Widespread evidence for elephant exploitation by Last Interglacial Neanderthals on the North European plain.

Neanderthals hunted and butchered straight-tusked elephants, the largest terrestrial mammals of the Pleistocene, in a lake landscape on the North European plain, 125,000 years ago, as recently shown by a study of the Last Interglacial elephant assemblage from Neumark-Nord (Germany). With evidence for a remarkable focus on adult males and on their extended utilization, the data from this location are thus far without parallel in the archaeological record. Given their relevance for our knowledge of the Neanderthal niche, we investigated whether the Neumark-Nord subsistence practices were more than a local phenomenon, possibly determined by local characteristics. Analyzing elephant remains from two other Last Interglacial archaeological sites on the North European plain, Gröbern and Taubach, we identified in both assemblages similar butchering patterns as at Neumark-Nord, demonstrating that extended elephant exploitation was a widespread Neanderthal practice during the (early part of the) Last Interglacial. The substantial efforts needed to process these animals, weighing up to 13 metric tons, and the large amounts of food generated suggest that Neanderthals either had ways of storing vast amounts of meat and fat and/or temporarily aggregated in larger groups than commonly acknowledged. The data do not allow us to rule out one of the two explanations, and furthermore both factors, short-term larger group sizes as well as some form of food preservation, may have played a role. What the data do show is that exploitation of large straight-tusked elephants was a widespread and recurring phenomenon amongst Last Interglacial Neanderthals on the North European plain.

Middle Pleistocene fire use: The first signal of widespread cultural diffusion in human evolution.

Control of fire is one of the most important technological innovations within the evolution of humankind. The archaeological signal of fire use becomes very visible from around 400,000 y ago onward. Interestingly, this occurs at a geologically similar time over major parts of the Old World, in Africa, as well as in western Eurasia, and in different subpopulations of the wider hominin metapopulation. We interpret this spatiotemporal pattern as the result of cultural diffusion, and as representing the earliest clear-cut case of widespread cultural change resulting from diffusion in human evolution. This fire-use pattern is followed slightly later by a similar spatiotemporal distribution of Levallois technology, at the beginning of the African Middle Stone Age and the western Eurasian Middle Paleolithic. These archaeological data, as well as studies of ancient genomes, lead us to hypothesize that at the latest by 400,000 y ago, hominin subpopulations encountered one another often enough and were sufficiently tolerant toward one another to transmit ideas and techniques over large regions within relatively short time periods. Furthermore, it is likely that the large-scale social networks necessary to transmit complicated skills were also in place. Most importantly, this suggests a form of cultural behavior significantly more similar to that of extant Homo sapiens than to our great ape relatives.

Reconstructing the provenance of the hominin fossils from Trinil (Java, Indonesia) through an integrated analysis of the historical and recent excavations.

In the early 1890s at Trinil, Eugène Dubois found a hominin skullcap (Trinil 2) and femur (Trinil 3, Femur I), situated at the same level ca. 10-15 m apart. He interpreted them as representing one species, Pithecanthropus erectus (now Homo erectus) which he inferred to be a transitional form between apes and humans. Ever since, this interpretation has been questioned-as the skullcap looked archaic and the femur surprisingly modern. From the 1950s onward, chemical and morphological analyses rekindled the debate. Concurrently, (bio)stratigraphic arguments gained importance, raising the stakes by extrapolating the consequences of potential mixing of hominin remains to the homogeneity of the complete Trinil fossil assemblage. However, conclusive evidence on the provenance and age of the hominin fossils remains absent. New Trinil fieldwork yielded unmanned aerial vehicle imagery, digital elevation models, and stratigraphic observations that have been integrated here with an analysis of the historical excavation documentation. Using a geographic information system and sightline analysis, the position of the historical excavation pits and the hominin fossils therein were reconstructed, and the historical stratigraphy was connected to that of new sections and test pits. This study documents five strata situated at low water level at the excavation site. Cutting into a lahar breccia are two similarly oriented, but asynchronous pre-terrace fluvial channels whose highly fossiliferous infills are identified as the primary targets of the historical excavations (Bone-Bearing Channel 1, 830-773 ka; Bone-Bearing Channel 2, 560-380 ka), providing evidence for a mixed faunal assemblage and yielding most of the hominin fossils. These channels were incised by younger terrace-related fluvial channels (terminal Middle or Late Pleistocene) that directly intersect the historical excavations and the reconstructed discovery location of Femur I, thereby providing an explanation for the relatively modern morphology of this 'bone of contention'. The paleoanthropological implications are discussed in light of the current framework of human evolution in Southeast Asia.

Evolution of Hominin Detoxification: Neanderthal and Modern Human Ah Receptor Respond Similarly to TCDD.

In studies of hominin adaptations to fire use, the role of the aryl hydrocarbon receptor (AHR) in the evolution of detoxification has been highlighted, including statements that the modern human AHR confers a significantly better capacity to deal with toxic smoke components than the Neanderthal AHR. To evaluate this, we compared the AHR-controlled induction of cytochrome P4501A1 (CYP1A1) mRNA in HeLa human cervix epithelial adenocarcinoma cells transfected with an Altai-Neanderthal or a modern human reference AHR expression construct, and exposed to 2,3,7,8-tetrachlorodibenzo-p-dioxin (TCDD). We compared the complete AHR mRNA sequences including the untranslated regions (UTRs), maintaining the original codon usage. We observe no significant difference in CYP1A1 induction by TCDD between Neanderthal and modern human AHR, whereas a 150-1,000 times difference was previously reported in a study of the AHR coding region optimized for mammalian codon usage and expressed in rat cells. Our study exemplifies that expression in a homologous cellular background is of major importance to determine (ancient) protein activity. The Neanderthal and modern human dose-response curves almost coincide, except for a slightly higher extrapolated maximum for the Neanderthal AHR, possibly caused by a 5'-UTR G-variant known from modern humans (rs7796976). Our results are strongly at odds with a major role of the modern human AHR in the evolution of hominin detoxification of smoke components and consistent with our previous study based on 18 relevant genes in addition to AHR, which concluded that efficient detoxification alleles are more dominant in ancient hominins, chimpanzees, and gorillas than in modern humans.

Homo erectus at Trinil on Java used shells for tool production and engraving.

The manufacture of geometric engravings is generally interpreted as indicative of modern cognition and behaviour. Key questions in the debate on the origin of such behaviour are whether this innovation is restricted to Homo sapiens, and whether it has a uniquely African origin. Here we report on a fossil freshwater shell assemblage from the Hauptknochenschicht ('main bone layer') of Trinil (Java, Indonesia), the type locality of Homo erectus discovered by Eugène Dubois in 1891 (refs 2 and 3). In the Dubois collection (in the Naturalis museum, Leiden, The Netherlands) we found evidence for freshwater shellfish consumption by hominins, one unambiguous shell tool, and a shell with a geometric engraving. We dated sediment contained in the shells with (40)Ar/(39)Ar and luminescence dating methods, obtaining a maximum age of 0.54 ± 0.10 million years and a minimum age of 0.43 ± 0.05 million years. This implies that the Trinil Hauptknochenschicht is younger than previously estimated. Together, our data indicate that the engraving was made by Homo erectus, and that it is considerably older than the oldest geometric engravings described so far. Although it is at present not possible to assess the function or meaning of the engraved shell, this discovery suggests that engraving abstract patterns was in the realm of Asian Homo erectus cognition and neuromotor control.

Neandertals revised.

The last decade has seen a significant growth of our knowledge of the Neandertals, a population of Pleistocene hunter-gatherers who lived in (western) Eurasia between ∼400,000 and 40,000 y ago. Starting from a source population deep in the Middle Pleistocene, the hundreds of thousands of years of relative separation between African and Eurasian groups led to the emergence of different phenotypes in Late Pleistocene Europe and Africa. Both recently obtained genetic evidence and archeological data show that the biological and cultural gaps between these populations were probably smaller than previously thought. These data, reviewed here, falsify inferences to the effect that, compared with their near-modern contemporaries in Africa, Neandertals were outliers in terms of behavioral complexity. It is only around 40,000 y ago, tens of thousands of years after anatomically modern humans first left Africa and thousands of years after documented interbreeding between modern humans, Neandertals and Denisovans, that we see major changes in the archeological record, from western Eurasia to Southeast Asia, e.g., the emergence of representational imagery and the colonization of arctic areas and of greater Australia (Sahul).

Population size does not explain past changes in cultural complexity.

Demography is increasingly being invoked to account for features of the archaeological record, such as the technological conservatism of the Lower and Middle Pleistocene, the Middle to Upper Paleolithic transition, and cultural loss in Holocene Tasmania. Such explanations are commonly justified in relation to population dynamic models developed by Henrich [Henrich J (2004)Am Antiq69:197-214] and Powell et al. [Powell A, et al. (2009)Science324(5932):1298-1301], which appear to demonstrate that population size is the crucial determinant of cultural complexity. Here, we show that these models fail in two important respects. First, they only support a relationship between demography and culture in implausible conditions. Second, their predictions conflict with the available archaeological and ethnographic evidence. We conclude that new theoretical and empirical research is required to identify the factors that drove the changes in cultural complexity that are documented by the archaeological record.

Use of red ochre by early Neandertals.

The use of manganese and iron oxides by late Neandertals is well documented in Europe, especially for the period 60-40 kya. Such finds often have been interpreted as pigments even though their exact function is largely unknown. Here we report significantly older iron oxide finds that constitute the earliest documented use of red ochre by Neandertals. These finds were small concentrates of red material retrieved during excavations at Maastricht-Belvédère, The Netherlands. The excavations exposed a series of well-preserved flint artifact (and occasionally bone) scatters, formed in a river valley setting during a late Middle Pleistocene full interglacial period. Samples of the reddish material were submitted to various forms of analyses to study their physical properties. All analyses identified the red material as hematite. This is a nonlocal material that was imported to the site, possibly over dozens of kilometers. Identification of the Maastricht-Belvédère finds as hematite pushes the use of red ochre by (early) Neandertals back in time significantly, to minimally 200-250 kya (i.e., to the same time range as the early ochre use in the African record).

On the earliest evidence for habitual use of fire in Europe.

The timing of the human control of fire is a hotly debated issue, with claims for regular fire use by early hominins in Africa at ∼ 1.6 million y ago. These claims are not uncontested, but most archaeologists would agree that the colonization of areas outside Africa, especially of regions such as Europe where temperatures at time dropped below freezing, was indeed tied to the use of fire. Our review of the European evidence suggests that early hominins moved into northern latitudes without the habitual use of fire. It was only much later, from ∼ 300,000 to 400,000 y ago onward, that fire became a significant part of the hominin technological repertoire. It is also from the second half of the Middle Pleistocene onward that we can observe spectacular cases of Neandertal pyrotechnological knowledge in the production of hafting materials. The increase in the number of sites with good evidence of fire throughout the Late Pleistocene shows that European Neandertals had fire management not unlike that documented for Upper Paleolithic groups.

The fragmented character of Middle Palaeolithic stone tool technology.

The importance of the transport of stone artefacts in structuring Neandertal lithic assemblages has often been addressed, but the degree to which this led to fragmentation of lithic reduction over Middle Palaeolithic landscapes has not been explicitly studied thus far. Large-scale excavations of Middle Palaeolithic open-air sites and refitting studies of the retrieved assemblages have yielded new, high-resolution data on the mobile aspects of Neandertal stone tool technology. In this paper, we integrate lithic technology and raw material data from recent studies of Middle Palaeolithic open-air and rock shelter sites in Western Europe. We demonstrate that the results of a variety of typological, technological (especially refitting), and lithological studies have important consequences for our knowledge of the acquisition of raw materials and subsequent production, usage and discard of stone artefacts in the Middle Palaeolithic. Neandertal production and use of stone tools was fragmented in three domains: the spatial, the temporal and the social domain. We show that this versatile segmentation of stone artefact handling strategies is a main determinant of the character of the Neandertal archaeological record. Our data testify to ubiquitous and continuous transport of stone artefacts of a wide variety of forms, picked by Neandertals using selection criteria that were sometimes far removed from what archaeologists have traditionally considered, and to some degree still consider, to be desired end products of knapping activities. The data presented here testify to the variability and versatility of Middle Palaeolithic stone tool technology, whose fragmented character created very heterogeneous archaeological assemblages, usually the product of a wide variety of independent import, use, discard and/or subsequent transport events.

Hunting and processing of straight-tusked elephants 125.000 years ago: Implications for Neanderthal behavior.

Straight-tusked elephants (Palaeoloxodon antiquus) were the largest terrestrial mammals of the Pleistocene, present in Eurasian landscapes between 800,000 and 100,000 years ago. The occasional co-occurrence of their skeletal remains with stone tools has generated rich speculation about the nature of interactions between these elephants and Pleistocene humans: Did hominins scavenge on elephants that died a natural death or maybe even hunt some individuals? Our archaeozoological study of the largest P. antiquus assemblage known, excavated from 125,000-year-old lake deposits in Germany, shows that hunting of elephants weighing up to 13 metric tons was part of the cultural repertoire of Last Interglacial Neanderthals there, over >2000 years, many dozens of generations. The intensity and nutritional yields of these well-documented butchering activities, combined with previously reported data from this Neumark-Nord site complex, suggest that Neanderthals were less mobile and operated within social units substantially larger than commonly envisaged.

Beaver exploitation, 400,000 years ago, testifies to prey choice diversity of Middle Pleistocene hominins.

Data regarding the subsistence base of early hominins are heavily biased in favor of the animal component of their diets, in particular the remains of large mammals, which are generally much better preserved at archaeological sites than the bones of smaller animals, let alone the remains of plant food. Exploitation of smaller game is very rarely documented before the latest phases of the Pleistocene, which is often taken to imply narrow diets of archaic Homo and interpreted as a striking economic difference between Late Pleistocene and earlier hominins. We present new data that contradict this view of Middle Pleistocene Lower Palaeolithic hominins: cut mark evidence demonstrating systematic exploitation of beavers, identified in the large faunal assemblage from the c. 400,000 years old hominin site Bilzingsleben, in central Germany. In combination with a prime-age dominated mortality profile, this cut mark record shows that the rich beaver assemblage resulted from repetitive human hunting activities, with a focus on young adult individuals. The Bilzingsleben beaver exploitation evidence demonstrates a greater diversity of prey choice by Middle Pleistocene hominins than commonly acknowledged, and a much deeper history of broad-spectrum subsistence than commonly assumed, already visible in prey choices 400,000 years ago.

An Asian perspective on early human dispersal from Africa.

The past decade has seen the Pliocene and Pleistocene fossil hominin record enriched by the addition of at least ten new taxa, including the Early Pleistocene, small-brained hominins from Dmanisi, Georgia, and the diminutive Late Pleistocene Homo floresiensis from Flores, Indonesia. At the same time, Asia's earliest hominin presence has been extended up to 1.8 Myr ago, hundreds of thousands of years earlier than previously envisaged. Nevertheless, the preferred explanation for the first appearance of hominins outside Africa has remained virtually unchanged. We show here that it is time to develop alternatives to one of palaeoanthropology's most basic paradigms: 'Out of Africa 1'.

Landscape modification by Last Interglacial Neanderthals.

Little is known about the antiquity, nature, and scale of Pleistocene hunter-gatherer impact on their ecosystems, despite the importance for studies of conservation and human evolution. Such impact is likely to be limited, mainly because of low population densities, and challenging to detect and interpret in terms of cause-effect dynamics. We present high-resolution paleoenvironmental and archaeological data from the Last Interglacial locality of Neumark-Nord (Germany). Among the factors that shaped vegetation structure and succession in this lake landscape, we identify a distinct ecological footprint of hominin activities, including fire use. We compare these data with evidence from archaeological and baseline sites from the same region. At Neumark-Nord, notably open vegetation coincides with a virtually continuous c. 2000-year-long hominin presence, and the comparative data strongly suggest that hominins were a contributing factor. With an age of c. 125,000 years, Neumark-Nord provides an early example of a hominin role in vegetation transformation.

Out of the North Sea: the Zeeland ridges Neandertal.

In 2001, a portion of human frontal bone was discovered in sediments extracted from the bottom of the North Sea, 15km off the coast of the Netherlands. The extraction zone is located in the so-called Zeeland Ridges area located at 51 degrees 40' northern latitude and 3 degrees 20' eastern longitude. The specimen was dredged up from sediments containing Late Pleistocene faunal remains and Middle Palaeolithic artefacts, including well-finished small handaxes and Levallois flakes. The details of the supraorbital morphology, as well as the quantitative assessment of the shape of the external surface of the squama using traditional and 3D geometric morphometrics, unambiguously assign the Zeeland Ridges frontal bone to Homo neanderthalensis. Carbon and nitrogen isotopic analysis indicate that the Zeeland Ridges hominin, like other Neandertals, was highly carnivorous and does not show evidence for the consumption of aquatic foods. A lesion on the outer table and diploic layer of the bone in the area of the supratoral sulcus can be interpreted as the result of an intradiploic epidermoid cyst, a type of neoplasm diagnosed for the first time in Neandertal remains. So far, the Zeeland Ridges Neandertal is the first Pleistocene fossil hominin found under seawater and the first recorded in the Netherlands.

Evidence for close-range hunting by last interglacial Neanderthals.

Animal resources have been part of hominin diets since around 2.5 million years ago, with sharp-edged stone tools facilitating access to carcasses. How exactly hominins acquired animal prey and how hunting strategies varied through time and space is far from clear. The oldest possible hunting weapons known from the archaeological record are 300,000 to 400,000-year-old sharpened wooden staves. These may have been used as throwing and/or close-range thrusting spears, but actual data on how such objects were used are lacking, as unambiguous lesions caused by such weapon-like objects are unknown for most of human prehistory. Here, we report perforations observed on two fallow deer skeletons from Neumark-Nord, Germany, retrieved during excavations of 120,000-year-old lake shore deposits with abundant traces of Neanderthal presence. Detailed studies of the perforations, including micro-computed tomography imaging and ballistic experiments, demonstrate that they resulted from the close-range use of thrusting spears. Such confrontational ways of hunting require close cooperation between participants, and over time may have shaped important aspects of hominin biology and behaviour.