PaleoAnthropology

Towards a Deeper Integration of ZooMS and Zooarchaeology at Palaeolithic Sites: Current Challenges and Future Directions

Geoffrey M. Smith, Karen Ruebens, Virginie Sinet-Mathiot, Frido Welker — Wed, 13 Nov 2024 00:00:00 +0100

Advances in biomolecular methods, in particular the study of ancient proteins (palaeoproteomics), have revolutionised 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 Palaeolithic 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 zooarchaeological data, especially within Palaeolithic 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 specimen at a site, but also provide novel insights into site formation histories, collection biases, carnivore behaviour, environmental conditions and past human subsistence, including site use, seasonality, carcass transport, prey preference and butchery practices.

Integrating Morphological and ZooMS-Based Approaches to Zooarchaeology at Vogelherd Cave in Southwestern Germany

Naihui Wang, Nicholas J. Conard, Katerina Douka — Wed, 13 Nov 2024 00:00:00 +0100

Zooarchaeology is an established subfield of archaeology that incorporates a variety of interdisciplinary tools. Advances in analytical methods like radiocarbon dating, stable isotope analysis, and ancient DNA analysis have added new dimensions to zooarchaeological research in the past century. In recent years, the addition of the ZooMS (Zooarchaeology by Mass Spectrometry) offers exciting new possibilities for studying faunal remains in archaeological contexts.
In this study, we use the Vogelherd Cave, a Palaeolithic site in the Swabian Jura of southwestern Germany, as a case study to showcase the advances in zooarchaeological analysis and changes in research focus. Gustav Riek from the University of Tübingen completely excavated the site’s rich deposits in 1931. In 2005-2012 and 2022-2023, Nicholas J. Conard’s team from the University of Tübingen excavated Riek’s backdirt using modern excavation techniques.
While Ulrich Lehmann published the first systematic analysis of the faunal assemblage from a paleontological perspective in 1954, it was not until the early 2000s that Laura Niven undertook a comprehensive zooarchaeological study. In 2014, Boger and colleagues analysed the fauna remains from the backdirt to gain a more complete view of the faunal assemblage. The current study adds the first ZooMS analysis of 287 fragmentary bones from the same site. Our focus lies on highly fragmented faunal remains obtained from water-screened sediment of the backdirt.
Here, we compile and compare our ZooMS results to previous faunal datasets from Vogelherd. The history of research at this site provides a representative example of how the research has expanded over time and how novel analytical methods contribute to the interpretation of a site. By juxtaposing traditional zooarchaeological data and ZooMS data, we explore the strengths and weaknesses of each approach and contemplate how to integrate these methods in future research.

Investigating Species Composition in the Early Aurignacian of Le Piage (France) Through Collagen Fingerprinting (ZooMS) of Screen-Recovered Small Bone Fragments

Wed, 13 Nov 2024 00:00:00 +0100

In the paleoproteomic field, ZooMS (Zooarchaeology by Mass Spectrometry) has been developed to identify morphologically non-diagnostic animal remains to taxon, offering insights into human subsistence practices. Here, we report new ZooMS analyses of 1,050 Early Aurignacian (ca. 37,000-34,000 cal BP) bone fragments from the site of Le Piage (Lot, France). The studied sample is heavily fragmented and was retrieved through water sieving. In our analysis, we compare the taxonomic identifications of bone remains using traditional morphological attributes with remains identified using ZooMS and discuss the implications of the taxonomic patterns. Our results indicate that the faunal spectrum identified through ZooMS differs from the species composition obtained through morphological analyses. While reindeer remains the dominant species, bovids and other cervids are better represented in the ZooMS fraction. Two rare taxa, a hare (Lepus sp.) and a previously unidentified carnivore (Pantherinae/Hyaenidae/Mustelidae), were also identified using ZooMS. We also observe an increase of Bos/Bison remains in the spongious assemblage that is possibly explained by the use of spongy portions of long bones as fuel. Our work adds new data on patterns of reindeer dominance during the Early Aurignacian and illustrates how ZooMS identifications of screen-recovered small bone fragments can enhance our understanding of Paleolithic site occupation and subsistence strategies.

Human Subsistence Before and After the 8.2 ka cal BP Event in Northern Iberia: Archaeozoology and Proteomic Data From the Macromammal Assemblage of El Mazo Rock Shelter

Wed, 13 Nov 2024 00:00:00 +0100

El Mazo rock shelter (Asturias, Spain) contains a long and well-dated Mesolithic sequence that shows an extended human presence between 9 and 7.4 ka cal BP, including the climatic 8.2 ka event. This is evident from fireplaces, abundant lithic tools, and a wide human diet, proved by the presence of marine and terrestrial specimens and several plant macroremains (charcoal, seeds, and fruits). This event continues to spark debates about the causes of changes in human behavior and subsistence strategies. Therefore, the El Mazo long sequence allows for identifying human subsistence patterns before, during, and after the 8.2 ka event. Regarding the use of terrestrial mammals, archeozoological and taphonomic analysis indicates the red deer as the primary prey, followed by wild boar and roe deer, with a high representation of adults and juveniles. Cut and percussion marks are associated with disarticulation and skinning on those prey. At the same time, thermoalteration is abundant along the sequence and was the most common alteration observed in the macromammal assemblage. The high representation of fragmented axial bones, including ribs and vertebrae, and fetal/newborn elements prevented more detailed information about human subsistence and prey transport, which drove those bones to be identifiable primarily to mammal size. As a result, these bones are often classified as indeterminate or identified only as size class. To tackle this morphological limitation, in this study, we applied ZooArchaeology by Mass Spectrometry (ZooMS) to determine the whole spectrum of axial elements. Proteomics results indicate those remains correspond mainly to adult and infant red deer and wild boar. Besides, it shows a broader herbivore and carnivore spectrum than archaeozoology showed, including humans, bovines, small carnivores, canids, or undetected species such as the Eurasian beaver. Finally, the mammal assemblage reveals how the sudden temperature decrease during the 8.2 ka cal BP event significantly impacted human subsistence and population decline. After the 8.2 ka event, intensification indices show that herbivore consumption increased significantly, overexploiting all available resources, while marine resources intensified.

Integrating ZooMS and Zooarchaeology to Assess the Châtelperronian and Carnivore Occupations at Cassenade (Dordogne, France)

Karen Ruebens, Geoff M. Smith, Emmanuel Discamps, Jean-Jacques Hublin — Wed, 13 Nov 2024 00:00:00 +0100

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 Palaeolithic 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 behaviour.
In this paper, we apply ZooMS to all morphologically unidentifiable bone fragments larger than 20 mm (n = 817) recovered from the 2012-2013 excavated Châtelperronian layer of Cassenade (France). Collagen was extracted using an ammonium-bicarbonate (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 (50.8% vs 16.7%). Conversely, Ursidae, the dominant taxa in the morphologically identifiable remains (36.6%), 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 none of the fragments show traces of human activity, which is most likely related to low bone surface readability. 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.
Three-dimensional data is available for all the ZooM-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 diet 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

Emmanuel Discamps, Karen Ruebens, Geoff Smith, Jean-Jacques Hublin — Wed, 13 Nov 2024 00:00:00 +0100

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 archaeological 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 Cassenade Palaeolithic site.

At the Châtelperronian site of Cassenade (France), all bone fragments larger than 20 mm (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.

Comparing Neanderthal and Modern Human Subsistence at Riparo Bombrini: An Integrated Archaeozoological, Multivariate Taphonomic, and ZooMS Analysis

Wed, 13 Nov 2024 00:00:00 +0100

The Liguro-Provençal arc yields unique deposits documenting the Middle-Upper Paleolithic transition. Yet, interpreting shifts in subsistence strategies in this region has remained challenging, mainly due to taphonomic processes and the lack of archaeological assemblages excavated with modern techniques. For instance, faunal assemblages from the Balzi Rossi Paleolithic site complex, dated 43-36 ky cal BP, are notoriously fragmented, impeding morphology-based taxonomic identification and the application of most conventional archaeozoological methods. Furthermore, collagen preservation is also often poor, making identification through proteomic techniques such as ZooMS difficult. This study analyzes three assemblages documenting the transition at one of Balzi Rossi’s sites, Riparo Bombrini, using an integrated approach that combines archaeozoological methods, multivariate taphonomic analysis, and FTIR-aided ZooMS.

The results indicate that the Proto-Aurignacian faunal assemblages were accumulated almost exclusively by anatomically modern human foragers, whereas the final Mousterian was accumulated as a result of brief, alternating site visits by Neanderthals and medium-sized carnivores. Continuous faunal exploitation strategies are observed through the final Mousterian and the Proto-Aurignacian levels, such as the predominance of prime-aged cervids hunted near the site, often brought back whole and processed at Riparo Bombrini. However, the faunal assemblages also document changes in the mortality profiles, carcass treatment, site function, and land-use, starting in the Proto-Aurignacian. These changes include increased events of hunting vulnerable large ungulates during prolonged occupations of the site, increased carcass curation for bone fuel, decreased carnivore activities on the site, and the diversification of bone tool types and raw materials to produce symbolic objects. In addition, the results align with previous hypotheses suggesting a hyperlocal adaptation of the very last Neanderthals to have occupied the site, followed by dense occupations of the site and shifting mobility strategies within a large territory associated with the overlying Proto-Aurignacian assemblages lasting through climatic instability. This study offers the first detailed view of human subsistence during the transition in the region and sets up test hypotheses about the changing nature of hominin behavioral ecology.

Integrating Morphology and ZooMS-Identified Fauna Provides Insights Into Species Diversity and Neanderthal - Carnivores Interactions in Shared Landscapes: Evidence from Picken's Hole, Britain

Wed, 13 Nov 2024 00:00:00 +0100

Reconstructing the faunal palaeoecology of landscapes occupied by Neanderthals and their competitors is essential for better understanding their ecological niche, decisions, and behaviors. Late Pleistocene faunal assemblages in Britain are highly fragmented with interpretations relying on the morphologically identifiable portion of the assemblage and the indeterminate bone fragments often dismissed. This paper applies two methodologies (taphonomic analysis and zooarchaeology by mass spectrometry (ZooMS)) to extract data from morphologically indeterminate bone fragments recovered from the late Middle Palaeolithic contexts of Picken’s Hole, Somerset, and integrate this new data with the extant zooarchaeological study of the morphologically identifiable faunal specimens. 2205 indeterminate bone fragments from Unit 3 were categorized to mammal class size and broad element type, taphonomic observations recorded (weathering, carnivore bone surface modification, etc.). Then 708 samples were selected for ZooMS. The ZooMS-identified faunal spectrum agrees with Scott’s (2018) study of the morphologically identifiable portion, indicating an open cool steppe tundra environment. The faunal proportions differ greatly between identification method, however, especially when dentition is removed. Woolly rhinoceros, mammoth and horse have the lowest ZooMS to Morph identification ratios (high ZooMS-NISP and low morph-NISP), while grey wolf/arctic fox and reindeer have the highest (low ZooMS-NISP and high morph-NISP). In addition, while the morphology-identified portion is dominated by dentition and foot bones, the ZooMS-identified portion includes more rib and long bone fragments. Weathering levels overall are low and do not appear to be a primary cause of fragmentation, while carnivore gnawing and evidence of digestion are observed across most species and fragment sizes, with carnivore digestion responsible for most fragmentation. The exception is grey wolf/arctic fox which display no evidence of predation activity, suggesting that they and Neanderthals may have used the site intermittently alongside repeated use by large carnivores. This paper underscores the significance of extracting and integrating information from indeterminate bone fragments to offer insight into Neanderthal behavior and interactions with prolific carnivores operating in a shared landscape.

Book Review of Handbook of Pleistocene Archaeology in Africa: Hominin Behavior, Geography, and Chronology

Geoffrey Clark — Wed, 13 Nov 2024 00:00:00 +0100

This is a review of an edited volume. It's about the Pleistocene archaeology of Africa.

14th Annual Meeting of the European Society for the study of Human Evolution Abstracts Zagreb, 11-15 September 2024

ESHE: European Society for the study of Human Evolution — Wed, 13 Nov 2024 00:00:00 +0100

ESHE abstracts 2024.