Connecting Middle Palaeolithic datasets: The interplay of zooarchaeological and lithic data for unravelling Neanderthal behaviour

The ongoing refinement of archaeological excavation and recording methods over the last decades has led to a significant increase in quantitative Middle Palaeolithic datasets that provide a record of past Neanderthal behaviour. Stone tools and butchered animal remains are the two main categories of Middle Palaeolithic archaeological remains and both provide distinctive insights into site formation and Neanderthal behaviour. However, the integration of these quantitative lithic and zooarchaeological datasets is key for achieving a full understanding of both site-specific and broader-scale patterns of Middle Palaeolithic subsistence. To explore novel ways to enhance the incorporation of these datasets, we organised a session at the 82nd annual meeting of the Society for American Archaeology in Vancouver. An underlying theme was the problem of linking lithic and faunal variability. Are variations in subsistence strategies reflected by changes in toolmaking decisions? This paper will briefly introduce the possible ways these Middle Palaeolithic datasets can be integrated, illustrated with the papers included in this special volume, and discuss its potential for understanding the variability and interconnectedness of Neanderthal technologies and subsistence strategies

Neanderthal subsistence, taphonomy and chronology at Salzgitter-Lebenstedt (Germany): a multifaceted analysis of morphologically unidentifiable bone

Abstract Pleistocene faunal assemblages are often highly fragmented, hindering taxonomic identifications and interpretive potentials. In this paper, we apply four different methodologies to morphologically unidentifiable bone fragments from the Neanderthal open-air site of Salzgitter-Lebenstedt (Germany). First, we recorded zooarchaeological attributes for all 1362 unidentifiable bones recovered in 1977. Second, we applied zooarchaeology by mass spectrometry (ZooMS) to 761 fragments, and calculated glutamine deamidation values. Third, we assessed the collagen preservation of 30 fragments by near-infrared spectroscopy (NIR) and, finally, we pretreated 10 bones with high predicted collagen values for radiocarbon dating. All returned dates at, or beyond, the limit of radiocarbon dating, indicating an age of older than 51 000 years ago. The ZooMS faunal spectrum confirms a cold environment, dominated by reindeer, alongside mammoth, horse and bison. The low occurrence of carnivore modifications (1%) contrasts with an abundance of human modifications (23%). Cut marks and marrow fractures were observed across reindeer, horse and bison. The mammoth remains are less well preserved and show a lower degree of human modifications, indicating, perhaps, a different taphonomic history. Overall, this study illustrates the importance of retaining, studying and incorporating the unidentifiable bone fraction to optimize interpretations of site formation and subsistence behaviour at Palaeolithic sites

When lithics hit bones: Evaluating the potential of a multifaceted experimental protocol to illuminate Middle Palaeolithic weapon technology

Recent zooarchaeological and isotope analyses have largely settled the debate surrounding Neanderthal hunting capacities, repeatedly demonstrating their successful acquisition of large ungulates. Nevertheless, the functional identification of individual tools as hunting weapons remains a methodological challenge. In-depth studies have focussed mainly on small subsets of lithic artefacts from selected assemblages assessing features of breakage patterns, retouch, shape and use wear. Studies focussing on associated hunting lesions are rarer and often focus on reconstructing very specific bone surface marks encountered in the archaeological record. This study aims to add to our understanding of the formation and characteristics of projectile impact marks (PIMs) on bone through a series of highly monitored, replicative experiments, using thrusting and throwing spears with replica Levallois points into two wild pig carcasses. In total, 152 shots were made, and for each a series of attributes was recorded, including velocity and location of impact. Subsequent quantitative analyses focussed on understanding the various factors underlying the formation of different types of projectile impact marks. These experiments demonstrate that PIM formation results from the properties of both the impacting projectile and bone element. PIMs can signal impacts caused by different delivery methods but only on some parts of the skeleton. These results are contextualised in relation to the occurrence and recognition of Palaeolithic PIMs and patterns of Neanderthal behaviour. These experiments are only a first step in improving the recognition of these signatures in the archaeological record and providing better insights into understanding of the mechanisms of Neanderthal hunting

Integrating ZooMS and Zooarchaeology to assess the Châtelperronian and carnivore occupations at Cassenade (Dordogne, France)

Archaeological animal bone assemblages are often highly fragmented, meaning that for over 70% of the recovered bone fragments we do not know what animal (or human) species they belonged to. This is especially problematic in Paleolithic contexts, when both humans and carnivores repeatedly occupied the same caves and rock shelters. Identifying bone fragments from these dual occupation contexts through Zooarchaeology by Mass Spectrometry (ZooMS) can provide additional insights into both carnivore and human behavior. In this paper, we apply ZooMS to the majority of morphologically unidentifiable bone fragments larger than 20mm (n=840) recovered from the 2012–2013 excavated Châtelperronian layer of Cassenade (France). Collagen was extracted using an ammoniumbicarbonate (AmBic) buffer and over 99% of the sampled bone fragments could be identified taxonomically. While the proportion of Equidae is similar in both ZooMS and zooarchaeological components, Bos/Bison is represented by a threefold increase in the ZooMS fraction (50.5% vs. 16.6%). Conversely, Ursidae, the dominant taxa in the morphologically identifiable remains (36.4%), only formed 7.3% of the ZooMS fragments. Carnivores are also present, but in low numbers (0–2%), and include Hyaenidae, Panthera and Canidae. In the ZooMS fraction, only few bones show traces of human activity (ca. 2%), which is most likely related to low bone surface readability. These show that human groups at Cassenade were processing Bovinae and Equidae, as well as Ursidae and rhinoceros. Conversely, traces of carnivore activity are abundant, and we were able to taxonomically identify 334 bone fragments that were digested by carnivores (as indicated by acid etched surfaces). While large proportions of the Rhinocerotidae (63.79%), Elephantidae (52%), Equidae (48%), and Bos/Bison (45%) remains were digested by carnivores, this is only the case for 1.7% of the Ursidae ZooMS fragments. Threedimensional data are available for all the ZooMS-identified fragments and confirm the near-exclusive presence of cave bear in the lower part of the sequence. Further, the ZooMS spatial data identified a restricted presence of mammoth in the middle part of the sequence and a diverging presence of reindeer and Cervid/saiga remains at the bottom and top. Overall, this study illustrates the added value of integrating zooarchaeological and ZooMS datasets to obtain additional insights into past ecologies, changing site use, carnivore diets, and human subsistence practices

Can ZooMS help assess species abundance in highly fragmented bone assemblages? Integrating morphological and proteomic identifications for the calculation of an adjusted ZooMS-eNISP

Zooarchaeology by Mass Spectrometry (ZooMS) is a rapid, low-cost, collagen-based method for the taxonomic identification of animal tissues. It is now increasingly applied to bone fragments from archeological contexts, creating large taxonomic datasets. How to integrate these ZooMS identifications within general zooarchaeological theoretical frameworks, such as estimates of species abundance and taxonomic richness, remains problematic. Past large-scale ZooMS analyses of Eurasian Paleolithic faunal assemblages have shown a general trend towards an increased representation of large ungulates (mainly Bos/Bison) in the ZooMS fraction, often coupled with a decrease in medium-sized taxa (e.g., reindeer). Here we propose several hypotheses to explain these identification discrepancies, involving identification biases and differential fragmentation patterns across various taxa, and test them using the case study of the Paleolithic site of Cassenade. At the Châtelperronian site of Cassenade (France), nearly all bone fragments larger than 20mm (n=1,119) have been identified to taxa, either through comparative morphology (n=364) or ZooMS (n=755). Each of these fragments was weighed and measured, creating a unique database to explore the relation between fragmentation and identification. Analysis shows that fragment size and mass distributions are distinct across taxa if only bones identified by morphology are considered, but, somehow counter-intuitively, extremely similar across taxa of various body sizes when all their bones are integrated. In particular, the bones of larger ungulates tend to be broken into a higher number of fragments, inducing an over-representation of larger taxa in ZooMS-NISP. Our dataset also shows that sorting long-bone shaft fragments by body size classes should be seen by zooarchaeologists as a process that, in addition to being prone to risks of misidentification, provides highly biased information of little use for estimating species abundance. To overcome this issue, we propose the calculation of an adjusted equivalent ZooMS NISP (ZooMS-eNISP) by dividing, for each taxon, the total ZooMS mass of identified bones (g) by the mean mass of morphologically identified bones for that taxon (g/NISP). The advantage of this method is that it considers site-specific characteristics of the faunal assemblage, notably bone preservation, which is especially important in Paleolithic contexts. Finally, we propose that ZooMS-eNISP, despite its limits, can facilitate the integration of both identification methods to produce a more refined picture of patterns of species representation, site formation, and human behavior at an archaeological site

Towards a deeper integration of ZooMS and zooarchaeology at Paleolithic sites: current challenges and future directions

Advances in biomolecular methods, in particular the study of ancient proteins (paleoproteomics), have revolutionized how we can taxonomically identify archaeological bone fragments. Alongside traditional zooarchaeological assignments based on the visual inspection of morphological criteria, variations in collagen type I amino acid sequences can now be used to distinguish which animal a bone fragment belonged to. Using MALDI-ToF mass spectrometry, this method, known as Zooarchaeology by Mass Spectrometry (ZooMS), is now being applied regularly to archaeological faunal assemblages and, often at a large-scale, at Paleolithic sites. However, detailed explorations of how these ZooMS datasets can best be integrated with zooarchaeological and taphonomic data are only in their infancy. To further advance this field, we hosted a workshop at the University of Kent in 2023, bringing together both zooarchaeologists and ZooMS specialists, to showcase and discuss various ways of integrating ZooMS and zoo-archaeological data, especially within Paleolithic contexts. This special issue results from the papers presented at this workshop. In this introductory paper we reflect on the open discussion sessions that formed an essential part of the workshop. First, we discuss a series of methodological challenges; this includes the recording of zooarchaeology and taphonomy on morphologically unidentifiable bone fragments, ZooMS study design and sample selection, pre-screening and sampling, pre-treatment and collagen extraction, and the acquisition, processing, and interpretation of MALDI data. Second, we delve deeper into the interpretive potential, and the wealth of future research directions, of a full contribution of ZooMS to a range of zooarchaeological research topics. In concordance with the seven research papers in this issue, this introduction illustrates how a well-designed study, integrating zooarchaeological and taphonomic observations across both the morphological and ZooMS-identified fractions, cannot only increase the number of identifiable specimens at a site, but also provide novel insights into site formation histories, collection biases, carnivore behavior, environmental conditions, and past human subsistence, including site use, seasonality, carcass transport, prey preference, and butchery practices

Ramification of lithic production and the search of small tools in Iberian Peninsula Middle Paleolithic

The notion of recycling and it relationship with ramified productions and small tool production in Late Middle Paleolithic from the Iberian Peninsula are investigated. Results from Amalda, Axlor, Pe~na Miel, and Quebrada show that the production of small tools is one of the principal objectives of lithic provisioning in these sites. Whereas in Axlor and Amalda, this is achieved through the ramification of production, due to the remoteness of flint sources, in Quebrada, where raw material sources are closer, small flakes are obtained at the end of Levallois production. The implications for Neandertal society organization of this small tool production is discussed, and its evolution is observed from a diachronic perspective