Early Pleistocene

The vegetation and the climatic context in which the first hominins entered and dispersed in Europe during the Early Pleistocene are reconstructed, using literature review and a new climatic simulation. Both in situ fauna and in situ pollen at the twelve early hominin sites under consideration indicate the occurrence of open landscapes: grasslands or forested steppes. The presence of ancient hominins (Homo of the erectus group) is only possible at the transition from glacial to interglacial periods, the full glacial being too cold for them and the transition interglacial to glacial too forested. Glacial-interglacial cycles forced by obliquity showed paralleled vegetation successions, which repeated c. 42 times during the course of the Early Pleistocene (2.58-0.78 Ma), providing 42 narrow windows of opportunity for hominins to disperse into Europe. The climatic conditions of this Early Pleistocene protocratic vegetation are compared with a climatic simulation for 9 ka ago without ice...

This paper reports the results of renewed fieldwork at the HWK EE site (Olduvai Gorge, Tanzania). HWK EE is positioned across the boundary between Lower and Middle Bed II, a crucial interval for studying the emergence of the Acheulean at Olduvai Gorge. Our excavations at HWK EE have produced one of the largest collections of fossils and artefacts from any Oldowan site, distributed across several archaeological units and a large excavation surface in four separate trenches that can be stratigraphically correlated. Here we present the main stratigraphic and archaeological units and discuss site formation processes. Results show a great density of fossils and stone tools vertically through two stratigraphic intervals (Lemuta and Lower Augitic Sandstone) and laterally across an area of around 300 m(2), and highlight the confluence of biotic and abiotic agents in the formation of the assemblage. The large size and diversity of the assemblage, as well as its good preservation, qualify HWK...

Recent functional and zooarchaeological studies conducted on the archeological finds of Pirro Nord (PN13) produced new, reliable data on early European hominid subsistence activities. The age of the site is estimated to be ~ 1.3–1.6 Ma, based on bio-chronological data, and the archeological excavation of the Pirro Nord 13 fissure led to the discovery of more than 300 lithic artifacts associated with thousands of vertebrate fossil remains of the final Villafranchian (Pirro Nord Faunal Unit). The analysis of the fossil faunal remains allowed for the identification of anthropogenic traces linked to the exploitation of different animal carcass (cut marks and intentional bone breakages). Use-wear traces were also observed on some flint artifacts and have been interpreted as the result of the exploitation of animal resources by early hominids and carnivores. It has not been possible to identify the type of access that hominins developed on the carcasses, although it has been established that the hominins competed with carnivores for animal resources. The stone tools and faunal remains with anthropogenic traces recovered in the PN13 fissure represent among the earliest evidence of hominin faunal exploitation in Europe.

The past geographic distribution of the genus Theropithecus Primates: Cercopithecidae) is mainly restricted to Africa. Outside that continent, the earliest reported records of this genus consist of a calcaneus of cf. Theropithecus sp. from ‘Ubeidiya (Israel, 1.6e1.2 Ma [millions of years ago]), as well as three associated cervical vertebrae from Pirro Nord (Italy, 1.7-1.3 Ma) attributed to Theropithecus sp. The attribution of the Pirro Nord vertebrae to this genus has been disputed on morphometric grounds, although their assignment to a large-bodied cercopithecid has remained undisputed. Here we report unpublished cervical vertebral specimens with a similar morphology and, given their significance for the paleobiogeography of Theropithecus (purportedly representing its earliest European record), we reevaluate their taxonomic attribution. In particular, we reconsider the possibility that they belong to another non-primate mammal recorded at this site. Based on both qualitative and metric morphological comparisons, we strongly favor an alternative attribution of the cervical vertebrae from Pirro Nord to the large porcupine Hystrix refossa, which is widely documented at the site by both dentognathic and other postcranial remains. We therefore conclude that the dispersal of Theropithecus out of Africa before ca. 1 Ma (when it is recorded by dental remains from Cueva Victoria, Spain) is currently based only on the calcaneus from ‘Ubeidiya tentatively attributed to this genus.

The equid material from the late Pliocene locality of Sésklo (Thessaly, Greece) is described and compared in this article. It belongs to a large and fairly stout Equus stenonis form, which shares many morphological characters with the species samples from Saint-Vallier, La Puebla de Valverde and Olivola, as well as with already known Equus stenonis samples from other Greek localities (Dafneró, Vólax). Its main features are the big skull and
limbs, the short protocones and the very simple enamel plication in the teeth. Large sized and relatively robust stenonid horses are common elements of the late Pliocene faunas of Greece.

The paper is devoted to a rare case of an encounter in the Early Pleistocene cultural layer of a bone with cut marks and a flint tool bearing use-wear traces. These artifacts were discovered in the context of Oldowan industry at the site Muhkai IIa, layer 2013-2, which dates back to no later than 1.95 million years ago. Complex analysis of the material coming from this layer allows to determine it as a butchery site. We suppose that there were sites with different specialization in this period of time in the Caucasus. Therefore, the functional type should be taken into account in the course of technological and typological generalizations on the most ancient lithic industries.

The earliest hominin occupation of Europe is one of the most debated topics in palaeoanthropology. However, the purportedly oldest of the Early Pleistocene sites in Eurasia lack precise age control and contain stone tools rather than human fossil remains. Here we report the discovery of a human mandible associated with an assemblage of Mode 1 lithic tools and faunal remains bearing traces of hominin processing, in stratigraphic level TE9 at the site of the Sima del Elefante, Atapuerca, Spain. Level TE9 has been dated to the Early Pleistocene (approximately 1.2-1.1 Myr), based on a combination of palaeomagnetism, cosmogenic nuclides and biostratigraphy. The Sima del Elefante site thus emerges as the oldest, most accurately dated record of human occupation in Europe, to our knowledge. The study of the human mandible suggests that the first settlement of Western Europe could be related to an early demographic expansion out of Africa. The new evidence, with previous findings in other Atapuerca sites (level TD6 from Gran Dolina), also suggests that a speciation event occurred in this extreme area of the Eurasian continent during the Early Pleistocene, initiating the hominin lineage represented by the TE9 and TD6 hominins

The dietary morphological methods of mesowear and microwear were applied to ungulates of the Late Pliocene fauna of Sésklo (Thessaly, Greece). The results provide evidence for the predominance of open grassland in the area, as the most common species, Equus stenonis, was a strict grazer. The rare cervid cf. Croizetoceros ramosus was the only browser. The antelopes (genera Gazella and Gazellospira) yielded discrepant microwear and mesowear results. This is interpreted as an indication of regional or seasonal dietary resource differentiation, inferring that the antelopes were probably mixed feeders that grazed occasionally or periodically.

Most of the research on fossil mammals from Umbria (central Italy) has been carried out in the southwestern branch of the Tiber basin, due to its paleontological richness. This portion of the basin extends from Perugia to Terni and corresponds to a well-defined half-graben filled with fluvial-lacustrine deposits. The paleontological sample presented here was discovered in a sand and gravel quarry at Podere San Lorenzo, East of the town of Deruta. The stratigraphic succession exposed in the quarry is no longer visible, but we describe here a new outcrop (Palazzone), which is not far from Podere San Lorenzo and shows comparable facies associations. The two successions were deposited in a fluvial environment characterized by an average reduction of the hydrodynamic energy from the bottom upwards. They are referred to the Early Pleistocene Santa Maria di Ciciliano Subsyntheme (Madonna dei Bagni Lithofacies). Large mammal remains are attributed to Mammuthus cf. meridionalis (Nesti, 1825)...

This paper reports the results of renewed fieldwork at the HWK EE site (Olduvai Gorge, Tanzania). HWK EE is positioned across the boundary between Lower and Middle Bed II, a crucial interval for studying the emergence of the Acheulean at Olduvai Gorge. Our excavations at HWK EE have produced one of the largest collections of fossils and artefacts from any Oldowan site, distributed across several archaeological units and a large excavation surface in four separate trenches that can be stratigraphically correlated. Here we present the main stratigraphic and archaeological units and discuss site formation processes. Results show a great density of fossils and stone tools vertically through two stratigraphic intervals (Lemuta and Lower Augitic Sandstone) and laterally across an area of around 300 m2, and highlight the confluence of biotic and abiotic agents in the formation of the assemblage. The large size and diversity of the assemblage, as well as its good preservation, qualify HWK E...

This paper reports the results of renewed fieldwork at the HWK EE site (Olduvai Gorge, Tanzania). HWK EE is positioned across the boundary between Lower and Middle Bed II, a crucial interval for studying the emergence of the Acheulean at Olduvai Gorge. Our excavations at HWK EE have produced one of the largest collections of fossils and artefacts from any Oldowan site, distributed across several archaeological units and a large excavation surface in four separate trenches that can be stratigraphically correlated. Here we present the main stratigraphic and archaeological units and discuss site formation processes. Results show a great density of fossils and stone tools vertically through two stratigraphic intervals (Lemuta and Lower Augitic Sandstone) and laterally across an area of around 300 m(2), and highlight the confluence of biotic and abiotic agents in the formation of the assemblage. The large size and diversity of the assemblage, as well as its good preservation, qualify HWK...

Fossil remains of Mammuthus meridionalis were found in 2016 at Tasso, near Terranuova Bracciolini (Province of Arezzo, Italy) in Early Pleistocene deposits of the second depositional phase of the Upper Valdarno. The interest in this new find derives from its clear stratigraphic position, at the top of the so-called “Sabbie e Limi del  Torrente Oreno” (sand and silt deposits typical of the Oreno Creek), close to the transition to the marginal alluvial fan deposits. The excavations yielded the remains of a skull of Mammuthus meridionalis still preserving part of its tusks, and an ulna, likely of the same individual. The fossils were found at the base of a fluvial channel infilled with sediments from the Pratomagno Ridge. Over time the channel was subjected to channel bank collapse and was infilled with sand deposited during small- to middle-scale flooding. The elephant skull, which is deformed due to sediment loading and partially eroded, belongs to a relatively large individual, presumably a male. The channel deposits provided also other small fossil remains of Equus stenonis, Pseudodama nestii and Pachy crocuta brevirostris. These late Villafranchian taxa suggest more or less open steppe/forested savannah landscapes in response to the glacial/interglacial cycles of that time period.

This article proposes a revision of the best preserved fossil cervid recovered from the Upper Siwaliks, which, thus far, has been insufficiently described in the literature. The combination of antler and craniodental characters place the deer under study in the genus Metacervocerus, an extinct lineage with a vast Eurasian distribution and a sister phylogenetic relationship with the modern genus Axis. Metacervocerus is distinguished from Axis by its relatively longer upper premolar series and by the form of its second antler bifurcation which has a second tine that is inserted on the anterior side of the main beam. The new examination of the cranial and dental morphology of the specimen from the Siwaliks revealed peculiarities of its cranial and dental morphology that clearly distinguish Metacervocerus punjabiensis from M. shansius from the Late Tertiary of China. The article provides a taxonomic and evolutionary evaluation of craniodental characters of the type specimen of M. punjabiensis, and an analysis of the evolutionary significance of specific antler characteristics, as well as the phylogenetic relationships of the genus Metacervocerus.

In this study, a new Early Pleistocene proximal hand phalanx (ATE9-2) from the Sima del Elefante cave site (TE - Sierra de Atapuerca, Spain), ascribed to Homo sp., is presented and comparatively described in the context of the evolution of the genus Homo. The ATE9-2 specimen is especially important because of the paucity of hand bones in the human fossil record during the Early Pleistocene. The morphological and metrical analyses of the phalanx ATE9-2 indicate that there are no essential differences between it and comparator fossil specimens for the genus Homo after 1.3 Ma (millions of years ago). Similar to Sima de los Huesos and Neandertal specimens, ATE9-2 is a robust proximal hand phalanx, probably reflecting greater overall body robusticity in these populations or a higher gracility in modern humans. The age of level TE9 from Sima del Elefante and morphological and metrical studies of ATE9-2 suggest that the morphology of the proximal hand phalanges and, thus, the morphology of...

The past geographic distribution of the genus Theropithecus (Primates: Cercopithecidae) is mainly restricted to Africa. Outside that continent, the earliest reported records of this genus consist of a calcaneus of cf. Theropithecus sp. from 'Ubeidiya (Israel, 1.6-1.2 Ma [millions of years ago]), as well as three associated cervical vertebrae from Pirro Nord (Italy, 1.7-1.3 Ma) attributed to Theropithecus sp. The attribution of the Pirro Nord vertebrae to this genus has been disputed on morphometric grounds, although their assignment to a large-bodied cercopithecid has remained undisputed. Here we report unpublished cervical vertebral specimens with a similar morphology and, given their significance for the paleobiogeography of Theropithecus (purportedly representing its earliest European record), we re-evaluate their taxonomic attribution. In particular, we reconsider the possibility that they belong to another non-primate mammal recorded at this site. Based on both qualitative ...

Over the last few decades, several types of evidence such as presence of hominin remains, lithic assemblages, and bones with anthropogenic surface modifications have demonstrated that early human communities inhabited the European subcontinent prior to the Jaramillo Subchron (1.07–0.98 Ma). While most studies have focused primarily on early European lithic technologies and raw material management, relatively little is known about food procurement strategies. While there is some evidence showing access to meat and other animal-based food resources, their mode of acquisition and associated butchery processes are still poorly understood. This paper presents a taphonomic and zooarchaeological analysis of the Fuente Nueva-3 (FN3) (Guadix-Baza, Spain) faunal assemblage, providing a more in-depth understanding of early hominin subsistence strategies in Europe. The present results show that hominins had access to the meat and marrow of a wide range of animal taxa, including elephants, hippo...

OBJECTIVES Here we describe the case of an ectopic maxillary third molar (M3 ), preventing the eruption of the M2 , in the individual H3 of the hominin hypodigm of level TD6.2 of the Early Pleistocene site of Gran Dolina (Sierra de Atapuerca, Spain). MATERIALS AND METHODS The fossil remains from the TD6.2 level of the Gran Dolina site (about 170 specimens) are assigned to Homo antecessor. Different geochronological methods place these hominins in the oxygen isotopic stage 21, between 0.8 and 0.85 million years ago (Ma). The immature individual H3 is represented by an almost complete midface (ATD6-69), preserving various teeth in situ. We used high-resolution microtomograhy (mCT) to investigate the abnormal position of the left M3 , virtually reconstruct M2 , and M3 as well as assessing the development stage of these. Finally, we compare this case with extinct and extant populations. RESULTS Based on the identified signs, we suggest that individual H3 suffered from a unilateral impac...

This paper reports the results of renewed fieldwork at the HWK EE site (Olduvai Gorge, Tanzania). HWK EE is positioned across the boundary between Lower and Middle Bed II, a crucial interval for studying the emergence of the Acheulean at Olduvai Gorge. Our excavations at HWK EE have produced one of the largest collections of fossils and artefacts from any Oldowan site, distributed across several archaeological units and a large excavation surface in four separate trenches that can be stratigraphically correlated. Here we present the main stratigraphic and archaeological units and discuss site formation processes. Results show a great density of fossils and stone tools vertically through two stratigraphic intervals (Lemuta and Lower Augitic Sandstone) and laterally across an area of around 300 m(2), and highlight the confluence of biotic and abiotic agents in the formation of the assemblage. The large size and diversity of the assemblage, as well as its good preservation, qualify HWK...

This paper reports the results of renewed fieldwork at the HWK EE site (Olduvai Gorge, Tanzania). HWK EE is positioned across the boundary between Lower and Middle Bed II, a crucial interval for studying the emergence of the Acheulean at Olduvai Gorge. Our excavations at HWK EE have produced one of the largest collections of fossils and artefacts from any Oldowan site, distributed across several archaeological units and a large excavation surface in four separate trenches that can be stratigraphically correlated. Here we present the main stratigraphic and archaeological units and discuss site formation processes. Results show a great density of fossils and stone tools vertically through two stratigraphic intervals (Lemuta and Lower Augitic Sandstone) and laterally across an area of around 300 m(2), and highlight the confluence of biotic and abiotic agents in the formation of the assemblage. The large size and diversity of the assemblage, as well as its good preservation, qualify HWK...

The early Pleistocene is the key period for understanding the evolutionary history and palaeobiogeography of rabbits (Genus Oryctolagus). In western Europe, many species were endemic, making them a reliable indicator of the evolution of the terrestrial ecosystems in which many species have evolved. However, the morphological variability of rabbit species is still poorly understood and their phylogeny remains a subject of debate. Through both qualitative (morphological description) and quantitative (linear measurements and two-dimensional geometric morphometrics) approaches, we address here the morphometric diversity of the third lower premolar (p3), a tooth commonly used to distinguish leporid species, in order to assess intra- and inter-regional morphological variations in several early Pleistocene rabbit populations. Our results suggest that the different approaches are complementary and allow, on different levels, a full characterization of the p3 variability of early Pleistocene rabbits and to imply relations between populations. The size and shape variations of this tooth reflect the taxonomic and phylogenetic signals of the different species but were probably also significantly impacted by geographical position and local climatic conditions. In view of the great morphometric variability highlighted in this work, we suggest a careful reconsideration of certain dental criteria previously considered ‘diagnostic’ in the characterization of these species. However, the overall results allowed us to discuss the phylogeny of the genus Oryctolagus and to hypothesize the ecological requirements and different phases of the dispersal of taxa in western Europe, probably associated with global climate changes.

This paper describes the first early Pleistocene freshwater fish assemblage from the Dacian Basin, Romania. The fossil sample, consisting mainly of isolated teeth, supports the presence of a rich and taxonomically diverse fish assemblage: 17 taxa were identified, belonging to 10 genera of the families Cyprinidae, Salmonidae, Siluridae, Esocidae, and Percidae. The ecological affinities of the identified taxa suggest the early Pleistocene fishes from Copăceni lived in a large lowland mesophitic river. The faunal composition is comparable to that of other southeast European assemblages, however, it includes more thermophilic taxa, suggesting the Dacian Basin represented an early Pleistocene refugium in a cooling post-Pliocene environment. The taxonomical composition of the fossil assemblage suggests a pre-Pleistocene dispersion of freshwater taxa took place in the areas surrounding the Euxinian Basin.

Early Pleistocene volcanic soil series exposed in archaeological sites of the Lori Plateau and the Upper Akhurian Lowland of the Armenian Upland were studied. The study considers the isotopic composition of organic carbon and the content of n-alkanes, not previously analyzed in this area. Both these parameters indicate the waterlogged conditions of Early Pleistocene soil formation as is also indicated by morphology and magnetic susceptibility values. The reconstructed landscapes were represented by herbaceous bio-cenoses on hydromorphic soils with prevalence of C3-grasses.

Reconstruction of early Pleistocene hominin carcass acquisition and processing behaviors are necessarily based at least in part on butchered fossil bones. This paper provides zooarchaeological and taphonomic analyses and behavioral interpretations of three 1.5 million-year-old archaeofaunas from areas 1A and 103 in the Okote Member of the Koobi Fora Formation, northern Kenya: FwJj14A, FwJj14B, and GaJi14. These sites are all located in similar paleoenvironmental contexts, near shallow water with swampy, seasonally flooded areas, and some evidence for more wooded or gallery forest settings. Both individual specimen-and assemblage-level analyses of butchery-marked bones indicate that the hominins appear to have practiced similar butchery strategies at all of these sites, with butchery (defleshing, disarticulation, and marrow extraction) of both high-and low-ranked skeletal elements with no apparent preference for prey size, skeletal region, limb class, or limb portion. Only four tooth-marked specimens, including one likely crocodile-tooth-marked bone, are preserved in all three archaeofaunas. A paucity of limb epiphyses suggests that bone-crunching hyenids may have deleted these portions subsequent to hominin butchery. Strangely, there are no stone tools preserved with the 292 cut-marked and 27 percussion-marked faunal specimens (out of a total of 6,039 specimens), suggesting that raw material availability may have conditioned hominin lithic discard patterns at these locales. These assemblages increase our knowledge of the dietary behavior and ecology of Homo erectus, and provide support for variability in early Pleistocene hominin carcass foraging patterns.

This paper reports the results of renewed fieldwork at the HWK EE site (Olduvai Gorge, Tanzania). HWK EE is positioned across the boundary between Lower and Middle Bed II, a crucial interval for studying the emergence of the Acheulean at Olduvai Gorge. Our excavations at HWK EE have produced one of the largest collections of fossils and artefacts from any Oldowan site, distributed across several archaeological units and a large excavation surface in four separate trenches that can be stratigraphically correlated. Here we present the main stratigraphic and archaeological units and discuss site formation processes. Results show a great density of fossils and stone tools vertically through two stratigraphic intervals (Lemuta and Lower Augitic Sandstone) and laterally across an area of around 300 m2, and highlight the confluence of biotic and abiotic agents in the formation of the assemblage. The large size and diversity of the assemblage, as well as its good preservation, qualify HWK EE as a reference site for the study of the late Oldowan at Olduvai Gorge and elsewhere in Africa. In addition, the description of the stratigraphic and archaeological sequence of HWK EE presented in this paper constitutes the foundation for further studies on hominin behavior and paleoecology in Lower and Middle Bed II.

The fossil remains of cervids from Trlica in Montenegro are described and assigned to the elk or moose Alces cf. carnutorum, the roe deer ?Capreolus sp., the red deer Cervus elaphus, and Eucladoceros. The new species Eucladoceros montenegrensis n. sp. is named on the basis of 169 bones and teeth. The Early and early Middle Pleistocene was a time with a great diversity in giant deer. Their features are discussed, where possible described by morphometrics, and compared to the material from Trlica. On this basis the new species Eucladoceros montenegrensis is named. At present, the oldest well dated record of a red deer (Cervus elaphus) in Europe is from 936 to 990 ka. Various ages have been proposed for Trlica, both younger and older than the Jaramillo Subchron. The age of the locality is discussed.

Since 2007, excavations at Barranc de la Boella (Tarragona, Catalonia, Spain) have revealed three localities
with rich archaeo-paleontological assemblages: La Mina, El Forn and Pit 1. Palaeontology, palaeomagnetism
and cosmogenic analyses have dated these localities to close to 1 Ma. The presence of
Mammuthus meridionalis, Hippopotamus antiquus, Stephanorhinus cf. hundsheimensis, Mimomys savini and
Victoriamys chalinei stand out in the sample of macro and micro-mammals.
The lithic assemblages from the three sites are made up of percussion cobbles, choppers, choppercores,
cores, simple flakes, and some retouched flakes: mainly denticulates and notches. In the case of
the El Forn and Pit 1 localities, two large cutting tools have been recovered: a cleaver-like tool and a pick
made of hard-wearing schist. The lithic assemblage of Pit 1, which includes several refitting lithic sets, is
closely associated with the remains of a young-adult Mammuthus meridionalis, in a clear butchering site
context.
This evidence suggests that Barranc de la Boella is the oldest European Early Acheulean site, and one of
the oldest butchering site on the subcontinent during the late Early Pleistocene. The study of the variability
among these three localities in similar environmental conditions, together with information from
other sites, are discussed in order to gain further knowledge about the appearance of the Acheulean in
Europe, and its continuity or discontinuity in relation to pre-existing technologies.

Homo is the only genus that responds to environmental pressures by adopting such strategies as non-standardized behaviour, a general diet, and technology. Considering that basic idea, this paper introduces the general hypothesis that the Pliocene and Early Pleistocene Homo dispersals within and out of Africa were consequences of that human characteristic behaviour. In particular, they were driven by a process starting with the emergence of technology, followed in a second phase by its socialization. From this point, social reorganization within communities (changing behaviours and social, cultural and economic strategies) occurred, driving demographic growth, and frequently geographic expansion.

In this paper a fossil fauna found in Mesozoic marble fissure fillings is described. The locality was found by Dr. G. Theodorou, who collected the first material. The fauna consists of Micromammals (Leporidae and Arvicolidae) and big Mammals (Elephantidae, Bovidae and representatives of the genera Equus, Canis and Panthera). A brief description of these findings is given, with emphasis on the carnivores. The locality is considered to be of Lower Pleistocene age.