

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.

First continuous pre-Jaramillo to Jaramillo terrestrial vertebrate succession from Europe.

In this paper, the early Pleistocene small vertebrate sequence of Quibas-Sima (Quibas karstic complex, Murcia, SE Spain) is presented. The available magnetostratigraphic information together with the small vertebrate association, allow to reliably constrain the age of the different units. The basal unit of the section has recorded a reversed polarity assigned to the pre-Jaramillo Matuyama (C1r.2r, i.e., between 1.2 and 1.07 Ma). The intermediate units have recorded a normal polarity correlated directly with the Jaramillo subchron (C1r.1n, between 1.07 and 0.99 Ma), while the upper units record the post-Jaramillo reverse polarity (C1r.1r, i.e., between 0.99 and 0.78). Jaramillo subchron is especially significant regarding the earliest hominin dispersal in Western Europe. However, vertebrate faunas unambiguously correlatable with Jaramillo subchron are extremely rare in Europe. Thereby, the study of the Quibas-Sima sequence allows to characterize the vertebrate association synchronous to this paleomagnetic episode in southern Iberian Peninsula, and contributes to increase knowledge of the biotic and climatic events that took place in southern Europe at the beginning of the Early-Middle Pleistocene transition, prior to the Matuyama-Brunhes boundary. A continuous small vertebrate succession has been reported, including representatives of the families Bufonidae, Pelodytidae, Testudinidae, Gekkonidae, Blanidae, Lacertidae, Colubridae, Viperidae, Soricidae, Erinaceidae, Rhinolophidae, Vespertilionidae, Muridae, Gliridae, Sciuridae, Leporidae and Ochotonidae The ecological affinities of the faunal association suggest a progressive reduction in forest cover in the onset of the Jaramillo subchron.

Correction: Dynamic Similarity in Titanosaur Sauropods: Ichnological Evidence from the Fumanya Dinosaur Tracksite (Southern Pyrenees).

[This corrects the article DOI: 10.1371/journal.pone.0057408.].

An archosauromorph dominated ichnoassemblage in fluvial settings from the late Early Triassic of the Catalan Pyrenees (NE Iberian Peninsula).

The vertebrate recovery after the end-Permian mass extinction can be approached through the ichnological record, which is much more abundant than body fossils. The late Olenekian (Early Triassic) tetrapod ichnoassemblage of the Catalan Pyrenean Basin is the most complete and diverse of this age from Western Tethys. This extensional basin, composed of several depocenters, was formed in the latest phases of the Variscan orogeny (Pangea breakup) and was infilled by braided and meandering fluvial systems of the red-beds Buntsandstein facies. Abundant and diverse tetrapod ichnites are recorded in these facies, including Prorotodactylus mesaxonichnus isp. nov. (tracks possibly produced by euparkeriids), cf. Rotodactylus, at least two large chirotheriid morphotypes (archosauriform trackmakers), Rhynchosauroides cf. schochardti, two other undetermined Rhynchosauroides forms, an undetermined Morphotype A (archosauromorph trackmakers) and two types of Characichnos isp. (swimming traces, here associated to archosauromorph trackmakers). The Pyrenean ichnoassemblage suggests a relatively homogeneous ichnofaunal composition through the late Early Triassic of Central Pangea, characterized by the presence of Prorotodactylus and Rotodactylus. Small archosauromorph tracks dominate and present a wide distribution through the different fluviatile facies of the Triassic Pyrenean Basin, with large archosaurian footprints being present in a lesser degree. Archosauromorphs radiated and diversified through the Triassic vertebrate recovery, which ultimately lead to the archosaur and dinosaur dominance of the Mesozoic.

Ichnological evidence of Megalosaurid Dinosaurs Crossing Middle Jurassic Tidal Flats.

A new dinosaur tracksite in the Vale de Meios quarry (Serra de Aire Formation, Bathonian, Portugal)preserves more than 700 theropod tracks. They are organized in at least 80 unidirectional trackways arranged in a bimodal orientation pattern (W/NW and E/SE). Quantitative and qualitative comparisons reveal that the large tridactyl, elongated and asymmetric tracks resemble the typical Late Jurassic-Early Cretaceous Megalosauripus ichnogenus in all morphometric parameters. Few of the numerous tracks are preserved as elite tracks while the rest are preserved as different gradients of modified true tracks according to water content, erosive factors, radial fractures and internal overtrack formations. Taphonomical determinations are consistent with paleoenvironmental observations that indicate an inter-tidal flat located at the margin of a coastal barrier. The Megalosauripus tracks represent the oldest occurrence of this ichnotaxon and are attributed to large megalosaurid dinosaurs. Their occurrence in Vale de Meios tidal flat represents the unique paleoethological evidence of megalosaurids moving towards the lagoon, most likley during the low tide periods with feeding purposes.

Dynamic similarity in titanosaur sauropods: ichnological evidence from the Fumanya dinosaur tracksite (southern Pyrenees).

The study of a small sauropod trackway from the Late Cretaceous Fumanya tracksite (southern Pyrenees, Catalonia) and further comparisons with larger trackways from the same locality suggest a causative relationship between gait, gauge, and body proportions of the respective titanosaur trackmakers. This analysis, conducted in the context of scaling predictions and using geometric similarity and dynamic similarity hypotheses, reveals similar Froude numbers and relative stride lengths for both small and large trackmakers from Fumanya. Evidence for geometric similarity in these trackways suggests that titanosaurs of different sizes moved in a dynamically similar way, probably using an amble gait. The wide gauge condition reported in trackways of small and large titanosaurs implies that they possessed similar body (trunk and limbs) proportions despite large differences in body size. These results strengthen the hypothesis that titanosaurs possessed a distinctive suite of anatomical characteristics that are well reflected in their tracks and trackways.

The latest succession of dinosaur tracksites in Europe: Hadrosaur ichnology, track production and palaeoenvironments.

A comprehensive review and study of the rich dinosaur track record of the Tremp Formation in the southern Pyrenees of Spain (Southwestern Europe) shows a unique succession of footprint localities prior to the end-Cretaceous mass extinction event. A description of some 30 new tracksites and data on sedimentary environments, track occurrence and preservation, ichnology and chronostratigraphy are provided. These new track localities represent various facies types within a diverse set of fluvial environments. The footprint discoveries mostly represent hadrosaurian and, less abundantly, to sauropod dinosaurs. The hadrosaur tracks are significantly smaller in size than, but morphologically similar to, those of North America and Asia and are attributable to the ichnogenus Hadrosauropodus. The track succession, with more than 40 distinct track levels, indicates that hadrosaur footprints in the Ibero-Armorican region occur predominantly in the late Maaastrichtian (at least above the early Maastrichtian-late Maastrichtian boundary). The highest abundance is found noticeably found in the late Maastrichtian, with tracks occurring in the C29r magnetochron, within about the latest 300,000 years of the Cretaceous.