Wil Roebroeks’s research while affiliated with Leiden University and other places

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Publications (87)


DRIVERS of CHANGE Climate Variability, Stone Technologies and Prey Species in Aboriginal Tasmania and Southwest France
  • Poster
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June 2023

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Megan O’Farrel

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Neumark-Nord 2 (Germany) HP 7 sequence, with lithological units and the archaeological find levels (Sier et al., 2011), the stratigraphical distribution of charcoal particles, carbonised seeds (Kuijper, 2014), arboreal (AP) and non-arboreal pollen (NAP) and data regarding vegetation openness (Pop & Bakels, 2015); correlation of archaeological layers containing fire-related findings with vegetation openness episodes shown in red
Pollen analysis (pollen percentage of trees, shrubs, upland herbs and Corylus avellana) from Weimar-Niederweimar II.2 profile and macrofossil evidence (percentage of wood, charcoal and remains from plants occupying open, disturbed and nutrient-rich areas) from different palaeochannel fills at Weimar-Niederweimar (Germany). The sequence shown here is dated to the Younger Dryas (11,640 BP, gravel layer), Preboreal (11,400–10,970 BP, gravel layer) and Boreal periods (10,420–9,510 BP, sand/gyttja and gyttja layers); phases of Early Mesolithic anthropogenic impact within the Lahn valley area are shown in red (after Bos & Urz, 2003)
Pollen analysis (pollen percentage of Corylus, Melampyrum, Succisa, Potentilla-type and microcharcoal) and NPP evidence (percentage of Gelasinospora, Neurospora, Sporormiella) from a profile at North Gill 5B (North York Moors within England and Wales). This evidence reflects post-disturbance phases after burning and intensive grazing during the Late Mesolithic at North Gill. The profile consists of amorphous peat resting on sand at 100 cm. The inferred age of the basal peat lies within the Late Mesolithic based on dates available for a section a few tens of metres away from North Gill 5 (5,270 BP) and higher section of this site (4,540 BP at 73 cm) (after Innes & Blackford, 2003). Red shows the phase with the highest herbivore concentrations; this follows a phase with intensive burning
Tracking Hunter-Gatherer Impact on Vegetation in Last Interglacial and Holocene Europe: Proxies and Challenges

September 2022

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1,038 Reads

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22 Citations

Journal of Archaeological Method and Theory

We review palaeoenvironmental proxies and combinations of these relevant for understanding hunter-gatherer niche construction activities in pre-agricultural Europe. Our approach consists of two steps: (1) identify the possible range of hunter-gatherer impacts on landscapes based on ethnographic studies; (2) evaluate proxies possibly reflecting these impacts for both the Eemian (Last Interglacial, Middle Palaeolithic) and the Early–Middle Holocene (Mesolithic). We found these paleoenvironmental proxies were not able to unequivocally establish clear-cut differences between specific anthropogenic, climatic and megafaunal impacts for either time period in this area. We discuss case studies for both periods and show that published evidence for Mesolithic manipulation of landscapes is based on the interpretation of comparable data as available for the Last Interglacial. If one applies the ‘Mesolithic’ interpretation schemes to the Neanderthal record, three common niche construction activities can be hypothesised: vegetation burning, plant manipulation and impact on animal species presence and abundance. Our review suggests that as strong a case can be made for a Neanderthal impact on landscapes as for anthropogenic landscape changes during the Mesolithic, even though the Neanderthal evidence comes from only one high-resolution site complex. Further research should include attempts (e.g. by means of modelling studies) to establish whether hunter-gatherer impact on landscapes played out at a local level only versus at a larger scale during both time periods, while we also need to obtain comparative data on the population sizes of Last Interglacial and Holocene hunter-gatherers, as these are usually inferred to have differed significantly.



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

August 2021

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481 Reads

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69 Citations

Proceedings of the National Academy of Sciences

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.


FIG. 1. Map of the modern human and ancient hominin AHR mRNA sequence variants studied here and by Hubbard et al. (2016). The modern human AHR variant tested was based on manually expert-curated sequence information from the Ensembl and RefSeq databases (details in Materials and Methods). The bar gives a scaled representation of this 6,277-bp sequence including the 5 0 -UTR, the AHR protein-coding region, and the 3 0 -UTR. The numbers above are the coordinates of the variant sites within the Ensembl AHR reference transcript ENST00000242057.8. Below it, the base, or codon plus encoded amino acid (V, valine; A, alanine; R, arginine; K, lysine) present in each AHR variant is indicated. Cross-hatching indicates optimization for mammalian codon use and minimal secondary mRNA structure; absence of hatching indicates that the codon usage from the human reference sequence is maintained. Codon bases in bold italics differ naturally, respectively, in the synthetic sequences tested by Hubbard et al. (2016), have been changed as compared with the modern human reference. Apart from the indicated variable sites the reported ancient AHR transcript sequences are identical to the modern reference sequence (see supplementary materials S1 and S10, Supplementary Material online).
FIG. 2. HeLa cells were transfected with a pcDNA3.1/Zeo(þ)-based expression construct for the complete modern human (panels a, c, e) or Neanderthal AHR mRNA (panels b, d, f). Panel a-f) Cotransfection (1:1 w/w) with the EGFP expression plasmid pcDNA3-EGFP to visualize the transfection efficiency; a, b) EGFP fluorescence; c, d) EGFP fluorescence þ phase-contrast transillumination; e, f) transillumination only; the black marker spot is applied for orientation. The pcDNA3.1/Zeo(þ) vector backbone is carrying a zeocin resistance gene expression cassette (ZEO). Therefore, ZEO mRNA expression is proportional to the number of functional construct molecules inside the cells upon transfection and allows to quantify the transfection efficiency by qRT-PCR. To correct for possible differences in cDNA input, the ZEO mRNA levels as quantified by qRT-PCR were normalized for b-actin (ACTB) mRNA expression and the mean and standard error of the mean (SEM) of the ZEO/ACTB mRNA ratio (n ¼ 13) was plotted (panel g). A t-test on the difference in mean ratio between the transfections with the modern human and Neanderthal AHR expression constructs showed no significant difference (P < 0.648).
FIG. 4. Effect of the G185A polymorphism (rs7796976) observed in the 5 0 -UTR of the AHR transcript on EGFP expression. A 620-bp 5 0 -UTR sequence carrying either the 185G or 185A variant was attached to the 5 0 -side of the EGFP coding region within pcDNA3-EGFP, a cytomegalovirus promoter-driven EGFP fusion protein expression vector, and transfected into AHR-competent HepG2 human liver hepatoma cells. EGFP reporter protein expression directed by the EGFP mRNA carrying the AHR-derived 5 0 -UTR was normalized to the luciferase expression directed by a cotransfected constitutive luciferase expression construct (see Materials and Methods). Data are from four replicate transfections carried out with two different HepG2 cell batches. To compensate for differences in responsivity between cell batches, all data are presented relative to the mean response observed for the 185A variant. This box and whisker graph is presenting the difference in distribution of the data, with the box showing the 25-75% percentile range, calculated including the median (horizontal line), and mean (x), and the whiskers indicating the minimum and maximum value.
Evolution of Hominin Detoxification: Neanderthal and Modern Human Ah Receptor Respond Similarly to TCDD

November 2020

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103 Reads

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9 Citations

Molecular Biology and Evolution

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.


Environmental conditions at the Last Interglacial (Eemian) site Neumark‐Nord 2, Germany inferred from stable isotope analysis of freshwater mollusc opercula

June 2020

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273 Reads

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4 Citations

Mollusc biogenic carbonates are valuable records of past environmental conditions. In particular, carbonate oxygen (δ¹⁸O) and carbon (δ¹³C) stable isotopes can be used to reconstruct different physical and chemical parameters, according to the different genera used (marine, freshwater or terrestrial). The Last Interglacial (early Eemian) palaeolake of Neumark‐Nord 2 (NN 2), Germany provides an excellent example of a Neanderthal archaeological site with abundant freshwater carbonate remains. As in other European contexts, one of the most abundant species is Bithynia tentaculata . In order to provide a robust regional baseline for the interpretation of the archaeological data, this study includes a calibration phase on modern B. tentaculata opercula. The results indicate that these calcitic structures are likely to be subjected to a growth slowdown/cessation during summer, which influences their geochemistry, reflecting mainly the water properties of the rest of the year. This modern calibration, together with the existing palaeoenvironmental reconstructions developed for NN 2 (e.g. pollen data), represents a valuable opportunity to establish B. tentaculata opercula as reliable environmental proxies applicable to several other freshwater contexts. The isotope data of the NN 2 opercula, in agreement with the pollen record, indicate that the major archaeological horizon was formed during a rather wet period and potentially in a semi‐forested environment. However, human occupation occurred also during drier phases at the site and within a wide temperature range, indicating the absence of restricted environmental preferences by the local Neanderthal groups.


Fig. 2 Location of the Eemian lake basin Neumark-Nord 1 (NN1) and the small pond Neumark-Nord 2 (NN2) within and along the recent Geiseltalsee, Saxony-Anhalt, Germany. The black dots indicate potential basin structures documented by field surveys. The box inset indicates the position of the Neumark-Nord lakeland on a larger regional scale.
Fig. 3 Schematic north-south cross section of the Neumark-Nord 2 basin and its infill including the stratigraphic position of the archaeological find horizon NN2/2. The box inset refers to the main profile HP7 section. Vertical axis: height in meters above sea level. Horizontal axis: position of the NN2 basin in the excavation grid (in meters).
Fig. 5 Topographic map of the large lake basin Neumark-Nord 1 (NN1) in the center and the small "pool" Neumark-Nord 2 (NN2) in the upper right corner, with spatial distribution of (partial) mammal skeletons in basin NN1 (colored dots) and archaeological excavation areas (black rectangles). The pie charts indicate species distribution in selected excavation areas, at NN2 for the main find horizon NN2/2B (expressed as %MNI = minimum number of individuals), at NN1 for the lower littoral zone (unit 6.1) and upper littoral zone (unit 6.2) (expressed as %NISP = number of identified specimens per taxon). - (The synopsis is based principally on figures and data presented in Mania et al. 2010a; Meller 2010.).
Fig. 6 Composition of the ungulate thanatocoenoses (based on estimates of MNI = minimum number of individuals) in the Neumark lakeland during the Eemian Interglacial Cycle.
Fig. 7 Composition of the carnivore thanatocoenosis (based on estimates of MNI = minimum number of individuals) in the Neumark lakeland during the first mesocratic phase of the Eemian Interglacial Cycle.
The last interglacial (Eemian) lakeland of Neumark-Nord (Saxony-Anhalt, Germany). Sequencing Neanderthal occupations, assessing subsistence opportunities and prey selection based on estimations of ungulate carrying capacities, biomass production and energy

April 2020

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512 Reads

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7 Citations

The high-resolution palaeoenvironmental record from the last interglacial (Eemian) lakeland at NeumarkNord (Saxony-Anhalt, Germany) holds the rare potential to study Neanderthal subsistence ca.125,000 years ago in remarkable detail. Using the palynological record from the large lake Neumark-Nord 1 (NN1) and the adjacent small “pool” Neumark-Nord 2 (NN2) a sequence of lithic and faunal assemblages has been established, indicating Neanderthal presence in the lakeland throughout the Eemian. Calculations of the herbivore carrying capacity for the Neumark area during the Eemian reveal high biomass estimates. Using these estimates it was possible to calculate the structure of the source population, based on the faunal record rich in species and individuals, from littoral deposits assigned to the frst mesocratic phase of the Eemian. The results can be used as a frame of reference for making inferences on subsistence opportunities and prey selection based on taphonomical and zooarchaeological studies of the Neumark-Nord faunal assemblages. In a broader perspective our study demonstrates the benefts from inferring energy values (kcal) and body mass (kg) as ancillary parameters to zooarchaeological analyses to understand the relationships between death assemblage and their habitats from which they originate.


The Last Interglacial (Eemian) lakeland of Neumark-Nord (Saxony-Anhalt, Germany): Sequencing Neanderthal occupations, assessing subsistence opportunities and prey selection based on estimations of ungulate carrying capacity, biomass production and energy values

April 2020

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195 Reads

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3 Citations

The high-resolution palaeoenvironmental record from the last interglacial (Eemian) lakeland at Neumark-Nord (Saxony-Anhalt, Germany) holds the rare potential to study Neanderthal subsistence ca. 125,000 years ago in remarkable detail. Using the palynological record from the large lake Neumark-Nord 1 (NN1) and the adjacent small "pool" Neumark-Nord 2 (NN2) a sequence of lithic and faunal assemblages has been established , indicating Neanderthal presence in the lakeland throughout the Eemian. Calculations of the herbi-vore carrying capacity for the Neumark area during the Eemian reveal high biomass estimates. Using these estimates it was possible to calculate the structure of the source population, based on the faunal record rich in species and individuals, from littoral deposits assigned to the first mesocratic phase of the Eemian. The results can be used as a frame of reference for making inferences on subsistence opportunities and prey selection based on taphonomical and zooarchaeological studies of the Neumark-Nord faunal assemblages. In a broader perspective our study demonstrates the benefits from inferring energy values (kcal) and body mass (kg) as ancillary parameters to zooarchaeological analyses to understand the relationships between death assemblage and their habitats from which they originate.


Oxygen isotope analyses of Equus teeth evidences early Eemian and early Weichselian palaeotemperatures at the Middle Palaeolithic site of Neumark-Nord 2, Saxony-Anhalt, Germany

November 2019

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403 Reads

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15 Citations

Quaternary Science Reviews

Here we present phosphate oxygen isotope (d 18 O PO4) data from horse (Equus sp.) tooth enamel (bio-apatite) from the early Eemian and early Weichselian find levels at the archaeological site of Neumark-Nord 2, Germany. Based on the relationship between d 18 O PO4 of bioapatite, body water, local precipitation and air temperature, these data are used to reconstruct palaeoclimatic conditions contemporary to the different phases of Neanderthal activity at the site. Bulk enamel samples representing one year of growth were taken from horse teeth from early Eemian (NN2/2b [~121 ± 5 ka], and NN2/1c) and early Weich-selian (NN2/0;~93 ± 7 ka) find levels, and d 18 O PO4 values were then utilised to calculate d 18 O of local environmental water and mean annual air temperature (MATs) during these phases of MIS5. Results indicate that during the early Eemian MAT was~9 C, with some evidence of variability through time. Although~3 C lower, the calculated early Weichselian MAT exceeds that indicated by other local and regional climate proxy datasets, suggesting that Neanderthal activity may have coincided with more ameliorate phases of the early Weichselian in this area.


Citations (66)


... While unquestionably robust, such approaches cannot answer all questions about landscape evolution, sometimes because there is insufficient high-resolution material (typically sedimentary) available to analyze, sometimes because it is uncertain what particular strata represent. But perhaps most profoundly, such approaches cannot humanize the past; they cannot tell us how particular events may have affected particular groups of humans, perhaps by either constraining or broadening their activities, and, conversely, they cannot tell us what role humans may have played in landscape evolution (perhaps through de-vegetation or the intentional modification of landforms) and their motivations for doing so (Nikulina et al. 2022;Prober et al. 2016;Rosen et al. 2017). ...

Reference:

Calibrating Holocene human-environment interactions using ancient narratives: The example of Ngurunderi in South Australia
Tracking Hunter-Gatherer Impact on Vegetation in Last Interglacial and Holocene Europe: Proxies and Challenges

Journal of Archaeological Method and Theory

... 用火 [12][13][14][15] 、人属物种首次移居更高纬度地区 [16][17][18][19] 等。通过对这些中更新世古人类类群的 研究,我们可以更好地审视现代人在体质特征、行为模式、社会组织形式等方面的起源 和演化细节。 目前已有的化石证据显示,这些新类型的古人类于约30万年前出现在东亚 [20] ,并 于约10万年前消失 [21] 。这一人群的典型代表有距今约30万-20万年的华龙洞人 [20] 、大 荔人 [22][23][24][25] 、金牛山人 [26][27][28] 、马坝人 [29][30][31] ,距今22.4万-16.1万年的许家窑人 [32] 和距今12.5 万-10.5万年的许昌人 ...

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

Proceedings of the National Academy of Sciences

... The white-vestigial mutant genotype is characterized by the white color of the eyes, generated by the presence of the recessive gene "w", located on chromosome I, and by the presence of reduced and rudimentary wings caused by the presence of the recessive gene "vg", located on chromosome II (Figure 1d). Studies showed the influence of the "vg" gene in the translation elongation process (TEF) [53], a process necessary for the generation of messenger RNA (mRNA) proteins, thus reducing the capacity of detoxification mechanisms [54]. ...

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

Molecular Biology and Evolution

... And when it comes to causal generics such as "smoking causes cancer" and "dogs bite people, " they function as guidelines for caution in actions (Sterken, 2015). Archaeologically and anthropologically, detached communication about the relations between animals, their tracks and their behavior is difficult without using some form of generics (MacDonald and Roebroeks, 2013). Although speculative, this connection could be an explanation of why generics are so central in language. ...

Neanderthal linguistic abilities: an alternative view
  • Citing Chapter
  • July 2013

... Malacological analysis of Eemian lake sediments was undertaken across Europe, e.g. Germany (Caspers et al., 2002;Brose et al., 2006;Meng et al., 2009a, b;Strahl et al., 2010;Menzel-Harloff and Meng, 2015;Kenzler et al., 2018;Milano et al., 2020), Netherlands (Cleveringa et al., 2000), Great Britain (Meijer and Preece, 1995;Preece, 1999;Langford et al., 2017), Lithuania (Sanko and Gaigalas, 2007;Sanko et al., 2011) and Belarus (Sanko et al., 2011). Mollusc shells are frequently found in deposits filling Eemian paleolakes, especially in lacustrine chalk and calcareous gyttja. ...

Environmental conditions at the Last Interglacial (Eemian) site Neumark‐Nord 2, Germany inferred from stable isotope analysis of freshwater mollusc opercula

... At the Eemian lakeshore site of Neumark-Nord in east-central Germany (ca 125 kya), evidence for Neanderthal presence coincides with pollen evidence for open landscape patches and substantially increased charcoal input [155]. This may have promoted the availability and accessibility of acorns, hazelnuts and perhaps sloe plums [159] and created suitable habitats for the close-distance hunting of elephants and deer [160,161], possibly framed by human-induced ecologies of fear [162], as herbivore biodiversity is extremely high in the lake basin while evidence for carnivore interference appears minimal [163]. In the Middle Stone Age (MSA) of Lake Malawi (ca 100-80 kya), another early human-environment feedback system with emergent biodiversity consequences is in evidence [22,142]. ...

The last interglacial (Eemian) lakeland of Neumark-Nord (Saxony-Anhalt, Germany). Sequencing Neanderthal occupations, assessing subsistence opportunities and prey selection based on estimations of ungulate carrying capacities, biomass production and energy

... Pleistocene foragers living in environments with rich, dense, and predictable resources, to some degree even cocreated by their landscape modification activities (43), may have coalesced into short-term larger groups. The repetitive food bonanzas documented on the north European plain, seen in the context of the rich Last Interglacial environments filled with other, less dangerous large mammals that at Neumark-Nord were hunted all year round and in large numbers (61), could have provided occasions to gather in larger groups or even have been carried out explicitly in the contexts of such short-term aggregations. ...

The Last Interglacial (Eemian) lakeland of Neumark-Nord (Saxony-Anhalt, Germany): Sequencing Neanderthal occupations, assessing subsistence opportunities and prey selection based on estimations of ungulate carrying capacity, biomass production and energy values
  • Citing Chapter
  • April 2020

... Stable oxygen isotopes from meteoric water (mainly derived from rainfall) have a strong relationship with 235 mean air temperatures in mid to high latitudes (Rozanski et al., 1992;Dansgaard, 1964) on a regional-to-236 local scale. Obligate drinkers, such as bovines and horses, acquire this water and record its isotopic 237 composition in their teeth and bones with a fixed, but species-specific offset (Pederzani and Britton, 2019 ...

Oxygen isotope analyses of Equus teeth evidences early Eemian and early Weichselian palaeotemperatures at the Middle Palaeolithic site of Neumark-Nord 2, Saxony-Anhalt, Germany
  • Citing Preprint
  • November 2019

Quaternary Science Reviews

... The main stratigraphic units are overbank deposits formed in a lowenergy watercourse during MIS 13, glacially tectonised in MIS 12 (Lewis, 1992). Although West et al. (2014) propose a different site formation theory, interpreting the clayeysilts as MIS 7 doline deposits, an MIS 13 age is supported by litho-stratigraphy (Lewis et al., 2019, as well as the mammalian taxa, Stephanorhinus hundsheimensis and Trogontherium sp., known to be extinct after MIS 12 and MIS 11, respectively (Preece & Parfitt, 2012;Stuart, 1992). Shear-planes within the High Lodge series are also symptomatic of englacial rafting (Lewis, 1992). ...

Human occupation of northern Europe in MIS 13: Happisburgh Site 1 (Norfolk, UK) and its European context
  • Citing Article
  • May 2019

Quaternary Science Reviews

... This stark contrast led Bradley (2002, 2012) and Bradley and Stanford (2004) to propose an alternative origin of Clovis (and a hypothesized precursor culture in the Mid-Atlantic US) in Western Europe, where the Solutrean industry has fine bipointed bifaces and macroblades, made using specific knapping techniques (e.g., overshot flaking) shared with Clovis. However, recent AMS radiocarbon dates show that the Solutrean dates to ca. 26,000-22,000 cal BP, ending about 9000 years before Clovis begins (Zilhao 2013;Cascalheira and Bicho 2015;Ducasse et al. 2019Ducasse et al. , 2020Verpoorte et al. 2019). Furthermore, the genomes of ancient and recent Native Americans show no early input from Upper Paleolithic Western Europeans. ...

Improving the chronological framework for Laugerie-Haute Ouest (Dordogne, France)
  • Citing Article
  • December 2018

Journal of Archaeological Science Reports