Citation
A Zooarchaeological analysis of neandertal [sic] cave site Arma Veirana in Liguria, Italy

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Title:
A Zooarchaeological analysis of neandertal [sic] cave site Arma Veirana in Liguria, Italy
Creator:
Charolla, Breeanna Chantel
Place of Publication:
Denver, CO
Publisher:
University of Colorado Denver
Publication Date:
Language:
English

Thesis/Dissertation Information

Degree:
Master's ( Master of arts)
Degree Grantor:
University of Colorado Denver
Degree Divisions:
Department of Anthropology, CU Denver
Degree Disciplines:
Anthropology
Committee Chair:
Hodgkins, Jamie M.
Committee Members:
Beekman, Christopher
Stone, Tammy

Notes

Abstract:
There are several theories regarding the extinction of Neandertals by 40,000 years ago. The climatic stress hypothesis posits environmental instability during the Late Pleistocene caused stress among already declining populations of Neandertals and other large cold-adapted mammals across Europe leading to their extinctions. If there was a negative climatic change causing stress for Neandertals, changes in butchering intensity as preserved on faunal remains should be present among assemblages. The purpose of this thesis is to provide a taphonomic and zooarchaeological analysis of four layers at newly excavated Italian cave site Arma Veirana (AV) located in the Ligurian Alps. Liguria may have served as a temperate area of refugia during global glacial events as fauna diversity is described as rich and continuous throughout the Late Pleistocene. This study sought to better understand how Neandertals reacted to changes in their environment and how they used the cave through time by reporting data on species representation, agents of bone accumulation, and the post-depositional destruction of the site. Tests were performed to evaluate changes in subsistence behaviors at AV and to explore the climatic stress hypothesis. This study specifically analyzed the presence of prey species (e.g. deer) in the cave and looked for evidence of carcass processing. Prey body size at AV fluctuates through time, with all layers preserving higher frequencies of medium-sized prey. Two layers preserves larger sizes which may signify a period of dense forest growth and more reliable resources. Transport pattern analysis results show Neandertals were making decisions at kill sites on what body parts to transport back to the difficult to access cave. Analysis of surface modification on both high and low utility bones shows an increase in percussion marks per fragment (after controlling for fragment size) between two layers. Carnivore activity increased as Neandertals utilized the cave less over time. Carnivores modified bones only in the layer with the highest frequency of carnivore remains (9.26%). Faunal assemblages from Arma Veirana reject the hypothesis that Neandertals were unable to adapt subsistence behaviors to deal with nutritional stress as there is evidence for a change in nutrient extraction intensity.

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A ZOOARCHAEOLOGICAL ANALYSIS OF NEANDERTAL CAVE SITE ARMA VEIRANA
IN LIGURIA, ITALY
by
BREEANNA CHANTEL CHAROLLA B.A. Arizona State University, 2015
A thesis submitted to the Faculty of the Graduate School of the University of Colorado in partial fulfillment of the requirements for the degree of Master of Arts Anthropology Program
2019


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This thesis for the Master of Arts degree by Breeanna Chantel Charolla has been approved for the Anthropology Program
by
Jamie M. Hodgkins, Chair Christopher Beekman Tammy Stone
Date: August 3, 2019


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Charolla, Breeanna Chantel (M.A., Anthropology Program)
A Zooarchaeological Analysis of Neandertal Cave Site Arma Veirana in Liguria, Italy Thesis directed by Assistant Professor Jamie M. Hodgkins
ABSTRACT
There are several theories regarding the extinction of Neandertals by 40,000 years ago. The climatic stress hypothesis posits environmental instability during the Late Pleistocene caused stress among already declining populations of Neandertals and other large cold-adapted mammals across Europe leading to their extinctions. If there was a negative climatic change causing stress for Neandertals, changes in butchering intensity as preserved on faunal remains should be present among assemblages. The purpose of this thesis is to provide a taphonomic and zooarchaeological analysis of four layers at newly excavated Italian cave site Arma Veirana (AV) located in the Ligurian Alps. Liguria may have served as a temperate area of refugia during global glacial events as fauna diversity is described as rich and continuous throughout the Late Pleistocene. This study sought to better understand how Neandertals reacted to changes in their environment and how they used the cave through time by reporting data on species representation, agents of bone accumulation, and the post-depositional destruction of the site. Tests were performed to evaluate changes in subsistence behaviors at AV and to explore the climatic stress hypothesis. This study specifically analyzed the presence of prey species (e.g. deer) in the cave and looked for evidence of carcass processing. Prey body size at AV fluctuates through time, with all layers preserving higher frequencies of medium-sized prey. Two layers preserves larger sizes which may signify a period of dense forest growth and more reliable resources. Transport pattern analysis results show Neandertals were making decisions at


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kill sites on what body parts to transport back to the difficult to access cave. Analysis of surface modification on both high and low utility bones shows an increase in percussion marks per fragment (after controlling for fragment size) between two layers. Carnivore activity increased as Neandertals utilized the cave less over time. Carnivores modified bones only in the layer with the highest frequency of carnivore remains (9.26%). Faunal assemblages from Arma Veirana reject the hypothesis that Neandertals were unable to adapt subsistence behaviors to deal with nutritional stress as there is evidence for a change in nutrient extraction intensity.
This form and content of this abstract are approved. I recommend its publication.
Approved: Jamie M. Hodgkins


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ACKNOWLEDGEMENTS
I would first like to thank committee, Jamie Hodgkins, Tammy Stone, and Chris Beekman for all their comments and guidance along this journey. Jamie, thank you for the support and lessons you provided about being a woman in science, I am a master because you pushed me to be my best. I would also like to thank Dr. Curtis Marean for the experience of a lifetime (several times over) which allowed me to fully explore my love of archaeology, it was a pleasure and honor to learn from you.
The biggest thank you goes to my wonderful parents for always supporting me and my dreams— even if they happen to involve me playing around in the dirt all over the world. You provided me the strength to keep going by showing me what hard work and dedication truly looks like my entire life. You gave me the lessons and tools to see this degree through and I’m the luckiest daughter in the world to have parents like you constantly cheering me on— and yelling at me when I’ve let stress take control for a little too long. You kept me grounded over the last three years and I will be forever grateful and anxiously waiting to return the generosity you incessantly deliver.
Erin Clark, I will always appreciate your listening, support, and solutions to problems... real or imaginary. Thank you for the laughs, tears, and trips whenever one was needed most—my favorite will always be laughing with the raccoons in Booth Bay. Saundra Malanowicz, thank you for always being the sweetest person and a telepathic lifeline whenever the going got rough, your voice can forever sooth me.
While you might not find it significant, I can’t image the end of this chapter without you by my side, EA. It is something that will never go forgotten or taken for granted but always cherished.


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Last but absolutely not least, Sarah Maureen Simeonoff, words cannot express the reality of failure without you. I’m so lucky to have met you all those years ago in that dusty old room. Thank you for your unrelenting love and helping me see the good in everything. I’m so glad to have had you laughing our way through some of life’s biggest journeys thus far. I’m insanely honored to have you by my side on this one. Here’s to the countless hours spent encouraging each other to read one more article, write one more paragraph, make one more figure—this thesis could not have been completed, and this degree would have been unattainable without you, suffering right alongside me—I had the most fun with you.


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TABLE OF CONTENTS
CHAPTER
I. INTRODUCTION.......................................................1
Neandertals and the Late Pleistocene...............................1
Oxygen Isotope Stage 3.............................................2
Climatic Stress....................................................3
Neandertal Morphology and Physiology...............................4
Liguria, Italy and Arma Veirana....................................7
II. BACKGROUND........................................................10
Hypotheses for Extinction.........................................10
Competitive Replacement.....................................10
Climatic Stress.............................................14
Nutritional Stress..........................................14
Theoretical Basis of This Study...................................16
Behavioral Ecology..........................................16
Optimal Foraging Theory.....................................17
Middle Range Theory.........................................18
Arma Veirana......................................................19
Excavation Methods..........................................22
Cave Setting................................................25
Black Mousterian............................................27
Granular....................................................28
Compact Strong Brown........................................28
Rocky Brown.................................................29


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III. METHODS.............................................................30
Sample Selection and Data Collection................................30
Anthropogenic vs. Taphonomic Marks............................30
Taphonomic Assessment.........................................31
Trampling Marks...............................................32
Taxonomic Assessment..........................................33
Surface Modification Analysis.......................................34
Breakage Patterns.............................................35
Agents of Accumulation........................................36
Cut Marks.....................................................37
Percussion Marks..............................................38
Carnivore Tooth Marks.........................................40
Transport Strategies................................................40
Skeletal Abundance............................................40
Food Utility and Transport....................................41
Testing the Climatic Stress Hypothesis..............................43
Nutritional Stress............................................43
Statistical Analysis..........................................44
IV. RESULTS.............................................................46
Taphonomic Analysis and Taxonomic Representation....................46
Post-depositional Damage......................................46
Evidence of Cooking by Neandertals............................53
Taxonomic Analysis
55


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Prey Species Represented at AV.............................56
Spatial Representation of Fauna at AV......................56
Skeletal Element Representation............................60
Surface Modification Analysis....................................61
Breakage Patterns..........................................62
Agents of Accumulation.....................................65
Transport Behaviors..............................................67
Analysis of Nutrient Extraction Behaviors........................70
Evaluating Nutritional Stress..............................71
Spatial Analysis of Percussion Marks.......................73
V. DISCUSSION AND CONCLUSION........................................82
Discussion.......................................................82
Black Mousterian...........................................82
Granular...................................................83
Compact Strong Brown.......................................85
Rocky Brown................................................86
Conclusion.......................................................88
REFERENCES....................................................................90
APPENDIX
A. Transport data.....................................................107
B. ANCOVAdata.........................................................109


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CHAPTERI INTRODUCTION
Neandertals and the Late Pleistocene
Homo neandertalensis was first discovered in 1856 at the cave site Feldhofer Grotte in the Neander Valley of Germany and has ever since captivated the imagination of archaeologists and the public alike (Papagianni and Morse, 2013; Wynn and Coolidge, 2012). Neandertal skeletal remains and associated material culture have been recovered from sites covering great geographical range across Eurasia (van Andel and Davies, 2003; Villa and Roebroeks, 2014; Zilhao, 2014) and are known to have had a temporal distribution from 350-39 kya (Higham et al, 2014; Villa and Roebroeks, 2014). Understanding Neandertal extinction during the Late Pleistocene has long been a highly provocative topic within paleoanthropological research (Higham et al, 2014). It is especially interesting as modern humans (Homo sapiens) outlived Neandertals to ultimately inhabit the entire globe. Causes for Neandertal extinction throughout the paleoanthropological literature are frequently divided between population changes brought on by the emergence of modern humans in Europe and the competitive replacement of Neandertals upon their arrival (Banks et al., 2008; d’Errico and Gobi, 2003; Marean, 2007; Mellars, 2004, 2007; Mellars and French, 2011) and climatic induced stress experienced by Neandertals due to the unstable environment during the Late Pleistocene (Dogandzic and McPherron, 2013; Higham et al., 2014; Finlayson et al., 2004; Villa and Roebroeks, 2014). This study contributes to this discussion by examining faunal material recovered from cave site Arma Veirana located in Liguria, Italy to better understand how


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Neandertals reacted to changes in their environment and investigate how the cave was used through time.
Oxygen Isotope Stage 3
The climate and environment of the Late Pleistocene is frequently discussed as chaotic and unstable through the oxygen isotope record 60-20 kya, also known as Oxygen Isotope Stage Three (OIS-3). Extensive research organized by van Andel and Davies (2003) has been done on OIS-3 to better understand the role climate change played in changes in Neandertal populations. This pair organized an interdisciplinary project which incorporated biological anthropology, archaeology, zooarchaeology, geology, palynology, paleoentomology, topography, and paleoclimatological variables from GISP2 Greenland ice cores (Grootes et al., 1993; van Andel and Tzedakis, 1996; Meese et al., 1997; Stuiver and Grootes, 2000; Delmotte et al., 2001; van Andel, 2002) and three pollen cores extracted from Lago Grande di Monticchio in Italy (van Andel and Tzedakis, 1996; Watts et al., 1996; Allen et al., 2000; van Andel and Davies, 2003). Ice cores yield alternating warm and cold phases on 100-to-1000-year times scales, termed Dansgaard-Oeschger (D-O) phases (Grootes et al., 1993; van Andel, 2002; Huntley et al., 2003; van Andel and Davies, 2003). To investigate the role quick climatic changes played on Neandertal extinction van Andel and Davies (2003) focused on 3 major phases: the Warm Transitional (57-38 ka) and Early Cold (37-28 ka) to investigate Neandertal and modern human population changes across Europe and the Last Glacial Maximum (LGM, 27-18 ka) to better understand environments and demographics at the OIS-3/OIS-2 boundary. These data were used to create rigorous and robust paleoclimatic and paleoenvironmental databases, models, and simulations of OIS-3 and its effects on Neandertal, modern human, and large mammalian population


3
distributions across Europe (Barron and Pollard, 2002; Pollard and Barron, 2003; Alfano et al„ 2003).
Climatic Stress
Climatic stress is recognized as a possible contributing factor for Neandertal extinction. The global climate of the OIS-3 was unstable, characterized by "rapid oscillations in temperatures, within a millennium”— these oscillations have been linked to a decline in megafauna populations across Europe (Dansgaard et al., 1993; Finlayson and Carrion, 2007; Stewart, 2003b:116; Stewart et al., 2004). Drastic changes in the climate and environment would have negatively impacted abundance of flora and fauna. Therefore, it has been hypothesized climatic fluctuations were stressful for Neandertal populations who lived as hunter gatherers and extracted resources from their local ecosystems (Finlayson and Giles Pacheco, 2000; Finlayson et al., 2001, 2004, 2008; Hodgkins et al., 2016; Hublin and Roebroeks, 2009, Stewart, 2003b). As explained by Hodgkins et al. (2016:1): "if cold climates stressed Neandertals, their subsistence behaviors may have changed, requiring intensified use of prey through more extensive nutrient extraction from faunal carcasses”.
To investigate the role climate change had on Neandertals and other large mammalian populations, the distribution of five large extant and extinct taxa ranging in spatial and temporal distribution across western and central Europe during the Warm Transitional, Early Cold, and LGM periods were compiled in the Stage 3 Mammalian Database (Stewart et al., 2003a; Stewart et al., 2003b; Stewart, 2005). Totaling 468 dated faunal remains from 294 sites, the database includes 1912 radiometric dates, ultimately assigning dates to 119 mammalian taxa and allowing for detailed climate-mammal model simulation comparisons (Stewart et al., 2003a). Results from these simulations indicate


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taxon, geographic range, and climatological variables such as temperature and precipitation tolerance were fundamental to mammalian distribution through OIS-3 (Stewart et al., 2003a) and that Neandertals indeed were among those fauna that saw massive declines in population and ultimately extinction at the end of OIS-3 (Stewart et al., 2003b). Population declines in megafauna taxa such as mammoths (Mammuthus primigenius) and woolly rhinos (Coelodonta antiquitatis] (Stewart et al., 2004; Finlayson and Carrion, 2007) and carnivores such as cave bears [Ursus spelaea] (Stewart et al.,
2003b) are frequently observed across Europe at the end of OIS-3. These extinctions support the climatic stress hypothesis as it appears various large cold-adapted mammals including Neandertals were negatively impacted by the changing climate.
Neandertal Morphology and Physiology
In comparison to modern humans, Neandertal skeletal morphology indicates they were robust: relatively shorter, squatter with stocky limbs, barrel-chests, and exhibited muscular hypertrophy (Finlayson, 2004; Holliday, 1997; Pearson etal., 2000; Weaver 2003, 2009). Neandertal physiology implies they were a species that evolved to be optimally adapted to the glacial climates of the Pleistocene (Holliday, 1997; Pearson et al., 2000; Weaver, 2003, 2009; van Andel and Davies, 2003; Finlayson, 2004, 2009; Hublin, 2009). Ecogeographic biological rules provide reasoning for animals within the same species appearing noticeably dissimilar in direct relation to maintaining optimal core body temperatures (Bergmann, 1847; James, 2018). Allen’s rule stipulates bodies with short, stocky limbs are better at conserving heat, therefore better adapted to life in cold temperatures (James, 2018). Bergmann’s rule (1847) was first put forth to detail a newly observed pattern among animal size and geographic distribution as "large-bodied animal


5
species tend to live further north than their small-bodied relatives” (Blackwell et al., 1999:165). Larger overall body sizes as compared to other species within the same genus can be linked to locations colder in temperature and that species’ ability for optimal body thermoregulation (Bergmann, 1847; Blackwell etah, 1999; James, 2018).
Although most features of Neandertal bodies support the idea they were optimally adapted to the cold (short limbs but stocky, robust trunks), the face appears to deviate from patterns of cold adaptation observed in modern humans (Franciscus, 2003; Holton and Franciscus, 2008; Rae et ah, 2011; Weaver, 2003, 2009). For example, the depressed nasal floors common in Neandertals are also common among modern human populations in sub-Saharan Africa (Franciscus, 2003) and the wide nasal aperture common of Neandertals is commonly found in modern humans in equatorial regions (Holton and Franciscus, 2008). Although these features are not indicative of cold adaptation, "the narrow superior internal nasal dimensions, tall nasal apertures, and projecting nasal bridges of Neandertals could be, because these features are typically found in high-latitude” modern humans (Weaver, 2009:16031). Interestingly, certain skeletal elements, for example "the low angle between the neck of the femur and the shaft is a characteristic of highly active individuals (Trinkaus, 1993),” signifying Neandertals were also adapted for a highly active lifestyle (Finlayson, 2004:82).
The inference Neandertals were highly active has been investigated through studies of energy expenditure in comparison to various modern human populations (Churchill, 2006; Froehle and Churchill, 2009; Mateos et al., 2014; Snodgrass and Leonard, 2009; Sorensen and Leonard, 2001). The assumed high body mass, high activity level, and basal metabolic rate (BMR) of Neandertals have been calculated using predictive equations and


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adjusted for the negative impact cold weather and stress possibly had on thermoregulation (Mateos et al., 2014). There exist several methods for calculating Neandertal Total Energy Expenditure (TEE) (Mateos etal., 2014). Sorensen and Leonard (2001) estimated Neandertal TEE "using sex-specific mean Neandertal masses and World Health Organization (WHO) equations to predict BMRs and eventually adjusting for the effects of cold climate on BMR” (Mateos et al., 2014). Other researchers have calculated Neandertal TEE using ethnographic data of caloric consumption of arctic hunter-gatherer populations while correcting for the more robust Neandertal (Steegmann etal., 2002); or using BMR calculations based on assumed skin surface area (Churchill, 2006). Results from these studies stipulate Neandertal TEE values ranged between 3500-5000 calories per day (Mateos et al., 2014)— a higher range than modern hunter-gatherer groups requiring 3000-4000 calories per day (Hublin and Roebroeks, 2009). Perhaps Neandertals were at an advantage over modern humans in conserving body heat; however, such a muscular and robust hominin would have required more calories. Did a higher caloric intake cause stress for Neandertals when faced with heightened nutritional demands brought on by ecological ramifications of OIS-3? Paleoanthropological research aims to better understand the reasons for Neandertal extinction as they appear to have been optimally adapted for life in glacial environments, thrived for more than 300 thousand years (Villa and Roebroeks,
2014), and were likely the sole hominin inhabitants of Europe for 160 thousand years (Stewart, 2003b). Thus it is interesting these hominins did not persevere through the end
of the Pleistocene like modern humans.


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Liguria, Italy and Arma Veirana
Neandertal skeletal remains or material culture do not appear in the archaeological record after 40 kya (Banks et al., 2008; d’Errico and Sanchez Goni, 2003; Marean, 2007; Mellars, 2007; Mellars and French, 2011), indicating they went extinct across Europe within the Warm Transitional phase of OIS-3 (van Andel and Davies, 2003). Interestingly, Neandertal sites during this phase proved to have had the widest dispersal and show a preference for coastal regions of France, Italy, Portugal, and Spain (van Andel and Davies, 2003); it is possible these locations had more stable and reliable environments than those further north and served as areas of refugia (Banks et ah, 2008). Italy serves as a critically important region for studying Neandertal behaviors as the archaeological record from Italy includes some of the longest lasting Mousterian lithic assemblages in Europe (Churchill and Smith, 2000; Higham et ah, 2014). The Mousterian is a Middle Paleolithic tool technology exclusively associated with Neandertals across Europe and is characterized as a "widespread flake-based technology with a variety of retouched implement types” with rare occurrences of bifacial tools such as hand axes (Finlayson and Carrion, 2007:215).
The purpose of this thesis is to provide a taphonomic and zooarchaeological analysis of faunal assemblages recovered from a newly excavated Middle-to-Upper Paleolithic alpine cave site Arma Veirana (AV) located in Liguria, Northwest Italy. AV is a promising site to better understand Neandertal subsistence behaviors as it preserves archaeological layers with numerous Mousterian lithics and copious faunal remains. Zooarchaeology is a subfield within archaeology that utilizes a host of methods to determine how hunter-gatherers were accessing food resources in the past. Taphonomy is crucial to zooarchaeological analyses as it is the study of post-depositional damage of faunal remains


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from the time an animal dies to when its bones are analyzed (Dibble et al., 2006; Gifford-Gonzalez, 1991; Fisher, 1995; Lyman, 1987). Taphonomy allows for investigation of post-depositional damage and destruction of whole sites and of individual archaeology remains overtime. Therefore, it is critical to zooarchaeology as it allows researchers avenues to determine between a myriad of natural processes and anthropogenic modification of bones (hunting and butchering), either scenario providing dissimilar and unique site narratives.
The mammalian fauna of the Warm Transitional phase in Liguria have been characterized as taxonomically diverse between the earlier OIS-6 and OIS-3 (Valensi and Psathi, 2004). Specifically, the abundance of carnivores is known to have been high, suggesting a "richness of this region in ungulates and small mammals,” an environment that would have been beneficial in terms of prey species for Neandertals as well as carnivores (Valensi and Psathi, 2004:258). Forest taxa such as Cervus elaphus are well represented in Liguria during OIS-3, signifying periods of dense forest growth (Valensi and Psathi, 2004). Additionally, alpine taxa like Capra ibex are well represented (Valensi and Psathi, 2004) which indicate periods of little snow cover (Grignolio et al., 2004). While the climate of OIS-3 is commonly characterized as oscillating "from as warm as present-day during the last interglacial to as cold as those of the last glacial maximum” (Hardy, 2009:662), Liguria appears not to have suffered as greatly as other locations further North in Europe (Valensi and Psathi, 2004). In fact, "the Ligurian region suggests that the climatic oscillations registered during the period between OIS 6 to OIS 3 were not very pronounced” (Valensi and Psathi, 2004:260).
Arma Veirana preserves four Neandertal-associated archaeological layers with abundant faunal remains, a number of which preserve evidence of butchering and cooking.


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Preliminary radiocarbon dates show that Neandertals were utilizing AV more than 45,000 years ago. Fauna diversity is described as rich and "both relatively high and fairly continuous between OIS 6 and OIS 3, suggesting temperate climatic conditions even during the coldest periods” (Valensi and Psathi, 2004:258). Thus it is possible Neandertal subsistence behaviors would have stayed relatively constant throughout stratigraphic layers at AV.
This study specifically analyzed the presence of prey-species in the cave and looked for evidence of carcass processing by Neandertals. Taphonomic patterns such as bone fragment size, type of bone breakage, and the frequency of epiphyses to shaft fragments were also recorded to assess the preservation of faunal material at the site to better understand site use through time. Methods were used to determine if Neandertals were the sole bone accumulators at the site or if carnivores also had access to the bones and modified them. This is an exploratory investigation of the cave regarding the intensity by which Neandertals or other agents such as carnivores accumulated and modified faunal
remains in the cave over time.


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CHAPTER II BACKGROUND
Hypotheses for Extinction
Reasons behind Neandertal extinction remain hotly debated among paleoanthropologists. Competitive replacement (Banks et al., 2008; Marean, 2007; Mellars, 2004, 2007; Mellars and French, 2011) and climatic induced stress serve as the main competing theories for Neandertal extinction 39 kya (d’Errico and Gobi, 2003; Dogandzic and McPherron, 2013; Higham et al., 2014; Finlayson etal., 2004; Villa and Roebroeks, 2014). Limited cognition is a reoccurring factor in the argument for the competitive replacement of Neandertals by modern humans, as they entered Europe roughly the same time Neandertals stop appearing in the archaeological record (Banks et al., 2008; d’Errico and Sanchez Goni, 2003; Marean, 2007; Mellars, 2007; Mellars and French, 2011). The climatic stress hypothesis states that the negative environmental impacts brought on by the instability of OIS-3 were stressful for dwindling Neandertal populations across Europe and ultimately contributed to their extinction (Finlayson and Carrion, 2007; Hodgkins et al., 2016; Hublin, 2009; Hublin and Roebroeks, 2009).
Competitive Replacement
New research proposes modern humans first evolved in Northern Africa as early as 300 kya (Callaway, 2017) before leaving Africa, entering the Near East and ultimately Europe by 45 kya (Villa and Roebroeks, 2014). Replacement of all archaic human populations (such as Neandertals) unable to adapt to climate changes through competition with cognitively superior modern humans serves as a popular avenue for understanding drastic demographic changes in Europe (Banks et al., 2008; d’Errico and Sanchez Goni,


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2003; Leroyer and Leroi-Gourhan, 1983; Leroyer, 1988; Marean, 2007; Mellars and French, 2011; Shea, 2003). Research proposes Neandertals and modern humans co-existed in Europe for roughly 5,000 years (Higham et al., 2011, 2014; Pinhasi et al., 2011) with evidence for site chronologies overlapping (Galvan et al., 2014; Higham et al., 2014; Wood et al., 2013). Poor cognitive ability (Marean, 2007; Mellars and French, 2011; Shea, 2003) and Neandertal inability to adapt to climate changes have long been used as factors in the competitive replacement of Neandertals (d’Errico and Sanchez Goni, 2003; Leroyer and Leroi-Gourhan, 1983; Leroyer, 1988; Marean, 2007).
It is claimed that as modern humans entered areas inhabited by Neandertals there was "direct competition for space and resources” (Mellars, 2004:4) as a result of climatic instability (Mellars, 2004; Mellars and French, 2011). Researchers also propose the initial modern human populations in Europe were a powerful factor in demographic and territorial competition (Mellars and French, 2011). It is claimed that modern human "expansion resulted in competition with which the Neanderthal adaptive system was unable to cope” (Banks et al., 2008:5). It is argued that Neandertal hunting and foraging abilities were lacking as compared to modern humans and served as a contributing factor for their replacement (Marean, 2007). However, predictive models show that competition for food resources would not have proved great enough for competition until 20 kya (Sprensen, 2011), long after Neandertals went extinct. Therefore, "if Neandertals were not... "disadvantaged”, how can we explain that they did not survive?” (Villa and Roebroeks, 2014:6).
The literature contends that Neandertals were indeed cognitively capable of adapting to climatic changes (Dogandzic and McPherron, 2013; Finlayson et al., 2004; Villa


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and Roebroeks, 2014). To say Neandertals were cognitively inferior to modern humans (Mellars and French, 2011; Wynn and Coolidge, 2008, 2011; Wynn et al., 2016) and thus the reason they were susceptible to competitive replacement is to say there is evidence for such inferiority within the archaeological record. There would need to be archaeological inference for direct competition, for example, evidence of modern human attack on Neandertals over resources (Marean, 2007). Yet there remains no archaeological evidence for direct competition between Neandertals and modern humans (Finlayson et al., 2004; Shea, 2003). In fact, the archaeological record supports strong cognitive abilities in terms of symbolic expression (Villa and Roebroeks, 2014; Zilhao, 2014), contradicting the persistent assumption of an inferior trait of Neandertals as compared to modern humans and a key cause for their demise (Mellars and French, 2011; Wynn and Coolidge, 2008, 2011; Wynn et al., 2016).
Perforated and pigment-stained marine shells from Africa and the Near East are "widely accepted as evidence for body ornamentation, implying behavioral modernity” of early modern humans throughout the Middle Stone Age in Africa and the Upper Paleolithic in the Near East (Zilhao et al., 2010:1023). Yet, there remains disagreement on what is considered complex symbolic behavior regarding Neandertal cognition (Villa and Roebroeks, 2014; Zilhao, 2014). Some researchers deem Neandertals cognitively inferior to modern humans (Mellars and French, 2011; Wynn and Coolidge, 2008, 2011; Wynn et al., 2016) yet there exists a surplus of evidence that supports Neandertals were behaviorally and cognitively indistinguishable from modern humans in terms of symbolic expression as displayed throughout the archaeological record (Villa and Roebroeks, 2014; Zilhao, 2014).


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New research (Hoffman etal., 2018) proposes Iberian Neandertals were using "marine shells such as beads and pigment containers... as early as 115 ka, predating any [modern human] expression of symbolism in Eurasia” (Rodriguez-Hidalgo at al., 2018:2). The Neandertal archaeological record includes cases of decorated bone tools (Caron etal., 2011; Morin and Laroulandie, 2012), marine shells (d’Errico etal., 2010; Hoffman etal., 2018; Peresani et al., 2013; Roebroeks et al., 2012; Zilhao et al., 2010) and raptor talons (Morin and Laroulandie, 2012; Radovcic etal., 2015; Rodriguez-Hidalgo atal., 2018; Romandini et al., 2014) altered for adornment, and the use of natural pigments as colorants (Caron etal., 2011; d’Errico etal., 2010; Hoffman etal., 2018; Roebroeks etal., 2012; Zilhao et al., 2010).
Zooarchaeological analyses of avifaunal assemblages across Europe have found Neandertals took care during the removal of raptor talons from birds of prey (Morin and Laroulandie, 2012; Rodriguez-Hidalgo atal., 2018; Romandini etal., 2014) as early as 130 kya (Radovcic et al., 2015). Eight white-tailed eagle talons have been recovered from Kirpina in Croatia (130 kya) and show minimal cut marks with polished facets in roughly the same places indicating the talons were part of a jewelry assemblage (Radovcic et al.,
2015). As talons contain zero nutritional value, it is inferred that the careful extraction techniques must have been implemented to remove talons to produce minimal damage by lithic tools (Romandini et al., 2014). Researchers suggest that as large raptors are some of the rarest birds in the world, "these large and powerful diurnal birds attracted hominins and stimulated their use as a symbolic media by Neanderthals” (Romandini et al., 2014:9). Whatever the reason, it appears Neandertals took inordinate care in the extraction of raptor talons for adornment—indicative of symbolic behavior and cognition. If Neandertals


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were symbolically indistinguishable from modern humans within the archaeological record (Villa and Roebroeks, 2014; Zilhao, 2014), there must be a reason other than competitive replacement to account for Neandertal extinction and modern human survival during unstable OIS-3.
Climatic Stress
It is argued that demographic changes which occurred after the arrival of modern humans in Europe were not uniquely due to Neandertal "cognitive and technological inferiority causing rapid and total replacement” (Villa and Roebroeks, 2014:7). In fact, competitive replacement research by d’Errico and Goni (2003) reinforced the role climate change played in Neandertal extinction (Finlayson et al., 2004), which indirectly influenced the climatic stress hypothesis. This hypothesis states that "Neandertal extinction was driven in part by the rapid succession of climate changes at the end of the Pleistocene [and] assumes that glacial cycles caused local environmental shifts that negatively impacted Neandertal populations” (Hodgkins et al., 2016:2). This stress is claimed to have been severe enough on Neandertal populations to not allow them to recover between climatic fluctuations (Finlayson and Giles Pacheco, 2000; Finlayson etal., 2001, 2004, 2008; Hodgkins et al., 2016; Hublin and Roebroeks, 2009) and is evaluated in the archaeological record via zooarchaeological analysis of nutritional stress displayed via changes in butchering intensity as preserved on faunal remains.
Nutritional Stress
Several zooarchaeological studies have exposed a pattern of Neandertals intensifying butchering of carcasses in a response to heightened nutritional stress (Castel et., 2017; Hodgkins et al., 2016; Sharon and Oron, 2013; Valensi et al., 2011). It is assumed


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that populations experiencing nutritional stress would not ignore bone marrow or grease as the caloric value of fat is nearly twice that of protein or carbohydrate (Outram, 2001). In France, Neandertal butchering strategies are known to include intensive processing of bones indicative of grease rendering (Castel et al., 2017). Intensification of butchering manifests in the zooarchaeological record via an increase in the frequency of cut and/or percussion marks (Binford, 1978; Jin and Mills, 2011; Nilssen, 2000; Outram, 1999, 2000). In the Levant, Neandertals appear to have intensified butchering practices indicated by the presence of a high frequency of three distinct lithic tools indicative of "slicing and removing large quantities of flesh”— and evident from the high frequency of cut marks displayed on fauna (Sharon and Oron, 2013:184).
A controversial topic within the literature on Neandertal social and subsistence behaviors is cannibalism observed at Moula-Guercy cave in southeast France (120-100 kya) (Valensi et al. 2011:48). Moula-Guercy has "yielded over a hundred Neandertal remains with evidence of cannibalism on six specimens” including two adults, 2 adolescents aged 15/16 years, and two youths aged 6/7 years (Valensi et al., 2011:53). The head, ribs, and vertebral columns show minimal cut marks, implying that the butchers "took great care in cutting up the corpses” (Valensi et al., 2011:53). A comparison between Neandertal remains and Red deer (the most abundant fauna) indicates similar processing practices: butchering marks are consistent with processing for nutritive extraction including skinning, disarticulation, and defleshing along with intensive breakage to extract the brain, bone marrow, or mandibular tissue (Valensi et al., 2011). This shows intensity in butchering and is indicative of elevated levels of nutritional stress (Binford, 1978; Jin and Mills, 2011; Nilssen, 2000; Outram, 1999, 2000). The nutritional stress of Neandertals at


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Moula-Guercy must have been great as it seems they were forced to eat their own yet practiced a higher level of care in the butchering of people by leaving fewer cut marks than the preserved on the hunted prey like Red deer.
These discussed intensified subsistence strategies employed by Neandertals follow behavioral ecological predictions of optimal foraging theory (OFT) (Binford, 1978) as "the extent to which bones are exploited for their fat depends upon the level of dietary stress the community is under” (Outram, 1991:104). This study was conducted under a behavioral ecology theoretical framework which allows zooarchaeological analysis and interpretation of hominin behavior over time.
Theoretical Basis of This Study Behavioral Ecology
Behavioral ecology is a robust theoretical perspective often implemented in the research of hominins as it assumes "behavioral diversity is largely the result of variability in specific socioecological settings” (Bird and O’Connell, 2006:146) and that organisms are subject to behavioral evolution relative to ecological conditions (Nettle etal., 2013). In other words, it is the study of how organisms adapt behaviorally to a wide array of social and ecological circumstances in fitness-enhancing ways (Boone and Smith, 1998). Behavioral ecology explores "why certain patterns of behavior have emerged and continue to persist and looks to their socioecological context in seeking answers” (Bird and O’Connell, 2006:144). Behavioral ecology is a valuable theoretical model in application of Neandertal research, including reasoning of changing subsistence behaviors to adapt to changing environments.


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Optimal Foraging Theory
OFT was originally developed in the late 1970s under an evolutionary or behavioral ecology theoretical framework (Krebs, 1978; Pyke et al., 1977) before being applied to archaeological research (Binford, 1979; Keene, 1983). OFT relies on an assumed intimate relationship between hunter-gatherers and their environment to explain different mobility patterns across time and space (Keene, 1983; Kelly, 1983; Kelly and Todd, 1988). In addition to evolutionary concepts like natural selection and adaptation, OFT further considers diet, group size, and foraging location to interpret prehistoric behavior (Keene, 1983). OFT is beneficial in Neandertal behavioral research as it has been applied archaeologically and ethnographically (Binford, 1978; Levin and Potpov, 1964; Morin, 2007) and is considered "a constructive approach to the study of Pleistocene foraging strategies” (Dusseldorp, 2012:2).
Drastic climatic changes alter the environmental carrying capacity, which is described as "a theoretical concept that represents the maximum amount of biomass an ecosystem may support” (Rodriguez et ah, 2014:122). In other words, carrying capacity is the level of subsistence readily available to both herbivores and carnivores across the landscape, determined by a variety of ecological factors such as temperature, rainfall, and solar radiation (McNaughton etah, 1989; Oesterheld etah, 1992; Nemani etah, 2003; Rodriguez et ah, 2014). Changes in these ecological factors directly influenced the carrying capacity of an environment— the fauna and flora available for hunting and gathering by Neandertals across Europe (van Andel and Davies, 2003). In contrast, if an environment and therefore carrying capacity is stable through time, zooarchaeological assemblages should not show a distinct change in subsistence behavior through time.


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Zooarchaeological analyses operate under an OFT framework to "isolate behavioural differences in the exploitation strategies represented” in faunal assemblages (Dusseldorp, 2012:2). In fact, OFT provides a link between the ease of acquisition of prey directly and the level of field processing of carcasses and the likelihood of preferential transport of certain skeletal elements in relation to the associated reward in terms of nutrition (Binford, 1978; Metcalfe and Barlow, 1992; Cannon, 2001, 2003; Faith, 2007). It is proposed that Neandertals required very high return rates from foraging (Snodgrass and Leonard, 2009); therefore, decreases in the environmental carrying capacity would have had severe impacts on Neandertal populations (Dussledorp, 2012). Intensified butchering strategies displayed by Neandertals in the zooarchaeological record (Castel et., 2017; Hodgkins etal., 2016; Sharon and Oron, 2013; Valensi etal., 2011) follow these expectations of OFT (Binford, 1980; Keene, 1983; Kelly, 1983; Pyke, 1984; Pyke et al.,
1977).
Middle Range Theory
Zooarchaeological studies of bone surface modification are predicated on middle range theory to provide testable experimental data to compare with archaeological data during analysis. First developed within sociology in the 1950s (Verhagen and Whitley, 2011:64), middle range theory sought to provide "a logical structure in which working hypotheses can be confirmed or negated... [and] inductive and deductive perspectives can be effective”. Within archaeology, middle range theory was spearheaded by Binford (1977) to promote methodological investigations of possible cause and effect processes of site formation (Kosso, 1991)—testing assumptions of uniformitarian principles, or the assumption that those processes occurring today also occurred in the past (Lyman, 1987).


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For example, uniformitarianism stipulates that because wind storms are common today, they too must have commonly occurred throughout prehistory. These assumptions are critical to zooarchaeological research as a wind storm has taphonomic implications for archaeological site interpretations—for example, cover and compaction of a site by animals can produce microstriations (called trampling marks) on faunal remains that closely resemble markings produces by butchering (Andrews and Cook, 1985; Behrensmeyer et al., 1986; Dominguez-Rodrigo etal., 2009; Gifford-Gonzalez, 1991; Fiorillo, 1989; Fisher, 1995; Olsen and Shipman, 1988; White, 1992). It is therefore imperative zooarchaeology play a role of "interpreting evidence and testing claims about the past” through middle range theory (Kosso, 1991:623).
This zooarchaeological study adheres to a behavioral ecology theoretical framework to investigate subsistence strategies employed by Neandertals at the cave site Arma Veirana roughly 45 kya in the Liguria region of Northwestern Italy. This thesis is interested in how Neandertals reacted to their environment and reports a taphonomic and zooarchaeological assessment of their behavior in the cave through time.
Arma Veirana
Arma Veirana (AV) is an archaeological cave site situated at the base of a steep cliff (441 m.a.s.l. elevation) in the Neva Valley montane region of the Ligurian Alps in Northwestern Italy (Figure 1). The cave is situated approximately 14 km inland from the modern Mediterranean coast (44°08’45.4020” N, 008°04’18.8508” E) and just outside the small town Erli where the field laboratory is located. Initial archaeological investigation began in 2015, followed again in 2016, 2017, and 2018.


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Radiocarbon dating has been performed on numerous charcoal and bone samples recovered from Neandertal-associated layers throughout the cave. Unfortunately, these results require caution as the bottom of the sequence is older than 45 kya (Hodgkins, 2018 Personal communication), pushing the accuracy boundaries of radiocarbon dating. Therefore, this study assumes the earliest time of Neandertal occupation atAV was at least 45 kya: during the Warm Transitional phase of OIS-3 (57-38 kya) (van Andel and Davies, 2003). Due to the current lack of reliable radiocarbon dates, this study relies on field observations to provide diagnostic changes in sediment and artifacts following the laws of stratigraphy and superposition to account for changes in time.
Figure 1: Google Earth map showing location of Arma Veirana in relation to modern day Italy.
Archaeological excavations follow two key laws initially put forth in the field of geology: 1) the law of superposition, and 2) the law of stratigraphy (Harris, 1979; Howe, 1970). Within archaeology, the law of superposition stipulates the oldest layer must always be the bottom-most layer in an archaeological sequence, as it must have been deposited


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before the layers covering it were deposited (Harris, 1979). The law of stratigraphy was first detailed in the late 1700s to explain the apparent changes in sediment composition not solely based on rocks and dirt, but also fossilized organic remains such as plants and animals: a "law of strata identified by fossils” (Harris, 1979:111). Within archaeology, stratigraphy considers archaeological artifacts deposited within each layer in that "each level or stratum is dated to a time after that of manufacture of the most recent artefact found in it” (Harris, 1979:111). In Europe, many Middle Paleolithic (MP) to Upper Paleolithic (UP) sites have lacked the ability for reliable site reconstructions to investigate stratigraphic contexts and processes of site formation, typically due to the lack of application of modern technological excavation methods as they were originally investigated prior to inception of current methods (Hublin, 2015).
Archaeological deposits at AV have not been previously excavated, allowing for application of modern methods such as three-dimensional (3D) surveying and mapping of all archaeological materials and stratigraphic boundaries and contexts using digital total stations since the initial investigation of the cave in 2015. Additional 3D imaging methods such as stereophotogrammetiy and LiDAR scanning have been employed at AV to further investigate stratigraphy, geological cave setting, and on-going excavation settings. Photo-capable drones have been commissioned to develop a 3D model of the cave and surrounding valley aiding in spatial analysis of hominin landscape use. While these 3D imaging methods provide highly detailed and impressive data, they were not utilized in this study; however, they promise to provide insight into further analysis of the cave and paleoenvironmental reconstructions of alpine Liguria.


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Excavation Methods
Excavations at AV follow methods put forth by researchers at the Middle Stone Age (early modern human) cave and rockshelter site complex Pinnacle Point outside Mossel Bay, South Africa (Brown et al., 2010; Bernatchez and Marean, 2011; Dibble et al., 2007; Marean et al., 2004, 2010; Oestmo and Marean, 2014). The recording system at Pinnacle Point was designed to ensure all measurements "made on-site of artifacts, features, sections, surfaces, and everything else is recorded by total station directly to handheld computer” as this "give[s] all finds a 3D provenience” (Oestmo and Marean, 2014:5956) allowing for "piece-plotting of finds and samples, section drawing, feature measurement and drawing, and grid lay-out” (Bernatchez and Marean, 2011:1) with virtually no human transcription error (Dibble et al., 2007).
AV uses prismless Topcon Pulse total stations operated by Spectra Precision handheld tablet computers running Carlson Survey Pro Software (Hodgkins, 2017 Personal communication). Total stations are electronic instruments with a built-in distance meter to measure angles and distances resulting in a specific point position in space—point-specific coordinate data, which allow for reconstructing spatial relationships and 3D distributions of materials for reference after excavation and ultimate destruction of stratigraphic boundaries (Bernatchez and Marean, 2011; Brown etal., 2012: Dibble etal., 2007; Marean et al., 2004, 2010; McPherron et al., 2005). Geographic Information Systems (GIS), such as ArcMap (employed in this study), use these data to digitally map spatial relationships of excavated materials and excavation processes (Marean et al., 2004, 2010; McPherron et al., 2005) including stratigraphy and geologic formations often used to produce detailed paleoenvironmental reconstructions (Anemone etal., 2011).


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At AV, field observations are made "first by noting and documenting the geological and geographical setting of the site, followed by detailed sedimentological and pedological description of stratigraphic units” (Miller and Czechowski, 2016:1) to determine and assign unique and descriptive names to such stratigraphic units, or layers. Changes in geogenic and anthropogenic factors such as sediment composition (texture, color, moisture, etc.),
Figure 2: Entrance of the cave facing South, showing location of the main excavation trench near the mouth. Photo taken in 2016, courtesy of Dominique Meyer.


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artifact density, and depositional disturbances (rootlets, rodent burrows, etc.) contribute to classification of these stratigraphic layers (Miller and Czechowski, 2016).
Layers at AV are easily distinguishable in the field based on clear sediment differences (Miller and Czechowski, 2016) and are assigned as Stratigraphic Aggregates (StratAggs) and named after diagnostic sediment characteristics such as color and texture (Brown etal., 2012; Marean et al., 2004, 2010; Oestmo and Marean, 2014). There exist four StratAggs of analytical relevance currently exposed in the main excavation trench near the mouth of the cave (Figure 4) from lowest to highest: Black Mousterian (BM), Granular (Gr), Compact Strong Brown (CSB), and Rocky Brown (RB) (Figure 3).
RB
SO cm
CSB
Gr
BM
east wall profile
Figure 3: Profile view of the east wall within the main excavation trench displaying the contacts for the four StratAggs analyzed. The red star corresponds with Figure 4.
Individual excavation units are organized in the following ranked order: Square, Quad, StratUnit, and Lot Number (Lot) (Brown et al., 2012; Marean et al., 2004, 2010; Oestmo and Marean, 2014). There are four excavation units divided within each 1 m Square— 50 cm quadrants (Quads) are assigned on their directional bearing (NE, NW, SE,


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SW) and are allocated a unique Lot Number (Marean et al., 2004, 2010; Oestmo and Marean, 2014). All artifacts and features (regardless of size) that become exposed within these units are shot in with total stations and collected in individual specimen bags with unique catalogue barcode numbers (Dibble et al., 2007) which have been "assigned to a square, quadrant, and stratigraphic unit” (Oestmo and Marean, 2014:5956). Digital recording of artifacts, stratigraphy, and features are supplemented by handwritten excavator notebooks which are digitally preserved for future reference (Marean et al.,
2004, 2010; Oestmo and Marean, 2014). This two-step method of artifact collection is employed in an attempt to mitigate human error during recording (Dibble et al., 2007; Bernatchez and Marean, 2011).
Cave Setting
Neandertals are assumed to be the main inhabitants of the cave at least 45 kya as numerous faunal remains and Mousterian lithics characterized by the contemporary use of Levallois and Discoid methods produced on mainly local raw materials have been recovered from all four StratAggs analyzed here (Hodgkins, 2017 Personal communication). Preliminary lithic analysis of AV highlights the dominant utilization of Discoid production with few Levallois elements (Negrino et al., 2017; Riel-Salvatore and Negrino, 2018). Exploited raw materials prove mainly to be locally sourced quartz, quartzite, and jasper which is believed to have been sourced near the modern town of Ortovero (Negrino et al., 2016) approximately 20 km away. Additionally, a preliminary geoarchaeological investigation of AV has begun to assess stratigraphy throughout the cave to develop a model for site formation (Miller and Czechowski, 2016) following current methods of micromorphological analysis (Courty et al., 1989; Mallol and Mentzer, 2015).


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Combined, these preliminary results provide fine-grained details about sediment composition within each StratAgg that are valuable in evaluating the relationship between stratigraphy and cultural remains throughout the sequence aiding in interpretations of occupation of the cave (Miller and Czechowski, 2016).
Arma Veirana is a limestone cave formed along a fault which eroded an offset portion ofbedrockto form the cave as it is today (Miller and Czechowski, 2016). The cave is 44 meters deep from front to back, with an erosion-produced sloped surface displaying younger sediments preserved towards the back and older sediments exposed near the front (Figure 4; Hodgkins, 2017 Personal communication).
Jn Cave
44 m long
' 10 m high x 11 m wide
v minimum of 5 m of sediment
Figure 4: Cave cross-section derived from 3D models established using photogrammetry of the cave and surrounding valley. The red star indicates location of the main excavation trench.


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Deposits within the cave consist of coarse and fine-grained clastic sediments and sedimentary rocks such as flowstones and speleothems; a tufa-cemented breccia at the cave entrance suggests there used to be more deposits than today (Miller and Czechowski,
2016). In fact, it appears deposits at the front of the cave were thicker than at present and have eroded to expose Middle-to-Upper Paleolithic deposits near the cave entrance (Miller and Czechowski, 2016). Excavations at AV are arranged in several trenches to investigate the stratigraphic sequences throughout the cave (Miller and Czechowski, 2016). While there are numerous StratAggs exposed throughout the cave, this study is solely concerned with the four Middle Paleolithic deposits exposed in the main excavation trench: BM, Gr, CSB, and RB (from oldest to youngest). These four are of analytical interest as none of the contacts between these StratAggs "show clear evidence of erosion or stasis of deposition”; however, "this does not conclusively demonstrate evidence of continuous deposition, it does suggest that major erosive processes are relatively absent” in the StratAggs studied here (Miller and Czechowski, 2016:5)—potentially providing a continuous archaeological sequence at AV. While out of scope for this study, this holds great analytical intrigue for the Arma Veirana Project.
Black Mousterlan
The lowest StratAgg excavated thus far, Black Mousterian (BM) was named after the dark sediment color seemingly resultant of a high density of charcoal and Mousterian lithic artifacts (Hodgkins, 2017 Personal communication). The sediment in BM is defined as a medium-fine grained silt with granules and sub-angular bedrock fragments up to 10cm (horizontally oriented) and overall dark in color (10YR 3/2 Munsell) with a high proportion of charcoal preserved throughout (Miller and Czechowski, 2016).


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Micromorphological thin sections show clear anthropogenic activity via high densities of charcoal, burned organic material, and burned fat (Miller and Czechowski, 2016). Faunal and Mousterian lithic artifact densities are greatest in the BM and decrease from the bottom to the top of the sequence (Hodgkins, 2017 Personal communication), possibly suggesting a change in site use over time. Minimal zooarchaeological analysis of fauna recovered from AV had been performed prior to this study; however, initial analysis found butchery marks are common on bone fragments throughout the BM assemblage (Hodgkins, 2017 Personal communication). High densities of burned organic material, Mousterian artifacts, and butchery marks on faunal remains suggests Neandertals were using the cave as a butchering and/or cooking site during the BM at least 45 kya.
Granular
Sediment in Granular (Gr) is characterized as a medium sandy silt with granules, gravel, and sub-angular strongly weathered bedrock fragments (up to 5-10 cm) with a slight color difference within the StratAgg between the upper portion (10YR 4/3 Munsell) and lower (10YR 4/4 Munsell) with high artifact density present in the upper and diffusing toward the lower portion (Miller and Czechowski, 2016), this could represent a change in occupation and site use. Mousterian lithics and faunal remains are common in Gr and micromorphological thin sections show a clear contact between BM and Gr StratAggs marked by a decrease in anthropogenic evidence (Miller and Czechowski, 2016).
Compact Strong Brown
Contact between the Gr and Compact Strong Brown (CSB) is sharp with a distinct change in sediment and rock: fine sand/silt with a similar clay content as Rocky Brown, smaller (5 cm) subangualr rock fragments with higher proportion of schist instead of


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limestone (Miller and Czechowski, 2016). Micromorphological thin sections fail to show a clearly expressed contact between Gr and CSB, however they reveal evidence suggestive of gentle downslope movement of sediment in CSB that is not present in Gr (Miller and Czechowski, 2016). While anthropogenic components are rare in CSB (Miller and Czechowski, 2016), Mousterian lithics and faunal remains with butchery marks are found throughout (Hodgkins, 2017 Personal communication).
Rocky Brown
Rocky Brown (RB), is the youngest StratAgg analyzed and consists of silt and clay with granules, gravel, and large angular to sub-angular rocks (10-15 cm) oriented to the North dipping Northeast and is similar in color to Gr (10YR 4/4 Munsell) (Miller and Czechowski, 2016). RB has a higher proportion of coarse sediment material compared to CSB and the contact between the two has been noted by a thin gravel lens (Miller and Czechowski, 2016). Mousterian lithic artifacts and fauna are common and more abundant than in CSB yet lower than Gr and BM.


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CHAPTER III METHODS
Sample Selection and Data Collection
Following current zooarchaeological methods (Abe etal., 2002; Grayson, 1984; Lyman, 1994,1995; Marean etal., 2000, 2001), bone surface modification analysis at Arma Veirana (AV) was performed on each faunal fragment regardless of size and/or taphonomy. Taxonomic and taphonomic assessments were performed in the field laboratory located in Erli, Italy. Of particular interest to this study are the Neandertal-associated StratAggs with Mousterian lithic technology and copious faunal remains: Black Mousterian (BM), Granular (Gr), Compact Strong Brown (CSB), Rocky Brown (RB). Curation methods at AV allowed for random sampling of faunal fragments from these four StratAggs. All artifacts are organized by excavation square (Strat) and Lot number which permitted a random selection of a minimum of 10% of faunal fragments from the four StratAggs for analysis: BM n=713, Gr n=295, CSB n= 54, RB n= 112. Data collection was performed using forms in Microsoft Access to provide a uniform system regardless of analyst (Marean et al., 2001). Dino Lite Microscopes (<40x) with digital photographic capabilities were used to inspect surface modifications of each specimen, and to record all marks with the assistance of incident lighting(Blumenschine etal., 1996).
Anthropogenic vs. Taphonomic Marks
Taphonomy, or the study of what "happens between the death of an animal and the arrival of its bones in the laboratory” (Dibble et al., 2006:3) is a critical component of zooarchaeological analysis. Taphonomic analysis evaluates bone surface modification marks preserved on archaeological faunal remains to interpret agents of accumulation of


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bones and the processes that potentially contributed to the modification of remains for hundreds, thousands, or millions of years (Gifford-Gonzalez, 1991; Fisher, 1995; Lyman, 1987). To adequately distinguish anthropogenic and taphonomic markings, the field of zooarchaeology has commissioned a number of experimental and naturalistic studies under a middle range theoretical framework to contribute critical insight into prehistoric diets (Behrensmeyer et al., 1986; Blumenschine, 1988,1995; Blumenschine and Selvaggio, 1988; Blumenschine etal., 1996; Egeland etal., 2004; Faith, 2007; Faith and Gordon, 2007; Marean and Spencer, 1991; Olsen and Shipman, 1988; Pickering and Egeland, 2006; Villa and Mahieu, 1991).
Experimental studies and inter-analyst blind tests have shown bone surface modification analysis benefits from the use of low-magnification and incident lighting to best expose the presence of any markings preserved on faunal fragments (Blumenschine et al., 1996; Dominguez-Rodrigo etal., 2009). In blind tests, Blumenschine and colleagues (1996) accomplished impressive accuracy in discerning between cutting/scraping marks, hammerstone percussion marks, and carnivore tooth marks. They "determined the presence or absence of conspicuous and inconspicuous marks with 97% three-way correspondence” and "diagnosed marks of known origin to actor and effector with 99% accuracy” (Blumenschine et al., 1996:493). These experimental data allow for rigorous comparisons against zooarchaeological assemblages to determine agents of accumulation (i.e., hominin, carnivore) and effector (i.e., hammerstone, carnivore ravaging).
Taphonomic Assessment
Manual calipers were used to record the maximum length and width (mm) of specimens to investigate issues of preservation of faunal remains per StratAgg. Additional


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data such as color and fossilization level (none, light, heavy) were collected on all specimens to further evaluate changes in preservation in the cave over time. The overall surface visibility of fragments and obstructions such as matrix-cover and dendritic etching were recorded in 10% increments (.00-1.0). All zooarchaeological tests were performed on fragments with at least 20% surface visibility. The maximum burning stage was assigned on a scale from 0-6, with zero representing no evidence of burning and 6 as fully calcined bone which turns completely white and chalky (Nicholson, 1993; Shipman et al., 1984; Stiner et al., 1995). Long bone shaft fragment completeness was recorded on a scale of 0-3 in terms of bone circumference where: 1 demonstrates less than half of the element’s original circumference, 2 is more than half, and 3 is a complete circumference in at least one portion of the fragment (Bunn, 1983; Villa and Mahieu, 1991).
Trampling Marks
Taphonomic marks produced by forces such as sediment compaction (Behrensmeyer etal., 1989; Fisher 1992; Marean etal., 2000) and trampling (Olsen and Shipman, 1988; Fiorillo, 1989; Dominguez-Rodrigo and Barba, 2006; Dominguez-Rodrigo et al., 2009) contribute to bone surface modification. It is imperative zooarchaeological analysis properly distinguish intentional hominin-produced butchery marks from marks left by sediment compaction as they share several diagnostic criteria (Behrensmeyer et al., 1986; Dominguez-Rodrigo et al., 2009; Olsen and Shipman, 1988) yet provide vastly different site narratives during analysis.
Most researchers have relied on microstriations to characterize both: a) the marked similarities between cut marks and trampling marks (Figure 5), and b) the presence and/or frequency of microstriations to distinguish between these marks (Dominguez-Rodrigo et


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al., 2009). It is now well understood that trampling marks are "much shallower (and frequently irregular)” than flaked cut marks (both simple and retouched) (Galan and Dominguez-Rodrigo, 2014:1064). At AV, trampling was recoded whenever present to explore variations in site use and post-depositional destruction over time.
Figure 5: Example of experimentally produced trampling marks. Measurements highlight the shallowness, frequency, and irregularity of marks on a single fragment.
Taxonomic Assessment
Skeletal element (e.g. femur, humerus, long bone fragment), side of bone (right, left, unidentifiable), and portion of bone (e.g. epiphysis, midshaft, shaft) were assigned for all fragments whenever possible. If taxon identification was unavailable, a general taxonomic grouping was made (e.g. bovid, carnivore, mammal, etc.). Identification on the species level was limited due to the lack of a comparative collection in the field laboratory; those that were considered identifiable were brought back to the zooarchaeology laboratory at the University of Colorado Denver where a number of fragments were further identified to bone element, side, taxon, and even species. The four StratAggs analyzed here preserve remains of medium-sized artiodactyl (bovids/cervids) prey species, including Capra ibex (goat), Cervus elaphus (Red deer), and Capreolus capreolus (Roe deer). This follows the


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taxonomic representation of Liguria which has reported deer as the most common prey species at the time of Neandertal occupation at AV (Valensi and Psathi, 2004).
Animal body size categories were assigned on a scale from 0-6, with 0 representing small fauna such as rodents and birds and 6 representing large fauna such elephants (Brain, 1981; Bunn etal., 1988). For example, of the identifiable species at AV, Capra ibex (50-100 kg, Aublet et al., 2008) and Capreolus capreolus (15-50 kg, Nowak and Wilson, 1999) are recorded as size 2 (20-113 kg) and Cervus elaphus (75-340 kg, Nowak and Wilson, 1999) are recorded as size 3 (113-340 kg) (Brain, 1981; Bunn etal., 1988). Animal body size is an important factor when considering transporting carcasses across the landscape as hunters favor smaller prey when travelling distances to butchering, cooking, and/or consuming sites (Binford, 1978; Schoville and Otarola-Castillo, 2014). Transporting heavy loads across tricky alpine terrain to a consumption site was undoubtedly a concern for Neandertals using the cave to butcher and consume their prey.
Surface Modification Analysis
Surface modification analyses at AV were conducted on "high survival” long bones including humeri, radii, ulnae, femora, tibiae, fibulae, metacarpals, and metatarsals, crania, and mandibles as they have thick cortical walls and are more likely to preserve under archaeological contexts (Faith and Gordon, 2007:873; Marean and Cleghorn, 2003). High survival long bones are used to quantify bone breakage patterns as they have "the highest probability of preservation at levels consistent with their discard” (Marean and Frey, 1997; Lam et al., 1999; Marean and Cleghorn, 2003; Cleghorn and Marean, 2004; Faith and Gordon, 2007) and minimize "the impacts of post-discard forces of bone destruction,


35
allowing for increased confidence in comparisons of abundance and modification” (Hodgkins et al., 2016:6).
The relative frequencies of long bone shaft fragments are compared to epiphyseal fragments in a critical first step to evaluate possible biases in excavation and artifact retention methods (Marean et al., 2000; Pickering et al., 2003). A higher frequency of epiphyseal fragments relative to shaft fragments has been found to be resultant of sampling biases (Marean, 1998; Marean and Kim, 1998; Marean et al., 2000) and "a good indication that shaft fragments were passed over for more identifiable epiphyseal fragments during excavation or curation” (Hodgkins and Marean, 2017:7). As this is the first zooarchaeological analysis of fauna at AV, it is important to maintain no biases in excavation, curation, or artifact sampling prior to analysis.
Breakage Patterns
Within zooarchaeology, long bone fragments (e.g. humeri, metapodials, etc.) are the most descriptive in terms of determining how and why faunal remains are present at a given site (Marean and Kim, 1998; Marean etal., 2000; Pickering etal., 2003; Villa and Mahieu, 1991). Following criteria outlined by Villa and Mahieu (1991), it is possible to distinguish between nutritive and non-nutritive bone breaks. Nutritive breaks occur when bones are broken in a fresh state— indicative of intentional agency by either hominins and/or carnivores with the intention of extracting bone grease and marrow from fresh bones which produces curved and/or v-shaped breakage outlines with oblique angles (Marean et al., 2000; Villa and Mahieu, 1991). In contrast, non-nutritive bone breaks are produced when bones are broken in a dry state— after a bone has lost all soft tissue, grease, or other nutrients, which produces transverse breakage outlines with right angles.


36
Marean etal. (2000) compiled experimental studies providing expected frequencies for zooarchaeological assemblages to compare data against. The breakage patterns of faunal remains at AV were compared against these expected frequencies where ‘Hominid Only’ and ‘Carnivore Only’ assemblages refer to those where humans and carnivores were the sole bone accumulators respectively while ‘Hominid to Carnivore’ depicts assemblages where humans had first access to bones for nutritional extraction, followed by carnivores. These expected frequencies serve as a baseline for the main ways in which bones are assumed to accumulate in archaeological settings.
Agents of Accumulation
Hominins have relied on hunting as a major source of nutrition since 1.76 mya (Fernandez-Jalvo et al., 1999; Thompson et al., 2019 In press), with a wide variety of hunting and butchering strategies that modify bone surfaces leaving behind different types of markings. Critical to zooarchaeology is the evaluation and assignment of diagnostic criteria for proper identification of bone surface modification marks to determine agents of bone accumulation. Caves and rockshelters were dominant residential locations for Neandertals in Western Europe (Marean, 2005). However, Neandertals were not alone in utilizing caves, carnivores such as canids are known to use them to den, thus contributing to accumulation of fauna (Brain, 1981) and/or post-depositional destruction of the site.
Agents of accumulation are determined through analysis of the relative frequencies (number of marks) of percussion marks and tooth marks preserved on bovid/cervid long bone fragments as these serve as proxies for hominin and carnivore activity respectively (Blumenschine, 1988,1995; Marean and Spencer, 1991; Marean etal., 1992, 2000; Marean and Bertino, 1994; Capaldo, 1995,1997). Comparing "the frequency and anatomical


37
distribution of marks, not simply their presence or absence on particular skeletal parts or taxa... are most informative about hominid behavioural ecology and site formation” (Blumenschine etal., 1996:504). At AV these frequencies are compared to expected frequencies as compiled and outlined by Marean et al. (2000) to establish the dominant bone accumulator and modifier for each StratAgg.
Butchery marks such as cut marks (Abe et al., 2002; Behrensmeyer et al., 1989, 1995; Blumenschine and Selvaggio, 1988,1991; Blumenschine et al., 1996; Bunn, 1981; Capaldo, 1995; Egeland, 2003; Fisher, 1992; Potts and Shipman, 1981) and percussion marks (Binford, 1981; Blumenschine, 1995; Blumenschine etal., 1996; Bunn, 1981,1989; Capaldo and Blumenschine, 1994; Dominguez-Rodrigo and Barba, 2006; Lyman, 1987; Outram, 2001; Pickering and Egeland, 2006) are produced in dissimilar ways and thus produce distinguishable marks. Both are recorded by frequency and location (i.e., epiphysis, near epiphysis, shaft) to investigate changes in butchering intensity between StratAggs (Outram, 1999, 2001).
Cut Marks
Experimental research began in the 1980s to produce diagnostic criteria for cut marks (Abe etal., 2002; Behrensmeyer etal., 1989,1995; Blumenschine and Selvaggio, 1988,1991; Blumenschine etal., 1996; Bunn, 1981; Capaldo, 1995; Egeland, 2003; Fisher, 1992; Potts and Shipman, 1981). Marks produced by cutting to remove flesh from bones can be produced by a variety of different tools such as flakes and bifaces or scrapers, which leave roughly similar mark morphologies (Blumenschine et al., 1996:496). Blumenschine and colleagues (1996) complied criteria from numerous studies to provide simple classification of marks to determine actor (human or carnivore) and effector through mark


38
morphologies. Cut marks produce deep V-shaped cross sections often with longitudinal microstriations (Figure 6) while scrapers produce more broad and shallow marks (Blumenschine etal., 1996:496). It is important to note that while these distinctions were often made during data collection at AV, the distinction between flake, biface, or scraper in this study was not taken into account and were all re-coded simply as cut marks during analysis.
Figure 6: Example of size 3 non-ID mammal specimen recovered from BM (Specimen ID: 1086). Measurements are included to show the possible frequency and size of cut marks preserved on a single fragment.
Percussion Marks
Extensive experimental research provides diagnostic criteria for identifying percussion marks (PMs; Binford, 1981; Blumenschine, 1995; Blumenschine and Selvaggio, 1988; Blumenschine etal., 1996; Bunn, 1981,1989; Capaldo and Blumenschine, 1994; Dominguez-Rodrigo and Barba, 2006; Johnson, 1985; Lyman, 1987; Outram, 2001; Pickering and Egeland, 2006). PMs are a suite of marks left from hominins using hammerstones to break open long bones exposing the medullary cavities and nutrient-rich marrow within (Blumenschine and Selvaggio, 1988; Blumenschine et al., 1996:496, White, 1992). They are classified as pits, grooves, isolated patches of microstriations (Blumenschine etal., 1996:496), and notches (Capaldo and Blumenschine, 1994).


39
Figure 7: Size 2 bovid/cervid metacarpal recovered from BM (Specimen ID: 2003). Measurements are included to show the possible frequency and size of percussion marks preserved on a single fragment.
Microstriations are very shallow "in and/or emanating from pits and grooves, oriented transverse to the long axis [of the bone] and occurring in dense superficial patches (Figure 7, Blumenschine etal., 1996:496). Percussion notches (Figure 8) are broad U-shaped bone scars resultant of the striking of a hammerstone on a fresh bone to break it open (Capaldo and Blumenschine, 1994). Similar to the cut mark re-coding, distinctions were made between percussion marks and notches during data collection, yet both were re-coded as percussion marks for simplicity in analysis.
Figure 8: Size 3 cervid metapodial recovered from BM (Specimen ID: 1967). Measurements are included to show the possible frequency and size of percussion notches preserved on a single fragment.


40
Carnivore Tooth Marks
Experimental studies have shown that while humans generally focus their subsistence strategies on extraction of meat and marrow from long bone shafts, carnivores focus their consumption around epiphyseal ends due to their high grease content which provides an excellent source of nutrients (Cleghorn and Marean, 2007; Marean and Spencer, 1991). Carnivores are known to seek out rapid access to bones once humans have discarded them (Binford, 1988; Blumenschine, 1988; Blumenschine and Marean, 1993; Capaldo, 1995,1998; Marean etal., 2000; Selvaggio, 1998). Carnivore tooth marks are known to be morphologically similar to hammerstone percussion marks as they both produce pits/notches on impact (Blumenschine and Selvaggio, 1988; Blumenschine etal., 1996). Binford (1981) first described tooth marks as "pits and scores”, which experimental studies have since critiqued to provide criteria to distinguish the two (Blumenschine et al., 1996:497; Capaldo and Blumenschine, 1994; Galen etal., 2008; Pickering, 2002). Transport Strategies Skeletal Abundance
Accurate skeletal element abundance data are imperative to reconstructions of transport strategies as they contribute to the calculation of the number of identifiable specimens (NISP), minimum number of individuals (MNI), minimum number of elements (MNE), and minimum number of animal units (MAU) within zooarchaeological assemblages. Skeletal element survivability plays a role in the likelihood of certain elements being present or absent from a zooarchaeological assemblage as low-survival skeletal elements with "thin cortical walls and low-density, grease-rich cancellous portions” including vertebra, ribs, pelves, scapulae, carpals, tarsals, and phalanges are


41
commonly absent from assemblages due to their weak nature (Faith and Gordon, 2007:873).
At AV, specific skeletal elements (e.g. femur, humerus), side of bone (right or left), and portion of bone (e.g. epiphysis, midshaft, shaft) were assigned whenever possible to calculate the MNE and MNI per StratAgg and investigate element abundance changes over time (Abe et al., 2002; Marean et al., 2001). To investigate food transport strategies, the MNE per StratAgg were used to calculate the MAU and the normed animal units (%MAU) to test the relationship between certain skeletal parts and their bone density values using the Standard Food Utility Index (SFUI) (Binford, 1978; Metcalfe and Jones, 1988). Low-utility skeletal elements do not hold much nutritional value and include metacarpals, metatarsals, crania, and mandibles; while high-utility elements yield plenty of flesh and are more attractive to hunters and include humeri, radii, femora, and tibiae (Faith and Gordon,
2007).
Food Utility and Transport
"The correlation between skeletal part abundance (%MAU) and bone density values allows us to estimate the differential preservation of the material” (Valensi and Psathi, 2004). The MAU was calculated using MNE values and diving by the number of times the bone would appear in a complete skeleton of the living animal (Binford, 1978; Grayson, 1984). For example, the MAU of an MNE of 6 Capreolus tibiae would be calculated by dividing the MNE (6) by the number of times tibiae appear in the body (2) (6/2 = MAU of 3). The %MAU was calculated using the highest valued MAU as a normed scale following Binford’s (1978) formula: dividing each MAU by the highest MAU value found at the site before being multiplied by 100. With the %MAU values calculated, they were compared to


42
the SFUI, or the expected amount of meat, marrow, and bone grease of each skeletal element. Comparing the %MAU of elements from each StratAgg to the SFUI allows for better understanding the relationship between bone utility and survivability (Metcalfe and Jones, 1988).
Animal body-part utility indices aid in the interpretation of zooarchaeological assemblages by determining levels of food utility via "meat, marrow, and bone grease associated with different body parts” and the likely ways in which hunters transported these parts for consumption (Binford, 1978; Faith and Gordon, 2007; Marean, 2005; Metcalfe and Jones, 1988:486; Schoville and Otarola-Castillo, 2014). As explained by Hodgkins et al. (2016:13), "as search time and/or distance to prey increases and/or encounter rates decrease, more field processing should take place so that lower utility parts will be left at the field processing sites, while higher utility parts will be transported to home base sites”. Typical patterns of bone accumulation in caves preserve higher frequencies of more complete transport of smaller animals (e.g. goats or small deer) and lower frequencies of more complete transport of larger animals as they would be too large to carry from a kill site to residential site (Marean, 2005). The "completeness of transport is a function of distance between the encounter site and the residential site. The farther the distance, the greater the filter on transport or anatomical units and bone” (Marean, 2005:372).
Within transport pattern analysis, zooarchaeological data are compared against four theoretical food utility curves. The three original utility curves, as proposed by Binford (1978) are: 1) bulk strategy—the entirety of a carcass except the lowest utility is transported; 2) gourmet strategy—only the highest utility portions of a carcass are


43
transported; and 3) unbiased strategy—portions of a carcass are transported in direct relation to their utility. Building off these, Faith and Gordon (2007) added a supplemental fourth utility curve: 4) the unconstrained strategy—where entire carcasses are transported. Zooarchaeological data are compared to these utility curves to better understand hunting behaviors ofhominins through time, contributing to interpretations of site use. This study is interested in understanding which specific prey transport strategies Neandertals at Arma Veirana utilized through time.
Testing the Climatic Stress Hypothesis Nutritional Stress
Nutritional stress is considered a key contributing factor for Neandertal extinction through the climatic stress hypothesis as the hypothesis is concerned with the heightened level of climatic instability during OIS-3 and considers the paleoenvironmentto better understand Neandertal subsistence strategies and adaptations to climate change (Finlayson, 2004; Hodgkins et al., 2016; Hublin, 2009; Sharon and Oron, 2013;). Stemming from Binford’s (1978) influential ethnoarchaeological work with the Nunamiut, numerous studies have supported the notion that there exists a relationship between heightened fat exploitation and intensified extraction methods (Bar-Oz and Munro, 2007; Outram, 1999; 2001), the size of bone fragmentation (Outram, 1999; Bar-Oz and Munro, 2007), and the transport of animal body parts and dietary significance (Faith and Gordon, 2007; Marean, 2005; Metcalf and Jones, 1988).
Within zooarchaeology, nutritional stress is evaluated as a heightened frequency of cut and/or percussion marks resulting from increased efforts of butchering and extracting marrow, fats, and bone grease (Bar-Oz and Munro, 2007; Outram, 1999, 2001; Hodgkins et


44
al., 2016). Binford’s (1978) work with the Nunamiut showed "that the intensity of bone marrow and grease exploitation is closely linked to levels of subsistence stress” (Outram, 2001:401). It is understood that "identifying bone marrow extraction strategies can provide valuable insight into the intensity of carcass use and the level of subsistence stress” (Bar-Oz and Munro, 2007:947). Therefore, investigating evidence for increased bone marrow extraction can provide information "about degrees of dietary stress” (Outram, 1999:104). In fact, nutritional stress has been documented throughout the ethnoarchaeological record via the extraction of bone fats such as marrow and bone grease (Binford, 1978; Levin and Potpov, 1964; Morin, 2007).
If Neandertals at Arma Veirana experienced nutritional stress, there should be evidence of intensive processing of prey by means of cut and percussion marks on low-utility bones with relatively little nutrients available for consumption such as phalanges (Bar-Oz and Munro, 2007), epiphyseal ends (Outram, 1999, 2001; Marean, 2005, 2007), and removing all nutrients from bones (Binford, 1978,1981; Nilssen, 2000). This study included low-utility skeletal elements including carpals, tarsals, and phalanges in its evaluations of butchering intensity to obtain a better understanding of the nutritional status of Neandertals atAV (Bar-Oz and Munro, 2007). Calculations of the frequency and distribution of cut marks (Binford, 1978; Nilssen, 2000) and percussion marks (Pickering and Egeland, 2006) were used to determine variations in butchering intensification by StratAgg.
Statistical Analysis
To test the hypothesis that cut mark and percussion mark frequencies remain relatively constant across time as a result of the reliable carrying capacity of Liguria


45
(Valensi and Psathi, 2004), a generalized linear model was applied in analyses of covariance (ANCOVA) (Miller and Chapman, 2001; Hodgkins etal., 2016). As this research is concerned with Neandertal behavioral ecology, the ANCOVAs allowed for testing butchering intensity with the frequencies of cut marks or percussion marks serving as proxies for nutritional stress (Bar-Oz and Munro, 2007; Outram, 1999, 2001; Hodgkins et al„ 2016).
The ANCOVA was designed following Hodgkins and colleagues’ analysis (2016:6) with individual StratAggs serving as independent variables, the number of cut marks or percussion marks as the dependent variables, and the maximum length and width of specimens was used to calculate the geometric mean of fragments (the covariates) (Jungers et al., 1995) "to control for the effects of fragments size on cut and percussion mark count across” StratAggs. The geometric mean can be understood as a "size proxy for each bone fragment” (Hodgkins et al., 2016:6). As measuring butchering intensity in this way results in count data (# of marks), and it is possible to have a count of zero, the ANCOVA "assumed a Poisson distribution (Zar, 2010), which is beneficially applied to ranges of values that include small numbers (Whitaker, 1914)” (Hodgkins etal., 2016:7). If a statistically meaningful relationship is exposed in analysis, a Tukey pairwise comparison is employed to establish where in the sequence there is a significant change in butchering frequencies. Statistical analyses were performed in SPSS software with a significance level of p<0.05.


46
CHAPTER IV RESULTS
Taphonomic Analysis and Taxonomic Representation
To interpret taphonomic post-depositional destruction of Arma Veirana (AV) through time, the extent of bone breakage was investigated. The average faunal fragment size, in terms of maximum length and width (mm), is overall small (<3 cm in length) and remains relatively consistent across the four StratAggs analyzed: Black Mousterian (BM), Granular (Gr), Compact Strong Brown (CSB), and Rocky Brown (RB). A one-way ANOVA found a significant difference in both the average length (F= 9.069, p= 0.000) and width (F= 2.767, p= 0.044) of fragments across layers. A Tukey HSD found CSB differs in average length from the other StratAggs (p= 0.000) while fragments are similar in size and differ slightly in average width in Gr (p= 0.057) and BM (p= 0.057).
Post-depositional damage
Trampling marks indicate movement of bones on or in sediment (Behrensmeyer et al., 1989; Fisher 1992; Marean etal., 2000), at AV this could have been due to changes in site use by Neandertals and/or other animals, disturbing bones within sediment during times of accumulation (Olsen and Shipman, 1988; Fiorillo, 1989; Dominguez-Rodrigo and Barba, 2006; Dominguez-Rodrigo etal., 2009). Table 1 shows all four StratAggs preserve bones with microstriations indicative of trampling marks.
Table 1: Number of Identifiable Specimens (NISP) and frequency (%) of fragments preserving microstriations indicative of trampling marks.


47
To interpret site use, spatial analyses were performed to evaluate changes in taphonomy throughout StratAggs. All spatial relationship maps were produced in ArcMap (a GIS program) to investigate the visual representation of artifacts in situ prior to destruction of archaeological and stratigraphic contexts following excavation (Marean et al., 2004, 2010; McPherron et al., 2005). As fauna data are recorded as 3D coordinates, a 2D representation of profile and plan views help visualize spatial relationships between artifacts in situ. Profile views provide a horizontal representation of the main excavation trench facing South (Figure 2)—as if the viewer was standing within the trench observing
Trampling vs. Non-Trampling
Profile View (South Wall)
-3 -2 -2 -1 -1 -0 0 0 1

9 !
•
RB Profile View • No Trampling * Trampling CSB Profile View ♦ No Trampling ♦ Trampling Gr Profile View a No Trampling a Trampling BM Profile View ■ No Trampling ■ Trampling
* u** ♦* * ► ** t • • m CL* i < r
i *â–  m

9 '
-3 -2 -2 -1 -1 -0 0 0 1 1
Figure 9: Profile view of the South wall of the main excavation trench showing the distribution of bones with microstriations indicative of trampling vs. non-trampled bones. Map produced in ArcMap.


48
stratigraphy straight on. Plan views provide an overview representation of artifacts as if the observer was standing directly above excavation units looking straight down.
Figure 9 displays the profile view, showing an overall even distribution of bones with observed trampling marks across StratAggs. Individual maps (plan views) were produced for each StratAgg to better understand the distribution of trampled vs. non-trampled bones through the cave over time: BM (Figure 10), Gr (Figure 11), CSB (Figure 12), and RB (Figure 13).
Black Mousterian Plan View Trampling
-2-1-1-0 0 0


.1 •* V •Mtfl ♦ mfm *9N9n2+
« * < 1
w < No Trampling
▼ N • Trampling =1
-2-1-1-0 0 0
Figure 10: Plan view ofBM. Map produced in ArcMap.
Trampled bones appear to be evenly distributed through most of the BM (Figure 9),
however trampling marks are not observed in the northeastern-most excavated area. This


49
is similar to changes observed in other StratAggs—it is possible the cave was not as heavily utilized in this area as it is close to the East cave wall, perhaps it was too close to the wall and an unfavored area of the cave. Figure 10 shows an even distribution of trampled bones in the excavated areas of BM, yet with notably less representation in the southwestern area.
Granular Plan View Trampling
-2 -1 -1 -0 0 0 1 1


* ♦ ♦ t * ♦ t ♦♦ ♦♦♦ * ♦♦ ***\ ♦ * ♦ M * ♦ .♦ ♦♦ * ♦ ♦ .♦ ♦
♦ **i M ♦ 0 .♦* % ♦ ♦♦ ♦ ♦ # 4 ♦ ')»***♦ > ?*• ♦' h/ u* V' ♦s.Jv ft * ♦v *♦ V * ♦ ♦ ♦ 0 ♦ ♦ ♦ f *A< ♦ >

V N
♦ No Trampling ♦ Trampling I
â– 2 -1 -1-0 0 0 1 1
Figure 11: Plan view ofGr. Map produced in ArcMap.
As demonstrated in Figure 9, Gr has a low representation of trampled bones, with
noticeably lower frequencies of trampling marks observed on fauna recovered from the western quads. Figure 11 supports this with no observed trampling marks along the western boundary; otherwise trampled bones are evenly distributed.


50
Compact Strong Brown Plan View Trampling
-2-1-1-0 0 0

<0 ♦
(0 ♦ ♦ ♦ ♦ ♦ ♦ ♦ ♦ » *♦ ♦ ♦ ♦ ♦
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10 w
♦ No Trampling ♦ Trampling i
T N ♦
-2 -1 -1 -0 0 0
Figure 12: Plan view ofCSB. Map produced in ArcMap.
Figure 9 reveals trampling is common in CSB, though slightly less frequent in the
middle of the excavation trench. Figure 12 shows the trampled bones are evenly distributed, indicating post-depositional damage was common and fairly consistent through time during deposition of CSB which is consistent with the high frequency of trampling marks reported in Table 1.


51
-3 -2 -2 -1 -1 â– locky -0 !rown Tramp 0 Plan V ing 0 iew 1 1 2 2 3
CO 00

# ♦ A 9 •# • » • t.
co • * •* * ♦ \* • CO
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lA ♦ CA
CA — *?•
♦ . «• * * •.:

CO Y • No Trampling • Trampling
ro i i
-3-3-2-2 -1 -1-000 1 1 2 23
Figure 13: Plan view of bones with microstriations indicative of trampling in RB. Map produced in ArcMap.
Figure 9 shows trampling marks are sparsely represented in RB with concentrations of trampled bones recovered from the quads located at the top of the sequence and becoming less concentrated lower in the trench. A higher frequency of trampled bones closer to the surface of RB is consistent with it be the youngest and therefore top-most layer, which was still susceptible to sediment compaction at the surface prior to excavation. Figure 13 shows that while sparse, trampled bones are fairly evenly distributed in the north and concentrated along the perimeter of the excavated area in the southeast.


52
Evidence of cooking by Neandertals
Coloration of fragments indicates carcass processing across levels included cooking by Neandertals, which appears to have remained relatively consistent at Arma Veirana through time (Figure 14). Analysis was performed on all specimens regardless of overall coloration with at least 20% visibility and evidence of burning (< 50% carbonized). The colors in Figure 14 relate to their level of burning, for reference: white represents fully calcined bones, or those having undergone the highest amount of burn time (delicate, dissolving bone).
Fully Calcined >50% Calcined
â–  <50% Calcined
â–  Fully Carbonized
â–  >50% Carbonized
â–  <50% Carbonized
â–  Total Burned
100%
Figure 14: The frequency of burned bones by maximum burning stages per StratAgg.
Figure 15 shows the profile view spatial relationship between burned and unburned
bones recovered from all StratAggs. Overall, burning appears sparse across StratAggs and there does not appear to be a pattern of a preferential burning area within the cave.
BURNING STAGES
0% 20% 40% 60% 80%


53
-2 -2 -1 Burned Profile Vie -1 vs. Unburn w (South Wall) -0 ed 0 0 1

9 * 9
< v- i- i- z- e- i i i i i
RB Profile View Unbumed ♦ Burned CSB Profile View Unbumed ♦ Burned Gr Profile View Unbumed a Burned BM Profile View Unbumed ■ Burned
1 ***** ♦ * ♦* **** a * * * *»■*** A A ♦ i .JF.f
â– 
Y
9
-2 -2 -1 -1-0 0 0 1
Figure 15: Spatial relationship between burned and unburned bones across StratAggs. Map produced in ArcMap.
As BM got its name from the burned nature of artifacts and sediment, it is unsurprising that it contains the densest concentration and most even distribution of burned bones of the StratAggs. CSB has more evidence of burning than RB, with a concentration of burned bones in eastern quads. Burned bones in Gr are infrequent but appear to be evenly distributed through time in the trench. RB preserves the lowest density of burned remains, with a small concentration at the top northwestern portion of the sequence and lacking any burned bones to the east.


54
Taxonomic Analysis
Table 2 displays the taxonomic representation at AV by the number of identifiable specimens (NISP) per StratAgg. The most representative taxon across StratAggs by far is non-ID mammal—most likely attributed to the small fragmentary nature of most specimens and the lack of a comparative collection in the field which made taxonomic classifications difficult. However, it is important to note that specimens were distinguished between mammal and carnivore whenever possible to better understand site use by Neandertals versus carnivores as they are well represented in caves throughout Liguria (Valensi and Psathi, 2004).
FAUNA ANALYZED
Taxon RB CSB Gr BM
Aves 2 0 0 1
Bovidae 0 2 0 10
Capra ibex 0 0 3 0
Bovid/Cervid 2 3 14 70
Canidae 1 0 0 0
Canis lupus 1 0 0 0
Non-ID Carnivore 4 1 6 6
Cervidae 1 1 0 7
Cervus 0 0 0 2
Cervus elaphus 0 0 0 3
Non-ID Mammal 97 41 249 555
Capreolous capreolous 2 0 0 1
Suidae 0 0 0 2
Ursus 1 4 7 3
Non-ID Other taxa 1 2 16 53
Total bones analyzed 112 54 295 713
Table 2: Number of identifiable specimens (NISP) and total number of specimens analyzed per StratAgg.


55
Prey species represented at AV
During data collection, faunal remains were assigned to specific taxon (e.g. Capreolous) whenever possible and when unidentifiable, they were assigned a classification level of bovid, cervid, or bovid/cervid depending on how distinguishable fragments were, shows the number of identifiable specimens (NISP) of artiodactyl taxa (bovidae and/or cervidae) are common in each layer with identifiable species including Capra (goat), Capreolous capreolous (Roe deer), Cervus (deer), and Cervus elaphus (Red deer). The specimens reported in also represent the bones used to perform zooarchaeological tests to determine agents of accumulation and bone modification.
BONES USED FOR TESTS
Taxon RB CSB Gr BM
Bovidae 0 2 0 10
Bovid/Cervid 2 3 14 70
Capra 0 0 3 0
Cervidae 1 1 0 7
Cervus 0 0 0 2
Cervus elaphus 0 0 0 3
Capreolous capreolous 2 0 0 1
Total analyzed 5 6 17 93
Table 3: NISP of identifiable bovid, cervid, or bovid/cervid long bones used for zooarchaeological tests.
Spatial representation of fauna at AV
Black Mousterian is by far the densest StratAgg in terms of faunal remains, Figure 16 shows the spatial relationship of fauna in this StratAgg. As previously discussed, non-identifiable mammal was the most commonly recorded fauna across StratAggs. It appears faunal remains are denser in the western excavated area than the eastern, this could be attributed to the East cave wall not serving as a preferred area to congregate within the


56
cave for Neandertals and/or other animals. Evenly distributed are copious bovid/cervid prey specimens including identifiable taxa like Cervus elaphus and Caprelous capreolous. Carnivores including non-identifiable taxa and Ursus are well represented and appear evenly distributed through BM, in both excavated areas. BM preserves interesting taxa including Suidae (pigs, n=2) andi4ves (bird, n=l).
-2 Black Moust -1“ erian Plan Vie 4)~ w gjn Legend Taxon Aves • Bovid • Bovid/Cervid ▲ Carnivore Cervid O Cervus elaphus • Mammal • Capreolous capreolous • Suid A Ursus

tO
to • • • * • * • * . •* •• •• * • V • ' •v •

1> • * • j . • • % » TT k A a 2

|
1> § w ■A §
r K
-2 ■2“ ■1 ■I® ■0 0“ 0a
Figure 16: Plan view spatial visualization of fauna from BM. Map produced in ArcMap.


57
Figure 17 shows the spatial relationship of fauna analyzed from Granular. Bovid/cervid prey specimens are sparsely distributed across this StratAgg in two loose clusters in the southeastern and western excavated portions; one Capra specimen is nestled within the western cluster. Identifiable Ursus remains are evenly distributed while non-identifiable carnivore remains appear in a loose cluster in the southeastern excavated region. It is worth noting there are no bovid/cervid yet four Ursus remains represented in the northeastern section of Gr, this might suggest a differential use of the cave between Ursus and Neandertals during deposition of this StratAgg.
Granular Plan View
.2 000000 .500000 _.| .000000 _0 500000 0003000 0500000 ^ OOOOOC


• . • 1 •• c *• V • • • •. \ . • • • . * • • • • • • ^
• •*: - ■yY: •% H * • «• • • u v*: %.y • • • " it ‘ • • • • • • • • • • :
Legend Taxon Bovid/Cervic O Capra ▲ Carnivore • Mammal ▲ Ursus
Y N
-2 .i« -oa hood 0oo 1 -ooooot
Figure 17: Plan view spatial visualization of fauna from Gr. Map produced in ArcMap.


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CSB preserves the lowest density of faunal remains across StratAggs yet Figure 18 shows mammals and bovid/cervid taxa are evenly distributed. The bovid and cervid specimens were not identifiable to species yet were distinct enough to distinguish between bovidae and cervidae families. Carnivore taxa including Ursus are solely represented in the southern region of CSB.
Compact Strong Brown Plan View
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Figure 18: Plan view spatial visualization of fauna from CSB. Map produced in ArcMap.
Figure 19 shows an even distribution of bovid/cervid and carnivore remains in Rocky Brown, including identifiable taxa like Aves, Canis lupis (wolf), Capreolous capreolous, and Ursus. In addition to the wolf remains there are non-identifiable Canidae remains.
Interestingly, the Capreolous specimens [n=2] are teeth identified as an incisor and third


59
pre-molar and are situated close to each other in the southeastern portion, likely representing one animal.
Rocky Brown Plan View
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Figure 19: Plan view spatial visualization of fauna in RB. Map produced in ArcMap.
Skeletal element representation
Table 4 lists the number of identifiable long bone skeletal elements assigned to bovid/cervids, either specific skeletal elements (e.g., femora), or simply long bone fragments when identification of specific skeletal elements was not possible. These bovid/cervid long bones were used in surface modification analyses to investigate evidence
of butchering, cooking, and transport strategies of Neandertals at AV through time.


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NISP SKELETAL ELMENTS
Skeletal Element RB CSB Gr BM
Femora 1 0 1 8
Fibulae 0 0 0 1
Humeri 1 0 0 8
Long bone fragments 0 1 2 8
Metacarpals 0 0 1 3
Metatarsals 0 0 0 5
Metapodials 0 2 1 6
Radii 0 0 0 4
Radioulnae 0 0 1 4
Tibiae 1 1 0 6
Ulnae 0 0 0 0
Total 3 4 6 53
Table 4: NISP skeletal elements (e.g. femora) or long bone fragments for all bovid/cervid remains.
Surface Modification Analysis
Before in-depth zooarchaeological tests are performed, it is paramount to ensure a lack of excavation and curation bias within faunal assemblages as non-identifiable fragments were regularly not analyzed in the past (Marean et al., 2000; Pickering et al., 2003). As AV began excavations in 2015, it was important to ensure a lack of biases prior to zooarchaeological test were performed; as such, all bone fragments were analyzed in this study. Figure 20 shows the relative abundance of shafts to epiphyses across StratAggs closely match expected frequencies taken from experimental data for sites where hominids and/or carnivores accumulated and modified faunal assemblages (Marean etal., 2000).
As all StratAggs closely match expected relative frequencies of shafts to epiphyses (Marean et al., 2000), it can be assumed they "retain their original frequencies of ends versus shafts at the time of breakage, are not biased through selective retention, and are consistent with either hominid and/or carnivore accumulation” (Hodgkins and Marean, 2017:10). Further taphonomic analysis was considered before several zooarchaeological


61
tests were employed to determine agents of bone accumulation and modification for each layer.
Rocky Brown Compact Strong Brown Granular Black Mousterian Carnivore Only Hominid to Carnivore Hominid Only
0% 20% 40% 60% 80% 100%
â–  Epiphyses â–  Shafts
Figure 20: The relative abundance of long bone shafts to epiphyses per layer as compared to experimental data compiled by Marean et al. (2000), experimental data are distinguished with dots.
Breakage patterns
Faunal assemblages per StratAgg were compared to expected frequencies (Marean et al., 2000) of nutritive (Figure 21) and non-nutritive breakage (Figure 22). Table 5 reports the raw zooarchaeological data used in this analysis of breakage patterns. Figure 21 shows all StratAggs have lower frequencies of nutritive bone breakage when compared to experimental data (Marean et al. 2000).
RB has the second highest frequency of nutritive breakage, though lower than expected in the literature. Likewise, BM and CSB preserve similar though low frequencies of nutritive bone breakage. This is interesting as BM and CSB have the densest (n= 713) and least dense (n= 54) faunal remains respectively yet so closely resemble one another in breakage analysis, possibly representing similar subsistence behaviors during deposition
Shafts vs. Epiphyses


62
of these StratAggs. While there appears to be drastic changes in bone accumulation across StratAggs, there is evidence for intentional extraction of nutrients from bones by either Neandertals and/or carnivores.
BONE BREAKAGE DATA
Nutritive Breakage
Layer Oblique % Curved % Total long bone ends
RB 112 73.68 105 69.08 152
CSB 48 55.81 48 55.81 86
GR 243 82.09 233 78.72 296
BM 265 57.21 468 57.21 818
Non-nutritive Breakage
Layer Right % Transverse % Total long bone ends
RB 36 23.68 33 21.71 152
CSB 50 58.14 52 60.47 86
GR 51 17.23 59 19.93 296
BM 279 34.11 265 32.40 818
Table 5: Long bones in each layer with nutritive breakage: oblique angles and curved outlines (top) and non-nutritive breakage: right angles and transverse outlines (bottom)—following methods outlined by Villa and Mahieu, 1991.
Figure 22 shows higher than expected non-nutritive breakage across StratAggs,
which suggests post-depositional forces strongly impacted the level of preservation of faunal remains, possibly creating a discrepancy in bone preservation between layers at Arma Veirana. If StratAggs preserve noticeably different breakage patterns, there must be reasons for these differences; changes in sediment composition and site use serve as possible contributions to changes in non-nutritive bone breakage.


63
Figure 161: The frequency ofbovid/cervid long bones broken with curved outlines and oblique angles following criteria outlined by Villa and Mahieu (1991). Data are plotted against experimental assemblages where nutrients were extracted by hominids only, hominids and carnivores, and carnivores only (Marean etal., 2000).


64
% TRANSVERSE BREAKS
Figure 22: The frequency ofbovid/cervid long bones broken with transverse outlines and right angles following criteria outlined by Villa and Mahieu (1991). Data are plotted against experimental assemblages where nutrients were extracted by hominids only, hominids and carnivores, and carnivores only (Marean et al., 2000).
Agents of accumulation
Bone surface modification data was utilized to better interpret bone accumulators and modifiers per StratAgg with the frequency of percussion marks and tooth marks serving as proxies for hominin and carnivore activity respectively and compared to expected frequency data outlined in various actualistic studies where humans and/or carnivores had access to the bones for nutritional extraction (Blumenschine, 1988,1995; Marean and Bertino, 1994; Capaldo, 1995,1997; Marean etal., 2000, 2004). While all four StratAggs preserve at least one percussion mark (PM) on bovid/cervid long bone fragments, CSB is the only layer that preserves a carnivore tooth mark (TM, Table 6).


65
Table 6: Raw data for the number ofbovid/cervid long bone fragments with at least one percussion mark (PM) or one tooth mark (TM). Note no specimens displayed both PM and TM.
Carnivores are known to favor the greasy epiphyseal ends of long bones (Marean and Spencer, 1991), thus preserving higher frequencies of tooth marks on epiphyses over midshafts (Faith, 2007). However, as epiphyses are spongy bone, they are less dense and more vulnerable to post-depositional damage than shaft fragments (Marean and Kim, 1998). Results from the breakage analysis (Figure 21 and Figure 22) and the frequency of shafts to epiphyses analysis (Figure 20) demonstrate epiphyseal ends were infrequently recorded at AV, suggesting they degraded over time through taphonomic forces and/or carnivore ravaging activity. However, it appears there was not a substantial amount of carnivore activity in any StratAgg (Figure 23), indicating substantial taphonomic destruction of bones overtime at AV.
Figure 23 shows a comparison of the percentage of PMs to TMs present on bovid/cervid long bone fragments per StratAgg. RB, Gr, and BM closely match expected frequencies of hominid only accumulation while CSB matches assemblages where humans first had access to bones before carnivores, further supporting the inference that


66
Neandertals were the main bone accumulators through time at AV while carnivores had intermittent access to the cave and bones within.
PERCUSSION VS. TOOTH MARKS
PC
<
s
0% 20% 40% 60% 80% 100%
% TOOTH MARKS
Figure 23: The percentage ofbovid/cervid/ long bone fragments with at least one tooth mark versus the percentage with at least one percussion mark. Data are plotted against experimental assemblages where nutrients were extracted by hominids only, hominids had first access before carnivores, and carnivores only (Blumenschine, 1988,1995; Marean and Spencer, 1991; Marean etal., 1992, 2000; Marean and Bertino, 1994; Capaldo, 1995,1997). Ellipses hold no statistical meaning but are used for visual aide.
Transport Behaviors
Figure 24 shows medium-sized (size 2-3) bovid/cervid fragments dominate through time. Ethnographic records show a high probability for hunters to exploit smaller game for easier transport from a kill site back to a butchering and/or consumption site (Binford, 1978; Schoville and Otarola-Castillo, 2014). As time went on at AV, it appears small to


67
medium prey were regularly transported to the cave for consumption. BM has the highest concentration of faunal remains of any layer it is the most diverse in prey size (sizes 2-5). Interestingly, as time goes on, bovid/cervid body sizes get relatively smaller: BM preserves sizes 2-5, Gr sizes 2-4, CSB only sizes 3 and 4, and RB only sizes 2 and 3— this is not a clear picture as well-represented prey size fluctuates through time. While more evidence is needed, these repetitive reductions in prey body size may suggest times of climate change and shifts in the environment’s carrying capacity as it is reported there were major declines in large mammalian populations across Europe during oscillating OIS-3 (Stewart et al., 2003b). If the flora available for fauna consumption became sparse and less reliable during climatic shifts, larger animals would be stressed as a result of limited food resources.
w
H 3
;x
OQ
0
BOVID/CERVID PREY SIZE
10
20 30
NISP
40
Size Distinction
2 (20-113 kg)
3 (113-340 kg)
4 (340-900 kg)
5 (900-2000 kg)
RB
CSB
Gr
BM
50
Figure 24: Number ofiden tifiable bovid/cervid by animal body size as detailed by Brain (1981). To investigate changes in food transport behavior at AV through time, only high
survival skeletal elements were used to calculate the minimum number of skeletal
elements (MNE) and the normed minimum animal units (%MAU) per StratAgg (Table 7).


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Unfortunately, the top three StratAggs (RB, CSB, and Gr) do not have sufficient data to compare against Metcalfe and Jones’ (1988) Standardized Food Utility Index (SFUI) and Binford’s (1978) and Faith and Gordon’s (2007) theoretical food utility curves; however, these data are reported for reference within the Appendix.
Table 7: MNE and MAU of all bovid/cervid fragments from Black Mousterian. Raw data are compared to the SFUI as published by Metcalfe and Jones (1988).
Figure 25 shows how the %MAU of each skeletal element within BM compared to the SFUI (Metcalf and Jones, 1988) of each element with its associated caloric value (Binford, 1978; Faith and Gordon, 2007; Metcalf and Jones, 1988) to interpret food transport strategies: bulk, gourmet, unbiased, or unconstrained. Figure 25 shows BM most closely falls between Binford’s (1978) bulk and unbiased transport strategies—suggesting a mixed strategy that fluctuated between transporting only high utility elements and transporting elements in direct correlation to their food utility respectively.


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Figure 25: The relative abundance of high survival skeletal elements (% minimum animal units) and their associated Standardized Food Utility Index (SFUl) [following Binford, 1978; Faith and Gordon, 2007; Metcalf and Jones, 1988). Data are plotted against utility curves (Binford, 1978; Faith and Gordon, 2007), curves hold no statistical meaning but are used for comparison. The Poly. Line holds no statistical meaning but shows the line of best fit for transport strategies in Black Mousterian.
Analysis of Nutrient Extraction Behaviors
Table 8 shows the low utility skeletal elements included in the analysis of butchering intensity at AV. These bones were analyzed regardless of taxon to obtain an understanding of the nutritional status of Neandertals over time. If Neandertals were indeed nutritionally stressed it should be assumed they would not leave any bones unprocessed—therefore intensifying butchering behaviors that manifest in the zooarchaeological record as an increased frequency of cut and/ or percussion marks. However, the environment of Liguria ~45kya is characterized as relatively stable with


70
sparse forests in steppe-like terrain (Holt et al., 2018) and an abundance of apline taxa like Capra ibex and Capreolous capreolous (Valensi and Psathi, 2004), both of which are frequently recorded at AV.
LOW UTILITY ELEMENTS
Skeletal Element RB CSB Gr BM
Astragalus 0 0 0 1
Calcaneus 1 0 1 1
Cranial 4 2 6 25
Cranial Fragment 0 2 0 8
Distal Phalanx 1 0 1 3
Intermediate Phalanx 0 0 0 5
Lunate 0 0 0 1
Magnum 0 0 1 0
Mandible 1 0 1 6
Maxilla 1 0 0 3
Metacarpal 2 0 2 4
Metacar pal III 1 0 0 0
Metapodial 1 4 3 17
Metatarsal 0 0 0 9
Non-ID Carpal 0 0 0 2
Phalanx 1 0 1 2
Pisiform 0 0 0 1
Proximal Phalanx 3 1 5 6
TOTALS 16 9 21 94
Table 8: NISP of all low utility skeletal elements (regardless of taxon) used to test butchering intensity and evaluate nutritional stress.
Evaluating Nutritional Stress
Two analyses of covariance (ANCOVA) were applied to test the hypothesis that cut mark (CM) and percussion mark (PM) frequencies remain relatively constant through time at AV as an effect of the reported reliable carrying capacity of Liguria (Valensi and Psathi, 2004). The ANCOVA testing for a difference in the frequency of CMs and controlling for fragments size (geometric mean of fragments) across StratAggs did not find a significant


71
difference in the frequency (y2= 0.709, df= 1, p=0.400): accepting the null hypothesis that there is not a significant change in the frequency of CMs in the different layers. However, Figure 26 shows a significant change in the frequency of PMs (y2= 5.846, df= 1, p= 0.016) across the layers (see Table 13,Table 14,Table 15Table 16 for raw data), namely in CSB.
The frequency of PMs per fragment increased in Gr, producing the biggest difference across StratAggs—suggesting an intensity in processing carcasses to more thoroughly extract marrow from within bones: signifying a time of nutritional stress at AV (Bar-Oz and Munro, 2007; Outram, 1999, 2001; Hodgkins et al., 2016). A Tukey pairwise comparison found a significant difference (p= 0.027, df= 3) in the frequency of percussion marks between BM and Gr. There is an increase of 0.31 percussion marks per fragment from BM to Gr.


72
Percussion Marks
(Wald Chi-Squarc= 5.846, df= 1, p=0.016)
3
2
S
e.
*
o 1
S?
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Os
0
-1
4 3 2 1
Strat (RB=4, CSB=3, Gi=2, BM=1)
Figure 26: ANCOVA results showing statistically significan t changes in the frequency of percussion marks (PM) on high and low utility skeletal elements through the layers. The central points are the mean number ofPMs preserved on skeletal elements. The error bars are the 95% confidence intervals for those means. The lines connecting the layers hold no statistical power but show the interaction effect between layers.
Spatial analysis of percussion marks
Utilization of plan and profile view GIS maps help illustrate the spatial and temporal
distribution of fauna analyzed at AV. Figure 27 and Figure 28 show a low density of fauna that preserve at least one percussion mark (PM) in BM. Percussed bones are evenly distributed through the StratAgg, yet with a lower concentration in the eastern excavated area in both plan and profile views.


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Black Mousterian Plan View
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Figure 27: Plan view spatial visualization of fauna with at least one percussion
mark (PM) in BM. Map created in ArcMap.


74
Black Mousterian Profile View
jSXMl J.OCOOOD _^J00000 ^ C Figure 28: Profile view (of the South wall) spatial visualization of fauna with at
least one percussion mark (PM) in BM. Map created in ArcMap.


75
Granular Plan View
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Figure 29: Plan view spatial visualization of fauna with at least one percussion mark (PM) in Gr. Map created in ArcMap.
Figure 29 shows a low density and fairly even distribution of fauna with PMs in Gr (plan view), with a lack of specimens with PMs in the eastern-most excavated area. Interestingly, Figure 30 shows an underrepresentation of PMs in the eastern section of the excavated section along the South wall (profile view). The absence of percussed bones in this area through time might suggest a differential use of space within the cave for percussing and processing carcasses. The profile view map also shows percussed bones roughly along the same horizon, possibly indicative of quick occupation events.


76
Granular Profile View
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Figure 30: Profile view (of the South wall) spatial visualization of fauna with at
least one percussion mark (PM) in Gr. Map created in ArcMap.


77
Compact Strong Brown Plan View
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Figure 31: Plan view spatial visualization of fauna with at least one percussion mark (PM) in CSB. Map created in ArcMap.
While there are few bones that have PMs in Compact Strong Brown (CSB), Figure 31 shows those that do are evenly distributed across the StratAgg. The profile view (Figure 32) shows an even distribution of percussed bones along two horizons in both the upper and lower excavated areas of CSB, this could also support the idea of quick occupation
events in Granular, possibly use of the cave for overnight camps.


78
Compact Strong Brown Profile View
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Figure 32: Profile view (of the South wall) spatial visualization of fauna with at
least one percussion mark (PM) in CSB. Map created in ArcMap.


79
s moo _£S ™ 00 _5»: 0000 4* moo moo .j.at F 0000 4.5C took y Br< 0000 Dwn 0000 j|.o Plan XXJOO .Q.S Vie nooo q W 0000 050 0000 15o 000 2 s0 0000 j.OO 0000 J-50 0000 40c
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Figure 33: Plan view spatial visualization of fauna with at least one percussion mark (PM) in RB. Map created in ArcMap.
Figure 33 shows few bones with PMs and only at the top of the southern portion of the excavated area of Rocky Brown (RB). The profile view (Figure 34) shows a very low frequency of percussed bones and only present in the lower excavated portion. It is interesting percussed bones are only found in selective areas within RB, this further supports quick use of the cave over time.


80
Rocky Brown Profile View
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Figure 34: Profile view (of the South wall) spatial visualization of fauna with at least one percussion mark (PM) in RB. Map created in ArcMap.


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CHAPTER V
DISCUSSION AND CONCLUSION
Discussion
Zooarchaeological analysis results suggest climate change influenced fluctuations in species abundance in Liguria through time as represented bovid/cervid prey body sizes oscillated at Arma Veirana. Interestingly, the most drastic shift in body size is observed between BM and the rest of the StratAggs when size 5 prey is not represented again. It is noteworthy that size 5 prey species occur only in this StratAgg as it could signify a time of dense forest growth and therefore not likely a time of nutritional stress amongst Neandertals if they had reliable food resources. Results from taphonomic and surface modification analyses support the inference that Neandertals used the cave for butchering and cooking prey while carnivores took advantage of the cave for scavenging or possibly even hibernating. There exists direct evidence for hominid to carnivore accumulation (Marean et al., 2000) in CSB. Carnivore remains represent a very small proportion of the assemblages from AV, namely Ursus (n= 15 total across StratAggs) and non-identiflable carnivores (n= 17 total across StratAggs) are found in every layer. Understanding the taphonomic setting of each StratAgg aids in the interpretations of site use over time.
Black Mousterian
It appears post-depositional damage was not substantial in BM: taphonomic analysis shows that BM preserves the highest number of individual trampled bones (n=85) yet the overall proportion of bones with trampling marks is lowest (11.9%). This is consistent with results from the breakage pattern analysis of non-nutritive bone breaks, with the overall second-lowest frequencies observed. BM also preserves the second-lowest


82
frequencies of nutritive bone breaks. The analysis of longbones of bovid/cervid prey species results show BM matches expected frequencies of hominid only accumulation with plenty percussion marks and no carnivore tooth marks. Additionally, BM preserves the overall highest proportion of burned bone (76.6%) and it is worth noting it is the only StratAgg that preserves fully calcined fragments [n=2), indicating Neandertals were burning to a further extent in BM. This is interesting as fully calcined bones are rarely recorded among zooarchaeological assemblages from Paleolithic sites across Italy (Stiner etal., 1995).
Results from the transport pattern analysis show BM falls between bulk and unbiased strategies (Binford, 1978). The bulk strategy "reflects the maximization of nutrient quantity returned to camp” and "should be practiced when demand for meat is high and hunters attempt to return as much mass as possible to camp” (Schoville and Otarola-Castillo, 2014:2). Neandertals transporting elements in direct relation to their utility follows the unbiased strategy, which Binford stipulates "should be the normative hunter-gatherer pattern of element transport” (Schoville and Otarola-Castillo, 2014:2). Together, these results do not suggest Neandertals were nutritionally stressed in the BM as they were clearly making decisions at kill sites on what body parts to transport back to the cave—not transporting entire carcasses for optimal caloric extraction.
Granular
Results suggest strong taphonomic forces were at play during deposition of Gr, such as sediment compaction by trampling, which contributed to the smallest observed fragmented bones of the StratAggs. Neandertals proved to be the main agents of accumulation in Gr as results closely match expected frequencies of hominid only


83
accumulation via analysis of percussion marks versus carnivore tooth marks on bovid/cervid long bones (Blumenschine, 1988,1995; Marean and Spencer, 1991; Marean etal., 1992, 2000; Marean and Bertino, 1994; Capaldo, 1995,1997). Further evidence supporting Neandertals were the main accumulators at AV during deposition of Gr comes from a high frequency of burned bones. In fact, 30% of the assemblage shows evidence for burning, indicating direct contact with fire either for cooking or warmth and may even signify a time of a warmer climate during Gr as fire use in by Neandertals in Europe is known to increase in temperate and decrease in colder climates yet this is proposed to demonstrate an opportunistic approach to fire use (Sandgate et al., 2011). This could be due to more trees growing in warmer environments and an overall easier acquisition of firewood to haul up to the cave.
As the carrying capacity of Liguria during OIS-3 is reported as reliable in terms of prey species (Valensi and Psathi, 2004) it is assumed butchering intensity was relatively constant through time at AV. Butchering behaviors should not fluctuate through StratAggs and there should not be a dramatic increase or decrease in the frequencies of cut marks and/or percussion marks through time. However, results from the ANCOVA show a statistically significant shift in the frequency of percussion marks preserved on both high and low utility skeletal elements (y2= 5.846, df= 1, p= 0.016), with an increase of 0.31 percussion marks per fragment (p= 0.027, df= 3) between BM and Gr which suggests greater efforts were put into extracting bone grease and marrow from within bones during this time. These results reject the null hypothesis that there is no change in nutrient extraction intensity. If Neandertals were adapting their subsistence behaviors to adapt to changes in their environment at AV, they were likely just as adaptive elsewhere in Europe


84
as well. This finding calls into question the rigidity of the climatic stress hypothesis, it is possible climate and environmental change were not as stressful for Neandertals as currently believed.
Compact Strong Brown
The sediment of CSB is defined as fine-grained sand and clay with small rocks (up to 5 cm) with micromorphological thin sections suggesting there was gentle downslope movement of sediment as well as a shift in geologic composition (Miller and Czechowski, 2016). These factors could signal a substantial shift in sediment composition modifying bones to a further extent in CSB than other StratAggs. As an effect of the sample size of CSB (as compared to other StratAggs) it preserves the highest proportion of trampled bones (31.5%) and the lowest number of bones with trampling marks (n= 17), suggesting infrequent but strong forces of site destruction.
Evidence for anthropogenic activity is sparse in CSB [n=54) and while every StratAgg analyzed contains bovid/cervid long bone fragments with at least one percussion mark, solely CSB preserves a carnivore tooth mark on these same bones, indicating Neandertals had initial access and carnivores scavenged the bones afterwards (Blumenschine, 1988,1995; Marean and Spencer, 1991; Marean etal., 1992, 2000; Marean and Bertino, 1994; Capaldo, 1995,1997). Interestingly, CSB is the only StratAgg with tooth marks on bones (n=5), both long bone and ribs. CSB likely represents dual occupation by Neandertals and carnivores of the cave over time as it preserves the highest proportion of carnivore remains (9.26%) including the second highest number of Ursus remains (n= 4). It is possible bears were utilizing the cave for hibernation through time, which could contribute to the elevated levels of post-depositional destruction in CSB. When Neandertals


85
were utilizing the cave, they were rarely using fire, or the cave in general as there is low evidence of burned bone (10%). These results together infer carnivores had increased access to the cave during deposition of CSB than in any other StratAgg while Neandertals maintained primary agents of accumulation over time.
Rocky Brown
RB preserves the second highest number (n= 25) and proportion of trampled bones (22.3%) likely suggesting strong post-depositional damage. This is consistent with every layer preserving higher than expected levels of non-nutritive bone breakage and lower than expected nutritive breaks as compared to experiments where bones were broken by humans and carnivores (Marean et al., 2000). The lower than expected nutritive bone breaks indicates post-depositional damage altered bones to remove nutritive breaks and replace them with non-nutritive breaks. RB preserves percussion marks on bovid/cervid long bones and there is evidence for cooking; although it is the lowest frequency (8%) of burned bones across StratAggs.
Bovid/cervid prey body sizes are smallest in RB (only 2 and 3) and though more data are needed this may suggest a time of decline in the carrying capacity or possibly a time of stress for Neandertals. It would be interesting to see results from transport pattern analysis of this layer to see if Neandertals were mostly hauling high yield bones back to the cave which could support a change in the environment. If they were mainly hunting smaller game, it would be expected for Neandertals to be more selective in what body parts to bring back to the cave for the highest caloric return. Unfortunately, RB lacked sufficient data to compare to the SFUI in this study. Results indicate Neandertals were not using the cave as a butchering site or camp site very often during deposition of RB.


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TEMPORAL TRENDS
Inference category RB CSB GR BM
Total bones analyzed n=112 n=54 n=295 n=713
Prey body sizes represented Sizes 2 and 3 Sizes 3 and 4 Sizes 2-4 Sizes 2-5: most diverse
Burned bones 8% 10% 30% 76.6% Only calcined fragments
Frequency of carnivore remains 6.25% Highest frequency: 9.26%. Only layer with tooth marks 4.41% Lowest frequency: 1.26%
Trampled bones 23.21% Least specimens yet highest frequency: 31.48% 17.63% Lowest frequency: 12.06%
Nutritive breakage 73.68% 55.81% 82.09% Most closely resembles hominid to carnivore accumulation. 57.21%
Non-nutritive breakage 23.68% 58.14% 17.23% 23.68%
Special attributes Most closely matches expected frequencies of non-nutritive breakage Carnivores had increased access to the cave. ANCOVA found a difference in the frequency of percussion marks. (PMs) (X2=5.846, df= 1, p= 0.016) and an increase of 0.31 PMs per fragment (p=0.027, df= 3). Only StratAgg with data for transport pattern analysis. Falls between bulk and unbiased strategies.
Table 9: Summary table of temporal trends observed atAV.


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Conclusion
This study used principles of Human Behavioral Ecology to evaluate changes in subsistence behaviors at Arma Veirana and to explore the climatic stress hypothesis. This taphonomic and zooarchaeological report has presented data on species representation, agents of bone accumulation, and the post-depositional destruction of the site through time. Notable differences between StratAggs are summarized in Table 9. The cave was used for processing bones and cooking prey as butchery marks and burned bones are common in all StratAggs. Bovid/cervid prey taxa are found in all layers: 12 bovid specimens and 17 cervid specimens including 3 Cervus elaphus, and 3 Capra were brought to the cave by Neandertals. Analysis of surface modifications indicates Neandertals intentionally processed bones to extract nutrients for consumption including evidence for disarticulation, defleshing, and breaking bones open for the marrow within. Cut marks and percussion marks are well represented across StratAggs, indicating Neandertals used the cave to butcher prey through time. Transport pattern analysis results show Neandertals were making decisions at kill sites on what body parts to transport back to the difficult to access cave.
Analysis of surface modification on both high and low utility bones shows an increase in the frequency of percussion marks preserved on fragments (0.31 marks per fragment, p= 0.027, df= 3) in Gr. This is direct evidence they were able to change their behaviors to adapt to change, yet what that change is remains uncertain. It is possible there was a drastic environmental change or that humans came onto the scene and caused subsistence stress for Neandertals. Whatever the reason, it is interesting Neandertals at AV


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modified their actions to increase the caloric extraction of prey during deposition of solely one layer.
AV promises to contribute to the ongoing discussion surrounding Neandertal subsistence behaviors and their ability to adapt to changing climates. The cave is of analytical value as it is unique for two reasons. First, it is difficult to access located in the mountains while most Neandertal cave sites in Italy are easily accesible along the coast. Second, occupation of the site occurred during seemingly temperate climates within a possible refugia area yet results from this study indicate a period of nutritional stress. Results indicate substantial shifts in site use over time: Neandertals used the cave less through time while carnivores took more advantage in their absence. The larger prey body sizes preserved in BM and Gr indicate a difference in the flora compared to smaller sizes in CSB and RB as larger cervids like Cervus elaphus prefer dense wooded areas (Valensi and Psathi, 2004). It is possible there was a change in the environment. It is also possible the cave was favored less through time as it was more physically demanding to access than the seemingly favored coastal caves across Italy. More secure dates from StratAggs at AV will contribute to an enhanced understanding of climate and environmental changes in Liguria.
Results from this preliminary analysis of faunal assemblages from AV reject the climatic stress hypothesis. This study shows Neandertals adapted subsistence behaviors to deal with nutritional stress. The climatic stress hypothesis fails to consider is how much behavior modification is sufficient for change to be successful. Perhaps Neandertals were more adaptive in their subsistence behaviors than they are currently given credit for. Further zooarchaeological analysis, combined with other methods briefly outlined in this report, will strengthen the understanding of Neandertal subsistence behaviors at AV


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through time. If Neandertals were indeed successful in adapting to survive environmental changes, it begs to ask how much adaptation would have saved the Neandertals? More finegrained zooarchaeological analyses of faunal assemblages across Europe will provide highly valuable and unique insight into subsistence behaviors and the role climate truly had in the extinction of Neandertals.


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A ZOOARCHAEOLOGICAL ANALYSIS OF NEANDERTAL CAVE SITE ARMA VEIRANA IN LIGURIA, ITALY by BREEANNA CHANTEL CHAROLLA B.A. Arizona State University, 2015 A thesis submitted to the Faculty of the Graduate School of the University of Colorado in partial fulfillment of the requirements for the degree of Master of Arts Anthropology Program 2019

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ii This thesis for the Master of Arts degree by Breeanna Chantel Charolla has been approved for the Anthropology Program by Jamie M. Hodgkins, Chair Christopher Beekman Tammy Stone Date: August 3, 2019

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iii Charolla, Breeanna Chantel (M.A., Anthropology Program) A Zooarchaeological Analysis of Neandertal Cave Site Arma Veirana in Liguria, Italy Thesis directed by Assistant Professor Jamie M. Hodgkins ABSTRACT There are several theories regarding the extinction of Neandertals by 40,000 years ago. The climatic stress hypothesis posits environmental instability during the Late Pleistocene caused stress among already declining populations of Neandertals and other large cold adapted mammals across Europe leading to their extinction s . If there was a negative climatic change causing stress for Neandertals, changes in butchering intensity as preserved on faunal remains should be present among assemblages . The purpose of this thesis is to provide a taphonomic and zooarchaeological analy sis of four layers at newly excavated Italian cave site Arma Veirana (AV ) located in the Liguria n Alps . Liguria may have served as a temperate area of refugia during global glacial events as fauna diversity is described as rich and continuous throughout the Late Pleistocene. This study sought to better understand how Neandertals reacted to changes in their environment and how they used the cave through time by reporting data on species representation, agents of bone accumulation, and the post d epositional destruction of the site . Tests were performed to evaluate changes in subsistence behaviors at AV and to explore the climatic stress hypothesis. This study specifically analyzed the presence of prey species ( e.g. deer ) in the cave and looked for evidence of carcass processing. Prey body size at AV fluctuates through time, with all layers preserving higher frequencies of medium sized prey. Two layer s preserv es larger sizes which may signify a period of dense forest growth and more reliable resourc es. Transport pattern analysis results show Neandertals were making decisions at

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iv kill sites on what body parts to transport back to the difficult to access cave. Analysis of surface modification on both high and low utility bones shows an increase in percu ssion marks per fragment (after controlling for fragment size) between two layers. Carnivore activity increased as Neandertals utilized the cave less over time . Carnivores modified bones only in the layer with t h e highest frequency of carnivore remains ( 9.26% ). F aunal assemblages from Arma Veirana reject the hypothesis that Neandertals were unable to adapt subsistence behaviors to deal with nutritional stress as there is evidence for a change in nutrient extraction intensity. This form and content of this abstract are approved. I recommend its publication. Approved: Jamie M. Hodgkins

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v ACKNOWLEDGEMENTS I would first like to thank committee, Jamie Hodgkins, Tammy Stone, and Chris Beekman for all their comm ents and guidance along this journey . Jamie, than k you for the support and lessons you provided about being a woman in science , I am a master because you pushed me to be my best. I would also like to thank Dr. Curtis Marean for the experience of a lifetime (several times over) which allowed me to fully explore my love of archaeo logy , i t was a pleasure and honor to learn from you. The biggest thank you goes to my wonderful parents for always supporting me and my dreams even if they happen to involve me playing around in the dirt a ll over the world. You provided me the strength to keep going by showing me what hard work and dedication truly looks like my entire life. You gave me the lessons and tools to see this degree t constantly cheering me on and too long. You kept me grounded over the last three years and I will be forever grateful a nd anxiously waiting to return the generosity you incessantly deliver. Erin Clark , I wi ll always appreciate your listening, support, and solutions to problems real or imaginary. Thank you for the laughs , tears , and trips when ever one w as needed most my favorite will always be laughing with the raccoons in Booth Bay. Saundr a Malanowicz , thank you for always being the sweetest person and a telepathic lifeline whenever the going got rough , your voice can forever sooth me. While you might not find it chapter without you by my side , EA . I t i s something that will never go forgotten or taken for granted but always cherished .

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vi Last but absolutely not least, S arah Maureen Simeonoff , words cannot express the reality of failure without you. old room. Thank you for your unrelenting love and helping me see the good in everything . biggest journeys thus far . insanely spent encouraging each other to read one more article, write one more paragraph, make one more figure this thesis could not have been complete d , and this degree would have been unattainable without you, suffering right alongside me I had the most fun with you.

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vii TABLE OF CONTENTS CHAPTER I. Oxygen Isotope Stage II. Hypotheses for E Theoretical Basis of This Excavati Rocky Bro

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viii III. Agents of Accumu IV. Post

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ix Transport V. DISCUSSION AND CON REFERENCES APPENDIX A. B.

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1 CHAPTER I INTRODUCTION Neandertals and the Late Pleistocene Homo neandertalensis was first discovered in 1856 at the cave site Feldhofer Grotte in the Neander Valley of Germany and has ever since captivated the imagination of archaeologists and the public alike (Papagianni and M orse, 2013; Wynn and Coolidge, 2012). Neandertal skeletal remains and associated material culture have been recovered from sites covering great geographical range across Eurasia (van Andel and Davies, 2003; Villa and Roebroeks, 2014; Zilhão, 2014) and are known to have had a temporal distribution from 350 39 kya ( Higham et al, 2014; Villa and Roebroeks, 2014). Understanding Neandertal extinction during the Late Pleistocene has long been a highly provocative topic within paleoanthropological research (Higham et al, 2014). It is especially interesting as modern humans ( Homo sapiens ) outlived Neandertals to ultimately inhabit the entire globe. Causes for Neandertal extinction throughout the paleoanthropological literature are frequently divided between populati on changes brought on by the emergence of modern humans in Europe and the competitive Marean, 2007; Mellars, 2004, 2007; Mellars and French, 2011) and climatic indu ced stress experienced by Neandertals due to the unstable environment during the Late Pleistocene Roebroeks, 2014). This study contributes to this discussion by examinin g faunal material recovered from cave site Arma Veirana located in Liguria, Italy to better understand how

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2 Neandertals reacted to changes in their environment and investigate how the cave was used through time. Oxygen Isotope Stage 3 The climate and enviro nment of the Late Pleistocene is frequently discussed as chaotic and unstable through the oxygen isotope record 60 20 kya, also known as Oxygen Isotope Stage Three (OIS 3). Extensive research organized by van Andel and Davies (2003) has been done on OIS 3 to better understand the role climate change played in changes in Neandertal populations. This pair organized an interdisciplinary project which incorporated biological anthropology, archaeology, zooarchaeology, geology, palynology, paleoentomology, topogr aphy, and paleoclimatological variables from GISP2 Greenland ice cores (Grootes et al., 1993; van Andel and Tzedakis, 1996; Meese et al., 1997; Stuiver and Grootes, 2000; Delmotte et al., 2001; van Andel, 2002) and three pollen cores extracted from Lago Gr ande di Monticchio in Italy (van Andel and Tzedakis, 1996; Watts et al., 1996; Allen et al., 2000; van Andel and Davies, 2003). Ice cores yield alternating warm and cold phases on 100 to 1000 year times scales, termed Dansgaard Oeschger (D O) phases (Groot es et al., 1993; van Andel, 2002; Huntley et al., 2003; van Andel and Davies, 2003). To investigate the role quick climatic changes played on Neandertal extinction van Andel and Davies (2003) focused on 3 major phases: the Warm Transitional (57 38 ka) and Early Cold (37 28 ka) to investigate Neandertal and modern human population changes across Europe and the Last Glacial Maximum (LGM, 27 18 ka) to better understand environments and demographics at the OIS 3/OIS 2 boundary. These data were used to create ri gorous and robust paleoclimatic and paleoenvironmental databases, models, and simulations of OIS 3 and its effects on Neandertal, modern human, and large mammalian population

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3 distributions across Europe (Barron and Pollard, 2002; Pollard and Barron, 2003; Alfano et al., 2003). Climatic Stress Climatic stress is recognized as a possible contributing factor for Neandertal extinction. The global climate of the OIS oscillations in temperatures, within a these oscillations have been linked to a decline in megafauna populations across Europe (Dansgaard et al., 1993; Finlayson and Carrion, 2007; Stewart, 2003b:116; Stewart et al., 2004 ). Drastic changes in the climate and environment would have negatively impacted abundance of flora and fauna. Therefore, it has been hypothesized climatic fluctuations were stressful for Neandertal populations who lived as hunter gatherers and extracted resources from their local ecosystems ( Finlayson and Giles Pac heco, 2000; Finlayson et al., 2001, 2004, 2008; Hodgkins et al., 2016; Hublin and Roebroeks, 2009 , Stewart, 2003b). climates stressed Neandertals, their subsistence behaviors may have changed, requiring in To investigate the role climate change had on Neandertal s and other large mammalian populations, the distribution of five large extant and extinct taxa ranging in spat ial and temporal distribution across western and central Europe during the Warm Transitional, Early Cold, and LGM periods were compiled in the Stage 3 Mammalian Database (Stewart et al., 2003a; Stewart et al., 2003b; Stewart, 2005). Totaling 468 dated faun al remains from 294 sites, the database includes 1912 radiometric dates, ultimately assigning dates to 119 mammalian taxa and allowing for detailed climate mammal model simulation comparisons (Stewart et al., 2003a). Results from these simulations indicate

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4 taxon, geographic range, and climatological variables such as temperature and precipitation tolerance were fundamental to mammalian distribution through OIS 3 (Stewart et al., 2003a) and that Neandertals indeed were among those fauna that saw massive decl ines in population and ultimately extinction at the end of OIS 3 (Stewart et al., 2003b). Population declines in megafauna taxa such as mammoths ( Mammuthus primigenius ) and woolly rhinos ( Coelodonta antiquitatis ) (Stewart et al., 2004; Finlayson and Carrio n, 2007) and carnivores such as cave bears ( Ursus spelaea ) (Stewart et al., 2003b) are frequently observed across Europe at the end of OIS 3. These extinctions support the climatic stress hypothesis as it appears various large cold adapted mammals includin g Neandertals were negatively impacted by the changing climate. Neandertal Morphology and Physiology In comparison to modern humans, Neandertal skeletal morphology indicates they were robust: relatively shorter, squatter with stocky limbs, barrel chests, and exhibited muscular hypertrophy (Finlayson, 2004; Holliday, 1997; Pearson et al., 2000; Weaver 2003 , 2009). Neandertal physiology implies they were a species that evolved to be optimally adapted to the glacial climates of the Pleistocene (Holliday, 1997; Pearson et al., 2000; Weaver, 2003, 2009; van Andel and Davies, 2003; Finlayson, 2004, 2009; Hublin, 2009). Ecogeographic biological rules provide reasoning for animals within the same species appearing noticeably dissimilar in direct relation to maintaining optimal core body temperatures (Bergmann, 1847; James, 2018). short, stocky limbs are better at conserving heat, therefore better adapted to life in cold b odied animal

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5 species tend to live further north than their small 1999:165). L arger overall body sizes as compared to other species within the same genus ability for optimal body thermoregulation ( Bergmann, 1847; Blackwell et al., 1999; James, 2018). Although most features of Neandertal bodies support the idea they were optimally adapted to the cold (short limbs but stocky, robust trunks), the face appear s to deviate from patterns of cold adaptation observed in modern humans (Franciscus, 2003; Holton and Franciscus, 2008; Rae et al., 2011; Weaver, 2003, 2009). For example, the depressed nasal floors common in Neandertals are also common among modern human populations in sub Saharan Africa (Franciscus, 2003) and the wide nasal aperture common of Neandertals is commonly found in modern humans in equatorial regions (Holton and he narrow superior internal nasal dimensions, tall nasal apertures, and projecting nasal bridges of Neandertals could be, because these features are typically found in high modern humans (Weaver, 2009:16031). Interestingly, certain skeletal eleme nts, for signifying Neandertals were also adapted for a highly active lifestyle (Finlayson, 2004:82). The inference Neandertals were highly active has been investigated through studies of energy expenditure in comparison to various modern human populations (Churchill, 2006; Froehle and Churchill, 2009; Mateos et al., 2014; Snodgrass and Leonard, 2009; Sore nsen and Leonard, 2001). The assumed high body mass, high activity level, and basal metabolic rate (BMR) of Neandertals have been calculated using predictive equations and

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6 adjusted for the negative impact cold weather and stress possibly had on thermoregul ation (Mateos et al., 2014). There exist several methods for calculating Neandertal Total Energy Expenditure (TEE) (Mateos et al., 2014). Sorensen and Leonard (2001) estimated specific mean Neandertal masses and World Health Organ ization (WHO) equations to predict BMRs and eventually adjusting for the effects of TEE using ethnographic data of caloric consumption of arctic hunter gatherer popula tions while correcting for the more robust Neandertal (Steegmann et al., 2002); or using BMR calculations based on assumed skin surface area (Churchill, 2006). Results from these studies stipulate Neandertal TEE values ranged between 3500 5000 calories per day (Mateos et al., 2014) a higher range than modern hunter gatherer groups requiring 3000 4000 calories per day (Hublin and Roebroeks, 2009). Perhaps Neandertals were at an advantage over modern humans in conserving body heat; however, such a muscular a nd robust hominin would have required more calories. Did a higher caloric intake cause stress for Neandertals when faced with heightened nutritional demands brought on by ecological ramifications of OIS 3? Paleoanthropological research aims to better under stand the reasons for Neandertal extinction as they appear to have been optimally adapted for life in glacial environments, thrived for more than 300 thousand years (Villa and Roebroeks, 2014), and were likely the sole hominin inhabitants of Europe for 160 thousand years (Stewart, 2003b). Thus i t is interesting these hominins did not persevere through the end of the Pleistocene like modern humans.

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7 Liguria, Italy and Arma Veirana Neandertal skeletal remains or material culture do not appear in the archa eological Mellars, 2007; Mellars and French, 2011), indicating they went extinct across Europe within the Warm Transitional phase of OIS 3 (van Andel and Davies, 2003). Interestingly, Neandertal sites during this phase proved to have had the widest dispersal and show a preference for coastal regions of France, Italy, Portugal, and Spain (van Andel and Davies, 2003); it is possible these locations had more stable and reli able environments than those further north and served as areas of refugia (Banks et al., 2008). Italy serves as a critically important region for studying Neandertal behaviors as the archaeological record from Italy includes some of the longest lasting Mou sterian lithic assemblages in Europe (Churchill and Smith, 2000; Higham et al., 2014). The Mousterian is a Middle Paleolithic tool technology exclusively associated with Neandertals across Europe and is characterized as a based technology rare occurrences of bifacial tools such as hand axes (Finlayson and Carrión, 2007:215). The purpose of this thesis is to provide a taphonomic and zooarchaeological analysis of faunal assemblages recovered from a newly excavated Middle to Upper Paleolithic alpine cave site Arma Veirana (AV) located in Liguria, Northwest Italy. AV is a promising site to better understand Neandertal subsistence behaviors as it preserves archaeological layers with numerous Mo usterian lithics and copious faunal remains. Zooarchaeology is a subfield within archaeology that utilizes a host of methods to determine how hunter gatherers were accessing food resources in the past. Taphonomy is crucial to zooarchaeological analyses as it is the study of post depositional damage of faunal remains

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8 from the time an animal dies to when its bones are analyzed (Dibble et al., 2006; Gifford Gonzalez, 1991; Fisher, 1995; Lyman, 1987). Taphonomy allows for investigation of post depositional dama ge and destruction of whole sites and of individual archaeology remains overtime. Therefore, it is critical to zooarchaeology as it allows researchers avenues to determine between a myriad of natural processes and anthropogenic modification of bones (hunti ng and butchering), either scenario providing dissimilar and unique site narratives. The mammalian fauna of the Warm Transitional phase in Liguria have been characterized as taxonomically diverse between the earlier OIS 6 and OIS 3 (Valensi and Psathi, 200 4). Specifically, the abundance of carnivores is known to have been high, that would have been beneficial in terms of prey species for Neandertals as well as carnivores ( Valensi and Psathi, 2004:258). Forest taxa such as Cervus elaphus are well represented in Liguria during OIS 3, signifying periods of dense forest growth (Valensi and Psathi, 2004). Additionally, alpine taxa like Capra ibex are well represented (Valensi an d Psathi, 2004) which indicate periods of little snow cover ( Grignolio et al., 2004). While the climate of OIS day 2009:662), Liguria appears not to have suffered as greatly as other loca tions further North oscillations registered during the period between OIS 6 to OIS 3 were not very Arma Veirana pres erves four Neandertal associated archaeological layers with abundant faunal remains, a number of which preserve evidence of butchering and cooking.

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9 Preliminary radiocarbon dates show that Neandertals were utilizing AV more than 45,000 years ago. Fauna dive continuous between OIS 6 and OIS 3, suggesting temperate climatic conditions even during subsistence behavio rs would have stayed relatively constant throughout stratigraphic layers at AV. This study specifically analyzed the presence of prey species in the cave and looked for evidence of carcass processing by Neandertals. Taphonomic patterns such as bone fragme nt size, type of bone breakage, and the frequency of epiphyses to shaft fragments were also recorded to assess the preservation of faunal material at the site to better understand site use through time. Methods were used to determine if Neandertals were th e sole bone accumulators at the site or if carnivores also had access to the bones and modified them. This is an exploratory investigation of the cave regarding the intensity by which Neandertals or other agents such as carnivores accumulated and modified faunal remains in the cave over time.

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10 CHAPTER II BACKGROUND Hypotheses for Extinction Reasons behind Neandertal extinction remain hotly debated among paleoanthropologists. Competitive replacement (Banks et al., 2008; Marean, 2007; Mellars, 2004, 200 7; Mellars and French, 2011) and climatic induced stress serve as the main and McPherron, 2013; Higham et al., 2014; Finlayson et al., 2004; Villa and Roebroeks, 2014). Limited cognition is a reoccurring factor in the argument for the competitive replacement of Neandertals by modern humans, as they entered Europe roughly the same a nd Sánchez Goñi, 2003; Marean, 2007; Mellars, 2007; Mellars and French, 2011). The climatic stress hypothesis states that the negative environmental impacts brought on by the instability of OIS 3 were stressful for dwindling Neandertal populations across E urope and ultimately contributed to their extinction (Finlayson and Carrion, 2007; Hodgkins et al., 2016; Hublin, 2009; Hublin and Roebroeks, 2009). Competitive Replacement New research proposes modern humans first evolved in Northern Africa as early as 300 kya (Callaway, 2017) before leaving Africa, entering the Near East and ultimately Europe by 45 kya (Villa and Roebroeks, 2014). Replacement of all archaic human population s (such as Neandertals) unable to adapt to climate changes through competition with cognitively superior modern humans serves as a popular avenue for understanding

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11 2003; Leroyer and Leroi Gourhan, 1983; Leroyer, 1988; Marean, 2007; Mellars and French, 2011; Shea, 2003). Research proposes Neandertals and modern humans co existed in Europe for roughly 5,000 years (Higham et al., 2011, 2014; Pinhasi et al., 2011) with eviden ce for site chronologies overlapping (Galván et al ., 2014; Higham et al ., 2014; Wood et al ., 2013). Poor cognitive ability (Marean, 2007; Mellars and French, 2011; Shea, 2003) and Neandertal inability to adapt to climate changes have long been used as fact ors in the Leroi Gourhan, 1983; Leroyer, 1988; Marean, 2007). It is claimed that as modern humans entered areas inhabited by Neandertals there instability (Mellars, 2004; Mellars and French, 2011) . Researchers also propose the initial modern human populations in Europe were a powerful factor in demographic and territorial competition (Mellars and French, 2011). It is claimed that modern human argued that Neandertal hunting and foraging abilities were lacking as compared to modern humans and served as a contributing factor for their replacement (Marean, 2007). However, predictive models show that competition for food resources would not have proved great enough for competition until 20 kya (Sørensen, 2011), lon Roebroeks, 2014:6). The literature contends that Neandertals were indeed cognitively capable of adapting to

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12 and Roebroeks, 2014). To say Neandertals were cognitively inferior to modern humans (Mellars and French, 2011; Wynn and Coolidge, 2008, 2011; Wynn et al., 2016) and thus the re ason they were susceptible to competitive replacement is to say there is evidence for such inferiority within the archaeological record. There would need to be archaeological inference for direct competition, for example, evidence of modern human attack on Neandertals over resources (Marean, 2007). Yet there remains no archaeological evidence for direct competition between Neandertals and modern humans (Finlayson et al., 2004; Shea, 2003). In fact, the archaeological record supports strong cognitive abiliti es in terms of symbolic expression (Villa and Roebroeks, 2014; Zilhão, 2014), contradicting the persistent assumption of an inferior trait of Neandertals as compared to modern humans and a key cause for their demise (Mellars and French, 2011; Wynn and Cool idge, 2008, 2011; Wynn et al., 2016). Perforated and pigment stained marine shells from Africa and the Near East are early modern humans throughout the Middle Stone Age in Africa and the Upper Paleolithic in the Near East (Zilhão et al., 2010:1023). Yet, there remains disagreement on what is considered complex symbolic behavior regarding Neandertal cognition (Villa and Roebroeks, 2014; Zilhão, 2014). Some researchers deem Neandertals cogni tively inferior to modern humans (Mellars and French, 2011; Wynn and Coolidge, 2008, 2011; Wynn et al., 2016) yet there exists a surplus of e vidence that supports Neandertals were behaviorally and cognitively indistinguishable from modern humans in terms o f symbolic expression as displayed throughout the archaeological record (Villa and Roebroeks, 2014; Zilhão, 2014).

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13 New research (Hoffman et al., 2018) proposes Iberian Neandertals were using 115 ka, predating any Rodríguez Hidalgo at al., 2018:2). The Neandertal archaeological record includes cases of decorated bone tools (Caron et al., rico et al., 2010; Hoffman et al., 2018; Peresani et al., 2013; Roebroeks et al., 2012; Zilhão et al., 2010) and raptor talons Hidalgo at al., 2018; Romandini et al., 2014) altered for adornmen t, and the use of natural pigments as colorants et al., 2010). Zooarchaeological analyses of avifaunal assemblages across Europe have found Neandertals took c are during the removal of raptor talons from birds of prey (Morin and Laroulandie, 2012; Rodríguez Hidalgo at al., 2018; Romandini et al., 2014) as early as 130 tailed eagle talons have been recovered from Kirpina i n Croatia (130 kya) and show minimal cut marks with polished facets in roughly 2015). As talons contain zero nutritional value, it is inferred that the careful extrac tion techniques must have been implemented to remove talons to produce minimal damage by lithic tools (Romandini et al., 2014). Researchers suggest that as large raptors are some of racted hominins Whatever the reason, it appears Neandertals took inordinate care in the extraction of raptor talons for adornment indicative of symbolic behavior and cognition. If Neandertals

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14 were symbolically indistinguishable from modern humans within the archaeological record (Villa and Roebroeks, 2014; Zilhão, 2014), there must be a reason other than competitive replacement to account for Neandertal extinction and modern human survival during unstable OIS 3. Climatic Stress It is argued that demographic changes which occurred after the arrival of modern inferiority causing rapid and to change played in Neandertal extinction (Finlayson et al., 2004), which indirectly influenced the climatic stre driven in part by the rapid succession of climate changes at the end of the Pleistocene [and] assumes that glacial cycles caused local environmental shifts that negatively impacted Neand severe enough on Neandertal populations to not allow them to recover between climatic fluctuations (Finlayson and Giles Pacheco, 2000; Finlayson et al., 2001, 2004, 2008; Hod gkins et al., 2016; Hublin and Roebroeks, 2009) and is evaluated in the archaeological record via zooarchaeological analysis of nutritional stress displayed via changes in butchering intensity as preserved on faunal remains. Nutritional Stress Several zooa rchaeological studies have exposed a pattern of Neandertals intensifying butchering of carcasses in a response to heightened nutritional stress (Castel et., 2017; Hodgkins et al., 2016; Sharon and Oron, 2013; Valensi et al., 2011). It is assumed

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15 that popul ations experiencing nutritional stress would not ignore bone marrow or grease as the caloric value of fat is nearly twice that of protein or carbohydrate (Outram, 2001). In France, Neandertal butchering strategies are known to include intensive processing of bones indicative of grease rendering (Castel et al., 2017). Intensification of butchering manifests in the zooarchaeological record via an increase in the frequency of cut and/or percussion marks (Binford, 1978; Jin and Mills, 2011; Nilssen, 2000; Outra m, 1999, 2000). In the Levant, Neandertals appear to have intensified butchering practices indicated by the and evident from the hig h frequency of cut marks displayed on fauna (Sharon and Oron, 2013:184). A controversial topic within the literature on Neandertal social and subsistence behaviors is cannibalism observed at Moula Guercy cave in southeast France (120 100 kya) (Valensi et al. 2011:48). Moula adolescents aged 15/16 years, and two youths aged 6/7 years (Valensi et al., 2011:53). The head, ribs, and vert ebral columns show minimal cut marks, implying that the butchers Neandertal remains and Red deer (the most abundant fauna) indicates similar processing practices: b utchering marks are consistent with processing for nutritive extraction including skinning, disarticulation, and defleshing along with intensive breakage to extract the brain, bone marrow, or mandibular tissue (Valensi et al., 2011). This shows intensity i n butchering and is indicative of elevated levels of nutritional stress (Binford, 1978; Jin and Mills, 2011; Nilssen, 2000; Outram, 1999, 2000). The nutritional stress of Neandertals at

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16 Moula Guercy must have been great as it seems they were forced to eat their own yet practiced a higher level of care in the butchering of people by leaving fewer cut marks than the preserved on the hunted prey like Red deer. These discussed intensified subsistence strategies employed by Neandertals follow behavioral ecologi extent to which bones are exploited for their fat depends upon the level of dietary stress behavioral ecol ogy theoretical framework which allows zooarchaeological analysis and interpretation of hominin behavior over time. Theoretical Basis of This Study Behavioral Ecology Behavioral ecology is a robust theoretical perspective often implemented in the rese subject to behavioral evolution relative to ecological conditions (Nettle et al., 2013). In other words, it is the study of how organisms adapt behaviorally to a wide array of social and ecological circumstances in fitness enhancing ways (Boone and Smith, 1998). emerged and continue Neandertal research, including reasoning of changing subsist ence behaviors to adapt to changing environments.

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17 Optimal Foraging Theory OFT was originally developed in the late 1970s under an evolutionary or behavioral ecology theoretical framework (Krebs, 1978; Pyke et al., 1977) before being applied to archaeolo gical research (Binford, 1979; Keene, 1983). OFT relies on an assumed intimate relationship between hunter gatherers and their environment to explain different mobility patterns across time and space (Keene, 1983; Kelly, 1983; Kelly and Todd, 1988). In add ition to evolutionary concepts like natural selection and adaptation, OFT further considers diet, group size, and foraging location to interpret prehistoric behavior (Keene, 1983). OFT is beneficial in Neandertal behavioral research as it has been applied archaeologically and ethnographically (Binford, 1978; Levin and Potpov, 1964; Morin, Drastic climatic changes alter the environmental c arrying capacity, which is the level of subsistence readily available to both her bivores and carnivores across the landscape, determined by a variety of ecological factors such as temperature, rainfall, and solar radiation (McNaughton et al., 1989; Oesterheld et al., 1992; Nemani et al., 2003; Rodríguez et al., 2014). Changes in these ecological factors directly influenced the carrying capacity of an environment the fauna and flora available for hunting and gathering by Neandertals across Europe (van Andel and Davies, 2003). In contrast, if an environment and therefore carrying capacit y is stable through time, zooarchaeological assemblages should not show a distinct change in subsistence behavior through time.

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18 behavioural differences in the exploitation strategies rep (Dusseldorp, 2012:2). In fact, OFT provides a link between the ease of acquisition of prey directly and the level of field processing of carcasses and the likelihood of preferential transport of certain skeletal elements in relation to the associated reward in terms of nutrition (Binford, 1978; Metcalfe and Barlow, 1992; Cannon, 2001, 2003; Faith, 2007). It is proposed that Neandertals required very high return rates from foraging (Snodgrass and Leonard, 2009); therefore, dec reases in the environmental carrying capacity would have had severe impacts on Neandertal populations (Dussledorp, 2012). Intensified butchering strategies displayed by Neandertals in the zooarchaeological record (Castel et., 2017; Hodgkins et al., 2016; S haron and Oron, 2013; Valensi et al., 2011) follow these expectations of OFT (Binford, 1980; Keene, 1983; Kelly, 1983; Pyke, 1984; Pyke et al., 1977). Middle Range Theory Zooarchaeological studies of bone surface modification are predicated on middle ran ge theory to provide testable experimental data to compare with archaeological data during analysis. First developed within sociology in the 1950s (Verhagen and Whitley, to promote methodological investigations of possible cause and effect processes of site formation (Kosso, 1991) testing assumptions of uniformitarian principles, or the assumption that those processes occurring today also occurred in the past (Lyman, 1987).

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19 For example, uniformitarianism stipulates that because wind storms are common today, they too must have commonly occurred throughout prehistory. These assumptions are critical to zooarchaeological research as a wind storm has taphonomic implications for archaeological site interpretations for example, cover and compaction of a site by animals can produce microstriations (called trampling marks) on faunal remains that closely resemble markings produces by butchering (Andrews and Cook, 1985; Behrensmeyer et al., 1986; Domínguez Rodrigo et al., 2009; Gifford Gonzalez, 1991; Fiorillo, 19 89; Fisher, 1995; Olsen and Shipman, 1988; White, 1992). It is therefore imperative zooarchaeology play a theory (Kosso, 1991:623). This zooarchaeological study adhere s to a behavioral ecology theoretical framework to investigate subsistence strategies employed by Neandertals at the cave site Arma Veirana roughly 45 kya in the Liguria region of Northwestern Italy. This thesis is interested in how Neandertals reacted to their environment and reports a taphonomic and zooarchaeological assessment of their behavior in the cave through time. Arma Veirana Arma Veirana (AV) is an archaeological cave site situated at the base of a steep cliff (441 m.a.s.l. elevation) in the Neva Valley montane region of the Ligurian Alps in Northwestern Italy ( Figure 1 ). The cave is situated approximately 14 km inland from the small town Erli where the field laboratory is located. Initial archaeological investigation began in 2015, followed again in 2016, 2017, and 2018.

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20 Radiocarbon dating has been performed on numerous charcoal and bone samples recovered from Neandertal associated layers throughout the cave. Unfo rtunately, these results require caution as the bottom of the sequence is older than 45 kya (Hodgkins, 2018 Personal communication), pushing the accuracy boundaries of radiocarbon dating. Therefore, this study assumes the earliest time of Neandertal occupa tion at AV was at least 45 kya: during the Warm Transitional phase of OIS 3 (57 38 kya) (van Andel and Davies, 2003). Due to the current lack of reliable radiocarbon dates, this study relies on field observations to provide diagnostic changes in sediment a nd artifacts following the laws of stratigraphy and superposition to account for changes in time. Archaeological excavations follow two key laws initially put forth in the field of geology: 1) the law of superposition, and 2) the law of stratigraphy (Harr is, 1979; Howe, 1970). Within archaeology, the law of superposition stipulates the oldest layer must always be the bottom most layer in an archaeological sequence, as it must have been deposited Figure 1 : Google Earth map showing location of Arma Veirana in relation to modern day Italy.

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21 before the layers covering it were deposited (Harris, 1979). The law of stratigraphy was first detailed in the late 1700s to explain the apparent changes in sediment composition not solely based on rocks and dirt, but also fossilized organic remains such as plants and (Harris, 1979:111). Within archaeology, level or stratum is dated to a time after that of manufacture of the most recent artefact n Europe, many Middle Paleolithic (MP) to Upper Paleolithic (UP) sites have lacked the ability for reliable site reconstructions to investigate stratigraphic contexts and processes of site formation, typically due to the lack of application of modern techn ological excavation methods as they were originally investigated prior to inception of current methods (Hublin, 2015). Archaeological deposits at AV have not been previously excavated, allowing for application of modern methods such as three dimensional ( 3D) surveying and mapping of all archaeological materials and stratigraphic boundaries and contexts using digital total stations since the initial investigation of the cave in 2015. Additional 3D imaging methods such as stereophotogrammetry and LiDAR scann ing have been employed at AV to further investigate stratigraphy, geological cave setting, and on going excavation settings. Photo capable drones have been commissioned to develop a 3D model of the cave and surrounding valley aiding in spatial analysis of hominin landscape use. While these 3D imaging methods provide highly detailed and impressive data, they were not utilized in this study; however, they promise to provide insight into further analysis of the cave and paleoenvironmental reconstructions of al pine Liguria.

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22 Excavation Methods Excavations at AV follow methods put forth by researchers at the Middle Stone Age (early modern human) cave and rockshelter site complex Pinnacle Point outside Mossel Bay, South Africa (Brown et al., 2010; Bernatchez an d Marean, 2011; Dibble et al., 2007; Marean et al., 2004, 2010; Oestmo and Marean, 2014). The recording system at Pinnacle site of artifacts, features, sections, surfaces, and everything else is record ed by total station directly to handheld plotting of finds and samples, section drawing, feature measurement and drawing, and grid lay (Bernatche z and Marean, 2011:1) with virtually no human transcription error (Dibble et al., 2007). AV uses prismless Topcon Pulse total stations operated by Spectra Precision handheld tablet computers running Carlson Survey Pro Software (Hodgkins, 2017 Personal communication). Total stations are electronic instruments with a built in distance meter to m easure angles and distances resulting in a specific point position in space point specific coordinate data, which allow for reconstructing spatial relationships and 3D distributions of materials for reference after excavation and ultimate destruction of st ratigraphic boundaries (Bernatchez and Marean, 2011; Brown et al., 2012: Dibble et al., 2007; Marean et al., 2004, 2010; McPherron et al., 2005). Geographic Information Systems (GIS), such as ArcMap (employed in this study), use these data to digitally map spatial relationships of excavated materials and excavation processes (Marean et al., 2004, 2010; McPherron et al., 2005) including stratigraphy and geologic formations often used to produce detailed paleoenvironmental reconstructions (Anemone et al., 201 1).

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23 ng the geological and geographical setting of the site, followed by detailed sedimentological and pedological unique and descriptive names to such stratigraphic uni ts, or layers. Changes in geogenic and anthropogenic factors such as sediment composition (texture, color, moisture, etc.), Figure 2 : Entrance of the cave facing South, showing location of the main excavation trench near the mouth. Photo taken in 2016, courtesy of Dominique Meyer.

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24 artifact density, and depositional disturbances (rootlets, rodent burrows, etc.) contribute to classification of these stratigraphic layers (Miller and Czechowski, 2016). Layers at AV are easily distinguishable in the field based on clear sediment differences (Miller and Czechowski, 2016) and are assigned as Stratigraphic Aggregates (StratAggs) and named after diagnostic sediment characteristics such as color and texture (Brown et al., 2012; Marean et al., 2004, 2010; Oestmo and Ma rean, 2014). There exist four StratAggs of analytical relevance currently exposed in the main excavation trench near the mouth of the cave ( Figure 4 ) from lowest to highest: Black Mousterian (BM), Granular (Gr), Compact Strong Brown (CSB), and Rocky Brown (RB) ( Figure 3 ). Individual excavation units are organized in the following ranked order: Square, Quad, StratUnit, and Lot Number (Lot) (Brown et al., 2012; Marean et al., 2004, 2010; Oestmo and Marean, 20 14). There are four excavation units divided within each 1 m Square 50 cm quadrants (Quads) are assigned on their directional bearing (NE, NW, SE, Figure 3 : Profile view of the east wall within the main excavation trench displaying the contacts for the four StratAggs analyzed. The red star corresponds with Figure 4 .

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25 SW) and are allocated a unique Lot Number (Marean et al., 2004, 2010; Oestmo and Marean, 2014). All artifac ts and features (regardless of size) that become exposed within these units are shot in with total stations and collected in individual specimen bags with square, quadran recording of artifacts, stratigraphy, and features are supplemented by handwritten excavator notebooks which are digitally preserved for future reference (Marean et al., 2004, 2010; Oestmo and Marean, 2014). This two step method of artifact collection is employed in an attempt to mitigate human error during recording (Dibble et al., 2007; Bernatchez and Marean, 2011). Cave Setting Neandertals are assumed to be the main inhabitants of the cave at least 45 kya as numerous fa unal remains and Mousterian lithics characterized by the contemporary use of Levallois and Discoid methods produced on mainly local raw materials have been recovered from all four StratAggs analyzed here (Hodgkins, 2017 Personal communication). Preliminary lithic analysis of AV highlights the dominant utilization of Discoid production with few Levallois elements (Negrino et al., 2017; Riel Salvatore and Negrino, 2018). Exploited raw materials prove mainly to be locally sourced quartz, quartzite, and jasper which is believed to have been sourced near the modern town of Ortovero (Negrino et al., 2016) approximately 20 km away. Additionally, a preliminary geoarchaeological investigation of AV has begun to assess stratigraphy throughout the cave to develop a mod el for site formation (Miller and Czechowski, 2016) following current methods of micromorphological analysis (Courty et al., 1989; Mallol and Mentzer, 2015).

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26 Combined, these preliminary results provide fine grained details about sediment composition within each StratAgg that are valuable in evaluating the relationship between stratigraphy and cultural remains throughout the sequence aiding in interpretations of occupation of the cave (Miller and Czechowski, 2016). Arma Veirana is a limestone cave formed alo ng a fault which eroded an offset portion of bedrock to form the cave as it is today (Miller and Czechowski, 2016). The cave is 44 meters deep from front to back, with an erosion produced sloped surface displaying younger sediments preserved towards the ba ck and older sediments exposed near the front ( Figure 4; Hodgkins, 2017 Personal communication). Figure 4 : Cave cross section derived from 3D models established using photogrammetry of the cave and surrounding valley. The red star indicates location of the main excavation trench.

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27 Deposits within the cave consist of coarse and fine grained clastic sediments and sedimentary rocks such as flowstones and speleothems; a tufa cemented breccia at the cave entrance suggests there used to be more deposits than today (Miller and Czechowski, 2016). In fact, it appears deposits at the front of the cave were thicker than at present and have eroded to expose Middle to Upper Paleolithic deposits near the cave entrance (Miller and Czechowski, 2016). Excavations at AV are arranged in several trenche s to investigate the stratigraphic sequences throughout the cave (Miller and Czechowski, 2016). While there are numerous StratAggs exposed throughout the cave, this study is solely concerned with the four Middle Paleolithic deposits exposed in the main exc avation trench: BM, Gr, CSB, and RB (from oldest to youngest). These four are of analytical interest as none of the evidence of continuous deposition, it here (Miller and Czechowski, 2016:5) potentially providing a continuous archaeological sequence at AV. While out of scope for th is study, this holds great analytical intrigue for the Arma Veirana Project. Black Mousterian The lowest StratAgg excavated thus far, Black Mousterian ( BM) was named after the dark sediment color seemingly resultant of a high density of charcoal and Mous terian lithic artifacts (Hodgkins, 2017 Personal communication). The sediment in BM is defined as a medium fine grained silt with granules and sub angular bedrock fragments up to 10cm (horizontally oriented) and overall dark in color (10YR 3/2 Munsell) with a high proportion of charcoal preserved throughout (Miller and Czechowski, 2016).

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28 Micromorphological thin sections show clear anthropogenic activity via high densities of charcoal, burned organic material, and burned fat (Miller and Czechowski, 2016). Faunal and Mousterian lithic artifact densities are greatest in th e BM and decrease from the bottom to the top of the sequence (Hodgkins, 2017 Personal communication), possibly suggesting a change in site use over time. Minimal zooarchaeological analysis of fauna recovered from AV had been performed prior to this study; however, initial analysis found butchery marks are common on bone fragments throughout the BM assemblage (Hodgkins, 2017 Personal communication). High densities of burned organic material, Mousterian artifacts, and butchery marks on faunal remains suggests Neandertals were using the cave as a butchering and/or cooking site during the BM at least 45 kya. Granular Sediment in Granular (Gr) is characterized as a medium sandy silt with granules, gravel, and sub angular strongly weathered bedrock fragments (up to 5 10 cm) with a slight color difference within the StratAgg between the upper portion (10YR 4/3 Munsell) and lower (10YR 4/4 Munsell) with high artifact density present in the upper and diffusing toward the lower portion (Miller and Czechowski, 2016), t his could represent a change in occupation and site use. Mousterian lithics and faunal remains are common in Gr and micromorphological thin sections show a clear contact between BM and Gr StratAggs marked by a decrease in anthropogenic evidence (Miller and Czechowski, 2016). Compact Strong Brown Contact between the Gr and Compact Strong Brown (CSB) is sharp with a distinct change in sediment and rock: fine sand/silt with a similar clay content as Rocky Brown, smaller (5 cm) subangualr rock fragments with hi gher proportion of schist instead of

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29 limestone (Miller and Czechowski, 2016). Micromorphological thin sections fail to show a clearly expressed contact between Gr and CSB, however they reveal evidence suggestive of gentle downslope movement of sediment in CSB that is not present in Gr (Miller and Czechowski, 2016). While anthropogenic components are rare in CSB (Miller and Czechowski, 2016), Mousterian lithics and faunal remains with butchery marks are found throughout (Hodgkins, 2017 Personal communication ) . Rocky Brown Rocky Brown (RB), is the youngest StratAgg analyzed and consists of silt and clay with granules, gravel, and large angular to sub angular rocks (10 15 cm) oriented to the North dipping Northeast and is similar in color to Gr (10YR 4/4 Munse ll) (Miller and Czechowski, 2016). RB has a higher proportion of coarse sediment material compared to CSB and the contact between the two has been noted by a thin gravel lens (Miller and Czechowski, 2016). Mousterian lithic artifacts and fauna are common a nd more abundant than in CSB yet lower than Gr and BM .

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30 CHAPTER III METHODS Sample Selection and Data Collection Following current zooarchaeological methods (Abe et al., 2002; Grayson, 1984; Lyman, 1994, 1995; Marean et al., 2000, 2001), bone surface modification analysis at Arma Veirana (AV) was performed on each faunal fragment regardless of size and/or taphonomy. Taxonomic and taphonomic assessments were performed in the field laboratory located in Erli, Italy. Of particular interest to this study are the Neandertal associated StratAggs with Mousterian lithic technology and copious faunal remains: Black Mousterian (BM), Granular (Gr), Compact Strong Brown (CSB), Rocky Brown (RB). Curation methods at AV allowed for random sampling of faunal fragments from these four StratAggs. All artifacts are organized by excavation square (Strat) and Lot number which permitted a r andom selection of a minimum of 10% of faunal fragments from the four StratAggs for analysis: BM n =713, Gr n= 295, CSB n= 54, RB n= 112. Data collection was performed using forms in Microsoft Access to provide a uniform system regardless of analyst (Marean et al., 2001). Dino Lite Microscopes ( 40x) with digital photographic capabilities were used to inspect surface modifications of each specimen, and to record all marks with the assistance of incident lighting (Blumenschine et al., 1996). Anthropogenic vs. Taphonomic Marks zooarchaeological analysis. Taphonomic analysis evaluates bone surf ace modification marks preserved on archaeological faunal remains to interpret agents of accumulation of

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31 bones and the processes that potentially contributed to the modification of remains for hundreds, thousands, or millions of years (Gifford Gonzalez, 19 91; Fisher, 1995; Lyman, 1987). To adequately distinguish anthropogenic and taphonomic markings, the field of zooarchaeology has commissioned a number of experimental and naturalistic studies under a middle range theoretical framework to contribute critica l insight into prehistoric diets (Behrensmeyer et al., 1986; Blumenschine, 1988, 1995; Blumenschine and Selvaggio, 1988; Blumenschine et al., 1996; Egeland et al., 2004; Faith, 2007; Faith and Gordon, 2007; Marean and Spencer, 1991; Olsen and Shipman, 1988 ; Pickering and Egeland, 2006; Villa and Mahieu, 1991). Experimental studies and inter analyst blind tests have shown bone surface modification analysis benefits from the use of low magnification and incident lighting to best expose the presence of any ma rkings preserved on faunal fragments (Blumenschine et al., 1996; Domínguez Rodrigo et al., 2009). In blind tests, Blumenschine and colleagues (1996) accomplished impressive accuracy in discerning between cutting/scraping marks, hammerstone percussion marks presence or absence of conspicuous and inconspicuous marks with 97% three way hese experimental data allow for rigorous comparisons against zooarchaeological assemblages to determine agents of accumulation (i.e., hominin, carnivore) and effector (i.e., hammerstone, carnivore ravaging). Taphonomic Assessment Manual calipers were use d to record the maximum length and width (mm) of specimens to investigate issues of preservation of faunal remains per StratAgg. Additional

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32 data such as color and fossilization level (none, light, heavy) were collected on all specimens to further evaluate changes in preservation in the cave over time. The overall surface visibility of fragments and obstructions such as matrix cover and dendritic etching were recorded in 10% increments (.00 1.0). All zooarchaeological tests were performed on fragments with a t least 20% surface visibility. The maximum burning stage was assigned on a scale from 0 6, with zero representing no evidence of burning and 6 as fully calcined bone which turns completely white and chalky (Nicholson, 1993; Shipman et al., 1984; Stiner et al., 1995). Long bone shaft fragment completeness was recorded on a scale of 0 3 original circumference, 2 is more than half, and 3 is a complete circumference in at leas t one portion of the fragment (Bunn, 1983; Villa and Mahieu, 1991). Trampling Marks Taphonomic marks produced by forces such as sediment compaction (Behrensmeyer et al., 1989; Fisher 1992; Marean et al., 2000) and trampling (Olsen and Shipman, 1988; Fiori llo, 1989; Dominguez Rodrigo and Barba, 2006; Domínguez Rodrigo et al., 2009) contribute to bone surface modification. It is imperative zooarchaeological analysis properly distinguish intentional hominin produced butchery marks from marks left by sediment compaction as they share several diagnostic criteria (Behrensmeyer et al., 1986; Domínguez Rodrigo et al., 2009; Olsen and Shipman, 1988) yet provide vastly different site narratives during analysis. Most researchers have relied on microstriations to characterize both: a) the marked similarities between cut marks and trampling marks ( Figure 5 ), and b) the presence and/or frequency of microstriations to distinguish between these marks (Domínguez Rodrigo et

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33 al., 2009). It is now well u Domínguez Rodrigo, 2014:1064). At AV, trampling was recoded whenever present to explore variations in site use and post depositional destruction over time. Taxonomic Assessment Skeletal element ( e.g. femur, humerus, long bone fragment), side of bone (right, left, unidentifiable), and portion of bone (e.g. epiphysis, midshaft, shaft) were assigned for all fragments whenever possible. If taxon identification was unavailable, a general taxonomic grou ping was made (e.g. bovid, carnivore, mammal, etc.). Identification on the species level was limited due to the lack of a comparative collection in the field laboratory; those that were considered identifiable were brought back to the zooarchaeology labora tory at the University of Colorado Denver where a number of fragments were further identified to bone element, side, taxon, and even species. The four StratAggs analyzed here preserve remains of medium sized artiodactyl (bovids/cervids) prey species, inclu ding Capra ibex (goat) , Cervus elaphus (Red deer) , and Capreolus capreolus (Roe deer). This follows the Figure 5 : Example of experimentally produced t rampling marks . Measurements highlight the shallowness, frequency, and irregularity of marks on a single fragment.

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34 taxonomic representation of Liguria which has reported deer as the most common prey species at the time of Neandertal occupation at AV (Valensi and Psat hi, 2004). Animal body size categories were assigned on a scale from 0 6, with 0 representing small fauna such as rodents and birds and 6 representing large fauna such elephants (Brain, 1981; Bunn et al., 1988). For example, of the identifiable species at AV, Capra ibex (50 100 kg, Aublet et al., 2008) and Capreolus capreolus (15 50 kg, Nowak and Wilson, 1999) are recorded as size 2 (20 113 kg) and Cervus elaphus (75 340 kg, Nowak and Wilson, 1999) are recorded as size 3 (113 340 kg) (Brain, 1981; Bunn et al., 1988). Animal body size is an important factor when considering transporting carcasses across the landscape as hunters favor smaller prey when travelling distances to butchering, cooking, and/or consuming sites (Binford, 1978; Schoville and Otárola Ca stillo, 2014). Transporting heavy loads across tricky alpine terrain to a consumption site was undoubtedly a concern for Neandertals using the cave to butcher and consume their prey. Surface Modification Analysis Surface modification analyses at AV we including humeri, radii, ulnae, femora, tibiae, fibulae, metacarpals, and metatarsals, crania, and mandibles as they have thick cortical walls and are more likely to preserve under archaeological contexts (Faith a nd Gordon, 2007:873; Marean and Cleghorn, 2003 ). High probability of preservation Lam et al., 1999; M arean and Cleghorn, 2003; Cleghorn and Marean, 2004; Faith and Gordon, 2007) and the impacts of post discard forces of bone destruction,

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35 (Hodgkins et al., 2016:6). T he relative frequencies of long bone shaft fragments are compared to epiphyseal fragments in a critical first step to evaluate possible biases in e xcavation and artifact retention methods (Marean et al., 2000; Pickering et al., 2003). A higher frequency of epiphyseal fragments relative to shaft fragments has been found to be resultant of sampling biases (Marean, 1998; Marean and Kim, 1998; Marean et that shaft fragments were passed over for more identifiable epiphyseal fragments during zooarchaeological analysis of fauna at AV, it is important to maintain no biases in excavation, curation, or artifact sampling prior to analysis. Breakage Patterns Within zooarchaeology, long bone fragments (e.g. humeri, metapodials, etc.) are the most descriptive in terms of determining how and why faunal remain s are present at a given site (Marean and Kim, 1998; Marean et al., 2000; Pickering et al., 2003; Villa and Mahieu, 1991). Following criteria outlined by Villa and Mahieu (1991), it is possible to distinguish between nutritive and non nutritive bone breaks . Nutritive breaks occur when bones are broken in a fresh state indicative of intentional agency by either hominins and/or carnivores with the intention of extracting bone grease and marrow from fresh bones which produces curved and/or v shaped breakage o utlines with oblique angles (Marean et al., 2000; Villa and Mahieu, 1991). In contrast, non nutritive bone breaks are produced when bones are broken in a dry state after a bone has lost all soft tissue, grease, or other nutrients, which produces transvers e breakage outlines with right angles.

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36 Marean et al. (2000) compiled experimental studies providing expected frequencies for zooarchaeological assemblages to compare data against. The breakage patterns of faunal remains at AV were compared against these e where humans had first access to bones for nu tritional extraction, followed by carnivores. These expected frequencies serve as a baseline for the main ways in which bones are assumed to accumulate in archaeological settings. Agents of Accumulation Hominins have relied on hunting as a major source of nutrition since 1.76 mya (Fernández Jalvo et al., 1999; Thompson et al., 2019 In press), with a wide variety of hunting and butchering strategies that modify bone surfaces leaving behind different types of markings. Critical to zooarchaeology is the evalua tion and assignment of diagnostic criteria for proper identification of bone surface modification marks to determine agents of bone accumulation. Caves and rockshelters were dominant residential locations for Neandertals in Western Europe (Marean, 2005). However, Neandertals were not alone in utilizing caves, carnivores such as canids are known to use them to den, thus contributing to accumulation of fauna (Brain, 1981) and/or post depositional destruction of the site. Agents of accumulation are determine d through analysis of the relative frequencies (number of marks) of percussion marks and tooth marks preserved on bovid/cervid long bone fragments as these serve as proxies for hominin and carnivore activity respectively (Blumenschine, 1988, 1995; Marean a nd Spencer, 1991; Marean et al., 1992, 2000; Marean and Bertino, 1994; Capaldo, 1995, 1997).

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37 distribution of marks, not simply their presence or absence on particular skeletal parts or (Blumenschine et al., 1996:504). At AV these frequencies are compared to expected frequencies as compiled and outlined by Marean et al. (2000) to establish the dominant bone accumulator and modifier for eac h StratAgg. Butchery marks such as cut marks (Abe et al., 2002; Behrensmeyer et al., 1989, 1995; Blumenschine and Selvaggio, 1988, 1991; Blumenschine et al., 1996; Bunn, 1981; Capaldo, 1995; Egeland, 2003; Fisher, 1992; Potts and Shipman, 1981) and percuss ion marks (Binford, 1981; Blumenschine, 1995; Blumenschine et al., 1996; Bunn, 1981, 1989; Capaldo and Blumenschine, 1994; Domínguez Rodrigo and Barba, 2006; Lyman, 1987; Outram, 2001; Pickering and Egeland, 2006) are produced in dissimilar ways and thus p roduce distinguishable marks. Both are recorded by frequency and location (i.e., epiphysis, near epiphysis, shaft) to investigate changes in butchering intensity between StratAggs (Outram, 1999, 2001). Cut Marks Experimental research began in the 1980s to produce diagnostic criteria for cut marks (Abe et al., 2002; Behrensmeyer et al., 1989, 1995; Blumenschine and Selvaggio, 1988, 1991; Blumenschine et al., 1996; Bunn, 1981; Capaldo, 1995; Egeland, 2003; Fisher, 1992; Potts and Shipman, 1981). Marks produce d by cutting to remove flesh from bones can be produced by a variety of different tools such as flakes and bifaces or scrapers, which leave roughly similar mark morphologies (Blumenschine et al., 1996:496). Blumenschine and colleagues (1996) complied crite ria from numerous studies to provide simple classification of marks to determine actor (human or carnivore) and effector through mark

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38 morphologies. Cut marks produce deep V shaped cross sections often with longitudinal microstriations ( Figure 6 ) while scrapers produce more broad and shallow marks (Blumenschine et al., 1996:496). It is important to note that while these distinctions were often made during da ta collection at AV, the distinction between flake, biface, or scraper in this study was not taken into account and were all re coded simply as cut marks during analysis. Percussion Marks Extensive experimental research provides diagnostic criteria for identifying percussion marks (PMs; Binford, 1981; Blumenschine, 1995; Blumenschine and Selvaggio, 1988; Blumenschine et al., 1996; Bunn, 1981, 1989; Capaldo and Blumenschine, 1994; Domínguez Rodrigo and Barba, 2006; Johnson, 1985; Lyman, 1987; Outram, 2001; Pickering and Egeland, 2006). PMs are a suite of marks left from hominins using hammerstones to break open long bones exposing the medullary cavities and nutrient rich marrow within (Blumenschine and Selvaggio, 1988; Blumenschine et al., 1996:496, White, 1992). They are classified as pits, grooves, isolated patches of microstriations (Blumenschine et al., 1996:496), and notches (Capaldo and Blumenschine, 1994). Figure 6 : Example of size 3 non ID mammal specimen recovered from BM (Specimen ID: 1086) . M easurements are included to show the possible frequency and size of cut marks preserved on a single fragment.

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39 oriented transverse to the long axis [ of the bone] and occurring in dense superficial patches ( Figure 7 , Blumenschine et al., 1996:496). Percussion notches ( Figure 8 ) are broad U shaped bone scars resultant of the striking of a hammerstone on a fresh bone to break it open (Capaldo and Blumenschine, 1994). Similar to the cut mark re coding, disti nctions were made between percussion marks and notches during data collection, yet both were re coded as percussion marks for simplicity in analysis. Figure 7 : S ize 2 bovid/cervid metacarpal recovered from BM (Specimen ID: 2003) . M easurements are included to show the possible frequency and size of percussion marks preserved on a single fragment . Figure 8 : S ize 3 cervid metapodial recovered from BM (Specimen ID: 1967) . M easurements are included to show the possible frequency and size of percussion notches preserved on a single fragment .

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40 Carnivore Tooth Marks Experimental studies have shown that while humans gener ally focus their subsistence strategies on extraction of meat and marrow from long bone shafts, carnivores focus their consumption around epiphyseal ends due to their high grease content which provides an excellent source of nutrients (Cleghorn and Marean, 2007; Marean and Spencer, 1991). Carnivores are known to seek out rapid access to bones once humans have discarded them (Binford, 1988; Blumenschine, 1988; Blumenschine and Marean, 1993; Capaldo, 1995, 1998; Marean et al., 2000; Selvaggio, 1998). Carnivor e tooth marks are known to be morphologically similar to hammerstone percussion marks as they both produce pits/notches on impact (Blumenschine and Selvaggio, 1988; Blumenschine et al., , which experimental studies have since critiqued to provide criteria to distinguish the two (Blumenschine et al., 1996:497; Capaldo and Blumenschine, 1994; Galen et al., 2008; Pickering, 2002 ). Transport Strategies Skeletal Abundance Accurate skeletal element abundance data are imperative to reconstructions of transport strategies as they contribute to the calculation of the number of identifiable specimens (NISP), minimum number of individuals (MNI), minimum number of elements (MNE), and minimum number of animal units (MAU) within zooarchaeological assemblages. Skeletal element survivability plays a role in the likelihood of certain elements being present or absent from a zooarchaeological assemblage as low survival skeletal elements w density, grease rich cancellous

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41 commonly absent from assemblages due to their weak nature (Faith and Gordon, 2007:873). At AV, speci fic skeletal elements (e.g. femur, humerus), side of bone (right or left), and portion of bone (e.g. epiphysis, midshaft, shaft) were assigned whenever possible to calculate the MNE and MNI per StratAgg and investigate element abundance changes over time ( Abe et al., 2002; Marean et al., 2001). To investigate food transport strategies, the MNE per StratAgg were used to calculate the MAU and the normed animal units (%MAU) to test the relationship between certain skeletal parts and their bone density values u sing the Standard Food Utility Index (SFUI) (Binford, 1978; Metcalfe and Jones, 1988). Low utility skeletal elements do not hold much nutritional value and include metacarpals, metatarsals, crania, and mandibles; while high utility elements yield plenty of flesh and are more attractive to hunters and include humeri, radii, femora, and tibiae (Faith and Gordon, 2007). Food Utility and Transport allows us to estimate the differe 2004). The MAU was calculated using MNE values and diving by the number of times the bone would appear in a complete skeleton of the living animal (Binford, 1978; Grayson, 1984). For example, the MAU of an MNE of 6 Capreolus tibiae would be calculated by dividing the MNE (6) by the number of times tibiae appear in the body (2) (6/2 = MAU of 3). The %MAU was calculated using the highest valued MAU as a normed scale following ividing each MAU by the highest MAU value found at the site before being multiplied by 100. With the %MAU values calculated, they were compared to

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42 the SFUI, or the expected amount of meat, marrow, and bone grease of each skeletal element. Comparing the %MA U of elements from each StratAgg to the SFUI allows for better understanding the relationship between bone utility and survivability (Metcalfe and Jones, 1988). Animal body part utility indices aid in the interpretation of zooarchaeological assemblages by these parts for consumption (Binford, 1978; Faith and Gordon, 2007; Marean, 2005; Metcalfe and Jone s, 1988:486; Schoville and Otárola Castillo, 2014 ). As explained by encounter rates decrease, more field processing should take place so that lower utility parts will be le ft at the field processing sites, while higher utility parts will be transported frequencies of more complete transport of smaller animals (e.g. goats or small deer) and lo wer frequencies of more complete transport of larger animals as they would be too large a function of distance between the encounter site and the residential si te. The farther the 2005:372). Within transport pattern analysis, zooarchaeological data are compared against four theoretical food utility curves. The three original uti lity curves, as proposed by Binford (1978) are: 1) bulk strategy the entirety of a carcass except the lowest utility is transported; 2) gourmet strategy only the highest utility portions of a carcass are

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43 transported; and 3) unbiased strategy portions of a carcass are transported in direct relation to their utility. Building off these, Faith and Gordon (2007) added a supplemental fourth utility curve: 4) the unconstrained strategy where entire carcasses are transported. Zooarchaeological data are compared to these utility curves to better understand hunting behaviors of hominins through time, contributing to interpretations of site use. This study is interested in understanding which specific prey transport strategies Neandertals at Arma Veirana utilized thro ugh time. Testing the Climatic Stress Hypothesis Nutritional Stress Nutritional stress is considered a key contributing factor for Neandertal extinction through the climatic stress hypothesis as the hypothesis is concerned with the heightened level of climatic instability during OIS 3 and considers the paleoenvironment to better understand Neandertal subsistence strategies and adaptations to climate change (Finlayson, 2004; Hodgkins et al., 2016; Hublin, 2009; Sharon and Oron, 2013;). Stemming from Binf studies have supported the notion that there exists a relationship between heightened fat exploitation and intensified extraction methods (Bar Oz and Munro, 2007; Outram, 1999; 2 001), the size of bone fragmentation (Outram, 1999; Bar Oz and Munro, 2007), and the transport of animal body parts and dietary significance (Faith and Gordon, 2007; Marean, 2005; Metcalf and Jones, 1988). Within zooarchaeology, nutritional stress is eval uated as a heightened frequency of cut and/or percussion marks resulting from increased efforts of butchering and extracting marrow, fats, and bone grease (Bar Oz and Munro, 2007; Outram, 1999, 2001; Hodgkins et

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44 provide valuable insight into the inte (Bar Oz and Munro, 2007:947). Therefore, investigating evidence for increased bone 1999:104). In fact, nutritiona l stress has been documented throughout the ethnoarchaeological record via the extraction of bone fats such as marrow and bone grease (Binford, 1978; Levin and Potpov, 1964; Morin, 2007). If Neandertals at Arma Veirana experienced nutritional stress, the re should be evidence of intensive processing of prey by means of cut and percussion marks on low utility bones with relatively little nutrients available for consumption such as phalanges (Bar Oz and Munro, 2007), epiphyseal ends (Outram, 1999, 2001; Mare an, 2005, 2007), and removing all nutrients from bones (Binford, 1978, 1981; Nilssen, 2000). This study included low utility skeletal elements including carpals, tarsals, and phalanges in its evaluations of butchering intensity to obtain a better understan ding of the nutritional status of Neandertals at AV (Bar Oz and Munro, 2007). Calculations of the frequency and distribution of cut marks (Binford, 1978; Nilssen, 2000) and percussion marks (Pickering and Egeland, 2006) were used to determine variations in butchering intensification by StratAgg. Statistical Analysis To test the hypothesis that cut mark and percussion mark frequencies remain relatively constant across time as a result of the reliable carrying capacity of Liguria

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45 (Valensi and Psathi, 2004), a generalized linear model was applied in analyses of covarianc e (ANCOVA) (Miller and Chapman, 2001; Hodgkins et al., 2016). As this research is concerned with Neandertal behavioral ecology, the ANCOVAs allowed for testing butchering intensity with the frequencies of cut marks or percussion marks serving as proxies f or nutritional stress (Bar Oz and Munro, 2007; Outram, 1999, 2001; Hodgkins et al., 2016). with individual StratAggs serving as independent variables, the number of cut marks or percussion marks as the dependent variables, and the maximum length and width of specimens was used to calculate the geometric mean of fragments (the covariates)(Jungers count in count data (# of marks), and it is possible to have a count of zero, a Poisson distribution (Zar, 2010), which is beneficially applied to ranges of values that meaningful relationship is exposed in analysis, a Tukey pa irwise comparison is employed to establish where in the sequence there is a significant change in butchering frequencies. Statistical analyses were performed in SPSS software with a significance level of p<0.05.

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46 CHAPTER IV RESULTS Taphonomic Analysis and Taxonomic Representation To interpret taphonomic post depositional destruction of Arma Veirana (AV) through time, the extent of bone breakage was investigated. The average faunal fragment size, in terms of maximum length and width (mm), is overall small (<3 cm in length) and remains relatively consistent across the four StratAggs analyzed: Black Mousterian (BM), Granular (Gr), Compact Strong Brown (CSB), and Rocky Brown (RB). A one way ANOVA found a significant difference in both the average length (F= 9.069, p= 0.000) and width (F= 2.767, p= 0.044) of fragments across layers. A Tukey HSD found CSB differs in average length from the other StratAggs (p= 0.000) while fragments are similar in size and differ slightly in average width in Gr (p= 0 .057) and BM (p= 0.057). Post depositional damage Trampling marks indicate movement of bones on or in sediment (Behrensmeyer et al., 1989; Fisher 1992; Marean et al., 2000), at AV this could have been due to changes in site use by Neandertals and/or other animals, disturbing bones within sediment during times of accumulation (Olsen and Shipman, 1988; Fiorillo, 1989; Dominguez Rodrigo and Barba, 2006; Domínguez Rodrigo et al., 2009). Table 1 shows all four StratAggs preserve bones with microstriations indicative of trampling marks. TRAMPLED BONES Layer: RB CSB Gr BM NISP % NISP % NISP % NISP % Trampled bone 25 22.3% 17 31.5% 52 17.6% 85 11.9% Table 1 : Number of Identifiable Specimens (NISP) and frequency (%) of fragments preserving microstriations indicative of trampling marks.

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47 To interpret site use, spatial analyses were performed to evaluate changes in taphonomy throughout StratAggs. All spatial relationship maps were produced in ArcMap (a GIS program) to investigate the visual representation of artifacts in situ prior to destruction of ar chaeological and stratigraphic contexts following excavation (Marean et al., 2004, 2010; McPherron et al., 2005). As fauna data are recorded as 3D coordinates, a 2D representation of profile and plan views help visualize spatial relationships between artif acts in situ. Profile views provide a horizontal representation of the main excavation trench facing South ( Figure 2 ) as if the viewer was standing within the trench observing Figure 9 : Profile view of the South wall of the main excavation trench showing the distribution of bones with microstriations indicative of trampling vs. non trampled bones. Map produced in ArcMap.

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48 stratigraphy straight on. Plan views provide an overview representation of artifacts as if the observer was standing directly above excavation units looking straight down. Figure 9 displays the profile view, showing an overall even distribution of bones with observed trampling marks across StratAggs. Individual maps (plan views) were produced for each StratAgg to better u nderstand the distribution of trampled vs. non trampled bones through the cave over time: BM ( Figure 10 ), Gr ( Figure 11 ), CSB (Figure 12 ), and RB ( Figure 13 ). Figure 10 : Plan view of BM. Map produced in ArcMap. Trampled bones appear to be evenly distributed through most of the BM ( Figure 9 ), however trampling marks are not observed in the northeastern most excavated area. This

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49 is similar to changes observed in other StratAggs it is possible the cave was not as heavily uti lized in this area as it is close to the East cave wall, perhaps it was too close to the wall and an unfavored area of the cave. Figure 10 shows an even d istribution of trampled bones in the excavated areas of BM, yet with notably less representation in the southwestern area. Figure 11 : Plan view of Gr. Map produced in ArcMap. As demonstrated in Figure 9 , Gr has a low representation of trampled bones, with noticeably lower frequencies of trampling marks observed on fauna recovered from the western quads. Figure 11 supports this with no observed trampling marks along the western boundary; otherwise trampled bones are evenly distributed.

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50 Figure 12 : Plan view of CSB. Map produced in ArcMap. Figure 9 reveals trampling is common in CSB, though slightly less frequent in the middle of the excavation trench. Figure 12 shows the trampled bones are evenly distributed, indicating post depositional damage was common and fairly consistent through time during deposition of CSB which is consistent with the high frequency of trampling marks reported in Table 1 .

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51 Figure 13 : Plan view of bones with microstriations indicative of trampling in RB. Map produced in ArcMap. Figure 9 shows trampling marks are sparsely represented in RB with concentrations of trampled bones recovered from the quads located at the top of the sequence and becoming less concentrated lower in the trench. A higher frequency of trampled bones closer to the su rface of RB is consistent with it be the youngest and therefore top most layer, which was still susceptible to sediment compaction at the surface prior to excavation. Figure 13 shows that while sparse, trampled bones are fairly evenly distributed in the north and concentrated along the perimeter of the excavated area in the southeast.

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52 Evidence of cooking by Neandertals Coloration of fragments indicates ca rcass processing across levels included cooking by Neandertals, which appears to have remained relatively consistent at Arma Veirana through time ( Figure 14 ). Analysis was performed on all specimens regardless of overall colors in Figure 14 relate to their level of burning, for reference: white represents fully calcined bones, or those having undergone the highest amount of burn time (delicate, dissolving bone). Figure 14 : The frequency of burned bones by maximum burning stages per StratAgg. Figure 15 shows the profile view spatial relationship between burned and unburned bones recovered from all StratAggs. Overall, burning appears sparse across StratAggs and there does not appear to be a pattern of a preferential burning area within the cave. N= 126 N= 32 N= 54 N= 7 0% 20% 40% 60% 80% 100% BM Gr CSB RB BURNING STAGES Fully Calcined >50% Calcined <50% Calcined Fully Carbonized >50% Carbonized <50% Carbonized Total Burned

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53 Figure 15 : Spatial relationship between burned and unburned bones across StratAggs. Map produced in ArcMap. As BM got its name from the burned nature of artifacts and sediment, it is unsurprising that it contains the densest concentrati on and most even distribution of burned bones of the StratAggs. CSB has more evidence of burning than RB, with a concentration of burned bones in eastern quads. Burned bones in Gr are infrequent but appear to be evenly distributed through time in the trenc h. RB preserves the lowest density of burned remains, with a small concentration at the top northwestern portion of the sequence and lacking any burned bones to the east.

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54 Taxonomic Analysis Table 2 displays the taxonomic representation at AV by the number of identifiable specimens (NISP) per StratAgg. The most representative taxon across StratAggs by far is non ID mammal most likely attributed to the small fragmentary nature of most specimens and the lack of a comparative collection in the field which made taxonomic classifications difficult. However, it is important to note that specimens were distinguished between mammal and carnivore whenever possible to better understand site use by Neandertals ve rsus carnivores as they are well represented in caves throughout Liguria (Valensi and Psathi, 2004). FAUNA ANALYZED Taxon RB CSB Gr BM Aves 2 0 0 1 Bovidae 0 2 0 10 Capra ibex 0 0 3 0 Bovid/Cervid 2 3 14 70 Canidae 1 0 0 0 Canis lupus 1 0 0 0 Non ID Carnivore 4 1 6 6 Cervidae 1 1 0 7 Cervus 0 0 0 2 Cervus elaphus 0 0 0 3 Non ID Mammal 97 41 249 555 Capreolous capreolous 2 0 0 1 Suidae 0 0 0 2 Ursus 1 4 7 3 Non ID Other taxa 1 2 16 53 Total bones analyzed 112 54 295 713 Table 2 : Number of identifiable specimens (NISP) and total number of specimens analyzed per StratAgg.

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55 Prey species represented at AV During data collection, faunal remains were assigned to specific taxon (e.g. Capreolous ) whenever possible and when unidentifiable, they were assigned a classification level of bovid, cervid, or bovid/cervid depending on how distinguishable fragments were. shows the number of identifiable specimens (NISP) of artiodactyl taxa (bovidae and/or cervidae) are common in each layer with identifiable species including Capra (goat), Capreolous capreolous (Roe deer) , Cervus (deer) , and Cervus el aphus (Red deer) . The specimens reported in also represent the bones used to perform zooarchaeological tests to determine agents of accumulation an d bone modification. BONES USED FOR TESTS Taxon RB CSB Gr BM Bovidae 0 2 0 10 Bovid/Cervid 2 3 14 70 Capra 0 0 3 0 Cervidae 1 1 0 7 Cervus 0 0 0 2 Cervus elaphus 0 0 0 3 Capreolous capreolous 2 0 0 1 Total analyzed 5 6 17 93 Table 3 : NISP of identifiable bovid, cervid, or bovid/cervid long bones used for zooarchaeological tests . Spatial representation of fauna at AV Black Mousterian is by far the densest StratAgg in terms of faunal remains, Figure 16 shows the spatial relationship of fauna in this StratAgg. As previously discussed, non identifiable mammal was the most commonly recorded fauna across StratAggs. It appears faunal remains are denser in the western excavated area than the eastern, this coul d be attributed to the East cave wall not serving as a preferred area to congregate within the

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56 cave for Neandertals and/or other animals. Evenly distributed are copious bovid/cervid prey specimens including identifiable taxa like Cervus elaphus and Caprelo us capreolous . Carnivores including non identifiable taxa and Ursus are well represented and appear evenly distributed through BM, in both excavated areas. BM preserves interesting taxa including Suidae (pigs, n=2) and Aves (bird, n=1). Figure 16 : Plan view spatial visualization of fauna from BM. Map produced in ArcMap.

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57 Figure 17 shows the spatial relationship of fauna analyzed from Granular. Bovid/cervid prey specimens are sparsely distributed across this StratAgg in two loose clusters in the southeastern and western excavated portions; one Capra specimen is nestled within the western cluster. Identifiable Ursus remains are evenly distributed while non identifiable carnivore remains appear in a loose cluster in the southeastern excavated region. It is worth noting the re are no bovid/cervid yet four Ursus remains represented in the northeastern section of Gr, this might suggest a differential use of the cave between Ursus and Neandertals during deposition of this StratAgg. Figure 17 : Plan view spatial visualization of fauna from Gr . Map produced in ArcMap.

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58 CSB preserves the lowest density of faunal remains across StratAggs yet Figure 18 shows mammals and bovid/cervid taxa are evenly distributed . The bovid and cervid specimens were not identifiable to species yet were distinct enough to distinguish between bovidae and cervidae families. Carnivore taxa including Ursus are solely represented in the southern r egion of CSB. Figure 19 shows an even distribution of bovid/cervid and carnivore remains in Rocky Brown, including identifiable taxa like Aves , Canis lupis (wolf) , Capreolous capreolous, and Ursus. In addition to the wolf remains there are non identifiable Canidae remains. Interestingly, the Capreolous specimens ( n =2) are teeth identified as an incisor and third Figure 18 : Plan view spatial visualization of fauna from CSB . Map produced in ArcMap.

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59 pre molar and are situated close to each other in the southeastern portion, likely representing one animal. Skeletal element representation Table 4 lists the number of identifiable long bone skeletal elements assigned to bovid/cervids, either specific skeletal elements (e.g., femora), or simply long bone fragments when id entification of specific skeletal elements was not possible. These bovid/cervid long bones were used in surface modification analyses to investigate evidence of butchering, cooking, and transport strategies of Neandertals at AV through time. Fig ure 19 : Plan view spatial visualization of fauna in R B. Map produced in ArcMap.

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60 NISP SKELETAL ELMENTS Skeletal Element RB CSB Gr BM Femora 1 0 1 8 Fibulae 0 0 0 1 Humer I 1 0 0 8 Long bone fragments 0 1 2 8 Metacarpals 0 0 1 3 Metatarsals 0 0 0 5 Metapodials 0 2 1 6 Radii 0 0 0 4 Radioulnae 0 0 1 4 Tibiae 1 1 0 6 Ulnae 0 0 0 0 Total 3 4 6 53 Table 4 : NISP skeletal elements (e.g. femora) or long bone fragments for all bovid/cervid remains. Surface Modification Analysis Before in depth zooarchaeological tests are performed, it is paramount to ensure a lack of excavation and curation bias within faunal assemblages as non identifiable fragments were regularly not analyzed in the past (Marean et al., 2000; Pickering et al., 2003). As AV began excavations in 2015, it was important to ensure a lack of biases prior to zooarchaeological test were performed; as such, all bone fragments were analyzed in this study. Figure 20 shows the relative abundance of shafts to epiphyses acros s StratAggs closely match expected frequencies taken from experimental data for sites where hominids and/or carnivores accumulated and modified faunal assemblages (Marean et al., 2000). As all StratAggs closely match expected relative frequencies of shafts to epiphyses versus shafts at the time of breakage, are not biased through selective retention, and are 2017:10). Further taphonomic analysis was considered before several zooarchaeological

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61 tests were employed to determine agents of bone accumulation and modification for each layer. Breakage patterns Faunal assemblages per StratAgg were compared to expected frequencies (Marean et al., 2000) of nutritive ( Figure 21 ) and non nutritive breakage ( Figure 22 ). Table 5 reports the raw zooarchaeological data used in this analysis of breakage patterns. Figure 21 shows all StratAggs have lower frequencies of nutritive bone breakage when compared to experimental data (Marean et al. 2000). RB has the second highest frequency of nutritive breakage, though lower than expected in the literature. Likewise, BM and CSB preserve similar though low frequencies of nutritive bone breakage. This is interesting as BM and CSB have the densest ( n = 713) and least dense ( n = 54) faunal remains respectively yet so closely resemble one another in breakage analysis, possibly representing similar subsistence behaviors during deposition N= 409 N= 148 N= 27 N= 76 N=21 N= 4 N=1 N= 1 0% 20% 40% 60% 80% 100% Hominid Only Hominid to Carnivore Carnivore Only Black Mousterian Granular Compact Strong Brown Rocky Brown Shafts vs. Epiphyses Epiphyses Shafts Figure 20 : The relative abundance of long bone shafts to epiphyses per layer as compared to experimental data compiled by Marean et al. (2000) , experimental data are distinguished with dots.

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62 of these StratAggs. While there appears to be drastic changes in bone accumulation across StratAggs, there is evidence for intentional extraction of nutrients from bones by either Neandertals and/or carnivores. BONE BREAKAGE DATA Nutritive Breakage Layer Oblique % Curved % Total long bone ends RB 112 73.68 105 69.08 152 CSB 48 55.81 48 55.81 86 GR 243 82.09 233 78.72 296 BM 265 57.21 468 57.21 818 Non nutritive Breakage Layer Right % Transverse % Total long bone ends RB 36 23.68 33 21.71 152 CSB 50 58.14 52 60.47 86 GR 51 17.23 59 19.93 296 BM 279 34.11 265 32.40 818 Table 5 : Long bones in each layer with nutritive breakage: oblique angles and curved outlines (top) and non nutritive breakage: right angles and transverse outlines (bottom) following methods outlined by Villa and Mahieu, 1991. Figure 22 s hows higher than expected non nutritive breakage across StratAggs, which suggests post depositional forces strongly impacted the level of preservation of faunal remains, possibly cr eating a discrepancy in bone preservation between layers at Arma Veirana. If StratAggs preserve noticeably different breakage patterns, there must be reasons for these differences; changes in sediment composition and site use serve as possible contribution s to changes in non nutritive bone breakage.

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63 Figure 16 1 : The frequency of bovid/cervid long bones broken with curved outlines and oblique angles following criteria outlined by Villa and Mahieu (1991). Data are plotted against experimental assemblages where nutrients were extracted by hominids only, hominids and carnivores, and carnivores only (Marean et al., 2000). RB CSB Hominid only Hominid to carnivore Carnivore only Gr BM 0% 10% 20% 30% 40% 50% 60% 70% 80% 90% 100% 0% 20% 40% 60% 80% 100% % OBLIQUE BREAKS % CURVED BREAKS NUTRITIVE BREAKAGE

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64 Figure 22 : The frequency of bovid/cervid long bones broken with transverse outlines and right angles following criteria outlined by Villa and Mahieu (1991). Data are plotted against experimental assemblages where nutrients were extracted by hominids only, hominids and carnivores, and carnivores only (Marean et al., 2000). Agents of accumulation Bone surface modification data was utilized to better interpret bone accumulators and modifiers per StratAgg with t he frequency of percussion marks and tooth marks serving as proxies for h ominin and carnivore activity respectively and compared to expected frequency data outlined in various actualistic studies where humans and/or carnivores had access to the bones for nutritional extraction (Blumenschine, 1988, 1995; Marean and Bertino, 1994 ; Capaldo, 1995, 1997; Marean et al., 2000, 2004). While all four StratAggs preserve at least one percussion mark (PM) on bovid/cervid long bone fragments, CSB is the only layer that preserves a carnivore tooth mark (TM, Table 6 ). RB CSB Hominid only Hominid to carnivore Carnivore only Gr BM 0% 10% 20% 30% 40% 50% 60% 70% 80% 90% 0% 20% 40% 60% 80% 100% % RIGHT BREAKS % TRANSVERSE BREAKS NON NUTRITIVE BREAKAGE

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65 PERCUSSION VS. TOOTH MARK DATA SKELETAL ELEMENT RB PM CSB PM Gr PM BM PM RB TM CSB TM Gr TM BM TM Humerus 0 0 0 6 0 0 0 0 Radius/radioulna 0 0 1 3 0 0 0 0 Metacarpal 0 0 1 1 0 0 0 0 Femur 1 0 0 2 0 0 0 0 Tibia 0 1 0 2 0 0 0 0 Metapodial 0 0 0 2 0 1 0 0 Metatarsal 0 0 0 1 0 0 0 0 Long bone fragments 0 1 1 2 0 0 0 0 Table 6 : Raw data for the number of bovid/cervid long bone fragments with at least one percussion mark (PM) or one tooth mark (TM). Note no specimens displayed both PM and TM. Carnivores are known to favor the greasy epiphyseal ends of long bones (Marean and Spencer, 1991), thus preserving higher frequencies of tooth marks on epiphyses over midshafts (Faith, 2007). However, as epiphyses are spongy bone, they are less dense and more vulnerable to post depositional damage than shaft fragments (Marean and Kim, 1998). Results from the breakage analysis ( Figure 21 and Figure 22) and the frequency of shafts to epiphyses analysis ( Figure 20) demonstrate epiphyseal ends were infrequently recorded at AV, suggesting they degraded over time through taphonomic forces and/or carnivore ravaging activity. However, it appears there was not a substantial amount of carnivore activity in any StratAgg ( Figure 23) , indicating substantial taphonomic destruction of bones over time at AV. Figure 2 3 shows a comparison of the percentage of PMs to TMs present on bovid/cervid long bone fragments per StratAgg. RB, Gr, and BM closely match expected frequencies of hominid only accumulation while CSB matches assemblages where humans first had access to bones before carnivores, further supporting the inference that

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66 Neandertals were the main bone accumulators through time at AV while carnivores had intermittent access to the cave and bones within. Figure 23 : The percentage of bovid/cervid/ long bone fragment s with at least one tooth mark versus the percentage with at least one percussion mark. Data are plotted against experimental assemblages where nutrients were extracted by hominids only, hominids had first access before carnivores, and carnivores only (Blu menschine, 1988, 1995; Marean and Spencer, 1991; Marean et al., 1992, 2000; Marean and Bertino, 1994; Capaldo, 1995, 1997). Ellipses hold no statistical meaning but are used for visual aide. Transport Behaviors Figure 24 s hows medium sized (size 2 3) bovid/cervid fragments dominate through time. Ethnographic records show a high probability for hunters to exploit smaller game for easier transport from a kill site back to a butchering and/or consumption site (Binford, 1978; S choville and Otárola Castillo, 2014). As time went on at AV, it appears small to RB CSB Gr BM 0% 10% 20% 30% 40% 50% 60% 0% 20% 40% 60% 80% 100% % PERCUSSION MARKS % TOOTH MARKS PERCUSSION VS. TOOTH MARKS Hominid only Homind to carnivore Carnivore only

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67 medium prey were regularly transported to the cave for consumption. BM has the highest concentration of faunal remains of any layer it is the most diverse in prey size (sizes 2 5). Interestingly, as time goes on, bovid/cervid body sizes get relatively smaller: BM preserves sizes 2 5, Gr sizes 2 4, CSB only sizes 3 and 4, and RB only sizes 2 and 3 this is not a clear picture as well represented prey size fluctuates through time . While more evidence is needed, these repetitive reductions in prey body size may suggest times of climate change declines in large mammalian populations across Europe du ring oscillating OIS 3 (Stewart et al., 2003b). If the flora available for fauna consumption became sparse and less reliable during climatic shifts, larger animals would be stressed as a result of limited food resources. Figure 24 : Number of identifiable bovid/cervid by animal body size as detailed by Brain (1981). To investigate changes in food transport behavior at AV through time, only high survival skeletal elements were used to calculate the minimum number of skeletal elements (MNE) and the normed minimum animal units (%MAU) per StratAgg ( Table 7 ). 0 10 20 30 40 50 5 4 3 2 NISP BODY SIZE BOVID/CERVID PREY SIZE RB CSB Gr BM Size Distinction 2 (20 113 kg) 3 (113 340 kg) 4 (340 900 kg) 5 (900 2000 kg)

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68 Unfortunately, the top three StratAggs (RB, CSB, and Gr) do not have suffi cient data to these data are reported for reference within the Appendix. TRANSPORT PATTERN DATA BLACK MOUSTERIAN Skeletal Element MNE L MNE R Side N/A MAU SFUI %MAU Cranial 1 0.5 9.1 6.7 Mandible 1 1 11.5 13.3 Humerus 1 7 7.5 36.8 100 Radius/radioulna 3 5 6.5 25.8 86.7 Metacarpal 3 1.5 5.2 20 Femur 1 2 5 7.5 100 100 Tibia 1 3 2 5.5 62.8 73.3 Metatarsal 1 4 4.5 37 60 Table 7 : MNE and MAU of all bovid/cervid fragments from Black Mousterian. Raw data are compared to the SFUI as published by Metcalfe and Jones (1988). Figure 25 shows how the %MAU of each skeletal element within BM compared to the SFUI (Metcalf and Jones, 1988) of each element with its associated caloric value (Binford, 1978; Faith and Gordon, 2007; Metcalf and Jones, 1988) to interpret foo d transport strategies: bulk, gourmet, unbiased, or unconstrained. Figure 25 shows BM most suggesting a mixed strategy that fluctuated between transporting only high utility elem ents and transporting elements in direct correlation to their food utility respectively.

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69 Figure 25 : The relative abundance of high survival skeletal elements (% minimum animal units) and their associated Standardized Food Utility Index (SFUI) (following Binford, 1978; Faith and Gordon, 2007; Metcalf and Jones, 1988). Data are plotted against utility curves (Binford, 1978; Faith and Gordon, 2007), curves hold no statistical meaning but are used for comparison. The Poly. Line holds no statistical meaning but shows the line of best fit for transport strategies in Black Mousterian. Analysis of Nutrient Extraction Behaviors Table 8 shows the low u tility skeletal elements included in the analysis of butchering intensity at AV. These bones were analyzed regardless of taxon to obtain a n understanding of the nutritional status of Neandertals over time. If Neandertals were indeed nutritionally stressed it should be assumed they would not leave any bones unprocessed therefore intensifying butchering behaviors that manifest in the zooarchaeological record as an increased frequency of cut and/ or percussion marks. However, the environment of Liguria ~45kya is characterized as relatively stable with Crania 0 20 40 60 80 100 0 20 40 60 80 100 % MAU SFUI BM Poly. (BM) Unconstrained Tibiae Metatarsals Mandibles Radioulnae Metacarpals Femora Humeri

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70 sparse forests in steppe like terrain (Holt et al., 2018) and an abundance of apline taxa like Capra ibex and Capreolous capreolous (Valensi and Psathi, 2004), both of which are frequently recorded at AV. LOW UTI LITY ELEMENTS Skeletal Element RB CSB Gr BM Astragalus 0 0 0 1 Calcaneus 1 0 1 1 Cranial 4 2 6 25 Cranial Fragment 0 2 0 8 Distal Phalanx 1 0 1 3 Intermediate Phalanx 0 0 0 5 Lunate 0 0 0 1 Magnum 0 0 1 0 Mandible 1 0 1 6 Maxilla 1 0 0 3 Metacarpal 2 0 2 4 Metacar pal III 1 0 0 0 Metapodial 1 4 3 17 Metatarsal 0 0 0 9 Non ID Carpal 0 0 0 2 Phalanx 1 0 1 2 Pisiform 0 0 0 1 Proximal Phalanx 3 1 5 6 TOTALS 16 9 21 94 Table 8 : NISP of all low utility skeletal elements (regardless of taxon) used to test butchering intensity and evaluate nutritional stress. Evaluating Nutritional Stress Two analyses of covariance (ANCOVA) were applied to test the hypothesis that cut mark (CM) and percussion mark (PM) frequencies remain relatively constant through time at AV as an effect of the reported reliable carrying capacity of Liguria (Valensi and Ps athi, 2004). The ANCOVA testing for a difference in the frequency of CMs and controlling for fragments size (geometric mean of fragments) across StratAggs did not find a significant

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71 00 ): accepting the nul l hypothesis that there is not a significant change in the frequency of CMs in the different layers. However, Figure 26 across the layers (see Table 13 , Table 14 , Table 15 Table 16 for raw data), namely in CSB. The frequency of PMs per fragment increased in Gr , producing the biggest difference across StratAggs suggesting an intensity in processing carcasses to more thoroughly extract marrow from within bones: signifying a time of nutritional stress at AV (Bar Oz and Munro, 2007; Outram, 1999, 2001; Hodgkins et al., 2016). A Tukey pairwise comparison found a significant difference (p= 0.027, df= 3) in the frequency of percussion marks between BM and Gr. There is an increase of 0.31 percussion marks per fragment from BM to Gr.

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72 Figure 26 : ANCOVA results showing statistically significant changes in the frequency of percussion marks (PM) on high and low utility skeletal elements through the layers. The central points are the mean number of PMs preserved on skeletal elements. The error bars are the 95% confidence intervals for those means. The lines connecting the layers hold no statistical power but show the interaction effect between layers. Spatial an alysis of percussion marks Utilization of plan and profile view GIS maps help illustrate the spatial and temporal distribution of fauna analyzed at AV. Figure 27 and Figure 28 show a low density of fauna that preserve at least one percussion mark (PM) in BM. Percussed bones are eve nly distributed through the StratAgg, yet with a lower concentration in the eastern excavated area in both plan and profile views.

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73 Figure 27 : Plan view spatial visualization of fauna with at least one percussion mark (PM) in BM. Map created in ArcMap.

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74 Figure 28 : Profile view (of the South wall) spatial visualization of fauna with at least one percussion mark (PM) in BM. Map created in ArcMap.

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75 Figure 29 : Plan view spatial visualization of fauna with at least one percussion mark (PM) in Gr. Map created in ArcMap. Figure 29 shows a low density and fairly even distribution of fauna with PMs in Gr (plan view), with a lack of specimens with PMs in the eastern most excavated area. Interestingly, Figure 30 shows an underrepresentation of PMs i n the eastern section of the excavated section along the South wall (profile view). The absence of percussed bones in this area through time might suggest a differential use of space within the cave for percussing and processing carcasses. The profile view map also shows percussed bones roughly along the same horizon, possibly indicative of quick occupation events.

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76 Figure 30 : Profile view (of the South wall) spatial visualization of fauna with at least one percussion mark (PM) in Gr . Map created in ArcMap.

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77 While there are few bones that have PMs in Compact Strong Brown (CSB), Figure 3 1 shows those that do are evenly distributed across the StratAgg. The profile view (Figure 3 2 ) shows an even distribution of percussed bones along two horizons in both the upper and lower excavated areas of CSB, this could also support the idea of quick occ upation events in Granular, possibly use of the cave for overnight camps. Figure 31 : Plan view spatial visualization of fauna with at least one percussion mark (PM) in CSB . Map created in ArcMap.

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78 Figure 32: Profile view (of the South wall) spatial visualization of fauna with at least one percussion mark (PM) in CSB. Map created in ArcMap.

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79 Figure 33 : Plan view spatial visualization of fauna with at least one percussion mark (PM) in RB. Map created in ArcMap. Figure 3 3 shows few bones with PMs and only at the top of the southern portion of the excavated area of Rocky Brown (RB). The profile view (Figure 3 4 ) shows a very low frequency of percussed bones and only present in the lower excavated portion. It is interesting pe rcussed bones are only found in selective areas within RB, this further supports quick use of the cave over time.

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80 Figure 34 : Profile view (of the South wall) spatial visualization of fauna with at least one percussion mark (PM) in RB. Map created in ArcM ap.

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81 CHAPTER V DISCUSSION AND CONCLUSION Discussion Zooarchaeological analysis results suggest climate change influenced fluctuations in species abundance in Liguria through time as represented bovid/cervid prey body sizes oscillated at A rma V eirana . Interestingly, the most drastic shift in body size is observed between BM and the rest of the StratAggs when size 5 prey is not represented again. It is noteworthy that size 5 prey species occur only in this StratAgg as it could signify a time of d ense forest growth and therefore not likely a time of nutritional stress amongst Neandertals if they had reliable food resources. Results from taphonomic and surface modification analyses support the inference that Neandertals used the cave for butchering and cooking prey while carnivores took advantage of the cave for scavenging or possibly even hibernating . T here exists direct evidence for hominid to carnivore accumulation (Marean et al., 2000) in CSB . C arnivore remains represent a very small proportion o f the assemblages from AV, namely Ursus ( n = 15 total across StratAggs) and non identifiable carnivores ( n = 17 total across StratAggs) are found in every layer . Understanding the taphonomic setting of each StratAgg aids in the interpretations of site use over time. Black Mousterian It appears post depositional damage was not substantial in BM: taphonomic analysis shows that BM preserves the highest number of individual trampled bones ( n =85) yet the overall proportion of bones with trampling marks is lowest (11.9%). This is consistent with results from the breakage pattern analysis of non nutritive bone breaks, with the overall second lowest frequenci es observed. BM also preserves the second lowest

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82 frequencies of nutritive bone breaks. The analysis of long bones of bovid/cervid prey species results show BM matches expected frequencies of hominid only accumulation with plenty percussion marks and no car nivore tooth marks. Additionally, BM preserves the overall highest proportion of burned bone (76.6%) and it is worth noting it is the only StratAgg that preserves fully calcined fragments ( n =2), indicating Neandertals were burning to a further extent in BM . This is interesting as fully calcined bones are rarely recorded among zooarchaeological assemblages from Paleolithic sites across Italy (Stiner et al., 1995). Results from the transport pattern analysis show BM falls between bulk and unbiased strategies Schoville and Otárola Castillo, 201 4:2). Neandertals transporting elements in direct relation to their utility follows should be the normative hunter Schoville and Otárola Castillo, 2014:2). Together, t hese results do not suggest Neandertals were nutritionally stressed in the BM as they were clearly making decisions at kill sites on what body parts to transport back to the cave not transporting entire carcasses for optimal caloric extraction. Granular Re sults suggest strong taphonomic forces were at play during deposition of Gr, such as sediment compaction by trampling, which contributed to the smallest observed fragmented bones of the StratAggs. Neandertals proved to be the main agents of accumulation in Gr as results closely match expected frequencies of hominid only

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83 accumulation via analysis of percussion marks versus carnivore tooth marks on bovid/cervid long bones (Blumenschine, 1988, 1995; Marean and Spencer, 1991; Marean et al., 1992, 2000; Marean a nd Bertino, 1994; Capaldo, 1995, 1997). Further evidence supporting Neandertals were the main accumulators at AV during deposition of Gr comes from a high frequency of burned bones. In fact, 30% of the assemblage shows evidence for burning, indicating dire ct contact with fire either for cooking or warmth and may even signify a time of a warmer climate during Gr as fire use in by Neandertals in Europe is known to increase in temperate and decrease in colder climates yet this is proposed to demonstrate a n opp ortunistic approach to fire use (Sandgate et al., 2011). This could be due to more trees growing in warmer environments and an overall easier acquisition of firewood to haul up to the cave. As t he carrying capacity of Liguria during OIS 3 is reported as reliable in terms of prey species (Valensi and Psathi, 2004) it is assumed butchering intensity was relatively constant through time at AV. B utchering behaviors should not fluctuate through StratAggs and there should not be a dramatic increase or decrease in the frequencies of cut marks and/or percussion marks through time. However , results from the ANCOVA show a statistically significant shift in the frequency of percussion marks preserved on both high df = 1, p= 0.016), with an increase of 0.31 percussion marks per fragment (p= 0.027, df = 3) between BM and Gr which suggests greater efforts were put into extracting bone grease and marrow from within bones during this time. These results reject the null hypothesis that there is no change in nutrient extraction intensity. If Neandertals were adapting their subsistence behaviors to adapt to changes in t heir environment at AV, they were likely just as adaptive elsewhere in Europe

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84 as well. This finding calls into question the rigidity of the climatic stress hypothesis, it is possible climate and environmental change were not as stressful for Neandertals as currently believed. Compact Strong Brown The sediment of CSB is defined as fine grained sand and clay with small rocks (up to 5 cm) with micromorphological thin sections suggesting there was gentle downslope movement of sediment as well as a shift in ge ologic composition (Miller and Czechowski, 2016). These factors could signal a substantial shift in sediment composition modifying bones to a further extent in CSB than other StratAggs. As an effect of the sample size of CSB ( as compared to other StratAggs ) it preserves the highest proportion of trampled bones (31.5%) and the lowest number of bones with trampling marks ( n = 17), suggesting infrequent but strong forces of site destruction . Evidence for anthropogenic activity is sparse in CSB ( n =54) and while every StratAgg analyzed contains bovid/cervid long bone fragments with at least one percussion mark, solely CSB preserves a carnivore tooth mark on these same bones, indicating Neandertals had initial access and carnivores scavenged the bones af terwards (Blumenschine, 1988, 1995; Marean and Spencer, 1991; Marean et al., 1992, 2000; Marean and Bertino, 1994; Capaldo, 1995, 1997). Interestingly, CSB is the only StratAgg with tooth marks on bones ( n =5), both long bone and ribs. CSB likely represents dual occupation by Neandertals and carnivores of the cave over time as it preserves the highest proportion of carnivore remains (9.26%) including the second highest number of Ursus remains ( n = 4) . It is possible bears were utilizing the cave for hibernati on through time, which could contribute to the elevated levels of post depositional destruction in CSB. When Neandertals

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85 were utilizing the cave, they were rarely using fire, or the cave in general as there is low evidence of burned bone (10%). These resul ts together infer carnivores had increased access to the cave during deposition of CSB than in any other StratAgg while Neandertals maintained primary agents of accumulation over time. Rocky Brown RB preserves the second highest number ( n = 25) and proport ion of trampled bones (22.3%) likely suggesting strong post depositional damage. This is consistent with every layer preserving higher than expected levels of non nutritive bone breakage and lower than expected nutritive breaks as compared to experiments w here bones were broken by humans and carnivores (Marean et al., 2000) . T he lower than expected nutritive bone breaks indicates post depositional damage altered bones to remove nutritive breaks and replace them with non nutritive breaks . RB preserves percus sion marks on bovid/cervid long bones and there is evidence for cooking; although it is the lowest frequency (8%) of burned bones across StratAggs. Bovid/cervid prey body sizes are smallest in RB (only 2 and 3) and though more data are needed this may suggest a time of decline in the carrying capacity or possibly a time of stress for Neandertals. It would be interesting to see results from transport pattern analysis of this layer to see if Neandertals were mostly hauling high yield bones back to the cav e which could support a change in the environment. If they were mainly hunting smaller game, it would be expected for Neandertals to be more selective in what body parts to bring back to the cave for the highest caloric return. Unfortunately, RB lacked suf ficient data to compare to the SFUI in this study . Results indicate Neandertals were not using the cave as a butchering site or camp site very often during deposition of RB.

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86 TEMPORAL TRENDS Inference category RB CSB GR BM Total bones analyzed n=112 n=54 n=295 n=713 Prey body sizes represented Sizes 2 and 3 Sizes 3 and 4 Sizes 2 4 Sizes 2 5: most diverse Burned bones 8% 10% 30% 76.6% Only calcined fragments Frequency of carnivore remains 6.25% Highest frequency: 9.26%. Only layer with tooth marks 4.41% Lowest frequency: 1.26% Trampled bones 23.21% Least specimens yet highest frequency: 31.48% 17.63% Lowest frequency: 12.06% Nutritive breakage 73.68% 55.81% 82.09% Most closely resembles hominid to carnivore accumulation. 57.21% Non nutritive breakage 23.68% 58.14% 17.23% 23.68% Special attributes Most closely matches expected frequencies of non nutritive breakage Carnivores had increased access to the cave. ANCOVA found a difference in the frequency of percussion marks. (PMs) 1, p= 0.016) and an increase of 0.31 PMs per fragment (p=0.027, df= 3). Only StratAgg with data for transport pattern analysis. Falls between bulk and unbiased strategies. Table 9 : Summary table of temporal trends observed at AV.

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87 Conclusion This study used principles of Human Behavioral Ecology to evaluate changes in subsistence behaviors at Arma Veirana and to explore the climatic stress hypothesis. This taphonomic and zooarchaeological report has presented data on species representation, a gents of bone accumulation, and the post depositional destruction of the site through time . Notable differences between StratAggs are summarized in Table 9. The cave was used for processing bones and cooking prey as butchery marks and burned bones are comm on in all StratAggs. Bovid/cervid prey taxa are found in all layers: 12 bovid specimens and 17 cervid specimens including 3 Cervus elaphus , and 3 Capra were brought to the cave by Neandertals. Analysis of surface modifications indicate s Neandertals intenti onally processed bones to extract nutrients for consumption including evidence for disarticulation, defleshing, and breaking bones open for the marrow within. Cut marks and percussion marks are well represented across StratAggs, indicating Neandertals used the cave to butcher prey through time. Transport pattern analysis results show Neandertals were making decisions at kill sites on what body parts to transport back to the difficult to access cave. Analysis of surface modification on both high and low ut ility bones shows an increase in the frequency of percussion marks preserved on fragments (0.31 marks per fragment, p= 0.027, df= 3) in Gr. This is direct evidence they were able to change their behaviors to adapt to change , yet w hat that change is remains uncertain . I t is possible there was a drastic environmental change or that humans came onto the scene and caused subsistence stress for Neandertals. Whatever the reason , it is interesting Neandertals at AV

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88 modified their actions to increase the cal oric extraction of prey during deposition of solely one layer. AV promises to contribute to the ongoing discussion surrounding Neandertal subsistence behaviors and their ability to adapt to changing climates. T he cave is of analytical value as it is unique for two reasons . First, it is difficult to access located in the mountains while most Neandertal cave sites in Italy are easily accesible along the co ast . Second, occupation of the site occurred during seemingly temperate climate s within a possible refugi a area yet results from this study indicate a period of nutritional stress . Res ults indicate substantial shifts in site use over time: Neandertals used the cave less through time while carnivores took more advantage in their absence . The larger prey body sizes preserved in BM and Gr indicate a difference in the flora compared to smaller sizes in CSB and RB as larger cervids like Cervus elaphus prefer dense wooded areas (Valensi and Psathi, 2004) . It is possible there was a change in the environm ent. It is also possible the cave was favored less through time as it was more physically demanding to access than the seemingly favored coastal caves across Italy . More secure dates from StratAggs at AV will contribute to an enhanced understanding of clim ate and environmental changes in Liguria. Results from this preliminary analysis of faunal assemblages from AV reject the climatic stress hypothesis . This study shows Neandertals adapted subsistence behaviors to deal with nutritional stress . The climatic s tress hypothesis fails to consider is how much behavior modification is sufficient for change to be successful. Perhaps Neandertals were more adaptive in their subsistence behaviors than they are currently given credit for. Further zooarchaeological analy sis, combined with other methods briefly outlined in this report, will strengthen the understanding of Neandertal subsistence behaviors at AV

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89 through time. If Neandertals were indeed successful in adapting to survive environmental changes, it begs to ask h ow much adaptation would have saved the Neandertals? More fine grained zooarchaeological analyses of faunal assemblages across Europe will provide highly valuable and unique insight into subsistence behaviors and the role climate truly had in the extinction of Neandertals.

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90 REFERENCES Abe, Y., Marean, C.W., Nilssen, P.J., Assefa, Z., Stone, E.C., 2002. The analysis of cutmarks on archaeofauna: A review and critique of quantification procedures, and a new image analysis GIS approach. American Antiquity 67, 643 663. Albon, S.D. and Langvatn, R., 1992. Plant phenology and the benefits of migration in a temperate ungulate. Oikos , pp.502 513. Alfano, M.J., Barron, E.J., Pollard, D., Huntley, B. and Allen, J.R., 2003. Comparison of climate model results with European vegetation and permafrost during oxygen isotope stage three. Quaternary Research, 59(1), pp.97 107. Allen, J.R., Watts, W.A. and Huntley, B., 2000. Weichselian palynostratigraphy, palaeovegetation and palaeoenvironment; the reco rd from Lago Grande di Monticchio, southern Italy. Quaternary International, 73, 91 110. Anemone, R.L., Conroy, G.C. and Emerson, C.W., 2011. GIS and paleoanthropology: Incorporating new approaches from the geospatial sciences in the analysis of primate an d human evolution. American journal of physical anthropology , 146 (S53), pp.19 46. Aublet, J.F., Festa Bianchet, M., Bergero, D. and Bassano, B., 2009. Temperature constraints on foraging behaviour of male Alpine ibex ( Capra ibex ) in summer. Oecologia , 159 ( 1), pp.237 247. Banks, W.E., d'Errico, F., Peterson, A.T., Kageyama, M., Sima, A. and Sánchez Goñi, M.F., 2008. Neanderthal extinction by competitive exclusion. PLoS One , 3 (12), p.e3972. Bar Oz, G. and Munro, N.D., 2007. Gazelle bone marrow yields and Epip alaeolithic carcass exploitation strategies in the southern Levant. Journal of Archaeological Science , 34 (6), pp.946 956. Barron, E.J. & D. Pollard, 2002. High resolution climate simulations of Oxygen Isotope Stage 3 in Europe. Quaternary Research 58 , 296 309. Behrensmeyer, A.K., Gordon, K.D. and Yanagi, G.T., 1986. Trampling as a cause of bone surface damage and pseudo cutmarks. Nature, 319(6056), p.768. Bergmann, K.G.L.C. (1847) Über die Verhältnisse der wärmeokönomie der Thiere zu ihrer Grösse. Göttinge r Studien, 3, 595 708. Benazzi, S., Douka, K., Fornai, C., Bauer, C.C., Kullmer, O., Svoboda, J., Pap, I., Mallegni, F., Bayle, P., Coquerelle, M. and Condemi, S., 2011. Early dispersal of modern humans in Europe and implications for Neanderthal behaviour . Nature, 479(7374), pp.525 528.

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102 Riel Salvatore, J., Ludeke, I.C., Negrino, F. and Holt, B .M., 2013. A spatial analysis of the late mousterian levels of Riparo Bombrini (Balzi Rossi, Italy). Canadian Journal of Archaeology/Journal Canadien d'Archéologie , pp.70 92. Riel Salvatore, J. and Negrino, F., 2018. Proto Aurignacian Lithic Technology, M obility, and Human Niche Construction: A Case Study from Riparo Bombrini, Italy. In Lithic Technological Organization and Paleoenvironmental Change (pp. 163 187). Springer, Cham. Rodríguez, J., Blain, H.A., Mateos, A., Martín González, J.A., Cuenca Bescós , G. and Rodriguez Gomez, G., 2014. Ungulate carrying capacity in Pleistocene Mediterranean ecosystems: Evidence from the Atapuerca sites. Palaeogeography, Palaeoclimatology, Palaeoecology, 393, pp.122 134. Rodríguez Hidalgo, A., Morales, J.I., Cebriá, A. , Courtenay, L.A., Fernández Marchena, J.L., García Argudo, G.G., Marín, J., Saladié, P., Soto, M., Tejero, J.M. and Fullola, J.M., 2018. The châtelperronian Neandertals of Cova Foradada (Calafell, Spain) used Iberian imperial eagle phalanges for symbolic purposes (No. e27133v1). PeerJ Preprints. Roebroeks, W., Sier, M.J., Nielsen, T.K., De Loecker, D., Parés, J.M., Arps, C.E. and Mücher, H.J., 2012. Use of red ochre by early Neandertals. Proceedings of the National Academy of Sciences, 109(6), pp.1889 189 4. Romandini, M., Peresani, M., Laroulandie, V., Metz, L., Pastoors, A., Vaquero, M. & Slimak, L. 2014, "Convergent evidence of eagle talons used by late Neanderthals in Europe: a further assessment on symbolism", PloS one, vol. 9, no. 7, pp. e101278. Sa ndgathe, D.M., Dibble, H.L., Goldberg, P., McPherron, S.P., Turq, A., Niven, L. and Hodgkins, J., 2011. On the role of fire in Neandertal adaptations in Western Europe: evidence from PaleoAnthropology , 2011 , pp.216 242. Sankararaman, S., Patterson, N., Li, H., Pääbo, S. and Reich, D., 2012. The date of interbreeding between Neandertals and modern humans. PLoS Genet , 8 (10), p.e1002947. Sankararaman, S., Mallick, S., Dannemann, M., Pr fer , K., Kelso, J., Pääbo, S., Patterson, N., Reich, D., Patterson, N., Reich, D., 2014. The genomic landscape of Neanderthal ancestry in present day humans. Nature 507, 354:7. Schoville, B.J. and Otárola Castillo, E., 2014. A model of hunter gatherer skelet al element transport: The effect of prey body size, carriers, and distance. Journal of human evolution , 73 , pp.1 14. Shea, J.J., 2003. Neandertals, competition, and the origin of modern human behavior in the Levant. Evolutionary Anthropology: Issues, News , and Reviews: Issues, News, and Reviews , 12 (4), pp.173 187.

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105 Whitaker, L., 1914. On the Poisson Law of Small Numbers. Biometrika 10, 36e71. Wynn, T. and Coolidge, F.L., 2011. How to think like a Neandertal . OUP USA. Wynn, T. and Coolidge, F.L., 2008. A Stone Age meeting of minds. American Scientist, 96(1), p.44. Wynn, T. and Coolidge, F.L., 2011. The implications of the working memory model for the evolution of modern cognition. International Jo urnal of Evolutionary Biology , 2011 . Wynn, T., Overmann, K.A. and Coolidge, F.L., 2016. The false dichotomy: a refutation of the Neandertal indistinguishability claim. Journal of Anthropological Sciences, 94, pp.1 22. Zar, J.H., 2010. Biostatistical Anal ysis, 5th ed. Prentice Hall Inc, Upper Saddle River. Aurignacian interstratification at the Châtelperronian type site and implications for the behavioral modernity of Neandertals. Proceedings of the National Academy of Sciences , 103 (33), pp.12643 12648. Zilhão, J., Angelucci, D.E., Badal Sanchez, M.J., Montes Bernardez, R ., Murcia Mascaros, S., Perez Sirvent, C., Roldan Symbolic use of marine shells and mineral pigments by Iberian Neandertals. Proc. Natl. Acad. Sci. 107, 023 1028.

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106 APPENDIX A. Transpor t Data ROCKY BROWN Skeletal element MNE L MNE R Side N/A MAU SFUI %MAU Cranial 9.1 Mandible 11.5 Humeru S 1 0.5 36.8 100 Radius/radioulna 25.8 Metacarpal 5.2 Femur 1 0.5 100 100 Tibia 1 0.5 62.8 100 Metatarsal 37 Table 10 : The minimum number of skeletal elements (MNE) and the minimum number of animal units (MAU) of all bovid/cervid fragments from Rocky Brown. Raw data are compared to the Standard Food Utility Index (SFUI) as published by Metcalfe and Jones (1988). COMPACT STRONG BROWN Skeletal element MNE L MNE R Side N/A MAU SFUI %MAU Cranial 9.1 Mandible 11.5 Humerus 36.8 Radius/radioulna 25.8 Metacarpal 5.2 Femur 100 Tibia 2 1 62.8 100 Metatarsal 37 Table 11 : MNE and MAU of all bovid/cervid fragments from Compact Strong Brown. Raw data are compared to the SFUI as published by Metcalfe and Jones (1988).

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107 GRANULAR Skeletal element MNE L MNE R Side N/A MAU SFUI %MAU Cranial 9.1 Mandible 11.5 Humerus 36.8 Radius/radioulna 1 0.5 25.8 100 Metacarpal 1 0.5 5.2 100 Femur 1 0.5 100 100 Tibia 62.8 Metatarsal 37 Table 12 : MNE and MAU of all bovid/cervid fragments from Granular. Raw data are compared to the SFUI as published by Metcalfe and Jones (1988).

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108 A. ANCOVA Data ROCKY BROWN Skeletal Element High/low utility? # CM # PM Length (mm) Width (mm) Geometric Mean (=SQRT(L*W) Femur High 0 1 62 19 34.3220046 Long Bone Fragment High 2 0 20 7 11.83215957 Long Bone Fragment High 0 2 51 18 30.29851482 Long Bone Fragment High 0 3 47 12 23.74868417 Long Bone Fragment High 2 0 38 17 25.41653005 Long Bone Fragment High 0 0 31 9 16.70329309 Long Bone Fragment High 2 0 16 13 14.4222051 Metacarpal Low 9 0 61 23 37.4566416 Proximal Phalanx Low 2 0 18 4 8.485281374 Proximal Phalanx Low 4 0 31 14 20.83266666 Radius High 0 0 37 18 25.8069758 Rib Low 2 0 41 9 19.20937271 Rib Low 0 0 33 3 9.949874371 Ulna High 3 0 66 17 33.49626845 Table 13 : Raw data for ANCOVA analyses. COMPACT STRONG BROWN Skeletal Element High/low utility? # CM # PM Length (mm) Width (mm) Geometric Mean (=SQRT(L*W) Long Bone Fragment High 4 6 160 36 75.89466384 Long Bone Fragment High 3 0 52 12 24.97999199 Long Bone Fragment High 0 4 49 10 22.13594362 Long Bone Fragment High 0 0 99 11 33 Tibia High 1 0 175 42 85.732141 Rib Low 1 20 10 14.14213562 Rib Low 0 5 38 8 17.43559577 Table 14 : Raw data for ANCOVA analyses.

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109 GRANULAR Skeletal Element High/low utility? # CM # PM Length (mm) Width (mm) Geometric Mean (=SQRT(L*W) Calcaneus Low 0 2 19 22 20.4450483 Long Bone Flake High 2 0 19 11 14.45683229 Long Bone Fragment High 1 0 22 5 10.48808848 Long Bone Fragment High 0 5 11 9 9.949874371 Long Bone Fragment High 0 0 27 5 11.61895004 Long Bone Fragment High 0 2 7 3 4.582575695 Long Bone Fragment High 2 0 17 5 9.219544457 Long Bone Fragment High 0 0 15 4 7.745966692 Long Bone Fragment High 2 0 31 10 17.60681686 Long Bone Fragment High 0 2 12 5 7.745966692 Long Bone Fragment High 0 3 26 14 19.07878403 Long Bone Fragment High 0 2 6 4 4.898979486 Long Bone Fragment High 0 1 15 17 15.96871942 Long Bone Fragment High 0 1 10 4 6.32455532 Long Bone Fragment High 0 1 55 10 23.4520788 Long Bone Fragment High 0 0 17 5 9.219544457 Long Bone Fragment High 0 3 49 8 19.79898987 Long Bone Fragment High 1 0 21 12 15.87450787 Long Bone Fragment High 1 0 19 8 12.32882801 Long Bone Fragment High 0 4 45 15 25.98076211 Long Bone Fragment High 0 1 27 15 20.1246118 Long Bone Fragment High 2 0 42 14 24.24871131 Long Bone Fragment High 0 1 65 17 33.24154028 Long Bone Fragment High 0 1 19 5 9.746794345 Long Bone Fragment High 0 1 22 9 14.07124728 Magnum Low 0 0 42 43 42.49705872 Metacarpal Low 1 2 72 20 37.94733192 Proximal Phalanx Low 0 1 42 18 27.49545417 Rib Low 0 2 15 7 10.24695077 Rib Low 0 1 27 9 15.58845727 Rib Low 0 3 51 10 22.58317958 Rib Low 0 10 61 11 25.90366769 Ulna Low 0 1 98 19 43.15089802 Table 15 : Raw data for ANCOVA analyses.

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110 BLACK MOUSTERIAN Skeletal Element High/low utility? # CM # PM Length (mm) Width (mm) Geometric Mean (=SQRT(L*W) Astragalus Low 0 1 48 32 39.19183588 Cranial Fragment Low 0 6 25 17 20.61552813 Femur High 4 0 33 17 23.68543856 Femur High 0 2 89 23 45.2437841 Femur High 3 0 77 35 51.91338941 Femur High 0 0 66 33 46.66904756 Femur High 1 0 74 22 40.348482 Humerus High 2 1 30 12 18.97366596 Humerus High 1 4 83 29 49.06118629 Humerus High 2 3 71 36 50.55689864 Humerus High 0 3 38 21 28.24889378 Humerus High 0 2 40 28 33.46640106 Humerus High 0 6 48 22 32.49615362 Intermediate Phalanx Low 1 0 42 20 28.98275349 Long Bone Fragment High 1 0 21 8 12.9614814 Long Bone Fragment High 2 0 56 21 34.2928564 Long Bone Fragment High 0 2 24 9 14.69693846 Long Bone Fragment High 0 3 36 13 21.63330765 Long Bone Fragment High 0 3 17 6 10.09950494 Long Bone Fragment High 0 1 45 21 30.7408523 Long Bone Fragment High 0 1 7 6 6.480740698 Long Bone Fragment High 0 3 11 7 8.774964387 Long Bone Fragment High 0 0 16 8 11.3137085 Long Bone Fragment High 1 0 51 19 31.12876483 Long Bone Fragment High 1 1 39 23 29.94995826 Long Bone Fragment High 0 1 54 24 36 Long Bone Fragment High 0 1 43 12 22.71563338 Long Bone Fragment High 0 7 24 5 10.95445115 Long Bone Fragment High 2 0 49 25 35 Long Bone Fragment High 0 0 24 15 18.97366596 Long Bone Fragment High 0 0 53 28 38.52272057 Long Bone Fragment High 0 0 21 12 15.87450787 Long Bone Fragment High 2 0 23 12 16.61324773 Long Bone Fragment High 0 3 27 18 22.04540769 Long Bone Fragment High 1 0 37 8 17.20465053 Long Bone Fragment High 1 0 41 23 30.70830507

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111 BLACK MOUSTERIAN CONTINUED Skeletal Element High/low utility? # CM # PM Length (mm) Width (mm) Geometric Mean (=SQRT(L*W) Long Bone Fragment High 1 0 28 12 18.33030278 Long Bone Fragment High 1 0 27 7 13.74772708 Long Bone Fragment High 0 2 12 10 10.95445115 Long Bone Fragment High 0 2 20 12 15.49193338 Long Bone Fragment High 1 0 18 11 14.07124728 Long Bone Fragment High 0 2 12 5 7.745966692 Long Bone Fragment High 1 0 9 2 4.242640687 Long Bone Fragment High 0 0 37 9 18.24828759 Long Bone Fragment High 1 0 19 7 11.53256259 Long Bone Fragment High 0 1 26 8 14.4222051 Long Bone Fragment High 0 1 17 11 13.67479433 Long Bone Fragment High 0 0 22 9 14.07124728 Long Bone Fragment High 2 0 37 21 27.87471973 Long Bone Fragment High 2 3 31 19 24.2693222 Long Bone Fragment High 0 1 21 8 12.9614814 Long Bone Fragment High 0 7 42 13 23.36664289 Long Bone Fragment High 0 1 34 24 28.56571371 Long Bone Fragment High 1 1 16 4 8 Long Bone Fragment High 2 0 96 29 52.76362383 Long Bone Fragment High 0 2 27 12 18 Long Bone Fragment High 2 0 30 15 21.21320344 Long Bone Fragment High 0 3 41 8 18.11077028 Long Bone Fragment High 2 1 36 9 18 Long Bone Fragment High 3 1 34 8 16.4924225 Long Bone Fragment High 0 1 25 9 15 Long Bone Fragment High 0 1 51 14 26.72077843 Long Bone Fragment High 0 5 29 10 17.02938637 Long Bone Fragment High 0 2 14 18 15.87450787 Long Bone Fragment High 1 0 41 19 27.91057147 Long Bone Fragment High 1 1 36 8 16.97056275 Long Bone Fragment High 1 7 23 5 10.72380529 Long Bone Fragment High 1 0 14 11 12.40967365 Long Bone Fragment High 2 2 31 20 24.8997992 Long Bone Fragment High 3 1 22 11 15.55634919 Long Bone Fragment High 0 4 28 12 18.33030278 Long Bone Fragment High 0 2 30 11 18.16590212 Long Bone Fragment High 2 0 45 17 27.65863337

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112 BLACK MOUSTERIAN CONTINUED Skeletal Element High/low utility? # CM # PM Length (mm) Width (mm) Geometric Mean (=SQRT(L*W) Long Bone Fragment High 0 4 37 28 32.18695388 Long Bone Fragment High 0 2 43 41 41.98809355 Long Bone Fragment High 1 0 32 14 21.16601049 Long Bone Fragment High 1 0 15 10 12.24744871 Long Bone Fragment High 2 0 27 7 13.74772708 Mandible Low 1 0 50 21 32.40370349 Mandible Low 0 1 31 15 21.56385865 Mandible Low 1 1 17 8 11.66190379 Metacarpal High 3 0 44 12 22.97825059 Metapodial High 1 0 71 30 46.15192304 Metapodial High 0 2 42 10 20.49390153 Metatarsal High 1 0 40 13 22.8035085 Metatarsal High 2 0 38 17 25.41653005 Metatarsal High 1 3 111 22 49.4165964 Proximal Phalanx Low 0 0 21 12 15.87450787 Proximal Phalanx Low 2 0 29 19 23.47338919 Proximal Phalanx Low 1 0 45 22 31.46426545 Radioulna High 0 1 20 7 11.83215957 Radius High 1 0 161 20 56.74504384 Radius High 0 1 35 7 15.65247584 Radius High 0 3 52 9 21.63330765 Rib Low 1 0 13 5 8.062257748 Rib Low 0 1 15 5 8.660254038 Rib Low 0 3 53 25 36.40054945 Rib Low 0 3 40 17 26.07680962 Rib Low 0 3 37 27 31.60696126 Rib Low 0 1 28 14 19.79898987 Rib Low 0 3 31 14 20.83266666 Rib Low 0 5 54 7 19.4422221 Tibia Low 1 3 84 47 62.83311229 Tibia Low 0 3 58 19 33.19638535 Table 16 : Raw data for ANCOVA analyses.