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.     

Early Pleistocene human mandible from Sima del Elefante (TE) cave site in Sierra de Atapuerca (Spain): a comparative morphological study

We present a detailed morphological comparative study of the hominin mandible ATE9-1 recovered in 2007 from the Sima del Elefante cave site in Sierra de Atapuerca, Burgos, northern Spain. Paleomagnetic analyses, biostratigraphical studies, and quantitative data obtained through nuclide cosmogenic methods, place this specimen in the Early Pleistocene (1.2-1.3 Ma). This finding, together with archaeological evidence from different European sites, suggests that Western Europe was colonised shortly after the first hominin expansion out of Africa around the Olduvai subchron. Our analysis of the ATE9-1 mandible includes a geometric morphometric analysis of the lower second premolar (LP₄), a combined and detailed external and internal assessment of ATE9-1 roots through CT and microCT techniques, as well as a comparative study of mandibular and other dental features. This analysis reveals some primitive Homo traits on the external aspect of the symphysis and the dentition shared with early African Homo and the Dmanisi hominins. In contrast, other mandibular traits on the internal aspect of the symphysis are derived with regard to African early Homo, indicating unexpectedly large departures from patterns observed in Africa. Reaching the most occidental part of the Eurasian continent implies that the first African emigrants had to cross narrow corridors and to overcome geographic barriers favouring genetic drift, long isolation periods, and adaptation to new climatic and seasonal conditions. Given these conditions and that we are dealing with a long time period, it is possible that one or more speciation events could have occurred in this extreme part of Eurasia during the Early Pleistocene, originating in the lineages represented by the Sima del Elefante-TE9 hominins and possibly by the Gran Dolina-TD6 hominins. In the absence of any additional evidence, we prefer not include the specimen ATE9-1 in any named taxon and refer to it as Homo sp.     

The fossil bat assemblages from the Grotta dei Pipistrelli in Pantalica (southeastern Sicily,Italy): Chronological and palaeoecological implications

Bat remains are useful for palaeoecological reconstructions, they provide independent information on palaeoenvironment and are good indicators for hypogean microclimates. Nine taxa of Chiroptera divided into three families (Rhinolophidae, Vespertilionidae and Miniopteridae) and four genera (Rhinolophus, Myotis, Plecotus and Miniopterus) were discovered in three fossil assemblages from the Grotta dei Pipistrelli, in Sicily, a key region for an understanding of Quaternary climates and environments. Bat remains were deposited during three distinct timespans: one late Pleistocene, MIS 2, around the Last Glacial Maximum, and two Holocene, both referable to the Atlantic chronozone. The presence of yearlings, adult and old individuals suggests hibernating colonies, but the cave was also utilized as a nursery. The diversity of taxa indicates a composite landscape with prevailing vegetation cover and relatively warm climatic conditions. The percentage variations of the frequencies of the recognized taxa suggest a progressive increase of open spaces in the middle Holocene.     

Early Pleistocene human mandible from Sima del Elefante (TE) cave site in Sierra de Atapuerca (Spain): a palaeopathological study

Here we present a detailed palaeopathological study of the hominin mandible ATE9-1 found at the Sima del Elefante site (TE), Sierra de Atapuerca, Spain. This fossil represents the earliest hominin remains from Western Europe with an age of ca. 1.3 Ma. The specimen displays several dento-gnathic lesions; the antiquity and geographic location of this fossil justifies a detailed palaeopathological study to determine if the pathologies have significantly altered taxonomically relevant features. Our study reveals severe dental attrition combined with generalized hypercementosis, alveolar root exposure, mild periodontal disease, tooth dislocation, and an anomalous occlusal plane. We have also observed calculus deposits, two cystic lesions and an anomalous wear facet compatible with tooth picking. The majority of these pathological signs can be explained by compensatory eruption. We propose that these lesions are associated as causes, consequences, and amplifiers of one another within the framework of heavy and even traumatic occlusion, masticatory habits, or both traumatic occlusion and masticatory habits. Despite the severity of these lesions, occlusion was at least partially functional so it was unlikely to influence the survival of this individual. In addition, the lesions do not prohibit the taxonomic assessment of the mandible.     

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.     

The Oldowan site of Muhkai II,layer 80 (northeastern Caucasus): Spatial structure and cultural and chronological attribution of the lithic assemblage

In 2004–2006 a group of Lower Palaeolithic sites was discovered in the northeastern Caucasus: Ainikab I–VI, Gegalashur I–III and Muhkai I–II. Since then, they have been the subject of comprehensive scientific investigations. One site which is undergoing active research is the multilayered site of Muhkai II. Multiple cultural layers at the site have been found in a 73 m thick sequence of Early Pleistocene sediments. The geological age of the layers encompasses the interval 2.1–0.8 million years ago, or the majority of the Early Pleistocene. Layer 80 of Muhkai II is located in the central part of the section at a depth of −33.6 to −34.0 m. Its significance lies in the fact that the site contains two cultural layers–the upper and the lower (main)–that are preserved in situ or have been subject to minimal fluvial processes. The finds consist of lithic artefacts and osseous faunal remains. According to field observations and microstratigraphic analyses, the cultural layers can be characterised as the ancient surfaces of an occupation area. Two stages of occupation can be defined in the lower cultural layer. Layer 80 of Muhkai II was a periodically visited location at the banks of a waterbody, where the acquisition and butchery of large mammals took place. The faunal collection contains 301 bones, deriving from four carnivorous and six herbivorous species of mammals. The total lithic collection contains 1094 finds. According to typological and technological criteria the lithic assemblage relates to the Oldowan and has analogies with Oldowan sites in Africa (particularly with sites of the Olduvai Gorge as well as others). At the same time, some specifics of the lithic assemblage reflect the uniqueness of the site and its functional type. Layer 80 of the Muhkai II site is estimated to date to within the chronological interval ∼2.1–1.7 mya.     

First occurrence of the extinct deer Haploidoceros in the Iberian Peninsula in the Upper Pleistocene of the Cova del Rinoceront (Castelldefels,Barcelona)

The Cova del Rinoceront is the first site in the Iberian Peninsula where the genus Haploidoceros has been documented. This discovery of abundant remains is also its first recorded occurrence in Europe during the Upper Pleistocene. The new fossil record strengthens claims that this genus was widely distributed during the Pleistocene, occupying a longer time span at least until MIS 5. Cranial and post-cranial skeletal remains ascribed to this species were recovered from the uppermost layers of the Cova del Rinoceront. The diagnostic features that allow these remains to be unequivocally assigned to H. mediterraneus, include their cranial morphology and antler shape, comprising two sickle-shaped beams, curving backwards and laterally, and a long frontal basal tine. This paper highlights the problems of deer systematics and identification given the high degree of fragmentation of the genus, their relatively homogeneous morphology, a virtual absence of reliable characters and the overlap of measurements between species.     

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.     

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).     

Human hyoid bones from the middle Pleistocene site of the Sima de los Huesos (Sierra de Atapuerca, Spain)

This study describes and compares two hyoid bones from the middle Pleistocene site of the Sima de los Huesos in the Sierra de Atapuerca (Spain). The Atapuerca SH hyoids are humanlike in both their morphology and dimensions, and they clearly differ from the hyoid bones of chimpanzees and Australopithecus afarensis. Their comparison with the Neandertal specimens Kebara 2 and SDR-034 makes it possible to begin to approach the question of temporal variation and sexual dimorphism in this bone in fossil humans. The results presented here show that the degree of metric and anatomical variation in the fossil sample was similar in magnitude and kind to living humans. Modern hyoid morphology was present by at least 530 kya and appears to represent a shared derived feature of the modern human and Neandertal evolutionary lineages inherited from their last common ancestor.     

A 400,000-year-old Acheulean assemblage associated with the Aroeira-3 human cranium (Gruta da Aroeira,Almonda karst system,Portugal)

Bifaces dominate the Acheulean stone tools recovered during the archaeological excavation of layer X of Gruta da Aroeira, dated to 389–436 ka. Faunal remains and a human cranium were found in association with this lithic assemblage. The raw materials used are mostly quartz and quartzite cobbles available in the vicinity of the site. Technological and systematic analysis shows that there are no Levallois elements and suggests that on-site knapping consisted of the reduction of centripetal cores. Flake cleavers are absent. Use-wear analysis indicates the processing of hard materials, mainly wood. Gruta da Aroeira represents one of the few Middle Pleistocene sites that provide securely dated diagnostic human remains and associated Acheulean lithics, thus representing a major step forward in our understanding of the variability of westernmost Europe's Acheulean and of the human populations that made it.     

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.     

Fauna,environment and human presence during MIS5 in the North of Spain: The new site of Valdavara 3

Valdavara 3 is a new early late Pleistocene paleontological and archeological cave site in northwestern Iberia. Over 1400 fossils have been collected, representing about 40 species. The fauna is of interglacial aspect and is in accordance with the OSL dates from the fossiliferous layer, which indicate an age of 103–113 ka. The great taxonomical diversity indicates a varied landscape. A small collection of lithic artifacts was found associated with the fossils, demostrating presence of humans and suggesting short non-residential visits to the cave. The fossiliferous site was predominantly formed by natural processes. Many fossil localities have short or biased faunal lists, but the fossil fauna recovered from Valdavara 3 is remarkably diverse and may reflect the fauna which once lived there.     

Spatial behaviours of Early Oldowan toolmakers in the Shungura Formation (Lower Omo Valley,Ethiopia): Proposal for an integrated approach

The spatial relationships of the Early Oldowan toolmakers with their environment have been so far addressed through raw material procurement analyses and the characterization of hominid habitat. This paper proposes to integrate these two approaches into a broader spatial analysis encompassing archaeological and environmental data (palaeontological, geological and isotopic data) from Member F and lower Member G of the Shungura Formation (Lower Omo Valley, Ethiopia). Heterogeneity in data resolution induces a multiscale approach with three levels of analysis. The level of occurrence complex allows focusing on the characterization of archaeological occurrences and on their environmental settings. The level of “study area” allows working on hominid habitats and on their raw material procurement behaviours. Finally, at the Shungura Formation scale, we can address temporal issues related to the evolution of spatial behaviours between Member F and the lower part of Member G, ca. 2.3 to 2 million years (Ma).     

Dental remains of Lower Miocene small rodents from Napak (Uganda): Afrocricetodontidae,Myophiomyidae, Kenyamyidae and Sciuridae

The Early Miocene sites of Napak (Uganda), which have been prospected by the Uganda Palaeontology Expedition since 1985, have yielded abundant fossil remains, including a rich and diverse rodent fauna. Ever since the work of Lavocat (1973) on the East African Miocene rodents, which focused mainly on the Kenyan remains (but included some material from Napak discovered by Bishop during the 1950s and 1960s), few studies have been made on the Ugandan specimens. This lack provides motivation for further study of Napak micro-mammals, especially the abundant rodents, which were collected at five sites, Napak IV, V, XV, XXX and XXXI. The fossils described here are attributed to six species representing four families: Afrocricetodontidae (Afrocricetodon songhori, Protarsomys macinnesi), Myophiomyidae (Myophiomys arambourgi), Kenyamyidae (Simonimys genovefae, Kenyamys mariae) and Sciuridae (Vulcanisciurus africanus). Comparison of the dentition of extant and extinct taxa allows us to propose dietary hypotheses (granivory, herbivory and omnivory) for these rodents. In addition, the fossil rodents associated with other fauna from the same localities suggest a more-or-less dense forest environment with clearings, and the probable presence of a humid climate at the time of deposition.     

The open-air Paleolithic site of Mirak,northern edge of the Iranian Central Desert (Semnan,Iran): Evidence of repeated human occupations during the late Pleistocene

The northern edge of the Iranian Central Desert has provided valuable evidence of terminal Pleistocene human settlements. Mirak constitutes one of the largest open-air lithic scatters in the region, consisting of eight natural mounds. Fieldwork was initiated in 2015 by the joint Iranian-French program at Mirak 8. Preliminary results have demonstrated at least three successive phases of human occupation during the MIS3: an upper layer with clear Upper Paleolithic affinities and a maximum age of 28 ky, a lower layer with clear Middle Paleolithic affinities that dates around 47 ky, and an intermediate layer with mixed characteristics that can be seen as an intermediate Paleolithic phase which dates between 28 ± 2 and 38 ± 2 ky. At the time when Upper Paleolithic cultures originated in the Zagros Mountains, cultures with clear Middle Paleolithic affinities persisted nearby along the northern edge of the Iranian Central Plateau.     

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.     

Metric and morphological study of the upper cervical spine from the Sima de los Huesos site (Sierra de Atapuerca, Burgos, Spain)

In this article, the upper cervical spine remains recovered from the Sima de los Huesos (SH) middle Pleistocene site in the Sierra de Atapuerca (Burgos, Spain) are described and analyzed. To date, this site has yielded more than 5000 human fossils belonging to a minimum of 28 individuals of the species Homo heidelbergensis. At least eleven individuals are represented by the upper cervical (C1 and C2) specimens: six adults and five subadults, one of which could represent an adolescent individual. The most complete adult vertebrae (three atlases and three axes) are described, measured, and compared with other fossil hominins and modern humans. These six specimens are associated with one another and represent three individuals. In addition, one of these sets of cervical vertebrae is associated with Cranium 5 (Individual XXI) from the site. The metric analysis demonstrates that the Sima de los Huesos atlases and axes are metrically more similar to Neandertals than to our modern human comparative sample. The SH atlases share with Neandertals a sagittally elongated canal. The most remarkable feature of the SH (and Neandertal) axes is that they are craniocaudally low and mediolaterally wide compared to our modern male sample. Morphologically, the SH sample shares with Neandertals a higher frequency of caudally projected anterior atlas arch, which could reflect greater development of the longus colli muscle. In other features, such as the frequency of weakly developed tubercles for the attachment of the transverse ligament of the atlas, the Sima de los Huesos fossils show intermediate frequencies between our modern comparative samples and the Neandertals, which could represent the primitive condition. Our results are consistent with the previous phylogenetic interpretation of H. heidelbergensis as an exclusively European species, ancestral only to H. neanderthalensis.     

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.     

Avian fossil assemblages at the onset of the LGM in the eastern Alps: A palaecological contribution from the Rio Secco Cave (Italy)

The avian fossil assemblages from the late Pleistocene deposits of the Rio Secco Cave (north-eastern Italy) is presented herein. We studied the layers that date back to the end of MIS3 and the beginning of MIS2, which also contain evidence of Gravettian frequentation dated to 33.5–30 ka cal BP. The systematic analysis revealed the presence of 18 species and other supraspecific taxa that supported palaeoenvironmental reconstructions. Taxa indicate that, at the onset of LGM, site surroundings were characterised by conifer or mixed forests, open grasslands, slow-flowing water bodies and mountain meadows with rocky outcrops, as indicated by Lagopus muta. Today, this environment is found above the tree line (beyond 1500–2000 m) and cannot be detected near the site, located at 580 m asl. Noteworthy, is also the finding of the second Italian late Pleistocene fossil record of Picus canus.