Small mammals (Insectivora, Rodentia, Lagomorpha) from the Early Pleistocene hominin-bearing site of Dmanisi (Georgia).

Small mammals (insectivores, rodents, and lagomorphs) from Dmanisi are here reviewed for the first time and used as a tool for paleoenvironmental proxies. The small mammal faunal list is composed of shrews (Beremendia fissidens, cf. Beremendia minor, Crocidura kornfeldi), hamsters (Cricetulus sp., Allocricetus bursae), gerbils (Parameriones aff. obeidiyensis), murids (Apodemus cf. atavus), arvicolids (Mimomys pliocaenicus, Mimomys aff. pusillus), and pikas (Ochotona sp.). A paleoenvironmental reconstruction based on the habitat weighting method has been applied to the rodent assemblage. According to this method, the most common elements indicate an open-dry habitat (36.5%), followed by water edge (25.7%) and rocky (21.0%) elements. Open-wet (15.5%) and woodland elements (1.3%) are rare. Therefore, the habitat occupied by the hominids of Dmanisi was characterized by the prevalence of arid conditions, from steppe or semi-desert to open Mediterranean forest, with stony or rocky substrate and bushy areas. The presence of permanent aquatic environments is also documented. From a biogeographic point of view, the small mammal community from Dmanisi is composed mainly by Western or Central Asian elements, with a poor representation of European elements (Mimomys, Apodemus). It is concluded that Dmanisi hominins most possibly had ecological requirements which were different from those of the Early Pleistocene hominins from Western Europe, which settled on wetter habitats. It could be also possible that Dmanisi hominins entered Southern Caucasus at an interglacial phase before the deposition of the Dmanisi site.

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.

Early Pleistocene enamel proteome from Dmanisi resolves Stephanorhinus phylogeny.

The sequencing of ancient DNA has enabled the reconstruction of speciation, migration and admixture events for extinct taxa1. However, the irreversible post-mortem degradation2 of ancient DNA has so far limited its recovery-outside permafrost areas-to specimens that are not older than approximately 0.5 million years (Myr)3. By contrast, tandem mass spectrometry has enabled the sequencing of approximately 1.5-Myr-old collagen type I4, and suggested the presence of protein residues in fossils of the Cretaceous period5-although with limited phylogenetic use6. In the absence of molecular evidence, the speciation of several extinct species of the Early and Middle Pleistocene epoch remains contentious. Here we address the phylogenetic relationships of the Eurasian Rhinocerotidae of the Pleistocene epoch7-9, using the proteome of dental enamel from a Stephanorhinus tooth that is approximately 1.77-Myr old, recovered from the archaeological site of Dmanisi (South Caucasus, Georgia)10. Molecular phylogenetic analyses place this Stephanorhinus as a sister group to the clade formed by the woolly rhinoceros (Coelodonta antiquitatis) and Merck's rhinoceros (Stephanorhinus kirchbergensis). We show that Coelodonta evolved from an early Stephanorhinus lineage, and that this latter genus includes at least two distinct evolutionary lines. The genus Stephanorhinus is therefore currently paraphyletic, and its systematic revision is needed. We demonstrate that sequencing the proteome of Early Pleistocene dental enamel overcomes the limitations of phylogenetic inference based on ancient collagen or DNA. Our approach also provides additional information about the sex and taxonomic assignment of other specimens from Dmanisi. Our findings reveal that proteomic investigation of ancient dental enamel-which is the hardest tissue in vertebrates11, and is highly abundant in the fossil record-can push the reconstruction of molecular evolution further back into the Early Pleistocene epoch, beyond the currently known limits of ancient DNA preservation.

Multiproxy evidence for leaf-browsing and closed habitats in extinct proboscideans (Mammalia, Proboscidea) from Central Chile.

Proboscideans are so-called ecosystem engineers and are considered key players in hypotheses about Late Pleistocene megafaunal extinctions. However, knowledge about the autoecology and chronology of the proboscideans in South America is still open to debate and raises controversial views. Here, we used a range of multiproxy approaches and new radiocarbon datings to study the autoecology of Chilean gomphotheres, the only group of proboscideans to reach South America during the Great American Biotic Interchange (∼3.1 to 2.7 million years before present). As part of this study, we analyzed stable isotopes, dental microwear, and dental calculus microfossils on gomphothere molars from 30 Late Pleistocene sites (31° to 42°S). These proxies provided different scales of temporal resolution, which were then combined to assess the dietary and habitat patterns of these proboscideans. The multiproxy study suggests that most foraging took place in relatively closed environments. In Central Chile, there is a positive correlation between lower δ13C values and an increasing consumption of arboreal/scrub elements. Analyses of dental microwear and calculus microfossils have verified these leaf-browsing feeding habits. From a comparative perspective, the dietary pattern of South American gomphotheres appears to be constrained more by resource availability than by the potential dietary range of the individual taxa. This multiproxy study is aimed at increasing knowledge of the life history of gomphotheres and thus follows an issue considered one of the greatest challenges for paleontology in South America, recently pointed out by the need to thoroughly understand the role of ecological engineers before making predictions about the consequences of ecosystem defaunation.

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.

Top-down silicon microcantilever with coupled bottom-up silicon nanowire for enhanced mass resolution.

A stepped cantilever composed of a bottom-up silicon nanowire coupled to a top-down silicon microcantilever electrostatically actuated and with capacitive or optical readout is fabricated and analyzed, both theoretically and experimentally, for mass sensing applications. The mass sensitivity at the nanowire free end and the frequency resolution considering thermomechanical noise are computed for different nanowire dimensions. The results obtained show that the coupled structure presents a very good mass sensitivity thanks to the nanowire, where the mass depositions take place, while also presenting a very good frequency resolution due to the microcantilever, where the transduction is carried out. A two-fold improvement in mass sensitivity with respect to that of the microcantilever standalone is experimentally demonstrated, and at least an order-of-magnitude improvement is theoretically predicted, only changing the nanowire length. Very close frequency resolutions are experimentally measured and theoretically predicted for a standalone microcantilever and for a microcantilever-nanowire coupled system. Thus, an improvement in mass sensing resolution of the microcantilever-nanowire stepped cantilever is demonstrated with respect to that of the microcantilever standalone.

Age and date for early arrival of the Acheulian in Europe (Barranc de la Boella, la Canonja, Spain).

The first arrivals of hominin populations into Eurasia during the Early Pleistocene are currently considered to have occurred as short and poorly dated biological dispersions. Questions as to the tempo and mode of these early prehistoric settlements have given rise to debates concerning the taxonomic significance of the lithic assemblages, as trace fossils, and the geographical distribution of the technological traditions found in the Lower Palaeolithic record. Here, we report on the Barranc de la Boella site which has yielded a lithic assemblage dating to ∼1 million years ago that includes large cutting tools (LCT). We argue that distinct technological traditions coexisted in the Iberian archaeological repertoires of the late Early Pleistocene age in a similar way to the earliest sub-Saharan African artefact assemblages. These differences between stone tool assemblages may be attributed to the different chronologies of hominin dispersal events. The archaeological record of Barranc de la Boella completes the geographical distribution of LCT assemblages across southern Eurasia during the EMPT (Early-Middle Pleistocene Transition, circa 942 to 641 kyr). Up to now, chronology of the earliest European LCT assemblages is based on the abundant Palaeolithic record found in terrace river sequences which have been dated to the end of the EMPT and later. However, the findings at Barranc de la Boella suggest that early LCT lithic assemblages appeared in the SW of Europe during earlier hominin dispersal episodes before the definitive colonization of temperate Eurasia took place.

Walk the line: 600000 years of molar evolution constrained by allometry in the fossil rodent Mimomys savini.

The allometric-constraint hypothesis states that evolutionary divergence of morphological traits is restricted by integrated growth regulation. In this study, we test this hypothesis on a time-calibrated and well-documented palaeontological sequence of dental measurements on the Pleistocene arvicoline rodent species Mimomys savini from the Iberian Peninsula. Based on 507 specimens representing nine populations regularly spaced over 600 000 years, we compare static (within-population) and evolutionary (among-population) allometric slopes between the width and the length of the first lower molar. We find that the static allometric slope remains evolutionary stable and predicts the evolutionary allometry quite well. These results support the hypothesis that the macroevolutionary divergence of molar traits is constrained by static allometric relationships.

Late Miocene/Early Pliocene vertebrate fauna from Mallorca (Balearic Islands, Western Mediterranean): an update.

The vertebrate fossil record from the Balearic Islands (western Mediterranean) has improved considerably over the past decade, especially in Mallorca and Menorca. In Menorca, the Pliocene terrestrial fauna was updated by the discovery and description of the large-sized leporid Nuralagus, several reptiles and an amphibian. In Mallorca, paleontological exploration yielded 2 deposits with a Late Miocene/Early Pliocene chronology, Caló den Rafelino (CdR) and Na Burguesa-1 (NB-1). So far, 4 new mammalian taxa and 2 new reptiles have been identified for the CdR deposit, whereas the faunal assemblage from the recently discovered deposit (Apr 2012) of NB-1 is currently composed of, at least, 6 terrestrial mammals, 8 reptiles and an amphibian. Its faunal composition and some primitive characteristics of the obtained taxa suggest that the chronology of this deposit is slightly earlier than the CdR. The terrestrial vertebrates recorded in these 2 Mallorcan deposits are changing the view of the paleofaunal assemblage previously known for the Plio-Pleistocene of the island. Morphological characteristics displayed by some of the taxa suggest that these faunas would be at the beginning of an isolated evolution. In this paper we present a preliminary report on the fossils recovered from the NB-1 deposit, as well as some unpublished data from CdR, and we analyze the whole fauna from both Mallorcan deposits, focusing on taxonomical and paleobiogeographical aspects.

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.

Human neoteny revisited: The case of synaptic plasticity.

The process of learning requires morphological changes in the neuronal connections and the formation of new synapses. Due to the importance of memory and learning in our species, it has been suggested that the synaptic plasticity in a number of association areas is higher in the human brain than in other primates. Cortical neurons in mammals are characterized by higher metabolism, activity, and synaptic plasticity during development and the juvenile stage than in the adult. In Homo sapiens, brain development is retarded compared with other primates, especially in some association areas. These areas are characterized by the presence of neurons, which remain structurally immature throughout their lifespans and show an increase in the expression of the genes, which deal with metabolism and the activity and synaptic plasticity in adulthood. The retention of juvenile features in some adult neurons in our species has occurred in areas, which are related to episodic memory, planning, and social navigation. The increase of the aerobic metabolism in these neurons may lead, however, to higher levels of oxidative stress, therefore, favoring the development of neurodegenerative diseases which are exclusive, or almost exclusive, to humans, such as Alzheimer's disease.

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.

A new Lower Pleistocene archeological site in Europe (Vallparadis, Barcelona, Spain).

Here we report the discovery of a new late Lower Pleistocene site named Vallparadís (Barcelona, Spain) that produced a rich archeological and paleontological sequence dated from the upper boundary of the Jaramillo subchron to the early Middle Pleistocene. This deposit contained a main archeological layer with numerous artifacts and a rich macromammalian assemblage, some of which bore cut marks, that could indicate that hominins had access to carcasses. Paleomagnetic analysis, electron spin resonance-uranium series (ESR-US), and the biostratigraphic chronological position of the macro- and micromammal and lithic assemblages of this layer reinforce the proposal that hominins inhabited Europe during the Lower Pleistocene. The archeological sequence provides key information on the successful adaptation of European hominins that preceded the well-known fossil population from Atapuerca and succeeded the finds from Orce basin. Hence, this discovery enables us to close a major chronological gap in the early prehistory of Iberia. According to the information in this paper and the available data from these other sites, we propose that Mediterranean Western Europe was repeatedly and perhaps continuously occupied during the late Matuyama chron.

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.

A fourth hominin skull from Dmanisi, Georgia.

Newly discovered Homo remains, stone artifacts, and animal fossils from Dmanisi, Republic of Georgia, provide a basis for better understanding patterns of hominin evolution and behavior in Eurasia ca. 1.77 million years ago. Here we describe a fourth skull that is nearly complete, lacking all but one of its teeth at the time of death. Both the maxillae and the mandible exhibit extensive bone loss due to resorption. This individual is similar to others from the site but supplies information about variation in brain size and craniofacial anatomy within the Dmanisi paleodeme. Although this assemblage presents numerous primitive characters, the Dmanisi skulls are best accommodated within the species H. erectus. On anatomical grounds, it is argued that the relatively small-brained and lightly built Dmanisi hominins may be ancestral to African and Far Eastern branches of H. erectus showing more derived morphology.

Anthropology: the earliest toothless hominin skull.

The site of Dmanisi in the Eurasian republic of Georgia has yielded striking hominin, faunal and archaeological material as evidence for the presence of early Homo outside Africa 1.77 million years ago, documenting an important episode in human evolution. Here we describe a beautifully preserved skull and jawbone from a Dmanisi hominin of this period who had lost all but one tooth several years before death. This specimen not only represents the earliest case of severe masticatory impairment in the hominin fossil record to be discovered so far, but also raises questions about alternative subsistence strategies in early Homo.