# Multivariate assessments of activity-related skeletal changes: Interpreting Bell Beaker specialized male archery and social organization in Central Europe

J Ryan-despraz, Sébastien Villotte, Jocelyne Desideri, M Besse

► To cite this version:

J Ryan-despraz, Sébastien Villotte, Jocelyne Desideri, M Besse. Multivariate assessments of activity-related skeletal changes: Interpreting Bell Beaker specialized male archery and social organization in Central Europe. American Journal of Biological Anthropology, 2023, 182, pp.237 - 263. 10.1002/ajpa.24817. hal-04228855

HAL Id: hal-04228855

https://hal.science/hal-04228855v1

Submitted on 4 Oct 2023

HAL is a multi-disciplinary open access
archive for the deposit and dissemination of sci-
entific research documents, whether they are pub-
lished or not. The documents may come from
teaching and research institutions in France or
abroad, or from public or private research centers.

L'archive ouverte pluridisciplinaire HAL, est
destinée au dépôt et à la diffusion de documents
scientifiques de niveau recherche, publiés ou non,
émanant des établissements d'enseignement et de
recherche français ou étrangers, des laboratoires
publics ou privés.

<figure><img src="image_1.png" /></figure>

Distributed under a Creative Commons CC BY 4.0 - Attribution - International License
## RESEARCH ARTICLE

AMERICAN JOURNAL OF
BIOLOGICAL ANTHROPOLOGY
The official journal of the American Association of Biological Anthropologists

WILEY

# Multivariate assessments of activity-related skeletal changes: Interpreting Bell Beaker specialized male archery and social organization in Central Europe

J. Ryan-Despraz<fnref n="1" /> ID | S. Villotte<fnref n="2" />,<fnref n="3" />,<fnref n="4" /> ID | J. Desideri<fnref n="5" /> ID | M. Besse<fnref n="1" /> ID

### Correspondence

J. Ryan-Despraz, Laboratory of Prehistoric Archaeology and Anthropology, Department F.-A. Forel for Environmental and Aquatic Sciences, University of Geneva, Switzerland.
Email: jessica.ryan@unibe.ch

### Funding information

Schweizerischer Nationalfonds zur Förderung der Wissenschaftlichen Forschung, Grant/Award Number: 175169; Université de Genève

## Abstract

**Objectives:** The Bell Beaker period witnessed the rise of individual inhumations with “wealthy” burial contexts containing archery-related grave goods, leading archaeologists to label the individuals in these tombs as “archers.” This study looks to (1) compare the skeletons from male “archer” burials with those from male “non-archer” burials—those not having archery-related grave goods—in order to assess a possible link between burial context and physical activity, and (2) apply a biomechanics profile to evaluate whether the individuals associated with these “archer” burials practiced specialized archer activity.

**Materials and Methods:** The corpus (males only) included 46 “archers” and 40 “non-archers” from Bell Beaker individual inhumations. Osteological data included measurements, scores of entheseal changes, and a diagnosis of certain pathologies. Data analyses involved visual observations, hypothesis tests, dimension reduction, and MANOVA, with approaches aimed at exploring the treatment of data missingness.

**Results:** Measurement data revealed no differences between the two groups. Evaluations of entheseal changes found that “non-archers” had consistently more instances of bone surface modifications than “archers.” Individual assessments of specialized archer occupation identified 11 possible specialized archers.

**Discussion:** These findings indicate a possible labor differentiation represented through the presence of a probably prestigious “archer” burial context. This suggests a link between grave good presence and labor, but not between a Bell Beaker archery occupation and an “archer” burial context. Data analyses support the application of biomechanics to osteological analyses in order to assess specialized activity on the skeleton.

## KEYWORDS

archery, Bell Beaker, biomechanics, entheseal changes

<footnote n="1">¹Laboratory of Prehistoric Archaeology and Anthropology, Department F.-A. Forel for Environmental and Aquatic Sciences, University of Geneva, Geneva, Switzerland</footnote>

<footnote n="2">²UMR7206 Éco-Anthropologie, CNRS, MNHN, Université Paris Cité. Musée de l'Homme, Paris, France</footnote>

<footnote n="3">³Quaternary environments & Humans, OD Earth and History of life, Royal Belgian Institute of Natural Sciences, Brussels, Belgium</footnote>

<footnote n="4">⁴Unité de Recherches Art, Archéologie Patrimoine, Université de Liège, Liège, Belgium</footnote>

<footnote n="5">⁵Laboratory of Archaeology of Africa and Anthropology, Section of Biology, University of Geneva, Geneva, Switzerland</footnote>
# 1 | INTRODUCTION

The Bell Beaker period (third millennium BCE) acts as a transition from the end of the Neolithic to the beginning of the Bronze Age in Europe and North Africa (Besse, 2014; Furholt, 2020; Gallay, 1979; Harrison, 1980; Müller & van Willigen, 2001; Nicolis, 2001). This period, so-called for its distinctive pottery, presents a gradual eastward expansion over a vast geographic area while maintaining a largely uniform material culture, in particular with regard to ceramics (Figure 1) (Bailly & Salanova, 1999; Czebreszuk, 2014; Salanova, 2016). These findings originally signaled the presence of a large migration of the Bell Beaker peoples; however, archaeological analyses have found subtle regional differences as well as evidence for local material production (Besse, 2003, 2014, 2015; Czebreszuk, 2014; Harrison, 1980). This evidence for a prominent local tradition combined with the vast geographic reach of the Bell Beakers has raised the question as to how these cultural traditions spread, even leading many archaeologists to refer to this period as a “phenomenon” (Besse, 2014; Czebreszuk, 2014; Lemercier, 1998). Recent analyses

examining aDNA have found clear genetic differences between the Bell Beaker peoples of the West and those of the East (modern-day Central Europe), indicating that material similarities were not the result of human migration but rather of ideas (Olalde et al., 2018). At the same time, Bell Beaker genome studies in Britain identified telling similarities with samples from Central Europe, showing that, at least for part of the expansion, migration did play a role (Olalde et al., 2018; Reich, 2019). This complex relationship between migration patterns, local production, and a vast geographic area at such a pivotal time of European prehistory has inspired researchers to explore the daily lives and functioning of the Bell Beaker peoples.

## 1.1 | Evidence for Bell Beaker social organization

Wealth inequality is a strong indicator for social stratification, and, in archaeology, this is commonly illustrated through funerary traditions, in particular grave goods and tomb architecture. Throughout the Bell Beaker period, burials exhibit skewed distributions of wealth between

<figure><img src="image_2.png" /><figcaption>FIGURE 1 A distribution map showing the spread and regional variation of bell beakers throughout Europe, from Besse (2014), Figure 1).</figcaption></figure>
individuals, both in terms of item quantity and quality (e.g., the Bell Beaker cemeteries of Hoštice, Pavlov, and Hulín) (Binford, 1972; Chapman, 2015; Drozdová et al., 2011; Heyd, 2001, 2007; Ryan-Despraz, 2022). Many such goods, for example ornaments and gold decoration, are purely symbolic and show evidence of trade, a practice strongly linked to a centralized economy reminiscent of a complex social organization (Abegg et al., 2022). In addition to grave goods, Bell Beaker monumental tombs also exist, such as the dolmens of Sion “Petit-Chasseur” (Switzerland) and “Saint-Martin-de-Corléans” (Italy), though they are rare (Bocksberger, 1976; Corboud, 2009; De Gattis et al., 2018). Most monumental tombs were only for select individuals; however, their construction would still have required the coordination of multiple people—a level of organization consistent with the presence of leadership roles. Isotopic and genetic analyses also contribute to the assumption of Bell Beaker social stratification. Numerous studies have found evidence for a reduced male geographic mobility (i.e., male sedentarism) together with high maternal genetic diversity, indicating a level of female exogamy demonstrative of patriarchal societies (Fitzpatrick, 2011; Grupe et al., 1997; Sjögren et al., 2019). Lastly, it is worthwhile to consider child burials presenting numerous and “rich” burial items (e.g., ornamental objects, especially those made of metal or carved stone). Children could not yet have held specialized roles in a society, such as artisan or warrior; however, the Bell Beaker period witnessed the presence of “wealthy” child burials, including some instances where these were the most prominent in the cemetery (e.g., Popůvky (Czechia)) (Bálek et al., 1999; Bedáňová, 2023; Heyd, 2000; Matějičková, 2007; Turek & Černý, 2001). Many of these cases also contained archery-related items, and these “prestigious” burials could indicate the presence of important traditions linked to inheritance (Bedáňová, 2023; Sjögren et al., 2019).

## 1.2 | Bell Beaker funerary tradition

Investigations classify two general Bell Beaker funerary traditions in continental Europe: the Western Complex (modern-day Western Europe—e.g., France, Spain, and Portugal) characterized most commonly by the presence of collective burials in tombs erected by preceding cultures, and the Eastern Complex (modern-day Central Europe—e.g., Czechia, Hungary, and Poland) marked by the widespread presence of individual inhumations (Heyd, 2001; Müller, 2001; Vander Linden, 2015). However, it is important to note that this classification represents trends rather than a strict dichotomy between funerary complexes; an undefined border and multiple exceptions attest to the complexity of the overall funerary traditions (Lemercier et al., 2022; Salanova, 2022; Tchérémissinoff et al., 2022).

During this time, stone wristguards associated with archery began to appear, most notably in select individual burials of the Eastern Complex, and this has often led archaeologists to interpret these graves as having belonged to archers (Heyd, 2007; Turek, 2017). Indeed, combined with the presence of arrowheads, bow-shaped pendants, and artistic depictions, the Bell Beaker period saw a rise in archery representations, which has prompted hypotheses asserting an

archery prominence in social functioning and organization (Corboud, 2009; Nicolas, 2019, 2020; Ryan-Despraz, 2022; Ryan-Despraz & Nicolas, 2022). Additionally, “archer” burials tend to be more wealthy, in terms of both grave good quality and item quantity, while also being relatively uncommon (Fitzpatrick, 2011; Heyd, 2007). The question then arises as to who were these “archers” and what their role could have been in Bell Beaker society.

## 1.3 | Activity-related skeletal changes

Since the beginning of the 20th century, biological anthropologists have attempted to study daily life and behavior through analyses of activity-related skeletal changes (Capasso et al., 1999; Kennedy, 1998; Larsen, 1997; Merbs, 1983; Villotte, 2023). The principle is based on the idea that bone can adapt, or sometimes alter pathologically, to its external environment, for example to biomechanical constraints (Acosta et al., 2017; Deymier et al., 2019; Nikita et al., 2019; Weiss, 2003; Wolff, 1892). The form and morphology of entheses, which are the insertions on bone for tendons, ligaments, and articular capsules, could reflect such changes (Benjamin et al., 2002, 2004; Benjamin & McGonagle, 2001), and modifications at these points, enthesal changes (EC), are commonly linked to human biomechanics (Cashmore & Zakrzewski, 2013; Dutour, 1992; Havelková et al., 2011; Pálfi & Dutour, 1996; Perreard Lopreno, 2007; Schrader, 2019). In addition to EC, other skeletal changes used to identify activity and behavior have included degenerative joint disease (DJD) and diaphyseal dimensions and geometry (Bridges, 1991; Hawkey & Merbs, 1995; Jones et al., 1977; Jurmain et al., 2012; Rhodes & Knüsel, 2005; Ruff et al., 1994; Walker & Holliman, 1989). Like EC, DJD is also linked to mechanics as the degeneration of the articular cartilage leads to direct bone-on-bone contact through movement, although other important factors such as age and genetics could exacerbate the issue (Chen et al., 2017; Goodman & Martin, 2002; Rogers & Waldron, 1995; Salter, 2002; White, 2000). While the etiologies for DJD are numerous, its appearance in younger individuals is more commonly associated with biomechanics, especially when expressed asymmetrically (Capasso et al., 1999; Mann & Hunt, 2005; Merbs, 1983; Schwartz, 2007; Tainter, 1980). Lastly, metric analyses can also be useful for assessing biomechanical loading. Traditionally, this has specifically included cross-sectional geometry (Becker, 2019; Cowgill, 2018; Honda et al., 2015; Lieberman et al., 2004; Macintosh et al., 2013; Maggiano et al., 2008; Niinimäki et al., 2016; Nikita et al., 2011; Ogilvie & Hilton, 2011), however external measurements could also serve as a cheaper proxy to such techniques. For example, the calculation of bone dimensions (e.g., a “robusticity index”) and overall size comparisons could provide developmental insight when comparing populations.

## 1.4 | Objectives

This study applied non-destructive, macro-observations of external measurements, EC, and DJD to assess a collection of Bell Beaker
individuals, organized into two groups based on the presence of archery-related grave goods, for indications of physical activity in general and of specialized archery in particular. While ideally this study would have included both males and females, a lack of female “archer” burials necessitated a males-only focus. The goals of this study were two-fold: (1) to observe any morphological differences between the two groups, which could draw a cultural link between burial context and manual activity, and (2) to test the theory that “archer” burials belonged to specialized archers. This study will additionally evaluate the dependability of activity determinations from the skeleton as well as how data analyses could help improve the reliability of interpretations.

While this study focuses on “archer” burials with respect to daily life and social organization, assessments as to the function of archery, such as for hunting or warfare, are outside the scope of this study; however, related works provide a thorough discussion of this question (see Ryan-Despraz, 2021, 2022). This study also controls for a possible link between archaeological identity (“archer” burial) and biological identity (archer occupation). While “non-archer” burials could have also belonged to specialized archers, this study relies on the archaeological context to drive the hypothesis of occupation. For this reason, individual analyses of specialization do not include “non-archers”.

# 2 | MATERIALS AND METHODS

## 2.1 | Skeletal collection

This study examined 86 male skeletons from individual inhumations of the Bell Beaker Eastern Complex (modern-day Central Europe), specifically the regions of Lower and Upper Austria, Bavaria (Germany), Bohemia (Czechia), and Moravia (Czechia) (Supplementary Information Figures S1–S3). This focus on individual burials was essential because of the direct relationship between the individual and the burial context. The decision to group individuals from such a large geographic area was based on three reasons: (1) Bell Beaker attribution—archaeological analyses all confidently classified these individuals as “Bell Beaker”, (2) distribution—many inhumations are single finds not belonging to a cemetery, and (3) adequate sample size—“archer” burials are not prevalent and this study requires a level of bone surface preservation not common in prehistoric collections, therefore the research required an expanded geographic scope in order to ensure an adequate sample size.

The skeletal sample was organized into two groups: “archers” (group A, *n* = 46) (Table 1) and “non-archers” (group N, *n* = 40) (Table 2). The “archer” burials contained at least one object related to archery (i.e., stone wristguard, arrowhead, or bow-shaped pendant) (Figure 2), whereas the “non-archers” did not have any goods associated with archery.

## 2.2 | Sex, age, and measurements

This study controlled each skeleton for sex and age. The primary method of sex determination was DSP 2 (Brůžek et al., 2017), which

uses the discriminant analysis principle to compare pelvis measurements to an established worldwide reference sample. When preservation did not allow for DSP 2 analyses, additional methods involved morphological observations of the coxal bone, skull, and burial orientation, two of which were required for a sex determination (Acsádi & Nemeskéri, 1970; Bruzek, 2002; Brůžek, 1991; Buikstra & Ubelaker, 1994; Murail et al., 2005; Stevenson et al., 2009; Walker, 2008). Burial orientation for the Bell Beaker period is exceedingly consistent and reliable, especially for male burials; indeed, a study by Müller (2001) only found inconsistencies between biological sex and archaeological grave orientation 4.6–5.6% of the time (Heyd, 2001; Vander Linden, 2015). This study was not able to determine the sex of one group A individual, Lochenice 3, and they were therefore not included in the population analyses. However, this individual's seemingly “rich” burial context (i.e., multiple items, with many examples of carved stone) remained of interest, therefore the skeleton was still included in individual analyses for specialized archer activity.

Age determinations included four methods adapted to specific age groups. For adolescents and young adults, fusion stages were classified for the ischial tuberosity (Coqueugniot & Weaver, 2007) as well as for the sternal epiphysis of the clavicle, annular epiphysis of the thoracic vertebrae, annular epiphysis of the lumbar vertebrae, the iliac crest, and the inferior angle of the scapula (Schaefer et al., 2009). For individuals >35 years, evaluations of the sacro-pelvic iliac surface, including development scores for the auricular surface and modifications of the apical surface and iliac tuberosity, were implemented (Schmitt, 2001, 2005). Lastly, the Brooks and Suchey (1990) method for the os pubis served to classify individuals into one of six age groups throughout adulthood. As a final verification, individuals demonstrating tooth wear consistent with advanced aging were excluded (Faillace et al., 2017; Mays, 1998, 2002). Any individual for whom age was not confidently <60 years was not included in the study.

An upper and lower age limit was necessary for two main reasons: (1) studies have shown that both EC and DJD appear much more commonly with age (Alves Cardoso & Henderson, 2010; Jurmain et al., 2012; Pearson & Lieberman, 2004), and (2) children could not yet have been “specialized” in terms of physical development. For these reasons, this study implemented an age range of ~15 to 60 years. However, analyses do not differentiate between age groups due to an already relatively low sample size and the fact that determinations are often quite broad.

When preservation allowed, this study additionally gathered 56 measurements (per side) on the clavicle (*n* = 4), scapula (*n* = 9), humerus (*n* = 12), radius (*n* = 6), ulna (*n* = 6), femur (*n* = 11), tibia (*n* = 6), and fibula (*n* = 2) and calculated two bone proportion index ratios. The two ratios, calculated from the measurements, include

Type 1:

$$ \frac{\text{Right} - \text{Left}}{\text{Average}} $$

The resulting value then further provides indications for left-versus right-handed bone-size dominance trends.
<table><caption>TABLE 1 The suspected “archers” (Group A) used in the study.</caption><thead><tr><th>Site</th><th>Region</th><th>Grave/Inventory Number</th><th>Sex</th><th>Age</th><th>Grave context</th></tr></thead><tbody><tr><td>Altenmarkt</td><td>Bavaria (DE)</td><td>gr. 6, obj. 25</td><td>Male<sup>d</sup></td><td>30–60</td><td>Wristguard (left), arrowheads, dagger, bell beaker, hematite, flint, and boar’s tooth (Schmotz, 1990)</td></tr><tr><td>Brandýsek</td><td>Bohemia (CZ)</td><td>71</td><td>Male</td><td>&lt;60</td><td>Four flint arrowheads, copper dagger, bowl, jug, and two flint fragments (Kytlicová, 1960; Hájek, 1968a, 1968b)</td></tr><tr><td>Franzhausen</td><td>Lower Austria (AT)</td><td>230</td><td>Male</td><td>30–40</td><td>Wristguard, spherical cup, bowl, bronze, copper, and pearl rings/beads near the neck (Neugebauer &amp; Gattringer, 1982)</td></tr><tr><td>Gemeinlebarn</td><td>Lower Austria (AT)</td><td>2071</td><td>Male<sup>d</sup></td><td>30–50</td><td>Wristguard, two bell beakers, cup, handled pot, copper awl, dagger, and two metal spiral rings (Neugebauer &amp; Neugebauer, 1997; Neugebauer &amp; Neugebauer-Maresch, 2001)</td></tr><tr><td>Hoštice 1, za Hanou</td><td>Moravia (CZ)</td><td>821</td><td>Male<sup>d</sup></td><td>20–27</td><td>Bow-shaped pendant, pitcher, bowl, small container, worked stone tip, animal bones (domesticated), and worked bone (Drozdová et al., 2011; Růžičková, 2008, 2009)</td></tr><tr><td>Hoštice 1, za Hanou</td><td>Moravia (CZ)</td><td>862</td><td>Male<sup>d</sup></td><td>30–50</td><td>Wristguard, copper dagger, bell beaker, pitcher, bowl, bone button (unsure if v-shaped), and animal bones (Olivik, 2009a, 2009b; Drozdová et al., 2011)</td></tr><tr><td>Hoštice 1, za Hanou</td><td>Moravia (CZ)</td><td>863</td><td>Male<sup>d</sup></td><td>30–50</td><td>Seven arrowheads, gold and silver fragments, and animal bones (Drozdová et al., 2011; Olivik, 2009a, 2009b; Peška, 2013a; Sosna, 2012)</td></tr><tr><td>Hoštice 1, za Hanou</td><td>Moravia (CZ)</td><td>864</td><td>Male<sup>d</sup></td><td>30–50</td><td>Bow-shaped pendant, four arrowheads, bell beaker, pitcher, and animal bones (domesticated) (Drozdová et al., 2011; Olivik, 2009a, 2009b; Růžičková, 2009; Sosna, 2012)</td></tr><tr><td>Hoštice 1, za Hanou</td><td>Moravia (CZ)</td><td>884</td><td>Male<sup>d</sup></td><td>&lt;50</td><td>Wristguard, copper dagger, three pitchers, bowl, and animal bones (Drozdová et al., 2011; Olivik, 2009a, 2009b)</td></tr><tr><td>Hoštice 1, za Hanou</td><td>Moravia (CZ)</td><td>915</td><td>Male<sup>d</sup></td><td>&lt;50</td><td>Wristguard, arrowheads, copper dagger, two bell beakers, lithic blade, polishing stone, and animal bones (domestic) (Olivik et al., 2009a, 2009b; Drozdová et al., 2011; Sosna, 2012)</td></tr><tr><td>Hoštice 1, za Hanou</td><td>Moravia (CZ)</td><td>949</td><td>Male</td><td>30–60</td><td>Wristguard, four arrowheads, two pitchers, bowl, two silver spirals, three gold spirals, bone object (ring?), animal bones (domestic) (Olivik et al., 2009a, 2009b; Drozdová et al., 2011; Sosna, 2012; Peška, 2013a)</td></tr><tr><td>Hulín1</td><td>Moravia (CZ)</td><td>66</td><td>Male<sup>d</sup></td><td>&lt;60</td><td>Wristguard, boar tusk (Peška, 2013b)</td></tr><tr><td>Hulín1</td><td>Moravia (CZ)</td><td>73</td><td>Male<sup>d</sup></td><td>&lt;60</td><td>Two arrowheads, anvils, hammers (smith equipment), bowl, and decorated bell beakers (Peška, 2013b)</td></tr><tr><td>Hulín1</td><td>Moravia (CZ)</td><td>85</td><td>Male<sup>d</sup></td><td>30–50</td><td>Wristguard (Peška, 2013b)</td></tr><tr><td>Ivanovice</td><td>Moravia (CZ)</td><td>812</td><td>Male</td><td>&lt;60</td><td>Wristguard, two arrowheads, copper dagger, two mugs, and bowl with animal bones (Tkáč, 2008)</td></tr><tr><td>Ivanovice</td><td>Moravia (CZ)</td><td>814</td><td>Male</td><td>&lt;60</td><td>Arrowheads (Tkáč, 2008)</td></tr><tr><td>Landau (Dingolfing, 1981)</td><td>Bavaria (DE)</td><td>2118/24, 22/1193</td><td>Male</td><td>&gt;19</td><td>Wristguard, two arrowheads, daggers, and amber pearls (Heyd, 2000)</td></tr><tr><td>Ledce II</td><td>Moravia (CZ)</td><td>1/52</td><td>Male<sup>d</sup></td><td>&lt;60</td><td>Wristguard (left association), stone arrowhead, two decorated beakers, boar tusk, bone clasp, and bone object (Dvořák, 1992)</td></tr><tr><td>Lhánice</td><td>Moravia (CZ)</td><td>8</td><td>Male<sup>d</sup></td><td>&lt;60</td><td>Wristguard, copper dagger, cup, pitcher (Moucha, 2005)</td></tr><tr><td>Lochenice</td><td>Bohemia (CZ)</td><td>3</td><td>ND<sup>a</sup></td><td>30–40</td><td>Two arrowheads, quartzite blade, bowl, jug, bone pendant, and flint fragments (Hájek, 1968a, 1968b)</td></tr><tr><td>Oberstimm II/1982</td><td>Bavaria (DE)</td><td>2</td><td>Male</td><td>&lt;60</td><td>Wristguard, arrowheads, copper dagger, lithics, bowl, and bell beaker (Rieder, 1983)</td></tr></tbody></table>

(Continues)
<table><caption>TABLE 1 (Continued)</caption><thead><tr><th>Site</th><th>Region</th><th>Grave/Inventory Number</th><th>Sex</th><th>Age</th><th>Grave context</th></tr></thead><tbody><tr><td>Osterhofen-Altenmarkt</td><td>Bavaria (DE)</td><td>2</td><td>Male<sup>d</sup></td><td>30–50</td><td>Five wristguards, three arrowheads, three daggers, four small cups, one large cup, and flint (Schmotz, 1994)</td></tr><tr><td>Osterhofen-Altenmarkt</td><td>Bavaria (DE)</td><td>10</td><td>Male</td><td>&lt;60</td><td>One wristguard, bow-shaped pendant fragments, two copper objects, several small pots, and five bowls (Schmotz, 1994)</td></tr><tr><td>Osterhofen-Altenmarkt</td><td>Bavaria (DE)</td><td>11</td><td>Male<sup>d</sup></td><td>&lt;60</td><td>Arrowhead, copper objects, decorated bow-shaped pendants, flint fragments, and nine cups (Kern, 2016a; Schmotz, 1994)</td></tr><tr><td>Ostopovice</td><td>Moravia (CZ)</td><td>14</td><td>Male<sup>d</sup></td><td>30–60</td><td>Five arrowheads, four-footed bowl, small jug, and animal bones (Dvořák, 1992)</td></tr><tr><td>Pavlov</td><td>Bohemia (CZ)</td><td>500</td><td>Male</td><td>17–21</td><td>Two bow-shaped pendants (boar's tusk), copper dagger, jug, and pottery shard (Dvořák et al., 1996)</td></tr><tr><td>Pavlov</td><td>Bohemia (CZ)</td><td>502</td><td>Male</td><td>16–34</td><td>Bow-shaped pendant (boar's tooth), two arrowheads, jug, bowl, and animal bones (Dvořák et al., 1996)</td></tr><tr><td>Pavlov</td><td>Bohemia (CZ)</td><td>519</td><td>Male</td><td>15–20</td><td>Wristguard, arrowheads, bowl and jug fragments, and animal bones (Dvořák et al., 1996)</td></tr><tr><td>Praha 8-Kobylisy</td><td>Bohemia (CZ)</td><td>36,749</td><td>Male<sup>d</sup></td><td>&lt;50</td><td>Arrowhead, bow-shaped pendant, copper dagger, and ceramics (Fridrichová et al., 1995; Soudský, 1954)</td></tr><tr><td>Radovesice</td><td>Bohemia (CZ)</td><td>53/80-I</td><td>Male<sup>d</sup></td><td>20–29</td><td>Found in the same burial group as cremated child 53/80-II and grave goods more associated with child than this adult male; however they include wristguard, arrowheads, bow-shaped pendant, bell beaker, jug, ceramic fragments, v-shaped buttons<sup>b</sup> (Muška, 1981; Turek & Černý, 2001); Turek, 2004, 2006; Shbat, 2013)</td></tr><tr><td>Radovesice</td><td>Bohemia (CZ)</td><td>59/80-II</td><td>Male</td><td>20–29</td><td>Found in the same burial group as cremated child 53/80-II; grave goods more associated with child than this adult male; however, they include wristguard, bow-shaped pendant, arrowheads, and v-shaped buttons<sup>c</sup> (Muška, 1981; Shbat, 2013)</td></tr><tr><td>Radovesice</td><td>Bohemia (CZ)</td><td>116/78</td><td>Male</td><td>&lt;34</td><td>Wristguard, arrowhead, copper dagger, bell beaker, jug, antler, boar's tooth, lithics, beaver tooth, and flint fragments (Muška, 1981; Shbat, 2013; Turek, 2003)</td></tr><tr><td>Rosnice</td><td>Bohemia (CZ)</td><td>1/59</td><td>Male<sup>d</sup></td><td>19–34</td><td>Wristguard (left association), copper dagger, flint dagger, bow-shaped pendant, bell beaker, small jug, and copper ring (Hájek, 1968a, 1968b; Vokolek, 1965)</td></tr><tr><td>Rousínov</td><td>Bohemia (CZ)</td><td>19/2</td><td>Male</td><td>20–29</td><td>Double burial, second individual a male child; wristguard, arrowheads, sharpening stone, bow-shaped pendant, bell beaker, bowl, cup, and flint (Geisler, 1990)</td></tr><tr><td>Stehelčeves</td><td>Bohemia (CZ)</td><td>1</td><td>Male</td><td>&lt;60</td><td>Two wristguards, two bell beakers, bell beaker shards, jug, copper dagger, copper awl, boar tooth fragment, grindstone, stone anvil, and stone hammer (Hájek, 1966, Hájek, 1968a, 1968b)</td></tr><tr><td>Stehelčeves</td><td>Bohemia (CZ)</td><td>2</td><td>Male</td><td>&lt;60</td><td>Wristguard, stone arrowheads, three bell beakers, copper dagger, metal hammer, flint scraper, antler ax, and flint bevels (Hájek, 1966, Hájek, 1968a, 1968b)</td></tr><tr><td>Straubing Stadttäcker</td><td>Bavaria (DE)</td><td>1</td><td>Male</td><td>30–50</td><td>Wristguard, bow-shaped pendant, and cup (Christlein, 1982; Prammer, 1981)</td></tr><tr><td>Sulejovice</td><td>Bohemia (CZ)</td><td>3</td><td>Male</td><td>&lt;60</td><td>Wristguard, three arrowheads, two bell beakers, and two jugs, and chunk of amber (Hájek, 1962; Hájek, 1968a, 1968b)</td></tr><tr><td>Tödling</td><td>Upper Austria (AT)</td><td>4</td><td>Male</td><td>17–20</td><td>Bowl with animal bones (pig), dagger tip, three bow-shaped pendants, bit (horse) cross-section, orientation ENE-WSW (Kern, 2016b; Kern & Wiltschke-Schrotta, in press)</td></tr></tbody></table>
<table><caption>TABLE 1 (Continued)</caption><thead><tr><th>Site</th><th>Region</th><th>Grave/Inventory Number</th><th>Sex</th><th>Age</th><th>Grave context</th></tr></thead><tbody><tr><td>Tödling</td><td>Upper Austria (AT)</td><td>5</td><td>Male</td><td>20-35</td><td>Two handled bowls, five bow-shaped pendants, flint (blade fragment?), pebble (worked), orientation ENE-WSW (Kern and Wiltschke-Schrotta, in press)</td></tr><tr><td>Trieching</td><td>Bavaria (DE)</td><td>670, 22/1193</td><td>Male</td><td>30-50</td><td>Excavation grave 1; wristguard, arrowheads, lithic scraper, copper dagger, amber bead, and beaker (Kreiner, 1991)</td></tr><tr><td>Velké Přilepy</td><td>Bohemia (CZ)</td><td>2</td><td>Male</td><td>16-24</td><td>Flint arrowhead, bowl, and pitcher (Skružný et al., 2000)</td></tr><tr><td>Veselí na Moravě</td><td>Moravia (CZ)</td><td>XVII, 158</td><td>Male<sup>d</sup></td><td>16-24</td><td>Wristguard (left), jug, bowl, pebbles, and boar's tooth (Staňa, 1960)</td></tr><tr><td>Vykáň</td><td>Bohemia (CZ)</td><td>1</td><td>Male<sup>d</sup></td><td>&lt;60</td><td>Wristguard, bow-shaped pendant, 2 jugs, bowl, flint dagger, pot, and leg device (Hájek, 1968a, 1968b)</td></tr><tr><td>Weichering</td><td>Bavaria (DE)</td><td>17</td><td>Male<sup>d</sup></td><td>30-60</td><td>Bow-shaped pendant, cup, bowl, and ceramic fragments (Weinig, 1992)</td></tr><tr><td>Žabovřesky</td><td>Bohemia (CZ)</td><td>32,293</td><td>Male</td><td>&lt;60</td><td>Wristguard, bell beaker, pot, jug, and bowl (Blajerová, 1962; Zápotocký, 1962)</td></tr></tbody></table>

Abbreviation: ND, not determined.

<sup>a</sup>The biological sex determinations were not consistent. Even with the DSP2 method, one os coxal identified as male and the other as female. The Walker method revealed rather female characteristics, but the burial orientation was male (lying on the left side).

<sup>b</sup>The grave goods are seemingly more associated with the child burial. Therefore this individual's classification as A is debatable; however because the general burial context still included archery-related items, 53/80-I maintains an A status.

<sup>c</sup>As with 53/80-II, the grave goods are seemingly more associated with the child burial, therefore this individual's classification as A is debatable.

<sup>d</sup>A sex determination using the DSP 2 method.

Type 2:

$$ \frac{\text{Circumference}}{\text{Maximum Length}} $$

This ratio could allow for a comparison of midshaft “robusticity” differences between the two groups.

Tables 3 and 4 provide a complete list of the measurements and ratios, all of which follow the criteria established in Martin and Sal-ler (1957).

## 2.3 | Biomechanics

This study examined each individual with respect to a biomechanics profile. This profile involved a literature review and had three main objectives: (1) determine the precise movements and corresponding muscles activated in archery, (2) examine modern archery reports for common injuries, and (3) assess previous anthropological research on the skeletons of suspected archers. Establishing the biomechanics and common injuries in archery comprised organizing the findings of multiple sources into one complete overview of archery science. This overview identified 39 primary muscles activated during archery, and these muscles served as the basis of this study's assessment of EC. Table 5 lists these muscles, their side distribution (draw arm, bow arm, or both arms), and the corresponding source. The literature review for injuries found that they most commonly occur from over-use and are located on the draw arm shoulder, with prominent secondary locations at the elbow (Table 6). Lastly, this study is not the

first to examine specialized archery in the archaeological record (though it is the first for the Bell Beaker period); therefore, findings from seven previous studies were also considered. Primary observations from those works include the presence of os acromiale (Stirland, 2013), DJD of the glenoid cavity and humeroradial joint (Merbs, 1983; Ryan et al., 2018), larger upper limb measurements (Stirland, 2013; Thomas, 2014), overall greater levels of EC in archer populations when compared with non-archer populations (Ryan et al., 2018; Thomas, 2014; Tihanyi et al., 2015), and upper limb asymmetries in likely archers (Dutour, 1986; Molnar, 2006; Ryan et al., 2018; Stirland, 2013; Thomas, 2014; Tihanyi et al., 2015).

## 2.4 | Analyses of enthesal changes (EC)

The entheses chosen for evaluation were those linked to the muscles established by the biomechanics review. EC analyses included four different methods according to the type of enthesis (mainly, fibrous or fibrocartilaginous), with each enthesis matched to the method best able to encapsulate its surface appearance. Those methods included the Coimbra method, the Mariotti et al. method, the Villotte method, and an absence/presence method (Henderson et al., 2013, 2016; Mariotti et al., 2004, 2007; Villotte, 2006). The Coimbra method requires eight different scores for a single enthesis as it differentiates between the contour and the center. The Mariotti et al. method provides a score based on whether an enthesis presents signs of erosion, proliferation, or robusticity. This study used Villotte's Group 2 method for one enthesis, which scores based on the absence or presence of enthesophytes or an erosion surface. Lastly, the absence/presence
<table><caption>TABLE 2 The “non-archers” (Group N) used in the study.</caption><thead><tr><th>Site</th><th>Region</th><th>Grave/inventory number</th><th>Sex</th><th>Age</th><th>Grave context</th></tr></thead><tbody><tr><td>Augsburg, Uni-Süd 1991</td><td>Bavaria (DE)</td><td>5</td><td>Male<sup>a</sup></td><td>20-35</td><td>No archery context<sup>b</sup> (Kociumaka &amp; Dietrich, 1992)</td></tr><tr><td>Brandýsek</td><td>Bohemia (CZ)</td><td>18</td><td>Male<sup>a</sup></td><td>16-24</td><td>Bowl (Kytlicová, 1960; Hájek, 1968a, 1968b)</td></tr><tr><td>Brandýsek</td><td>Bohemia (CZ)</td><td>26</td><td>Male</td><td>&lt;50</td><td>Bowl, bone pendant (Kytlicová, 1960; Hájek, 1968a, 1968b)</td></tr><tr><td>Brno-Juliánov</td><td>Moravia (CZ)</td><td>1/67, 242</td><td>Male</td><td>&lt;60</td><td>Jug (Dvořák, 1992; Peškař, 1968; Stloukal, 1968)</td></tr><tr><td>Franzhausen II</td><td>Lower Austria (AT)</td><td>3382</td><td>Male</td><td>30-50</td><td>2 cups, oriented N-S facing East, surrounded by circular ditch (Kern, 2011)</td></tr><tr><td>Gemeinlebarn</td><td>Lower Austria (AT)</td><td>3559</td><td>Male</td><td>30-50</td><td>Beaker, two jugs, and two metal rings (Neugebauer &amp; Neugebauer, 1997; Neugebauer &amp; Neugebauer-Maresch, 2001)</td></tr><tr><td>Henzing</td><td>Lower Austria (AT)</td><td>3</td><td>Male<sup>a</sup></td><td>15-20</td><td>Grave group with Henzing grave 1, 2, animal bones, oriented N-S facing east (Friesinger, 1976; Jungwirth, 1976)</td></tr><tr><td>Holubice IV</td><td>Bohemia (CZ)</td><td>7</td><td>Male</td><td>&lt;50</td><td>Bell beaker (Rakovský, 1985)</td></tr><tr><td>Hoštice 1, za Hanou</td><td>Moravia (CZ)</td><td>818</td><td>Male</td><td>&lt;50</td><td>Ceramics, fragments of a copper plate (Drozdová et al., 2011)</td></tr><tr><td>Hoštice 1, za Hanou</td><td>Moravia (CZ)</td><td>859</td><td>Male<sup>a</sup></td><td>&lt;60</td><td>Ceramics (Drozdová et al., 2011)</td></tr><tr><td>Hoštice 1, za Hanou</td><td>Moravia (CZ)</td><td>931</td><td>Male</td><td>&lt;50</td><td>Ceramics, stone tool, and animal bones (Drozdová et al., 2011)</td></tr><tr><td>Hulín 1</td><td>Moravia (CZ)</td><td>72</td><td>Male</td><td>&lt;60</td><td>No archery context (Peška, 2013b)</td></tr><tr><td>Kněževes</td><td>Bohemia (CZ)</td><td>12</td><td>Male<sup>a</sup></td><td>30-60</td><td>Bowl, jug (Kytlicová, 1956; Hájek, 1968a, 1968b)</td></tr><tr><td>Laa Thaya</td><td>Lower Austria (AT)</td><td>NL 46</td><td>Male</td><td>~23</td><td>No archery context (Kern (2003), and personal communication)</td></tr><tr><td>Landau SO 1992</td><td>Bavaria (DE)</td><td>7</td><td>Male</td><td>&lt;60</td><td>Beaker, cup, jug, v-shaped buttons, and dagger (Heyd, 2000; Husty, 1993)</td></tr><tr><td>Leopoldsdorf</td><td>Lower Austria (AT)</td><td>5</td><td>Male<sup>a</sup></td><td>25-35</td><td>Grave group of five: several cups (some decorated), bowls, other ceramic forms, two gold burl rings, two simple gold rings, two faience pearls, amber jewelry, and animal bones (Willvonseder, 1937)</td></tr><tr><td>Lhánice</td><td>Moravia (CZ)</td><td>1</td><td>Male<sup>a</sup></td><td>&lt;60</td><td>Pot, cup, sheep bones (Moucha, 2005)</td></tr><tr><td>Libochovice</td><td>Bohemia (CZ)</td><td>32,244</td><td>Male</td><td>&lt;60</td><td>Bowl, cup, animal bones (Zápotocký, 1962)</td></tr><tr><td>Lochenice</td><td>Bohemia (CZ)</td><td>8</td><td>Male<sup>a</sup></td><td>&lt;60</td><td>None (Buchvaldek, 1990)</td></tr><tr><td>Lochenice</td><td>Bohemia (CZ)</td><td>15</td><td>Male</td><td>&lt;60</td><td>Jug, animal bones (Buchvaldek, 1990)</td></tr><tr><td>Morkůvky</td><td>Moravia (CZ)</td><td>9544</td><td>Male<sup>a</sup></td><td>40-50</td><td>Jugs, bowl, and animal bones (domestic) (Dvořák et al., 1996; Unger, 1984)</td></tr><tr><td>Ostopovice</td><td>Moravia (CZ)</td><td>20</td><td>Male<sup>a</sup></td><td>25-40</td><td>None (Dvořák, 1992)</td></tr><tr><td>Pavlov</td><td>Bohemia (CZ)</td><td>516</td><td>Male<sup>a</sup></td><td>&lt;50</td><td>Jug (Dvořák et al., 1996)</td></tr><tr><td>Pavlov</td><td>Bohemia (CZ)</td><td>517</td><td>Male</td><td>&lt;50</td><td>Bowl (Dvořák et al., 1996)</td></tr><tr><td>Pavlov</td><td>Bohemia (CZ)</td><td>525</td><td>Male<sup>a</sup></td><td>&lt;50</td><td>None (Dvořák et al., 1996)</td></tr><tr><td>Pavlov</td><td>Bohemia (CZ)</td><td>566</td><td>Male</td><td>14-26</td><td>Bowl (Dvořák et al., 1996)</td></tr><tr><td>Pavlov</td><td>Bohemia (CZ)</td><td>570</td><td>Male</td><td>&lt;60</td><td>Two bell beakers, copper dagger, and lithic industry (Dvořák et al., 1996)</td></tr><tr><td>Plotiště nad Labem</td><td>Bohemia (CZ)</td><td>36,100</td><td>Male<sup>a</sup></td><td>20-29</td><td>Bone awl, flint blade, and bone clasps (Vokolek, 1981)</td></tr><tr><td>Radovesice</td><td>Bohemia (CZ)</td><td>2/80</td><td>Male</td><td>&lt;60</td><td>Ceramic fragment (Turek, 2006)</td></tr><tr><td>Roštín</td><td>Moravia (CZ)</td><td>1</td><td>Male<sup>a</sup></td><td>&lt;60</td><td>Jug (Spurný, 1957)</td></tr><tr><td>Straubing-Lerchenhaid</td><td>Bavaria (DE)</td><td>2</td><td>Male<sup>a</sup></td><td>30-60</td><td>No archery context (personal contact with archaeologist Prof. Volker Heyd; Sjögren et al., 2019)</td></tr><tr><td>Straubing-Lerchenhaid</td><td>Bavaria (DE)</td><td>13</td><td>Male</td><td>20-39</td><td>No archery context (personal contact with archaeologist Prof. Volker Heyd; Sjögren et al., 2019)</td></tr></tbody></table>
<table><caption>TABLE 2 (Continued)</caption><thead><tr><th>Site</th><th>Region</th><th>Grave/inventory number</th><th>Sex</th><th>Age</th><th>Grave context</th></tr></thead><tbody><tr><td>Sulejovice</td><td>Bohemia (CZ)</td><td>2</td><td>Male</td><td>30-40</td><td>Found with two small children, bell beaker, flint blade, three pearls, two belt plates (Hájek, 1962, 1968a, 1968b)</td></tr><tr><td>Tišice</td><td>Bohemia (CZ)</td><td>30,634</td><td>Male</td><td>&lt;60</td><td>Bell beaker, bowl, and jug (Hájek, 1968a, 1968b)</td></tr><tr><td>Tišice</td><td>Bohemia (CZ)</td><td>30,635</td><td>Male</td><td>&lt;60</td><td>Bell beaker, bowl (Hájek, 1968a, 1968b)</td></tr><tr><td>Tödling</td><td>Upper Austria (AT)</td><td>3</td><td>Male</td><td>25-40</td><td>Bowl with animal bones, handled cup, and orientation NNW-SSE (Kern and Wiltschke-Schrotta, in press)</td></tr><tr><td>Tödling</td><td>Upper Austria (AT)</td><td>6</td><td>Male<sup>a</sup></td><td>15-21</td><td>Likely meat offerings (goat and sheep), orientation NE-SW (Kern and Wiltschke-Schrotta, in press)</td></tr><tr><td>Vyškov</td><td>Moravia (CZ)</td><td>2/1958, 136</td><td>Male</td><td>20-29</td><td>Four jugs, copper pin, three bone buttons (Ondráček, 1961; Stloukal, 1961)</td></tr><tr><td>Želešice</td><td>Moravia (CZ)</td><td>1/81</td><td>Male</td><td>&lt;60</td><td>None (Dvořák, 1992)</td></tr><tr><td>Zwingendorf<br>Alicenhof</td><td>Lower Austria (AT)</td><td>16</td><td>Male</td><td>35-45</td><td>None (Wiltschke-Schrotta et al., 2001)</td></tr></tbody></table>

<figure><img src="image_3.png" /><figcaption>FIGURE 2 The appearance of stone wristguards, arrowheads, and bow-shaped pendants in 46 suspected “archer” burials (males only).</figcaption></figure>

method existed primarily for certain muscle origins and the small insertions of the wrist and hand that are rarely preserved, and the presence classification had two further subcategories specifying a color or texture change versus a pathological irregular surface. Table 7 summarizes these scoring methods and Table 5 lists the method used to score each origin and insertion.

## 2.5 | Analyses of pathologies

Lastly, this study controlled for the absence or presence of DJD, rheumatoid arthritis (RA), diffuse idiopathic skeletal hyperostosis (DISH), and ankylosing spondylitis (AS). DJD is of interest because while it is linked to biomechanics, it can also be influenced by factors such as

age and genetic predisposition (Rogers & Waldron, 1995; Steckel et al., 2005; Waldron, 2009). RA, DISH, and AS required controls because they are examples of pathologies that can lead to EC, but do not have a biomechanical origin (Aufderheide et al., 1998; Mann & Hunt, 2005; Rothschild et al., 1990; Steckel et al., 2005; Waldron, 2009). For this reason, analyses must exclude individuals with these three pathologies; however, due to its links to biomechanics and the exclusion of senior individuals, analyses included skeletons with DJD, noting the specific locations as well as how it manifested on bone. Table 8 contains a summary of the criteria used to identify each pathology. Lastly, it is important to note that preservation did not always allow for a diagnosis of each pathology. Absence of a diagnosis therefore does not necessarily mean absence of a pathology; for example, a DISH diagnosis requires a preserved vertebral column, and

<footnote><sup>a</sup>A sex determination using the DSP 2 method.</footnote>

<footnote><sup>b</sup>The source only discusses grave 3, implying that all other graves have no additional burial context apart from a few ceramics.</footnote>
<table><caption>TABLE 3 The measurements taken for each individual when preservation allowed; all measurements and codes are based on the standards from Martin and Saller (1957); from Ryan-Despraz (2021, Table 34).</caption><thead><tr><th>Bone</th><th>Measurement</th><th>Code</th></tr></thead><tbody><tr><td rowspan="4">Clavicle</td><td>Maximum length</td><td>δ1</td></tr><tr><td>Vertical midshaft diameter</td><td>δ4</td></tr><tr><td>Sagittal midshaft diameter</td><td>δ5</td></tr><tr><td>Midshaft circumference</td><td>δ6</td></tr><tr><td rowspan="10">Scapula</td><td>Anatomical width</td><td>ε1</td></tr><tr><td>Anatomical length</td><td>ε2</td></tr><tr><td>Margo axillaris length</td><td>ε3</td></tr><tr><td>Supraspinatus fossa width</td><td>ε6</td></tr><tr><td>Length of scapular spine</td><td>ε7</td></tr><tr><td>Largest acromion width</td><td>ε9</td></tr><tr><td>Glenoid height</td><td>ε12</td></tr><tr><td>Transverse diameter of glenoid cavity</td><td>ε13</td></tr><tr><td>Maximum thickness of spinal crest</td><td>-</td></tr><tr><td rowspan="12">Humerus</td><td>Maximum length</td><td>ζ1</td></tr><tr><td>Breadth of proximal epiphysis</td><td>ζ3</td></tr><tr><td>Maximum distal breadth</td><td>ζ4</td></tr><tr><td>Maximum midshaft diameter</td><td>ζ5</td></tr><tr><td>Minimum midshaft diameter</td><td>ζ6</td></tr><tr><td>Midshaft circumference (at deltoid tuberosity)</td><td>ζ7a</td></tr><tr><td>Head circumference</td><td>ζ8</td></tr><tr><td>Transverse head diameter</td><td>ζ9</td></tr><tr><td>Vertical head diameter</td><td>ζ10</td></tr><tr><td>Breadth of trochlea</td><td>ζ11</td></tr><tr><td>Breadth of capitulum</td><td>ζ12</td></tr><tr><td>Breadth of olecranon fossa</td><td>ζ14</td></tr><tr><td rowspan="5">Radius</td><td>Maximum length</td><td>η1</td></tr><tr><td>Minimum circumference</td><td>η3</td></tr><tr><td>Maximum transverse shaft diameter</td><td>η4a</td></tr><tr><td>Minimum sagittal shaft diameter</td><td>η5a</td></tr><tr><td>Capitulum circumference</td><td>η5(3)</td></tr><tr><td rowspan="6">Ulna</td><td>Maximum length</td><td>θ1</td></tr><tr><td>Minimum circumference</td><td>θ3</td></tr><tr><td>Olecranon breadth</td><td>θ6</td></tr><tr><td>Olecranon depth</td><td>θ7</td></tr><tr><td>Dorso-volar diameter</td><td>θ11</td></tr><tr><td>Transverse diameter</td><td>θ12</td></tr><tr><td rowspan="6">Femur</td><td>Maximum length</td><td>λ1</td></tr><tr><td>Sagittal midshaft diameter</td><td>λ6</td></tr><tr><td>Transverse midshaft diameter</td><td>λ7</td></tr><tr><td>Circumference of midshaft</td><td>λ8</td></tr><tr><td>Upper transverse diaphysis diameter</td><td>λ9</td></tr><tr><td>Upper sagittal diaphysis diameter</td><td>λ10</td></tr></tbody></table>

(Continues)

<table><caption>TABLE 3 (Continued)</caption><thead><tr><th>Bone</th><th>Measurement</th><th>Code</th></tr></thead><tbody><tr><td rowspan="5">Tibia</td><td>Circumference of femoral neck</td><td>λ17</td></tr><tr><td>Vertical head diameter</td><td>λ18</td></tr><tr><td>Transverse head diameter</td><td>λ19</td></tr><tr><td>Circumference of the head</td><td>λ20</td></tr><tr><td>Epicondyle width</td><td>λ21</td></tr><tr><td rowspan="6">Fibula</td><td>Maximum length</td><td>ν1</td></tr><tr><td>Maximum proximal breadth</td><td>ν3</td></tr><tr><td>Maximum distal breadth</td><td>ν6</td></tr><tr><td>Maximum midshaft diameter</td><td>ν8</td></tr><tr><td>Transverse midshaft diameter</td><td>ν9</td></tr><tr><td>Midshaft circumference</td><td>ν10</td></tr><tr><td rowspan="2">Fibula</td><td>Maximum length</td><td>ξ1</td></tr><tr><td>Midshaft circumference</td><td>ξ4</td></tr></tbody></table>

not all individuals had this. This study excluded individuals with a positive diagnosis, but included individuals where a diagnosis was not possible. While this study acknowledges that such decisions are less than ideal, not including those individuals would have drastically reduced the sample size, even making many analyses impossible. This was additionally considered possible due to the very low prevalence of these pathologies—out of 48 individuals for whom a diagnosis was possible, only one case of DISH and zero cases of AS and RA were found (thus leaving 39 undiagnosed individuals).

## 2.6 | Data analysis

This study performed all data analyses in R (version 3.6.3) and these included hypothesis tests, dimension reduction, and MANOVA. The basic hypothesis tests aimed to signal possible variations between the measurements and EC of groups A and N and consisted of a t-test (Student's or Welch's depending on variance), Wilcoxon rank-sum test, Fisher's Exact test, and a chi-squared test. This study attempted to improve reliability by additionally performing bootstrapped t-tests, bootstrapped tests for the absolute value of the difference in median measurements, a Bonferroni correction, and controlling for each instance of a data outlier. Such tests applied to all datasets with an *n* ≥ 5 and an alpha value set at 0.05. Dimension reduction models involved principal component analysis (PCA) for the continuous data (measurements) and multiple correspondence analysis (MCA) for the categorical data (EC). MANOVA tests then further evaluated the resulting dimensions of each PCA and MCA.

Bootstrapped calculations aimed to improve reliability and compensate for unequal datasets, and non-bootstrapped tests were included to compare the two techniques. The bootstrapped tests created a data matrix for the given dataset, then ran 10,000 tests with a seed set at 99, and used resampling to obtain a final p-value.
<table><caption>**TABLE 4** Each of the calculated ratios, with all original measurements defined by Martin and Saller (1957); adapted from Ryan-Despraz (2021, Table 54).</caption><thead><tr><th colspan="2">Ratios</th></tr></thead><tbody><tr><td rowspan="3">Scapula</td><td>ε9R – ε9L/AVG</td></tr><tr><td>ε12R – ε12L/AVG</td></tr><tr><td>ε13R – ε13L/AVG</td></tr><tr><td rowspan="6">Clavicle</td><td>δ4R – δ4L/AVG</td></tr><tr><td>δ5R – δ5L/AVG</td></tr><tr><td>δ6R – δ6L/AVG</td></tr><tr><td>δ4R/δ1R</td></tr><tr><td>δ4L/δ1L</td></tr><tr><td>δ5R/δ1R</td></tr><tr><td rowspan="6">Humerus</td><td>ζ1R – ζ1L/AVG</td></tr><tr><td>ζ3R – ζ3L/AVG</td></tr><tr><td>ζ4R – ζ4L/AVG</td></tr><tr><td>ζ5R – ζ5L/AVG</td></tr><tr><td>ζ6R – ζ6L/AVG</td></tr><tr><td>ζ7aR – ζ7aL/AVG</td></tr><tr><td rowspan="6">Radius</td><td>ζ10R – ζ10L/AVG</td></tr><tr><td>ζ11R – ζ11L/AVG</td></tr><tr><td>ζ12R – ζ12L/AVG</td></tr><tr><td>ζ14R – ζ14L/AVG</td></tr><tr><td>ζ5R/ζ1R</td></tr><tr><td>ζ5L/ζ1L</td></tr><tr><td rowspan="6">Ulna</td><td>ζ6R/ζ1R</td></tr><tr><td>ζ6L/ζ1L</td></tr><tr><td>ζ7aR/ζ1R</td></tr><tr><td>ζ7aL/ζ1L</td></tr><tr><td>η1R – η1L/AVG</td></tr><tr><td>η3R – η3L/AVG</td></tr><tr><td rowspan="6">Femur</td><td>η4aR – η4aL/AVG</td></tr><tr><td>η5aR – η5aL/AVG</td></tr><tr><td>η5(6)R – η5(6)L/AVG</td></tr><tr><td>θ1R – θ1L/AVG</td></tr><tr><td>θ3R – θ3L/AVG</td></tr><tr><td>θ6R – θ6L/AVG</td></tr><tr><td>θ7R – θ7L/AVG</td></tr><tr><td rowspan="6">Femur</td><td>θ11R – θ11L/AVG</td></tr><tr><td>θ12R – θ12L/AVG</td></tr><tr><td>θ11R/θ1R</td></tr><tr><td>θ11L/θ1L</td></tr><tr><td>λ1R – λ1L/AVG</td></tr><tr><td>λ6R – λ6L/AVG</td></tr><tr><td>λ7R – λ7L/AVG</td></tr><tr><td>λ8R – λ8L/AVG</td></tr><tr><td>λ17R – λ17L/AVG</td></tr></tbody></table>

(Continues)

<table><caption>**TABLE 4** (Continued)</caption><thead><tr><th colspan="2">Ratios</th></tr></thead><tbody><tr><td rowspan="3">Tibia</td><td>λ18R – λ18L/AVG</td></tr><tr><td>λ19R – λ19L/AVG</td></tr><tr><td>λ20R – λ20L/AVG</td></tr><tr><td rowspan="4">Fibula</td><td>v6R – v6L/AVG</td></tr><tr><td>v8R – v8L/AVG</td></tr><tr><td>v9R – v9L/AVG</td></tr><tr><td>v10R – v10L/AVG</td></tr><tr><td></td><td>ξ4R – ξ4L/AVG</td></tr></tbody></table>

PCA and MCA are not possible on datasets containing missingness, which is a common problem with archaeological data. In order to address this issue, this study also tested multiple imputation models. For the PCA, each dataset required listwise deletion to lower the missingness to ≤30% before applying the MICE package (multivariate imputation by chained equations) (Buuren & Groothuis-Oudshoorn, 2011; Katitas, 2019; Lee & Simpson, 2014; Serneels & Verdonck, 2008; Zhang et al., 2019). This package imputes the missing data with plausible values in order to estimate regression coefficients not affected by the missing values. This follows a Conditional Multiple Imputation approach, which allows for increased flexibility as the distribution is calculated for each variable as opposed to each entire dataset (Buuren & Groothuis-Oudshoorn, 2011; Katitas, 2019). The imputation parameters include: number of multiple imputations based on percent missingness (e.g., 17% missingness applied 17 imputations), the predictive mean matching imputation model, 50 maximum iterations, and a seed set to 99. Imputations for the MCA applied a similar approach with the adapted package missMDA and the function imputeMCA, which uses regularized iterative MCA (Husson & Josse, 2014; Josse et al., 2012). Such methods have been tested in biological anthropology and identified as potential solutions to analyses with missing data (Wissler et al., 2022).

## 2.7 | Analyses at the individual level

Analyses at the individual level involved symmetry scores following the biomechanics profile. This was a two-step process: (1) organize the entheses according to draw arm, bow arm, or both arm activation (Table 5) and (2) note the symmetry of each. In other words, when an enthesis was visible for both the right and left sides, a comparison was made as to which side exhibited greater surface modification. Because this study assumes an association between enthesis appearance and muscle activation, this symmetry score could indicate whether a muscle was more physically developed on the right or left side. Therefore, the idea is that by organizing these symmetry scores with respect to the biomechanics of muscle activation in archery, it could be possible to identify specialized archers. When applicable, these individual analyses also considered the presence of trauma and DJD.
<table><caption>TABLE 5 The list of muscles associated with the draw arm, bow arm, and both arms in archery, scored for origins and insertions, according to various researchers (Ryan-Despraz, 2021, 2023).</caption><thead><tr><th></th><th>Activated muscle</th><th>Source and location distribution (draw arm/both arms/bow arm)</th><th>Sources</th><th>Scoring method</th></tr></thead><tbody><tr><td rowspan="11">Draw arm</td><td>m. extensor indicis</td><td>1/0/0</td><td>Kosar and Demirel (2004)</td><td>A/P—Origin, A/P—Insertion</td></tr><tr><td>m. extensor digiti minimi</td><td>1/0/0</td><td>Kosar and Demirel (2004)</td><td>A/P—Insertion</td></tr><tr><td>m. extensor carpi radialis longus</td><td>2/0/0</td><td>Kosar and Demirel (2004) and Sessa (1994)</td><td>A/P—Insertion</td></tr><tr><td>m. extensor carpi radialis brevis</td><td>2/0/0</td><td>Kosar and Demirel (2004), Sessa (1994)</td><td>A/P—Insertion</td></tr><tr><td>m. brachialis</td><td>3/0/0</td><td>Kosar and Demirel (2004), Lapostolle (2004), and Sessa (1994)</td><td>Mariotti et al.—Insertion</td></tr><tr><td>m. extensor digitorum</td><td>5/1/0</td><td>Açıkada et al. (2004), Clarys et al. (1990), Ertan et al. (2003), Kosar and Demirel (2004), Martin et al. (1990), and Tinazci (2011)</td><td>A/P—Insertion</td></tr><tr><td>m. flexor digitorum profundus</td><td>2/1/0</td><td>Ahmad et al. (2014), Kaymak et al. (2012), and Kosar and Demirel (2004)</td><td>A/P—Origin, A/P—Insertion</td></tr><tr><td>m. flexor digitorum superficialis</td><td>5/1/0</td><td>Ahmad et al. (2014), Clarys et al. (1990), Ertan et al. (2003), Kosar and Demirel (2004), Martin et al. (1990), and Tinazci (2011)</td><td>Mariotti et al.—Origin, A/P—Insertion</td></tr><tr><td>m. biceps brachii</td><td>5/0/1</td><td>Açıkada et al. (2004), Clarys et al. (1990), Kosar and Demirel (2004), Lapostolle (2004), Nishizono et al. (1987), and Sessa (1994)</td><td>Mariotti et al.—Origin, Coimbra—Insertion</td></tr><tr><td>m. trapezius</td><td>3/1/0</td><td>Clarys et al. (1990), Hu and Tang (2005), Tinazci (2011), and Zipp (1979)</td><td>Mariotti et al.—Insertion, A/P—Insertion</td></tr><tr><td>Superior m. trapezius</td><td>1/0/0</td><td>Nishizono et al. (1987)</td><td>A/P—Insertion</td></tr><tr><td>Middle m. trapezius</td><td>3/2/0</td><td>Kosar and Demirel (2004), Lapostolle (2004), Littke (2004), Nishizono et al. (1987), and Sessa (1994)</td><td>Mariotti et al.—Insertion</td></tr><tr><td>Inferior m. trapezius</td><td>1/1/0</td><td>Kosar and Demirel (2004) and Littke (2004)</td><td>Mariotti et al.—Insertion</td></tr><tr><td>m. rhomboid major</td><td>2/1/0</td><td>Kosar and Demirel (2004), Lapostolle (2004), and Littke (2004)</td><td>Mariotti et al.—Insertion</td></tr><tr><td>m. rhomboid minor</td><td>2/1/0</td><td>Kosar and Demirel (2004), Lapostolle (2004), and Littke (2004)</td><td>Mariotti et al.—Insertion</td></tr><tr><td>Posterior m. deltoid</td><td>3/1/0</td><td>Kosar and Demirel (2004), Lapostolle (2004), Sessa (1994), and Tinazci (2011)</td><td>Mariotti et al.—Origin</td></tr><tr><td>m. infraspinatus</td><td>1/1/0</td><td>Kosar and Demirel (2004) and Littke (2004)</td><td>A/P—Origin, Coimbra—Insertion</td></tr><tr><td>m. teres major</td><td>1/1/0</td><td>Kosar and Demirel (2004) and Lapostolle (2004)</td><td>Mariotti et al.—Origin, Mariotti et al.—Insertion</td></tr><tr><td>m. brachioradialis</td><td>1/1/0</td><td>Clarys et al. (1990) and Kosar and Demirel (2004)</td><td>Mariotti et al.—Origin, A/P—Insertion</td></tr><tr><td>lig. costoclavicular</td><td>2/0/0</td><td>Park et al. (2013) and Tihanyi et al. (2015)</td><td>Coimbra—Insertion</td></tr></tbody></table>
<table><caption>TABLE 5 (Continued)</caption><thead><tr><th></th><th>Activated muscle</th><th>Source and location distribution (draw arm/both arms/bow arm)</th><th>Sources</th><th>Scoring method</th></tr></thead><tbody><tr><td rowspan="10">Both arms</td><td>m. palmar interossei</td><td>1/0/1</td><td>Kosar and Demirel (2004) and Sessa (1994)</td><td>A/P—Origin, A/P—Insertion</td></tr><tr><td>m. pronator teres</td><td>0/1/0</td><td>Kosar and Demirel (2004)</td><td>Mariotti et al.—Origin, Mariotti et al.—Insertion</td></tr><tr><td>m. pronator quadratus</td><td>0/1/0</td><td>Kosar and Demirel (2004)</td><td>Mariotti et al.—Origin, A/P—Insertion</td></tr><tr><td>m. supinator</td><td>0/1/0</td><td>Lapostolle (2004)</td><td>Mariotti et al.—Origin, A/P—Insertion</td></tr><tr><td>m. latissimus dorsi</td><td>0/1/0</td><td>Kosar and Demirel (2004)</td><td>Mariotti et al.—Origin, Mariotti et al.—Insertion</td></tr><tr><td>m. pectoralis major</td><td>1/0/1</td><td>Açıkada et al. (2004) and Kosar and Demirel (2004)</td><td>Mariotti et al.—Origin, Mariotti et al.—Insertion</td></tr><tr><td>m. subscapularis</td><td>0/1/0</td><td>Lapostolle (2004)</td><td>Mariotti et al.—Origin, Coimbra—Insertion</td></tr><tr><td>m. supraspinatus</td><td>1/1/1</td><td>Kosar and Demirel (2004), Lapostolle (2004), and Littke (2004)</td><td>A/P—Origin, Coimbra—Insertion</td></tr><tr><td>m. teres minor</td><td>0/1/0</td><td>Kosar and Demirel (2004) and Littke (2004)</td><td>Mariotti et al.—Origin, Coimbra—Insertion</td></tr><tr><td>m. deltoids</td><td>3/1/3</td><td>Açıkada et al. (2004), Ahmad et al. (2014), Hu and Tang (2005), Kosar and Demirel (2004), Littke (2004), and Stirland, 2013, Zipp</td><td>Mariotti et al.—Insertion</td></tr><tr><td rowspan="3">Lateral m. deltoid</td><td></td><td>1/0/1</td><td>Clarys et al. (1990) and Nishizono et al. (1987)</td><td>Mariotti et al.—Origin</td></tr><tr><td>m. serratus anterior</td><td>0/1/0</td><td>Kosar and Demirel (2004)</td><td>Mariotti et al.—Insertion</td></tr><tr><td>m. extensor carpi ulnaris</td><td>1/0/1</td><td>Kosar and Demirel (2004) and Sessa (1994)</td><td>A/P—Insertion</td></tr><tr><td rowspan="10">Bow arm</td><td>m. flexor digiti minimi brevis</td><td>0/0/1</td><td>Kosar and Demirel (2004)</td><td>A/P—Insertion</td></tr><tr><td>m. opponens digiti minimi</td><td>0/0/1</td><td>Kosar and Demirel (2004)</td><td>A/P—Insertion</td></tr><tr><td>m. dorsal interossei</td><td>0/0/1</td><td>Kosar and Demirel (2004)</td><td>A/P—Origin, A/P—Insertion</td></tr><tr><td>m. lumbricales</td><td>0/0/1</td><td>Kosar and Demirel (2004)</td><td>A/P—Insertion</td></tr><tr><td>m. anconeus</td><td>0/0/1</td><td>Kosar and Demirel (2004)</td><td>Coimbra—Origin, Mariotti et al.—Insertion</td></tr><tr><td>m. flexor carpi ulnaris</td><td>0/0/2</td><td>Kosar and Demirel (2004) and Sessa (1994)</td><td>A/P—Insertion (pisiform, hamate)</td></tr><tr><td>m. flexor carpi radialis</td><td>0/0/1</td><td>Kosar and Demirel (2004)</td><td>A/P—Insertion</td></tr><tr><td>m. triceps brachii</td><td>0/1/5</td><td>Açıkada et al. (2004), Clarys et al. (1990), Kosar and Demirel (2004), Lapostolle (2004), Nishizono et al. (1987), and Sessa (1994)</td><td>Coimbra—Origin (long hd.), Mariotti et al.—Origin (lateral), A/P—Origin (medial), Villotte—Insertion</td></tr><tr><td>Anterior m. deltoid</td><td>0/1/1</td><td>Kosar and Demirel (2004) and Sessa (1994)</td><td>Mariotti et al.—Origin</td></tr></tbody></table>

Abbreviations: A/P, Absence/presence method.

Note: “Source and Location Distribution” indicates the number of sources citing each location (x/y/z = draw arm/both arms/bow arm), for example 2/1/0 signifies that 2 sources link this muscle only to the draw arm, 1 source links it to both arms, and no sources link it only to the bow arm, from Ryan-Despraz (2021, Table 28). The last column indicates the method used to score each enthesis as well as if it was applied to a muscle origin or insertion.
<table><caption>TABLE 6 Common injuries associated with specialized archery (Ryan-Despraz, 2021, 2023).</caption><thead><tr><th></th><th>Common Injuries</th><th>Sources</th></tr></thead><tbody><tr><td rowspan="3">Draw arm</td><td>Shoulder tendinitis— mainly the m. <i>supraspinatus</i>, but also the m. <i>infraspinatus</i>, m. <i>teres minor</i>, and m. <i>subscapularis</i></td><td>Gopal Adkitte et al. (2016), Hildenbrand and Rayan (2010), Kaynaroğlu and Kiliç (2012), and Mann and Littke (1989)</td></tr><tr><td>Shoulder impingement</td><td>Kaynaroğlu and Kiliç (2012), Littke (2004), Mann and Littke (1989), Naraen et al. (1999), Niestroj et al. (2018), Prine et al. (2016), and Singh and Lhee (2016)</td></tr><tr><td>Biceps tendinitis</td><td>Kaynaroğlu and Kiliç (2012) and Mann and Littke (1989)</td></tr><tr><td rowspan="4">Bow arm</td><td>Epicondylitis</td><td>Frostick et al. (1999), Lapostolle et al. (2004), Niestroj et al. (2018), Rayan (1992), Sessa (1994), Singh and Lhee (2016), and Whiteside and Andrews (1989)</td></tr><tr><td>Osteochondritis dissecans</td><td>Andrews and Whiteside (1993)</td></tr><tr><td>Osseous Hypertrophy at the elbow</td><td>Andrews and Whiteside (1993)</td></tr><tr><td>De Quervain's Tenosynovitis</td><td>Kaynaroğlu and Kiliç (2012) and Rayan (1992)</td></tr></tbody></table>

### 3 | RESULTS

All data is available through the original PhD thesis, which is available in open access through the University of Geneva's Archive Ouverte website (https://archive-ouverte.unige.ch/unige:151360).

#### 3.1 | Population Analyses—“Archer” Burials (A) vs. “Non-archer” Burials (N)

139 hypothesis tests looking at individual measurements revealed no significant differences between groups. PCA analyses also found no clustering in the data, a finding that was additionally supported through a MANOVA test for the principal components, with $F(32,2) = 1.0433$, $p = 0.3651$, Pillai's Trace = 0.0671, partial $\eta^2 = 0.07$. This finding remained true when group A was further sub-categorized according to wristguard and arrowhead presence, with no visible patterns in the data. A comparison between the regular t-test and the bootstrapped t-test results found the similar values, indicating no significant difference in overall reliability.

Assessments of each individual score yielded few significant differences in EC manifestation frequencies between groups, with only

<table><caption>TABLE 7 The four methods used to classify the enthesis, their basic criteria, and notations, with thorough definitions and references available in the corresponding publications (Henderson et al., 2013, 2016; Mariotti et al., 2004, 2007; Villotte, 2006, 2009), an increasing notation (number or letter) denotes a larger or more prominent surface modification (e.g., a stage 2 will cover more of the enthesis surface than stage 1).</caption><thead><tr><th></th><th>Criteria—appearance on bone</th><th>Notation/EC Score</th></tr></thead><tbody><tr><td rowspan="6">Coimbra method (Henderson et al., 2013, 2016)</td><td>Contour: Bone formation (OP)</td><td>0, 1, or 2</td></tr><tr><td>Contour: Bone erosion (OL)</td><td>0, 1, or 2</td></tr><tr><td>Center: Texture change</td><td>0 or 1</td></tr><tr><td>Center: Bone formation (OP)</td><td>0, 1, or 2</td></tr><tr><td>Center: Erosion (OL)</td><td>0, 1, or 2</td></tr><tr><td>Center: Fine porosity (OL)</td><td>0, 1, or 2</td></tr><tr><td rowspan="2">Mariotti et al.'s method (Mariotti et al., 2004, 2007)</td><td>Center: Macro-porosity (OL)</td><td>0, 1, or 2</td></tr><tr><td>Center: Cavitation (OL)</td><td>0, 1, or 2</td></tr><tr><td rowspan="3">Villotte.'s Method (Villotte, 2006; 2009)</td><td>Robusticity</td><td>1a, 1b, 1c, 2, or 3</td></tr><tr><td>Osteophytic formation (OP)</td><td>0, 1, 2, or 3</td></tr><tr><td>Osteolytic formation (OL)</td><td>0, 1, 2, 3a, or 3b</td></tr><tr><td rowspan="3">Absence/presence method</td><td>Absence of EC</td><td>A</td></tr><tr><td>Enthesophyte presence (OP)</td><td>Ba or Ca</td></tr><tr><td>Erosion surface (OL)</td><td>Bb or Cb</td></tr><tr><td rowspan="3">Absence/presence method</td><td>Absence (no EC, no color/texture change)</td><td>A</td></tr><tr><td>Color/texture change</td><td>B</td></tr><tr><td>Presence (irregular surface)</td><td>C</td></tr></tbody></table>

Abbreviations: OP, osteophytic development; OL, osteolytic development.

one passing the Bonferroni correction (the robusticity score for the right *m. pronator teres*), and none between right and left sides. Further analyses aiming to broaden the scope created two binary sub-classifications of the EC: (1) general osteophytic and osteolytic modifications (i.e., bone proliferation vs. bone surface erosion) (Table 7), and (2) general presence or absence of EC (i.e., scores of 0 or A vs. all other scores). These examinations revealed that groups A and N have similar types of surface modifications (i.e., osteophytic and osteolytic development), but not similar levels of overall EC presence (Table 9). Overall, group N exhibited significantly more EC than group A.

The general MCA results illustrated similar patterns as the EC hypothesis test data, with a first dimension separating fairly well
<table><caption>**TABLE 8** The pathologies controlled for in this study and the diagnosis criteria.</caption><thead><tr><th></th><th>Criteria</th><th>Source</th></tr></thead><tbody><tr><td>Degenerative joint disease (DJD)</td><td>Absence or presence of eburnation If no eburnation, then must have two of the following:<ul><li>Marginal osteophyte</li><li>New bone on joint surface</li><li>Pitting on joint surface</li><li>Alteration in joint contour</li></ul></td><td>Rogers and Waldron (1995)<br>Waldron (2009)</td></tr><tr><td>Rheumatoid arthritis (RA)</td><td>Any combination of:<ul><li>Symmetrical, marginal erosions of small joints in the hands and/or feet</li><li>Sparing of sacroiliac joints</li><li>Minimal new bone formation</li><li>Absence of spinal fusion</li></ul></td><td>Waldron (2009)<br>Biehler-Gomez and Cattaneo (2018)</td></tr><tr><td>Diffuse idiopathic skeletal hyperostosis (DISH)</td><td>Absence or presence of paraspinal ligament ossification between at least two vertebrae<br>Classification stages:<ol><li>Vertebrae not available for observation</li><li>No evidence of DISH</li><li>Features of DISH are present</li></ol></td><td>Belanger and Rowe (2001)<br>Maat et al. (1995)<br>Mann and Hunt (2005)<br>Resnick and Niwayama (1978)<br>Steckel et al. (2005)<br>Vaishya et al. (2017)<br>Weinfeld et al. (1997)</td></tr><tr><td>Ankylosing spondylitis (AS)</td><td>Scoring classification:<ol><li>Vertebral and sacral bodies not available for observation</li><li>No AS visible in the preserved bones</li><li>Symmetrical fusion of both sacroiliac joints AND spinal fusion with no skip lesions</li></ol></td><td>Khan (2002)<br>Mann and Hunt (2005)<br>Šlaus et al. (2012)<br>Waldron (2009)</td></tr></tbody></table>

groups A and N, including slightly more clustering in group A and slightly more variation in group N (Figure 3). The variables most responsible for the dimension 1 variation include the *m. teres major*, *m. biceps brachii*, *m. deltoid*, *m. palmar interossei*, *m. pronator quadratus*, *m. pectoralis major*, *m. brachialis*, *t. common extensor*, and the *m. anconeus*. A MANOVA also signaled trends in the data, with a significant *p*-value but a mid-value Pillai's trace, with $F(49,2) = 9.9261$, $p = 0.0003$, Pillai's Trace = 0.3015, partial $\eta^2 = 0.30$. This discrepancy indicates that while the independent variables might not have an extremely strong effect on the dependent variables, there is still evidence of a statistically significant effect. A focused MCA controlling for wristguard and arrowhead presence revealed no additional trends in the data.

## 3.2 | Individual analyses—assessing the individuals of Group A for signs of specialized archery

These results examined each individual of group A for EC symmetry scores linked to specialized archery according to the previously established biomechanics profile. However, due to a lack of complete bone preservation, a symmetry score was not possible for all entheses. A “specialized archer” assessment required each individual to have this data at $n \ge 5$, and a lack of bone preservation excluded 20 individuals from this analysis. Of the remaining 26 individuals (including Lochenice 3, who was not included in the population analyses due to an undetermined sex), observations identified 11 as “confirmed” archers (C) and 15 as “improbable” archers (IA) based on biomechanic consistency (Table 10). Those individuals include possible right-handed archers: Hoštice 1–862, Hoštice 1–821, Oberstimm II/1982–2, and Vykáň–1, and possible left-handed archers: Altenmarkt–11, Hoštice 1–915, Hoštice 1–884, Hulín 1–73, Lochenice–3, Radovesice–59/80-II, and Rosnice–I/59. Analyses using Fisher's Exact test found that these two new datasets (right-handed and left-handed “C” archers) are indeed statistically unusual (Table 11). Tests revealed significant differences for the symmetry frequencies of the draw arm vs. the bow arm in right-handed archers, the draw arm vs. the bow arm in left-handed archers, and the draw arms of right-handed archers vs. left-handed archers. The MCA examining these individuals also revealed trends in the data, with the first dimension managing to cluster individuals according to suspected archery handedness (Figure 4). There were two outliers, one right-handed (Vykáň 1) and one left-handed suspected archer (Lochenice 3), separated from the others by both dimensions, and from each other by the second dimension. The variables most responsible for the dimension 1 variation include: the *m. common extensor*, *l. costoclavicular*, *m. supraspinatus*, *m. pectoralis major*, left *m. anconeous*, *m. biceps brachii*, *m. teres minor*, *m. pronator quadratus*, *m. latissimus dorsi*, *m. teres major*, and the *m. deltoids*. A MANOVA was not possible due to the low sample size. PCA models and hypothesis tests examining the measurements revealed no patterns in the data. A distribution of the archery-related items showed no link between grave goods and a “confirmed” archer classification (Figure 5).

Six individuals from group A had confirmed DJD, two of whom classified as “C” (Lochenice 3 and Osterhofen 11), two as “IA”, and two did not present enough data for a classification. Both “C” individuals and one “IA” individual presented asymmetrical manifestations in the upper limb (Table 10).

## 4 | DISCUSSION

The first aim of this study was to analyze the Bell Beaker sample at a “population” level by comparing the skeletons of “archers” (Group A) and “non-archers” (Group N). Because “archer” burials tended to be more “wealthy”, the purpose of this analysis was to assess a potential link between burial wealth and skeletal morphology, in particular with regard to physical activity.
<table><caption>TABLE 9 The results of population analyses (“archers” (A) and “non-archers” (N)) of EC for osteophytic (OP) vs. osteolytic (OL) surface changes and overall presence of any surface modification vs. absence.</caption><thead><tr><th rowspan="2"></th><th colspan="2">OP vs. OL</th><th colspan="2">Surface modification presence vs. absence</th></tr><tr><th>Test result</th><th>95% CI prop. test</th><th>Test result</th><th>95% CI prop.test</th></tr></thead><tbody><tr><td>Coimbra method—Origins (χ² test)</td><td>χ²(1, N<sub>a</sub> = 977, N<sub>n</sub> = 941) = 1.73, p = 0.19</td><td>[-0.0313, 0.1461]</td><td>χ²(1, N<sub>a</sub> = 977, N<sub>n</sub> = 941) = 11.7, p < 0.001<br>Group N tended to have more instances of surface modifications</td><td>[-0.1278, -0.0335]</td></tr><tr><td>Coimbra method—insertions (χ² test)</td><td>χ²(1, N<sub>a</sub> = 1652, N<sub>n</sub> = 1342) = 0.20, p = 0.65</td><td>[-0.0832, 0.0529]</td><td>χ²(1, N<sub>a</sub> = 1652, N<sub>n</sub> = 1342) = 12.07, p < 0.001<br>Group N tended to have more instances of surface modifications</td><td>[-0.1013, -0.0273]</td></tr><tr><td>Villotte method—Insertion (Fisher's Exact test)</td><td>(N<sub>a</sub> = 41, N<sub>n</sub> = 29), p = 0.15</td><td>N/A</td><td>(N<sub>a</sub> = 41, N<sub>n</sub> = 29), p = 1</td><td>N/A</td></tr><tr><td>Mariotti et al. method—origins (χ² test)</td><td>χ²(1, N<sub>a</sub> = 535, N<sub>n</sub> = 590) = 5.48, p = 0.02<br>Group N tended to have more OL than group A</td><td>[-0.0459, 0.4267]</td><td>χ²(1, N<sub>a</sub> = 535, N<sub>n</sub> = 590) = 0.77, p = 0.38</td><td>[-0.1352, 0.0542]</td></tr><tr><td>Mariotti et al. Method—Insertions (χ² test)</td><td>χ²(1, N<sub>a</sub> = 757, N<sub>n</sub> = 779) = 0.53, p = 0.47</td><td>[-0.0637, 0.1337]</td><td>χ²(1, N<sub>a</sub> = 757, N<sub>n</sub> = 779) = 13.33, p < 0.001<br>Group N tended to have more instances of surface modifications</td><td>[-0.1490, -0.0434]</td></tr><tr><td>Absence/presence method—origins (χ² test)</td><td>N/A</td><td>N/A</td><td>χ²(1, N<sub>a</sub> = 120, N<sub>n</sub> = 169) = 16, p < 0.001<br>Group N tended to have more instances of surface modifications</td><td>[0.1381, 0.4300]</td></tr><tr><td>Absence/presence method—Insertions (χ² test)</td><td>N/A</td><td>N/A</td><td>χ²(1, N<sub>a</sub> = 206, N<sub>n</sub> = 217) = 26.14, p < 0.001<br>Group N tended to have more instances of surface modifications</td><td>[0.224, 0.4694]</td></tr><tr><td>All methods combined (χ² test)</td><td>χ²(1, N<sub>a</sub> = 3962, N<sub>n</sub> = 3681) = 1.08, p = 0.3</td><td>[-0.0205, 0.0649]</td><td>χ²(1, N<sub>a</sub> = 3962, N<sub>n</sub> = 3681) = 65.26, p < 0.001<br>Group N tended to have more instances of surface modifications</td><td>[-0.1150, -0.0698]</td></tr></tbody></table>

Note: Each test also included a proportion test (R function “prop.test”) and this provides a confidence interval for the difference in proportions; the alpha value was set to 0.05.

<figure><img src="image_4.png" /><figcaption>FIGURE 3 The MCA plot controlling for entheunal changes scores between the “archers” (A) and “non-archers” (N).</figcaption></figure>
<table><caption>TABLE 10 All suspected “archers” (n = 26) that provided at least five symmetry data points (“right,” “left,” or “symmetric” entheseseal modification prominence comparison), the associated muscles for each arm can be found in Table 3.</caption><thead><tr><th rowspan="2">Individual</th><th colspan="3">Draw Arm (20 associated muscles)</th><th colspan="3">Both Arms (13 associated muscles)</th><th colspan="3">Bow Arm (9 associated muscles)</th><th colspan="3">DJD</th><th rowspan="2">Assessment Result</th></tr><tr><th>Right</th><th>Left</th><th>Symmetric</th><th>Right</th><th>Left</th><th>Symmetric</th><th>Right</th><th>Left</th><th>Symmetric</th><th>Manifestation</th><th>Location</th><th>Assessment Result</th></tr></thead><tbody><tr><td>Altenmarkt, 2 (M)</td><td>5</td><td>0</td><td>3</td><td>10</td><td>2</td><td>4</td><td>7</td><td>1</td><td>3</td><td>2, 3, 4, and 5</td><td>Vertebrae, right distal clavicle, right distal ulna, and both proximal ulnas</td><td>IA</td></tr><tr><td>Altenmarkt, 6 (M)</td><td>2</td><td>1</td><td>4</td><td>3</td><td>2</td><td>5</td><td>2</td><td>1</td><td>2</td><td></td><td></td><td>IA</td></tr><tr><td>Altenmarkt, 10 (M)</td><td>0</td><td>0</td><td>1</td><td>1</td><td>0</td><td>2</td><td>2</td><td>0</td><td>1</td><td></td><td></td><td>IA</td></tr><tr><td>Altenmarkt, 11 (M)</td><td>1</td><td>3</td><td>7</td><td>6</td><td>1</td><td>8</td><td>2</td><td>0</td><td>6</td><td>2, 4</td><td>Right glenoid cavity, both acromions, both proximal clavicles, and right distal ulna</td><td>C, left-handed</td></tr><tr><td>Gemeinlebarn, 2071 (M)</td><td>2</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td><td>1</td><td>0</td><td>0</td><td></td><td></td><td>IA</td></tr><tr><td>Hošstice 1, 863 (M)</td><td>1</td><td>0</td><td>1</td><td>3</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td></td><td></td><td>IA</td></tr><tr><td>Hošstice 1, 862 (M)</td><td>1</td><td>0</td><td>1</td><td>1</td><td>0</td><td>2</td><td>0</td><td>1</td><td>0</td><td></td><td></td><td>C, right-handed</td></tr><tr><td>Hošstice 1, 821 (M)</td><td>1</td><td>0</td><td>1</td><td>1</td><td>0</td><td>4</td><td>0</td><td>0</td><td>0</td><td></td><td></td><td>C, right-handed</td></tr><tr><td>Hošstice 1, 949 (M)</td><td>1</td><td>1</td><td>3</td><td>1</td><td>0</td><td>2</td><td>1</td><td>0</td><td>2</td><td>2, 3</td><td>Vertebrae</td><td>IA</td></tr><tr><td>Hošstice 1, 915 (M)</td><td>1</td><td>2</td><td>0</td><td>2</td><td>0</td><td>5</td><td>1</td><td>0</td><td>0</td><td></td><td></td><td>C, left-handed</td></tr><tr><td>Hošstice 1, 884 (M)</td><td>0</td><td>1</td><td>4</td><td>1</td><td>1</td><td>2</td><td>0</td><td>0</td><td>3</td><td></td><td></td><td>C, left-handed</td></tr><tr><td>Hulín 1, 66 (M)</td><td>1</td><td>0</td><td>3</td><td>5</td><td>2</td><td>4</td><td>1</td><td>1</td><td>2</td><td></td><td></td><td>IA</td></tr><tr><td>Hulín 1, 73 (M)</td><td>3</td><td>0</td><td>3</td><td>2</td><td>0</td><td>6</td><td>0</td><td>0</td><td>2</td><td></td><td></td><td>C, left-handed</td></tr><tr><td>Hulín 1, 85 (M)</td><td>0</td><td>0</td><td>3</td><td>0</td><td>1</td><td>3</td><td>0</td><td>1</td><td>4</td><td></td><td></td><td>IA</td></tr><tr><td>Lochenice, 3 (ND)</td><td>0</td><td>2</td><td>5</td><td>0</td><td>1</td><td>7</td><td>2</td><td>0</td><td>1</td><td>2, 3, and 4</td><td>Right thumb, both glenoid cavities</td><td>C, left-handed</td></tr><tr><td>Oberstimm II/1982, 2 (M)</td><td>3</td><td>0</td><td>0</td><td>1</td><td>0</td><td>3</td><td>0</td><td>0</td><td>1</td><td></td><td></td><td>C, right-handed</td></tr><tr><td>Praha 8-Kobylysy, 36,749, obj. 13 (M)</td><td>1</td><td>1</td><td>5</td><td>3</td><td>0</td><td>5</td><td>0</td><td>0</td><td>3</td><td></td><td></td><td>IA</td></tr><tr><td>Radovesice, 59/80-II (M)</td><td>2</td><td>3</td><td>1</td><td>2</td><td>4</td><td>5</td><td>4</td><td>0</td><td>0</td><td></td><td></td><td>C, left-handed</td></tr><tr><td>Radovesice, 53/80-I (M)</td><td>2</td><td>1</td><td>4</td><td>3</td><td>2</td><td>6</td><td>1</td><td>1</td><td>3</td><td></td><td></td><td>IA</td></tr><tr><td>Rosnice, I/59 (M)</td><td>0</td><td>2</td><td>1</td><td>1</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td></td><td></td><td>C, left-handed</td></tr><tr><td>Stehelčěves, 2 (M)</td><td>1</td><td>0</td><td>3</td><td>2</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td><td></td><td></td><td>IA</td></tr><tr><td>Straubing Stadtäcker, 1 (M)</td><td>2</td><td>0</td><td>0</td><td>0</td><td>1</td><td>1</td><td>0</td><td>0</td><td>2</td><td></td><td></td><td>IA</td></tr><tr><td>Velké Prílepy, 2 (M)</td><td>2</td><td>2</td><td>2</td><td>5</td><td>0</td><td>3</td><td>0</td><td>0</td><td>1</td><td></td><td></td><td>IA</td></tr><tr><td>Vykán, 1 (M)</td><td>4</td><td>0</td><td>1</td><td>3</td><td>2</td><td>7</td><td>1</td><td>0</td><td>4</td><td></td><td></td><td>C, right-handed</td></tr><tr><td>Weichering, 17 (M)</td><td>2</td><td>0</td><td>5</td><td>3</td><td>1</td><td>5</td><td>2</td><td>0</td><td>0</td><td></td><td></td><td>IA</td></tr><tr><td>Žábovřesky (P7A 32,293) (M)</td><td>0</td><td>0</td><td>4</td><td>0</td><td>0</td><td>4</td><td>0</td><td>0</td><td>1</td><td></td><td></td><td>IA</td></tr></tbody></table>

Note: Estimated sex is indicated in parentheses, with M = male and ND = not determined; C = “confirmed” archer with the handedness assessment and IA = improbable archer; DJD manifestations include: 1 (eburnation), 2 (marginal osteophyte), 3 (new bone on joint surface), 4 (pitting on joint surface), and 5 (alteration in joint contour) (Rogers & Waldron, 1995; Waldron, 2009); from Ryan-Despraz (2021, Tables 109 and 116).
<table><caption>TABLE 11 The results of frequency tests controlling archery draw and bow arm activation for group C; R-H = right-handed, L-H = left-handed; from Ryan-Despraz (2021, Table 11).</caption><thead><tr><th colspan="3">Comparisons (Fisher's exact test) for asymmetric frequencies between "confirmed" archers</th></tr><tr><th>Test groups</th><th>Test parameters</th><th>Test result</th></tr></thead><tbody><tr><td>R-H Archers</td><td>Draw arm (D) vs. Bow arm (B)</td><td>(n<sub>D</sub> = 10, n<sub>B</sub> = 5), p = 0.04</td></tr><tr><td>L-H Archers</td><td>Draw arm (D) vs. Bow arm (B)</td><td>(n<sub>D</sub> = 31, n<sub>B</sub> = 19), p = 0.006</td></tr><tr><td>R-H vs. L-H Archers</td><td>Draw arm</td><td>(n<sub>Right</sub> = 10, n<sub>Left</sub> = 31), p = 0.003</td></tr><tr><td>R-H vs. L-H Archers</td><td>Bow arm</td><td>(n<sub>Right</sub> = 5, n<sub>Left</sub> = 19), p = 0.08</td></tr><tr><td>R-H vs. L-H Archers</td><td>Both arms</td><td>(n<sub>Right</sub> = 27, n<sub>Left</sub> = 64), p = 0.68</td></tr></tbody></table>

## 4.1 | The population analyses

Looking first at the measurement data, the lack of differentiation between A and N indicates that the two groups were not variable in terms of physical size and proportion. This is not necessarily surprising since these individuals were part of the same population. However, the measurement data, including values linked to “robusticity,” does not seem to function as a proxy for more internalized measurements. The lack of significance for these values contrasts with the EC findings, which did note differences in surface morphology between groups A and N. While more data are needed, this study cannot attest to the reliability of such analyses as a potential stand-in for analyses of diaphyseal geometry.

The EC results of the population analyses emphasized a difference between the type of surface modification and the overall amount of EC. In particular, the finding that groups A and N had similar occurrences of osteophytic and osteolytic modifications could demonstrate that the two groups shared a similar type of biomechanical stress—meaning that they lived in similar conditions with similar daily constraints. However, the finding that group N consistently had more instances of overall EC than group A could indicate the level of biomechanical stress. That is to say, while these two groups were part of the same general population, group N possibly had a more physically strenuous daily life. This could exhibit the presence of labor differentiation that is additionally identifiable through cultural burial practice. If “archer” burials were indeed more prestigious or wealthy (either socially, monetarily, or both), this could draw a direct link between that and lower levels of physical activity. Notably, the burials of group N were generally less rich, in both quality and quantity of grave goods (e.g., fewer instances of metal items or carved stone). This correlation between fewer EC and an archer burial could indicate a relationship in Bell Beaker society between labor and grave goods. At the same time, this study was not able to differentiate “archer” burials further through the type of archery-related item. There was no statistical significance when tests sub-categorized the burials

according to wristguard or arrowhead presence (a lack of samples excluded analyses controlling only for bow-shaped pendants). Yet, while wristguards and arrowheads are associated with the same activity, they do not appear together consistently in the same burial context—only 16 out of the 46 “archer” burials had both items. It is therefore possible that they did not have the same cultural function for the Bell Beaker peoples.

## 4.2 | The individual analyses

The second primary aim of this study was to assess the “archers” at an “individual” level for osteological indications of specialized archer activity. In other words, the goal was to see if the burial-item profile (“archery”) matched the biological profile (“archer” occupation). However, a lack of adequate preservation meant individual analyses were not possible for 20 skeletons. For the 11 possibly “confirmed” archers, four followed the expected patterns for a right-handed archer and seven for a left-handed archer. Initially, this result is surprising because the vast majority of *homo sapiens* throughout time and space are right-handed (74%–96%) (Faurie & Raymond, 2004; Llaurens et al., 2009; Uomini, 2009). Further investigation found that such a finding could be linked to eye laterality rather than hand laterality. Just as humans have a preferred hand, they also have a preferred eye, and while the two are often on the same side, this is not always the case. Bourassa et al. (1996) found that while on average 10% of people are left-handed, 33% are left-eye dominant, with additional studies reporting similar and even higher levels of cross dominance (Puri & Sethi, 2017; Robison et al., 1995). Most instructors advise cross-dominant archers to choose a side and adapt either the hand or the eye, and because precision is so vital to success, most archers choose to adapt their hand (Axford, 1995; Laborde et al., 2009; Porac & Coren, 1981). However, while these findings are important to consider, they are not sufficient for explaining this study's higher level of possible left-handed archers. At the same time, while this finding is noteworthy, statistically speaking the difference in frequencies is not significant.

Hypothesis tests for group C found a *p* < 0.05 for three out of four tests comparing the possible left- and right-handed archers. This signals something unusual in the data that merits further exploration and indeed, MCA analyses revealed similar trends. Examining the variables responsible for this variation could help improve osteological analyses of specialized archer activity. These findings indicate that this study's biomechanical theory managed to identify separate groups of individuals whose muscular development is somehow linked. This includes comparisons between not only draw and bow arms, but also the general comparison between possible right- and left-handed archers, which exhibits the potential efficacy of applying a biomechanics model when attempting to interpret specialized archery, including on datasets containing missingness. However, this final dataset of “C” individuals remains small, so additional approaches on other individuals would improve interpretations.
<figure><img src="image_5.png" /><figcaption>**FIGURE 5** The relationship between archery-related grave goods and individual classifications of specialized archery occupation (“confirmed” archers and improbably archers); adapted from Ryan-Despraz (2021, Figure 90).</figcaption></figure>

<figure><img src="image_6.png" /></figure>

Assessing group C for patterns in grave good appearance and distribution revealed no relationship between the archaeological context and a “confirmed” archer classification (Figure 5). This contributes to the conclusion that there are no black-and-white rules linking burial context to a likely occupation—not all of group A were likely specialized archers just as some members of group N could have been specialized archers. From an archaeological perspective, this could mean that grave goods, such as stone wristguards, represented a social function rather than an occupational function in Bell Beaker society. Therefore, rather than labeling an individual buried with archery-related items as an “archer”, it could be more accurate to identify this individual as someone of social importance. In terms of specialized occupation, it would be more reliable to examine each individual according to his or her biological attributes, including those individuals without the corresponding archaeological context (i.e., group N). A direct link between cultural identity (as exhibited through grave

goods) and occupational identity emphasizes the individual (i.e., an archer); however a lack of association, such as this study found, shifts this focus to the concept of archery as an ideal within Bell Beaker society.

## 4.3 | Limitations

As mentioned throughout this article, data missingness is a constant issue, and prehistoric skeletons are rarely fully preserved. Studies such as this rely on comparisons, and this becomes complicated when each individual presents different levels of preservation. For analyses at the population level, measurement and EC evaluations did not necessarily contain data from the same individuals, which is of course less than ideal, though it is also unavoidable. The question will always remain as to how the absence or presence of certain individuals may have
influenced the results. A controlled evaluation of each individual from group A helped to remove this bias, however even those individual analyses did not necessarily include symmetry assumptions for the same enthesis. One EC will probably not indicate an activity, however the ensemble could help to paint a clearer overview—this is why the application of the biomechanics profile was essential. Furthermore, due to small sample sizes and broad age determinations, this study was not able to further assess patterns linked to age groups, which could also be an important consideration in future studies. Overall, this study aimed to reduce the effects of data missingness and bias by examining several skeletal characteristics, applying several methods of data analysis, and evaluating the skeletons both as a broad population and as individuals.

A primary objective of this study was to test whether or not the individuals buried with archery-related items practiced a specialized archery occupation—meaning individual analyses only focused on group A. While this was a measured decision based on the specific research question, it would also have been interesting to consider individual archery analyses of “non-archers” (group N). This would have provided an additional layer of comparison with group A, especially in terms of interpreting social organization. It could therefore prove beneficial for any future studies to expand individual analyses in order to promote more complex population assessments.

Many researchers also question the reliability, repeatability, and efficacy of scoring enthesal changes, which remains a valid concern. This study attempted to address these limits by applying multiple methods best adapted to each enthesis, as well as scoring simply according to the absence or presence, thus removing possible biases linked to subjectivity. The individual analyses also applied a binary comparison between right and left sides, which limits the risk of error due to a possibly arbitrary classification. However, these methods lack quantification. More accurate evaluations will require the development of methods capable of quantifying bone surface morphology. Additionally, determining activity from the skeleton remains an unconfirmed field because researchers have still not been able to define exactly how biomechanics influences bone morphology or how to differentiate it from other etiologies. As such, many researchers have also begun to study bone microarchitecture with regard to physical activity (Agarwal et al., 2004; Berthon et al., 2015; Bouvard et al., 2012; Courteix et al., 1998; Ding et al., 2002; Griffin et al., 2010; Huiskes et al., 2000; T. M. Ryan & Shaw, 2012; Wallace et al., 2017). Ultimately, studies such as this assume a direct association between muscle activation and enthesal morphology, yet this is likely an oversimplification of the underlying biological processes. In particular, this study was based on a biomechanics profile for archery and therefore studied the EC for this specific group of entheses, which can be considered heterogeneous (see for instance Villotte & Santos, 2023). Some of these recorded EC are likely alterations caused by micro-trauma while others are possibly more adaptations (normal physiological responses to loading) (see Villotte, 2023). With this in mind, further research is required to better understand the relationship between the patterns identified in the biomechanics-based EC presence and the underlying processes that led to them. This is

also noteworthy considering that this study combined analyses of fibrous and fibrocartilaginous EC; however when considering what is currently known about these attachment sites, it is perhaps not appropriate to assume that they would respond the same way to stimulations. Lastly, the clinical data on modern archers describe soft tissue injuries and pain locations, however without clearer definitions as to the underlying biology, this information is difficult to transfer to analyses of dry bone. While there is obvious room for improvement in the field, the overall concept of linking activity-related skeletal changes and biomechanics to evaluate specialized activity remains promising.

## 5 | CONCLUSION

This study examined activity-related skeletal changes as a means for understanding Bell Beaker daily life and social organization with respect to archery. At a population level, differences in enthesal modifications between “archers” and “non-archers” exhibit a possible link between burial context and physical activity, such as manual labor. “Non-archer” burials were seemingly less prestigious, both in terms of quality and quantity of goods, and this corresponded to greater levels of enthesal changes potentially linked to physical activity. At an individual level, the data found that not all “archers” were likely specialized archers and that there is no correlation between a probable archer occupation and an archery burial context. This suggests that male Bell Beaker burials containing archery-related goods tended to reflect an individual’s social identity or role in society rather than their specialized, daily activities. Lastly, these findings support the premise that biological anthropologists can apply methods in osteology, biomechanics, and data analysis as a means for evaluating possible specialized activity, daily life, and behavior.

## AUTHOR CONTRIBUTIONS

**J. Despraz:** Conceptualization (lead); data curation (lead); formal analysis (lead); funding acquisition (lead); investigation (lead); methodology (lead); project administration (lead); visualization (lead); writing – original draft (lead); writing – review and editing (lead). **S. Villotte:** Formal analysis (supporting); methodology (supporting); supervision (lead); validation (supporting); writing – review and editing (supporting). **J. Desideri:** Conceptualization (supporting); project administration (supporting); supervision (lead); validation (supporting). **M. Besse:** Conceptualization (supporting); funding acquisition (supporting); project administration (lead); resources (supporting); supervision (lead); validation (equal); writing – review and editing (supporting).

## ACKNOWLEDGMENTS

This research was part of a PhD project entitled “The Application of Biomechanics and Bone Morphology to Interpret Specialized Activity and Social Stratification: The Case of Bell Beaker Archery” and financed by the Swiss National Science Foundation Doc.ch grant number P0GEP1_175169/1 (P.I. J. Ryan-Despraz). The University of Geneva Laboratory of prehistoric archaeology and anthropology also
provided additional funding. The authors would like to thank Karin Wiltschke-Schrotta (Natural History Museum of Vienna), Petr Veleminský (Prague National Museum), Zdeněk Tvrdý (Anthropos Institute of the Moravian Museum), and Mike Schweissing and George McGlynn (Bavarian State Collection of Anthropology and Palaeoanatomy) for providing access to the Bell Beaker skeletal collections. Clément Nicolas, Jan Turek, Volker Heyd, Lubomír Šebela, and Daniela Kern† provided additional assistance with the archaeological contexts. Lastly, the authors are grateful to Sylvain Sardy and Estella Poloni for their help with the data analysis. Open access funding provided by Universite de Geneve.

## DATA AVAILABILITY STATEMENT

Three additional Figures are submitted for the Supplementary Information. All data are available in open access on the University of Geneva's Archive Ouverte website for the original PhD thesis: https://archive-ouverte.unige.ch/unige:151360.

## ORCID

J. Ryan-Despraz https://orcid.org/0000-0002-0794-4317

S. Villotte https://orcid.org/0000-0002-2958-8034

J. Desideri https://orcid.org/0000-0003-1372-7420

M. Besse https://orcid.org/0000-0002-4752-9070

## REFERENCES

Abegg, C., Carloni, D., Cousseau, F., Derenne, E., & Ryan-Despraz, J. (2022). *The Bell Beaker culture in all its forms: Proceedings of the 22nd Meeting of 'Archéologie et Gobelets' 2021* (Geneva, Switzerland). *Archaeopress Archaeology*.

Açıkada, C., Ertan, H., & Tınazcı, C. (2004). Shooting dynamics in archery. In *sports medicine and science in archery* (pp. 15–36). Fédération Internationale de Tir à l'Arc.

Acosta, M. A., Henderson, C. Y., & Cunha, E. (2017). The effect of terrain on enthesal changes in the lower limbs. *International Journal of Osteoarchaeology*, 27, 828–838.

Acsádi, G., & Nemeskéri, J. (1970). *History of human life span and mortality*. Akadémiai Kiadó.

Agarwal, S. C., Dumitriu, M., Tomlinson, G. A., & Grynpas, M. D. (2004). Medieval trabecular bone architecture: The influence of age, sex, and lifestyle. *American Journal of Physical Anthropology*, 124(1), 33–44.

Alves Cardoso, F., & Henderson, C. Y. (2010). Enthesopathy formation in the humerus: data from known age-at-death and known occupation skeletal collections. *American Journal of Physical Anthropology*, 141, 550–560.

Andrews, J. R., & Whiteside, J. A. (1993). Common elbow problems in the athlete. *Journal of Orthopaedic & Sports Physical Therapy*, 17(6), 289–295.

Aufderheide, A., Rodríguez-Martin, C., & Langsjoen, O. (1998). *The Cambridge encyclopedia of human paleopathology* (Vol. 478). Cambridge University Press.

Axford, R. (1995). *Archery anatomy*. Souvenir Press.

Bailly, M., & Salanova, L. (1999). Les dates radiocarbone du Campaniforme en Europe occidentale: Analyse critique des principales séries de dates. In J. Evin (Ed.), *Actes du 3ème Congrès International 14C et Archéologie (Lyon, avril 1998)*, XXVI (pp. 219–224). Mémoires de la Société Préhistoriques Française.

Bálek, M., Dvořák, P., Kovárník, J., & Matějičková, A. (1999). Pohřebiště Kultury Zvoncovitých Pohárů v Tvoříhrázi (okr. Znojmo). In *Pravěk Supplementum* 4. Ústav Archeologiché Památkové Pěče Brno.

Becker, S. K. (2019). Osteoarthritis, entheses, and long bone cross-sectional geometry in the Andes: Usage, history, and future directions. *International Journal of Paleopathology*, 29, 45–53.

Bedáňová, A. (2023). Child archer-warrior and his ‘toys’ from the Bell Beaker cemetery in Popůvky. *Préhled Výzkumů*, 64(1), 85–94.

Belanger, T. A., & Rowe, D. E. (2001). Diffuse idiopathic skeletal hyperostosis: Musculoskeletal manifestations. *Journal of the American Academy of Orphopaedic Surgeons*, 9, 258–267.

Benjamin, M., Kumai, T., Milz, S., Boszczyk, B. M., Boszczyk, A. A., & Ralphs, J. R. (2002). The skeletal attachment of tendons–Tendon ‘entheses.’ *Comparative Biochemistry and Physiology Part A*, 133, 931–945.

Benjamin, M., & McGonagle, D. (2001). The anatomical basis for disease localisation in seronegative spondyloarthropathy at entheses and related sites. *The Journal of Anatomy*, 199, 503–526.

Benjamin, M., Moriggl, B., Brenner, E., Emery, P., McGonagle, D., & Redman, S. (2004). The “Enthesis Organ” Concept: Why Enthesopathies May Not Present as Focal Insertional Disorders. *Arthritis & Rheumatism*, 50(10), 3306–3313.

Berthon, W., Rittemard, C., Tihanyi, B., Pálfi, G., Coqueugniot, H., & Dutour, O. (2015). Three-dimensional microarchitecture of enthesal changes: Preliminary study of human radial tuberosity. *Acta Biologica Szegediensis*, 59, 79–90.

Besse, M. (2003). *L'Europe du 3e millénaire avant notre ère. Les céramiques communes au Campaniforme*. Cahiers d'archéologie romande.

Besse, M. (2014). Bell Beaker Common Ware During the Third Millennium BC in Europe. In J. Czebreszuk (Ed.), *Similar but different: Bell Beaker in Europe* (pp. 127–148). Sidestone Press.

Besse, M. (2015). Territorialité, transferts, interculturalités dans les contextes de la diffusion du Campaniforme en Europe. In N. Naudinot, L. Meignen, D. Binder, & G. Querré (Eds.), *Les Systèmes de Mobilité de la Préhistoire au Moyen Âge* (pp. 419–430). APDCA.

Biehler-Gomez, L., & Cattaneo, C. (2018). The diagnostic implications of two cases of known rheumatoid arthritis from the CAL Milano cemetery skeletal collection. *Journal of Forensic Sciences*, 63(6), 1880–1887.

Binford, L. R. (1972). Mortuary Practices: Their Study and Their Potential. In L. R. Binford (Ed.), *An archaeological perspective* (pp. 208–243). Seminar Press.

Blajerová, M. (1962). Anthropologické Zhodnocení Mladoeneolitických Kostrových Pohřbů z Libochovic, Žabovřesk a Štětí. *Sborník Severočeského Musea-Historia*, 3, 163–166.

Bocksberger, O. J. (1976). Le site préhistorique du Petit-Chasseur (Sion, Valais). 1 et 2. Le dolmen MVI, texte, catalogue et planches. *Cahiers d'Anthropologie Romande* 6.

Bourassa, D. C., McManus, I. C., & Bryden, M. P. (1996). Handedness and eye-dominance: A meta-analysis of their relationship. *Laterality*, 1(1), 5–34.

Bouvard, B., Mabilleau, G., Legrand, E., Audran, M., & Chappard, D. (2012). Micro and macroarchitectural changes at the tibia after botulinum toxin injection in the growing rat. *Bone*, 50(4), 858–864.

Bridges, P. S. (1991). Degenerative joint disease in hunter-gatherers and agriculturists from the Southeastern United States. *American Journal of Physical Anthropology*, 85, 379–391.

Brooks, S., & Suchey, J. M. (1990). Skeletal age determination based on the os pubis: A comparison of the Acsádi-Nemeskéri and Suchey-Brooks methods. *Human Evolution*, 5(3), 227–238.

Brůžek, J. (1991). *Fiabilité des procédés de détermination du sexe à partir de l'os coxal: Implications à l'étude du dimorphisme sexuel de l'homme fosible*. Muséum national d'histoire naturelle.

Bruzek, J. (2002). A Method for Visual Determination of Sex, Using the Human Hip Bone. *American Journal of Physical Anthropology*, 117, 157–168.

Brůžek, J., Santos, F., Dutailly, B., Murail, P., & Cunha, E. (2017). Validation and reliability of the sex estimation of the human os coxae using freely
available DSP2 software for bioarchaeology and forensic anthropology. *American Journal of Physical Anthropology*, 164(2), 440–449.

Buchvaldek, M. (1990). Pohřebiště lidu se zvoncovitými poháry. *Praehistorica*, 16, 29–49.

Buikstra, J., & Ubelaker, J. (1994). *Standard for data collection from human skeletal remains*. Arkansas Archaeological Survey Research Series No. 44.

Buuren, S. v., & Groothuis-Oudshoorn, K. (2011). mice: Multivariate impulation by chained equations in R. *Journal of Statistical Software*, 45(1) Article 1, 1–67.

Capasso, L., Kennedy, K. A. R., & Wilczak, C. A. (1999). *Atlas of occupational markers on human remains*. Edigrafital S.p.A.

Cashmore, L. A., & Zakrzewski, S. R. (2013). Assessment of musculoskeletal stress marker development in the hand. *International Journal of Osteoarchaeology*, 23, 334–347.

Chapman, J. (2015). The Balkan neolithic and chalcolithic. In C. Fowler, J. Harding, & D. Hofmann (Eds.), *The Oxford handbook of neolithic Europe* (pp. 157–174). Oxford University Press.

Chen, D., Shen, J., Zhao, W., Wang, T., Han, L., Hamilton, J. L., & Im, H.-J. (2017). Osteoarthritis: Toward a comprehensive understanding of pathological mechanism. *Bone Research*, 5, 16044.

Christlein, R. (1982). Landesarchäologie in Bayern 1981–Ein Überblick. In R. Christlein (Ed.), *Das archäologie Jahr in Bayern 1981* (pp. 13–38). Konrad Theiss Verlag Stuttgart.

Clarys, J. P., Cabri, J., Bollens, E., Sleeckx, R., Taeymans, J., Vermeiren, M., Van Reeth, G., & Voss, G. (1990). Muscular activity of different shooting distances, different release techniques, and different performance levels, with and without stabilizers, in target archery. *Journal of Sports Sciences*, 9, 235–257.

Coqueugniot, H., & Weaver, T. (2007). Brief communication: infracranial maturation in the skeletal collection From Coimbra, Portugal: New aging standards for epiphyseal union. *American Journal of Physical Anthropology*, 134, 424–437.

Corboud, P. (2009). Les stèles anthropomorphes de la nécropole néolithique du Petit-Chasseur à Sion. *Bulletin d'études Préhistoriques et Archéologiques Alpines*, 20, 1–89.

Courteix, D., Lespessailles, E., Peres, S. L., Obert, P., Germain, P., & Benhamou, C. L. (1998). Effect of physical training on bone mineral density in prepubertal girls: A comparative study between impact-loading and non-impact-loading sports. *Osteoporosis International*, 8(2), 152–158.

Cowgill, L. (2018). Cross-sectional geometry. In *The international encyclopaedia of biological anthropology* (pp. 1–3). John Wiley & Sons, Ltd.

Czebreszuk, J. (Ed.). (2014). *Similar but different. Bell Beakers in Europe*. Sidestone Press.

De Gattis, G., Curdy, P., Ferroni, A. M., Martinet, F., Poggiani Keller, R., Raiteri, L., Sarti, L., Zidda, G., & Mezzena, F. (2018). *Area megalitica di Saint-Martin-de-Corléans. Una visione aggiornata*. Le Château Edizioni (Documenti 13).

Deymier, A. C., Schwartz, A. G., Cai, Z., Daulton, T. L., Pasteris, J. D., Genin, G. M., & Thomopoulos, S. (2019). The multiscale structural and mechanical effects of mouse supraspinatus muscle unloading on the mature enthesis. *Acta Biomaterialia*, 83, 302–313.

Ding, M., Odgaard, A., Danielsen, C. C., & Hvid, I. (2002). Mutual associations among microstructural, physical and mechanical properties of human cancellous bone. *The Journal of Bone and Joint Surgery*, 84(6), 8.

Drozdová, E., Balueva, T. S., Veselovskaya, E., Smrčka, V., Benešová, J., Bůzek, F., Erban, V., Kanický, V., Ovesná, P., Nejechzebová, H., Vaňharová, M., Zocová, J., & Matějičková, A. (2011). *Hoštice I za Hanou*. Masarykova univerzita.

Dutour, O. (1986). Enthesopathies (Lesions of Muscular Insertions) as Indicators of the Activities of Neolithic Saharan Populations. *American Journal of Physical Anthropology*, 71(2), 221–224.

Dutour, O. (1992). Activités physiques et squelette humain: Le difficile passage de l'actuel au fossile. *Bulletins et Mémoires de La Société d'Anthropologie de Paris*, 4(3-4), 233-241.

Dvořák, P. (1992). *Die Gräberfelder der Glockenbecherkultur in Mähren I (Bez Blansko, Brno-město, Brno-venkov)*. Katalog der Funde (pp. 1-124). Mährische Archäologische Quellen.

Dvořák, P., Matějičková, A., Peška, J., & Rakovský, I. (1996). Gräberfelder Der Glockenbecherkultur in Mähren II: Katalog der Funde. Petr Dvořák Verlag Brno.

Ertan, H., Kentel, B., Tümer, S. T., & Korkusuz, F. (2003). Activation patterns in forearm muscles during archery shooting. *Human Movement Science*, 22, 37-45.

Faillace, K. E., Bethard, J. D., & Marks, M. K. (2017). The applicability of dental wear in age estimation for a modern American population. *American Journal of Physical Anthropology*, 164(4), 776–787.

Faurie, C., & Raymond, M. (2004). Handedness frequency over more than ten thousand years. *Proceedings of the Royal Society of London B: Biological Sciences*, 271, S43-S45.

Fitzpatrick, A. P. (2011). *The Amesbury Archer and the Boscombe Bowmen: Bell Beaker burials at Boscombe Down, Amesbury. Wiltshire*.

Fridrichová, M., Fridrich, J., Havel, J., & Kovářík, J. (1995). Praha v Pravěku. Archaeologica Pragensia Supplementum (2).

Friesinger, I. (1976). Glockenbecherzeitliche Grabfunde aus Henzing, Gemeinde Siegharts-kirchen, pol. Bez. Tulln, Niederösterreich. *Annalen Des Naturhistorischen Museums in Wien*, 80, 823–828.

Frostick, S. P., Mohammad, M., & Ritchie, D. A. (1999). Sports injuries of the elbow. *British Journal of Sports Medicine*, 33, 301–311.

Furholt, M. (2020). Mondes sociaux et communautés de pratique: Un modèle culturel polythétique pour l'Europe du IIe millénaire avant notreère à la lumière des débats actuels sur la migration. *Préhistoire Méditerranéennes*, 8, Article 8, 1–26.

Gallay, A. (1979). Le phénomène campaniforme: Une nouvelle hypothèse historique. *Archives suisses d'anthropologie générale*, 43(2), 231-258.

Geisler, M. (1990). Hrob Kultury se Zvoncovitými Poháhy z Rousínovce (pp. 119–127). *Pravěké a Slovanské Osílení Moravy*.

Goodman, A. H., & Martin, D. L. (2002). Reconstructing health profiles from skeletal remains. In R. H. Steckel & J. C. Rose (Eds.), *The Backbone of History. Health and Nutrition in the Western Hemisphere* (pp. 11–60). Cambridge University Press.

Gopal Adkitte, R., Shah, S., Jain, S., Walla, S., Chopra, N., & Kumar, H. (2016). Common injuries amongst Indian elite archers: A prospective study. *Saudi Journal of Sports Medicine*, 16(3), 210–213.

Griffin, N. L., D'Août, K., Ryan, T. M., Richmond, B. G., Ketcham, R. A., & Postnov, A. (2010). Comparative forefoot trabecular bone architecture in extant hominids. *Journal of Human Evolution*, 59(2), 202–213.

Grupe, G., Price, T. D., Schröter, P., Söllner, F., Johnson, C. M., & Beard, B. L. (1997). Mobility of Bell Beaker people revealed by strontium isotope ratios of tooth and bone: A study of southern Bavarian skeletal remains. *Applied Geochemistry*, 12(4), 517–525.

Hájek, L. (1962). Die Glockenbecherkultur in Böhmen und Mähren. In M.-E. Mariën (Ed.), *Inventaria Archaeologica: Corpus des Ensembles Archologiques* (Vol. 2, p. ČS 11–ČS 20). Rudolf Habelt Verlag/Bonn.

Hájek, J. (1968a). Archeologické studijní materiály: Die Glockenbecherkultur in Böhmen. *Archeologický Ústav ČSAV*.

Hájek, J. (1968b). Kultura zvoncovitých pohárů v (p. 5). Die Glockenbecherkultur in Bohmen. *Archeologické Studijní Materiály*.

Hájek, L. (1966). Die älteste Phase der Glockenbecherkultur in Böhmen und Mähren. *Pámátky Archeologické*, 1, 210–241.

Harrison, R. J. (1980). *The Beaker Folk: Copper Age archaeology in Western Europe* (Vol. 97). Thames and Hudson.

Havelková, P., Villotte, S., Velemínský, P., Poláček, L., & Dobisíková, M. (2011). Enthesopathies and activity patterns in the early medieval great moravian population: Evidence of division of labour. *International Journal of Osteoarchaeology*, 21, 487–504.
Hawkey, D. E., & Merbs, C. F. (1995). Activity-induced musculoskeletal stress markers (MSM) and subsistence strategy changes among ancient Hudson Bay Eskimos. *International Journal of Osteoarchaeology*, 5, 324-338.

Henderson, C. Y., Mariotti, V., Pany-Kucera, D., Villotte, S., & Wilczak, C. (2013). Recording specific enthesal changes of fibrocartilaginous entheses: Initial tests using the Coimbra method. *International Journal of Osteoarchaeology*, 23, 152-162.

Henderson, C. Y., Mariotti, V., Pany-Kucera, D., Villotte, S., & Wilczak, C. (2016). The New “Coimbra Method”: A biologically appropriate method for recording specific features of fibrocartilaginous enthesal changes. *International Journal of Osteoarchaeology*, 26, 925-932.

Heyd, V. (2000). *Die Spätkupferzeit in Süddeutschland* (Vol. 73). Rudolph Habelt GmbH.

Heyd, V. (2001). On the earliest Bell Beakers along the Danube. In F. Nicoslis (Ed.), *Bell Beakers today: Pottery, people, culture, symbols in prehistoric Europe* (pp. 387-410). Provincia Autonoma di Trento Servizio Beni Culturali Ufficio Beni Archeologici.

Heyd, V. (2007). Families, Prestige Goods, Warriors and Complex Societies: Beaker Groups of the 3rd Millennium cal BC Along the Upper and Middle Danube. *Prehistoric Society London. Proceedings*, 73, 327-379.

Hildenbrand, J. C., & Rayan, G. M. (2010). Archery. In D. Caine, P. Harmer, & M. A. Schiff (Eds.), *Epidemiology of Injury in olympic sports* (Vol. XVI, pp. 18-25). Wiley-Blackwell.

Honda, A., Matsumoto, M., Kato, T., & Umemura, Y. (2015). Exercise characteristics influence femoral cross-sectional geometry: A magnetic resonance imaging study in elite female athletes. *Osteoporos International*, 26, 1093-1098.

Hu, C. M., & Tang, W. T. (2005). Muscle activation by olympic female archers at different releasing rhythms. In *20th International Society of Biomechanics Congress, Poster*. 20th International Society of Biomechanics Congress, Cleveland.

Huiskes, R., Ruimerman, R., van Lenthe, G. H., & Janssen, J. D. (2000). Effects of mechanical forces on maintenance and adaptation of form in trabecular bone. *Nature*, 405(6787), 704-706.

Husson, F., & Josse, J. (2014). Multiple correspondence analysis. In J. Blasius & M. Greenacre (Eds.), *The Visualization and Verbalization of Data*. CRC Press.

Husty, L. (1993). Ein Gräberfeld der Glockenbecherkultur in Landau a.d. *Isar. Das Archäologische Jahr in Bayern*, 1992, 48-49.

Jones, H. H., Priest, J. D., Hayes, W. C., Chinn Tichenor, C., & Nagel, D. A. (1977). Humeral hypertrophy in response to exercise. *The Journal of Bone and Joint Surgery*, 59A, 204-208.

Josse, J., Chavent, M., Liquet, B., & Husson, F. (2012). Handling missing values with regularized iterative multiple correspondence analysis. *Journal of Classification*, 29(1), 91-116.

Jungwirth, J. (1976). Vier spätneolithische Skelette aus Henzing, Gemeinde Sieghartskirchen, Niederösterreich. *Annalen Des Naturhistorischen Museums in Wien*, 80, 829-842.

Jurmain, R., Alves Cardoso, F., Henderson, C., & Villotte, S. (2012). *Bioarchaeology's holy grail: The reconstruction of activity*. In A. L. Grauer (Ed.), *A Companion to Paleopathology* ((1st ed.). ed.). Blackwell Publishing Ltd.

Katitas, A. (2019). *Getting Started with Multiple Imputation in R*. University of Virginia Library- Statlab Articles https://uvastatlab.github.io/2019/05/01/getting-started-with-multiple-imputation-in-r/.

Kaymak, B., Özçakar, L., Ertan, H., Karabulut, I., & Akinci, A. (2012). Sono-graphic assessment of finger flexor tendons in olympic archers. *Turkish Journal of Physical Medicine and Rehabilitation*, 58, 85-87.

Kaynaroğlu, V., & Kiliç, Y. A. (2012). Archery-related sports injuries. In M. N. Doral & J. Karlsson (Eds.), *Sports injuries: Prevention, diagnosis, treatment and rehabilitation* (pp. 1081-1086). Springer-Verlag.

Kennedy, K. (1998). Markers of occupational stress: conspectus and prognosis of research. *International Journal of Osteoarchaeology*, 8, 305-310.

Kern, D. (2003). Überlegungen zum Nachweis von Mobilität und Migration in Ostösterreich zur Zeit der Glockenbecher. *Mitteilungen Der Anthropologischen Gesellschaft in Wien*, 133, 73-83.

Kern, D. (2011). Äneolithische und frühbronzezeitliche Hügelgräber in Ostösterreich. In H. Kowalewska-Marszałek & P. Włodarczak (Eds.), *Kurhany i obrządek pogrzebowy w IV-II tysiącleciu p.n.e* (pp. 161-168). Inst. Archeologii i Etnologii Polskiej Akad. Nauk.

Kern, D. (2016a). About bow-shaped and rod-shaped pendants. *Musaica Archaeologica*, 1, 95-104.

Kern, D. (2016b). Von stäbchenförmigen Anhängern und Knebeln (in Mitteleuropa). In M. Bartelheim, B. Horejs, & R. Krauss (Eds.), *Von Baden bis Troia: Ressourcennutzung, Metallurgie und Wissenstransfer. Verlag Marie Leidorf GMBH*.

Kern, D., & Wiltschke-Schrotta, K. (in press). *Einespätestendneolithische Gräbergruppe aus Tödling, Oberösterreich*. Unpublished manuscript.

Khan, M. A. (2002). Ankylosing spondylitis: Introductory comments on its diagnosis and treatment. *Annals of Rheumatic Diseases*, 61(Suppl III), iii3-iii7.

Kociumaka, C., & Dietrich, H. (1992). Ein Gräberfeld der Glockenbecherkultur vom Sportgelände der Universität Augsburg. *Das Archäologische Jahr in Bayern*, 1991, 67-68.

Kosar, N. S., & Demirel, H. A. (2004). Kinesiological analysis of archery. In E. Ergen & K. Hibner (Eds.), *Sports medicine and science in archery* (pp. 3-12). Fédération Internationale de Tir à l'Arc.

Kreiner, L. (1991). Drei neue Gräber der Glockenbecherkultur aus Trieching, Gde. Pilsting, Lkr. Dingolfing-Landau. *Vorträge Des 9. Niederbayerischen Archäologentages*, 1991, 131-136.

Kytlicová, O. (1956). Pohřebiště kultury zvoncovitých pohárů vKněževsi. In J. Filip (Ed.), *Archeologické rozhledy* (Vol. 3, pp. 328-349). Československá Akademie Věd.

Kytlicová, O. (1960). Eneolitické Pohřebiště v Brandýsku. *Památky Archeologické*, 51(2), 442-474.

Laborde, S., Dosseville, F. E. M., Leconte, P., & Margas, N. (2009). Interaction of Hand Preference with Eye Dominance on Accuracy in Archery. *Perceptual and Motor Skills*, 108(2), 558-564.

Lapostolle, J. C. (2004). Elbow pathologies in archery. In E. Ergen & K. Hibner (Eds.), *Sports medicine and science in archery* (pp. 70-81). Fédération Internationale de Tir à l'Arc.

Lapostolle, J. C., Hibner, K., & Ergen, E. (2004). FITA medical committee archery injuries survey (Juniors). In E. Ergen & K. Hibner (Eds.), *Sports medicine and science in Archery* (pp. 65-69). Fédération Internationale de Tir à l'Arc.

Larsen, C. S. (1997). *Bioarchaeology: Interpreting behavior from the human skeleton*. Cambridge University Press.

Lee, K. J., & Simpson, J. A. (2014). Introduction to multiple imputation for dealing with missing data. *Respirology*, 19(2), 162-167.

Lemercier, O. (1998). The Bell Beaker phenomenon in the southeast of France: The state of research and preliminary remarks about the TGV excavations and some other sites of the Provence. *Proceedings of the 2nd Meeting of the "Association Archéologie et Gobelets"* 23-41.

Lemercier, O., Furestier, R., Müller, A., Blaise, E., Bouville, C., Convertini, F., & Salanova, L. (2022). Chapitre XI. La sépulture individuelle campaniforme de La Fare, Forcalquier (Alpes-de-Haute-Provence). In Y. Tcheremissinoff (Ed.), *Les sépultures individuelles campaniformes en France* (pp. 145-159). CNRS Éditions.

Lieberman, D. E., Polk, J. D., & Demes, B. (2004). Predicting long bone loading from cross-sectional geometry. *American Journal of Physical Anthropology*, 123(2), 156-171.

Littke, N. (2004). Shoulder injuries: A rehab perspective. In E. Ergen & K. Hibner (Eds.), *Sports medicine and science in archery* (pp. 82-86). Fédération Internationale de Tir à l'Arc.

Llaurens, V., Raymond, M., & Faurie, C. (2009). Why are some people left-handed? An evolutionary perspective. *Philosophical Transactions of the Royal Society B*, 364, 881-894.
Maat, G. J. R., Mastwijk, R. W., & van der Velde, E. A. (1995). Skeletal distribution of degenerative changes in vertebral osteophytosis, vertebral osteoarthritis and DISH. *International Journal of Osteoarchaeology*, 5(3), 289-298.

Macintosh, A., Davies, T., Ryan, T., Shaw, C., & Stock, J. (2013). Periosteal versus true cross-sectional geometry: A comparison along humeral, femoral, and tibial diaphyses. *American Journal of Physical Anthropology*, 150, 442-452.

Maggiano, I., Schultz, M., Kierdorf, H., Sierra Sosa, T., Maggiano, C., & Tiesler Blos, V. (2008). Cross-sectional analysis of long bones, occupational activities and long-distance trade of the classic Maya from Xcambó-archaeological and osteological evidence. *American Journal of Physical Anthropology*, 136, 470-477.

Mann, R. W., & Hunt, D. R. (2005). *Photographic regional atlas of bone disease: A guide to pathologic and normal variation in the human skeleton* (3rd ed.). Charles C. Thomas.

Mann, D., & Littke, N. (1989). Shoulder injuries in archery. *Canadian Journal of Sports Science*, 14(2), 85-92.

Martin, P. E., Siler, W. L., & Hoffman, D. (1990). Electromyographic analysis of bow string release in highly skilled archers. *Journal of Sports Sciences*, 8(3), 215-221.

Mariotti, V., Facchini, F., & Giovanna Belcastro, M. (2004). Enthesopathies – proposal of a standardized scoring method and applications. *Collegium Antropologicum*, 28(1), 145-159.

Mariotti, V., Facchini, F., & Giovanna Belcastro, M. (2007). The study of entheses: Proposal of a standardised scoring method for twenty-three entheses of the postcranial skeleton. *Collegium Antropologicum*, 31(1), 291-313.

Martin, R., & Saller, K. (1957). Lehrbuch der Anthropologie. In *Systematischer Darstellung mit Besonderer Berücksichtigung der Anthropologischen Methoden* (Vol. 1). Gustav Fischer Verlag.

Matějíčková, A. (2007). Archaeological and anthropological analysis of the Bell beaker cemetery at Hoštice 1. In *Beaker Days 2007–Sicily [Lecture]*. Beaker Days.

Mays, S. (1998). *The archaeology of human bones*. Taylor & Francis e-Library.

Mays, S. (2002). The relationship between molar wear and age in an early 19th century AD archaeological human skeletal series of documented age at death. *Journal of Archaeological Science*, 29(8), 861-871.

Merbs, C. F. (1983). Patterns of activity-induced pathology in a Canadian Inuit population. *Archaeological Survey Canada*, 119, 120-128.

Molnar, P. (2006). Tracing prehistoric activities: Musculoskeletal stress marker analysis of a stone-age population on the island of gotland in the Baltic Sea. *American Journal of Physical Anthropology*, 129, 12-23.

Moucha, V. (2005). Pohřebišťé Lidu s Kulturou se Zvoncovitými Poháry ve Lhánicích (okr. Třebíč). *Pravěk NŘ*, 15, 25-58.

Müller, A. (2001). Gender Differentiation in burial rites and grave-goods in the Eastern or Bohemian-Moravian Group of the Bell Beaker Culture. In F. Nicolis (Ed.), *Bell Beakers today: Pottery, people, culture, symbols in prehistoric Europe* (pp. 589-599). Provincia Autonoma di Trento Servizio Beni Culturali Ufficio Beni Archeologici.

Müller, J., & van Willigen, S. (2001). New radiocarbon evidence for european Bell Beakers and the consequences for the diffusion of the Bell Beaker Phenomenon. In F. Nicolis (Ed.), *Bell Beaker today: Pottery, people, culture, symbols in prehistoric Europe* (pp. 59-80). Provincia Autonoma di Trento Servizio Beni Culturali Ufficio Beni Archeologici.

Murail, P., Brůžek, J., Houët, F., & Cunha, E. (2005). DSP: A Tool for Probabilistic Sex Diagnosis Using Worldwide Variability in Hip-Bone Measurements. *Bulletins et Mémoires de La Société d'Anthropologie de Paris*, 17, 167-176.

Muška, J. (1981). The Settlement and Cemetery Sites of the Bell Beaker Culture at Radovesice by Bílina. In *Archaeological News in The Czech Socialist Republic, Xe Congrès International des Sciences Préhistoriques et Protohistoriques, Mexico 1981* (p. 51). Institut d'archéologie de l'Académie des sciences tchécoslovaque.

Neugebauer, C., & Neugebauer, J.-W. (1997). Zum Forschungsstand der Glockenbecherkultur in Ostösterreich. In B. Fritsch, M. Maute, J. Müller, & C. Wolf (Eds.), *Tradition and Innovation, Prähistorische Archäologieals historische Wissenschaft, Festschrift für Christian Strahm* (pp. 307-324). Internationale Archäologie, Studia Honoria 3. Rahden.

Neugebauer, J., & Neugebauer-Maresch, C. (2001). Bell Beaker Culture in Austria. In F. Nicolis (Ed.), *Bell Beakers Today: Pottery, People, Culture, Symbols in Prehistoric Europe* (Vol. II, pp. 429-437). Provincia Autonoma di Trento Servizio Beni Culturali Ufficio Beni Archeologici.

Neugebauer, J.-W., & Gattringer, A. (1982). Die Kremser Schnellstrasse S 33: Zweiter Vorbericht über Ergebnisse der archäologischen Überwachung des Grossbauvorhabens durch die Abt. F. Bodendenkmale des Bundesdenkmalamtes im Jahre 1982. In H. Adler (Ed.), *Fundberichte aus Österreich: Herausgegeben vom Bundesdenkmalamt* (Vol. 21, pp. 63-95). Verlag Ferdinand Berger & Söhne.

Nicolas, C. (2019). The production and use of archery related items as a reflection of social changes during the Late Neolithic and the Early Bronze Age in Europe. In S. Kadrow & J. Müller (Eds.), *Habitus: The social dimension of technology and transformation* (pp. 115-139). Sidestone Press.

Nicolas, C. (2020). Bracer ornaments! An investigation of Bell Beaker stone "wrist-guards" from Central Europe. *Journal of Neolithic Archaeology*, 22, 15-107.

Nicolis, F. (Ed.). (2001). *Bell Beakers today. Pottery, people, cultures, symbols in prehistoric Europe*. Edizioni All'insegna del Giglio.

Niinimäki, S., Härkönen, L., Nikander, R., Abe, S., Knüsel, C., & Sievänen, H. (2016). The cross-sectional area of the gluteus maximus muscle varies according to habitual exercise loading: Implications for activity-related and evolutionary studies. *HOMO-Journal of Comparative Human Biology*, 67, 125-137.

Nikita, E., Siew, Y. Y., Stock, J., Mattingly, D., & Mirazón Lahr, M. (2011). Activity patterns in the Sahara Desert: An interpretation based on cross-sectional geometric properties. *American Journal of Physical Anthropology*, 146, 423-434.

Nikita, E., Xanthopoulou, P., Bertsatos, A., Chovalopoulou, M.-E., & Hafez, I. (2019). A three-dimensional digital microscopic investigation of enthesal changes as skeletal activity markers. *American Journal of Physical Anthropology*, 169(4), 704-713.

Ogilvie, M., & Hilton, C. (2011). Cross-sectional geometry in the Humeri of foragers and farmers from the Prehispanic American Southwest: Exploring patterns in the sexual division of labor. *American Journal of Physical Anthropology*, 144, 11-21.

Naraen, A., Giannikas, K. A., & Livesley, P. J. (1999). Overuse epiphyseal injury of the coracoid process as a result of archery. *International Journal of Sports Medicine*, 20(1), 53-55.

Niestroj, C. K., Schöffl, V., & Küpper, T. (2018). Acute and overuse injuries in elite archers. *The Journal of Sports Medicine and Physical Fitness*, 58 (7-8), 1063-1070.

Nishizono, H., Shibayama, H., Izuta, T., & Saito, K. (1987). Analysis of archery shooting techniques by means of electromyography. *International Society of Biomechanics in Sports, Symposium V*, 364-372.

Oladie, I., Brace, S., Allentoft, M. E., Armit, I., Kristiansen, K., Booth, T., Rohland, N., Mallick, S., Szécsényi-Nagy, A., Mittnik, A., Altena, E., Lipson, M., Lazaridis, I., Harper, T. K., Patterson, N., Broomandkhoshbacht, N., Diekmann, Y., Faltyskova, Z., Fernandes, D., ... Reich, D. (2018). The Beaker phenomenon and the genomic transformation of northwest Europe. *Nature*, 555(7695) Article 7695, 190-196.

Olivik, J. (2009a). *Sillicitové šipky a nátepní destíčky – jejich typologie a postavení v pohřebním ritu kultury zvoncovitých pohárů na Moravě* [Master's]. Masarykova Univerzita.

Olivik, J. (2009b). Studies on Bell Beaker lithic industry in Moravia. Arrowheads. In J. Kopacz, A. Přichystal, & L. Šebela (Eds.), *Lithic chipped industry of the Bell Beaker culture in Moravia*. Annex A. Polish Academy of Arts and Sciences.
Ondráček, J. (1961). Příspěvky k Poznání Kultury Zvoncovitých Poháhů na Moravě. PA, 52, 149–157.

Pálfi, G., & Dutour, O. (1996). Activity-induced skeletal markers in historical anthropological material. *International Journal of Anthropology*, *11*(1), 41–55.

Pearson, O. M., & Lieberman, D. E. (2004). The aging of Wolff's "Law": Ontogeny and Responses to mechanical loading in cortical bone. *Yearbook of Physical Anthropology*, 47, 63–99.

Perreard Lopreno, G. (2007). *Adaptation structurelle des os du membre supérieur et de la clavicule à l'activité* [PhD, University of Geneva]. https://doi.org/10.13097/archive-ouverte/unige:18785

Peška, J. (2013a). Morava na konci eneolitu. Olomouc.

Peška, J. (2013b). Two new burial sites of Bell Beaker Culture with an exceptional find from Eastern Moravia/Czech Republic. In M. P. P. Marínez & L. Salanova (Eds.), *Current Researches on Bell Beakers: Proceedings of the 15th International Bell Beaker Conference: From Atlantic to Ural. 5th-9th May 2011 Poio (Ponteverde, Galicia, Spain)* (pp. 61–72). Copynino-Centro de Impresión Digital.

Peškař, I. (1968). Hrob kultury zvoncovitých pohárů v Bně-Julianově (pp. 19–20). Ein Grab der Glockenbecherkultur in Brno-Julianov. *Přehled Výzkumů*.

Porac, C., & Coren, S. (1981). *Lateral Preferences and human behavior*. Springer-Verlag.

Prammer, J. (1981). Ausgrabungen und Funde im Raum Straubing 1981. *Jahresbericht Des Historischen Vereins Für Straubing Und Umgebung*, 83, 7–20.

Prine, B., Prine, A., Leavitt, T., Wasser, J., & Vincent, H. (2016). Prevalence and characteristics of archery-related injuries in archers: 3083 Board #148 June 3. *Medicine & Science in Sports & Exercise*, 48, 874.

Puri, V., & Sethi, A. (2017). Determination of ocular dominance and its association with handedness. *Indian Journal of Clinical and Experimental Ophthalmology*, 3(4), 504–506.

Rakovský, I. (1985). Pohřebiště Kultury Zvoncovitých Pohárů v Holubicích. *Archeologické Rozhledy*, XXXVII, 393–402.

Rayan, G. M. (1992). Archery-related injuries of the hand, forearm, and elbow. *Southern Medical Journal*, 85, 961–964.

Reich, D. (2019). Who We Are and How We Got Here: Ancient DNA and the new science of the human past. Oxford University Press.

Resnick, D., & Niwayama, G. (1978). Radiographic and pathologic features of spinal involvement in diffuse idiopathic skeletal hyperostosis (DISH). *Diagnostic Radiology*, 119(3), 559–568.

Rhodes, J., & Knüsel, C. (2005). Activity-related skeletal change in medieval humeri: Cross-sectional and architectural alterations. *American Journal of Physical Anthropology*, 128(3), 536–546.

Rieder, K. H. (1983). Ein Bestattungsplatz der Glockenbecherzeit aus Oberstimm, Gemeinde Manching, Landkreis Pfaffenhofen, Oberbayern. *Das Archäologische Jahr in Bayern*, 1982, 40–42.

Robison, S. E., Jacobsen, S., & Heintz, B. (1995). Hand-eye cross dominance: Poster # 38 (BV-334). *Optometry and Vision Science*, 72(12), 210.

Rogers, J., & Waldron, T. (1995). *A Field Guide to Joint Disease in Archaeology*. John Wiley & Sons.

Rothschild, B. M., Woods, R. J., & Ortel, W. (1990). Rheumatoid arthritis "in the buff": Erosive arthritis in defleshed bones. *American Journal of Physical Anthropology*, 82(4), 441–449.

Ruff, C. B., Walker, A., & Trinkaus, E. (1994). Postcranial robusticity in Homo. III: ontogeny. *American Journal of Physical Anthropology*, 93, 35–54.

Růžičková, P. (2008). Lukovité závěsky kultury zvoncovitých pohárů ve střední Evropě [Bachelor Thesis]. Masaryk University.

Růžičková, P. (2009). Bow-Shaped Pendants of the Bell Beaker Culture. In *Acta Archaeologica Carpathica* (Vol. XLIV, pp. 37–72). Polish Academy of Sciences Publishing House in Cracow.

Ryan, J., Desideri, J., & Besse, M. (2018). Bell Beaker Archers: Warriors or an Ideology? *Journal of Neolithic Archaeology*, 20(2018S), 97–122.

Ryan, T. M., & Shaw, C. N. (2012). Unique suites of trabecular bone features characterize locomotor behavior in human and non-human anthropoid primates. *PLOS ONE*, 7(7), e41037.

Ryan-Despraz, J. (2021). *The application of biomechanics and bone morphology to interpret specialized activity and social stratification: The case of bell beaker archery* [PhD thesis]. University of Geneva.

Ryan-Despraz, J. (2022). *Practice and prestige: An exploration of neolithic warfare, Bell beaker archery, and social stratification from an anthropological perspective*. Archaeopress Archaeology.

Ryan-Despraz, J. (2023). Archery and the arm. In C. S. Hirst, R. Gilmour, K. Plomp, & F. Alves Cardoso (Eds.), *Behaviour in our bones* (pp. 109–134). Elsevier.

Ryan-Despraz, J., & Nicolas, C. (2022). Examining the role of Bell Beaker archery through techno-functional and osteological analyses: reinvestigating the hoštice cemetery (Czech Republic). In C. Abegg, D. Carloni, F. Cousseau, E. Derenne, & J. Ryan-Despraz (Eds.), *The Bell Beaker culture in all its forms: Proceedings of the 22nd Meeting of 'Archéologie et Gobelets' 2021 (Geneva, Switzerland)* (pp. 191–207). Archaeopress Archaeology.

Salanova, L. (2016). Behind the warriors: Bell Beakers and identities in Atlantic Europe (Third Millennium B.C.). In *Celtic from the West 3: Atlantic Europe in the Metal Ages questions of shared language* (pp. 13–34). Oxbow Books.

Salanova, L. (2022). Chapitre X. Chronologie et facteurs d'évolution des sépultures individuelles campaniformes dans le Nord de la France. In Y. Tcheremissinoff (Ed.), *Les sépultures individuelles campaniformes en France* (pp. 125–142). CNRS Éditions.

Salter, D. M. (2002). Degenerative joint disease. *Current Diagnostic Pathology*, 8, 11–18.

Schaefer, M., Black, S., & Scheuer, L. (2009). *Juvenile osteology: A laboratory field manual*. Elsevier Inc.

Schmitt, A. (2001). *Variabilité de la sénescence du squelette humain. Réflexions sur les indicateurs de l'âge au décès: À la recherche d'un outil performant.* [PhD, Université Sciences et Technologies- Bordeaux I]. https://tel.archives-ouvertes.fr/tel-00255753

Schmitt, A. (2005). Une nouvelle méthode pour estimer l'âge au décès des adultes à partir de la surface sacro-pelvienne iliaque. *Bulletins et Mémorières de La Société d'Anthropologie de Paris*, 17(1-2), 89-101.

Schmotz, K. (1990). Eine Gräbergruppe der Glockenbecherkultur von Altenmarkt. *Arch. Jahr Bayern*, 1989, 59–60.

Schmotz, K. (1994). Eine Gräbergruppe der Glockenbecherkultur von Osterhofen-Altenmarkt. *Archäologische Denkmaler im Landkreis Degendorf*, 9, 5–28.

Schrader, S. (2019). Bioarchaeological approaches to activity reconstruction. In S. Schrader (Ed.), *Activity, diet and social practice: Addressing everyday life in human skeletal remains* (pp. 55–126). Springer International Publishing.

Schwartz, J. H. (2007). *Skeleton keys: An introduction to human skeletal morphology, development, and analysis* (2nd ed.). Oxford University Press.

Serneels, S., & Verdonck, T. (2008). Principal component analysis for data containing outliers and missing elements. *Computational Statistics & Data Analysis*, 52(3), 1712–1727.

Sessa, E. (1994). Etude de la biomécanique du tir à l'arc: Incidence sur la pathologie et la rééducation. *Annales de Kinésithérapie*, 21(8), 435–442.
Shbat, A. (2013). Zdravotní stav populací neolitu a eneolitu na základě studia kosterního materiálu z Čech [PhD, Charles University]. https://dspace.cuni.cz/handle/20.500.11956/59871

Singh, A., & Lhee, S. H. (2016). Injuries in archers. *Saudi Journal of Sports Medicine*, 16(3), 168.

Sjögren, K.-G., Olalde, I., Carver, S., Allentoft, M. E., Knowles, T., Kroonen, G., Pike, A. W. G., Schröter, P., Brown, K. A., Robson-Brown, K., Harrison, R. J., Bertemes, F., Reich, D., Kristiansen, K., & Heyd, V. (2019). Kinship and social organization in Copper Age Europe. A cross-disciplinary analysis of archaeology, DNA, isotopes, and anthropology from two Bell Beaker cemeteries. *BioRxiv*, 863944.

Skružný, L., Turek, J., & Vojtěchovská, I. (2000). Pohřrebiště z Období Zvoncovitých Pohárů ve Velkých Přílepech—Kamýku, okr. Praha-Západ. *Archeologie ve Středních Čechách*, 4, 59–78.

Šlaus, M., Novak, M., & Čavka, M. (2012). Four cases of ankylosing spondylitis in medieval skeletal series from Croatia. *Rheumatology International*, 32, 3985–3992.

Sosna, D. (2012). Stone arrowheads from Hoštice-I: Use-wear analysis. *Supplementum*, 24(1), 323–338.

Souský, B. (1954). Eneolitická hrazená osada v Praze-Kobylišis. *Archeologické Rozhledy*, VI, 142–147.

Spurný, V. (1957). Hroby lidu se zvoncovitými poháry v Roštíně u Kroměříže. *Archeologické Rozhledy*, IX, 415–416.

Staňa, Č. (1960). Hrob kultury zvoncovitých pohárů ve Veselí n.Mor. In *Přehled výzkumů* 1959 (pp. 32–35). Archeoogický Ústav Československé Akademie Věd.

Steckel, R. H., Larsen, C. S., Sciulli, P. W., & Walker, P. L. (2005). *Data collection Codebook*. Cambridge University Press.

Stevenson, J., Mahoney, E., Walker, P., & Everson, P. (2009). Technical Note: Prediction of sex based on five skull traits using decision analysis (CHAID). *American Journal of Physical Anthropology*, 139, 434–441.

Stirland, A. J. (2013). *The men of the Mary Rose: Raising the dead*. The History Press.

Stloukal, M. (1961). Antropologický posudek. *Památky Archeologické*, LII, 157–158.

Stloukal, M. (1968). *Kostra z hrobu s keramikou zvoncovitých pohárů z Brna-Juliánova* (pp. 20–21). Das Skelett aus dem Glockenbechergrab von Brno-Juliánov. *Přehled Výzkumů*.

Taintier, J. A. (1980). Behavior and Status in a Middle Woodland Mortuary Population from the Illinois Valley. *American Antiquity*, 45(2), 308–313.

Tchéremissinoff, Y., Convertini, F., Fouéré, P., & Salanova, L. (2022). Chaptre premier. La sépulture campaniforme de La Folie, Poitiers (Vienne). In Y. Tcheremissinoff (Ed.), *Les sépultures individuelles campaniformes en France* (pp. 11–19). CNRS Éditions.

Thomas, A. (2014). Bioarchaeology of the middle neolithic: Evidence for archery among early European farmers. *American Journal of Physical Anthropology*, 154, 279–290.

Tihanyi, B., Bereczki, Z., Molnár, E., Berthon, W., Révész, L., Dutour, O., & Pálfi, G. (2015). Investigation of Hungarian Conquest Period (10th c. AD) archery on the basis of activity-induced stress markers on the skeleton—Preliminary results. *Acta Biologica Szegediensis*, 59(1), 65–77.

Tinazci, C. (2011). Shooting dynamics in archery: A multidimensional analysis from drawing to releasing in male archers. *Procedia Engineering*, 13, 290–296.

Tkáč, P. (2008). Pohrebisko Kultúry Zvoncovitých Pohárov v Ivanoviciach Hané, Trať Borůvka (predbežná správa). In I. Cheben & I. Kuzma (Eds.), *Otázky Neolitu a Eneolitu Našich Krajín*—2007 (Vol. IX, pp. 287–294). Archaeologica Slovaca Monographie.

Turek, J. (2003). Řemeslná symbolika v pohřebním ritu období zvoncovitých pohárů: Suroviny, výroba a struktura společnosti v závěru eneolitu. In L. Šmejda & P. Vařeka (Eds.), *Sedmdesát neustupných let. Plzeň* (pp. 201–220). Aleš Čeněk.

Turek, J. (2004). Nátepnídestičky z období zvoncovitých pohárů, jejich suroviny, technologie aspočěnský význam. In J. Turek (Ed.), *K poctě Vládimíru Podborskému. Přátelé a žáci k sedmdesátým narozeninám*. = Zu Ehrenvon Vladimír Podborský. Von Freunden und Schülern zum siebzigsten Geburtstag (pp. 207–226). Ústav archeologie a muzeologie. Filozofická fakulta Masarykovy univerzity v Brně.

Turek, J. (2006). Období zvoncovitých pohárů v Evropě. *Archeologie ve Středních Čechách*, 10, 275–368.

Turek, J. (2017). Sex, transsexuality and archaeological perception of gender identities. *Archaeologies: Journal of the World Archaeological Congress*, 12(3), 340–358.

Turek, J., & Černý, V. (2001). Society, gender and sexual dimorphism of the Corded Ware and Bell Beaker populations. In F. Nicolis (Ed.), *Bell Beakers today: Potter, people, culture, symbolism in prehistoric Europe* (pp. 601–612). Provincia Autonoma di Trento Servizio Beni Culturali Ufficio Beni Archeologici.

Unger, J. (1984). Nové Nálezy Kultury Zvoncovitých Pohárů na Kloboucku u Brna. *Archeologické Rozhledy*, 36, 204–207.

Uomini, N. T. (2009). The prehistory of handedness: Archaeological data and comparative ethology. *Journal of Human Evolution*, 57, 411–419.

Vaishya, R., Vijay, V., Nwagbara, I. C., & Agarwal, A. K. (2017). Diffuse idiopathic skeletal hyperostosis (DISH) – A common but less known cause of back pain. *Journal of Clinical Orrthopaedics and Trauma*, 8, 191–196.

Vander Linden, M. (2015). Bell Beaker Pottery and Society. In C. Fowler, J. Harding, & D. Hofmann (Eds.), *The Oxford handbook of neolithic Europe* (pp. 605–620). Oxford University Press.

Villotte, S. (2006). Connaissances Médicales Actuelles, Cotation des Enthésopathies: Nouvelle Méthode. *Bulletins et Mémoires de La Société d'Anthropologie de Paris*, 18(1-2), 65–85.

Villotte, S. (2009). Enthésopathies et activités des hommes préhistoriques—Recherche méthodologique et application aux fossiles européens du Paléolithique supérieur et du Mésolithique. *BAR International Series* 1992. *Archaeopress*.

Villotte, S. (2023). Activity-related skeletal changes. In *Reference Module in Social Sciences* (pp. 1–9). Elsevier.

Villotte, S., & Santos, F. (2023). The effect of age on enthestral changes: A study of modifications at appendicular attachment sites in a large sample of identified human skeletons. *International Journal of Osteoarchaeology*, 33, 389–401.

Vokolek, V. (1965). Výzkum Východočeských Pohřebišť Lidu Popelnicových Polí V L. 1963-1964. *Archeologické Rozhledy*, 17, 644–647.

Vokolek, V. (1981). Eneolitický Hrob z Plotišť N.L. a Pohřebiště Zvoncovitých Pohárů z Předměřic nad Labem. *Archeologické Rozhledy*, XXXIII, 481–485.

Waldron, T. (2009). *Palaeopathology*. Cambridge University Press.

Walker, P. (2008). Sexing skulls using discriminant function analysis of visually assessed traits. *American Journal of Physical Anthropology*, 136, 39–50.

Walker, P., & Holliman, S. E. (1989). Changes in osteoarthritis associated with the development of a maritime economy among Southern California Indians. *International Journal of Anthropology*, 4, 171–183.

Wallace, I. J., Winchester, J. M., Su, A., Boyer, D. M., & Konow, N. (2017). Physical activity alters limb bone structure but not enthestral morphology. *Journal of Human Evolution*, 107, 14–18.

Weinfeld, R. M., Olson, P. N., Maki, D. D., & Griffiths, H. J. (1997). The prevalence of diffuse idiopathic skeletal hyperostosis (DISH) in two large American midwest metropolitan hospital populations. *Skeletal Radiology*, 26(4), 222–225.

Weinig, J. (1992). Ein neues Gräberfeld der Kupfer-und Frühbronzezeit bei Weichering. *Das Archäologische Jahr in Bayern*, 1991, 64–67.
Weiss, E. (2003). Understanding muscle markers: Aggregation and construct validity. *American Journal of Physical Anthropology*, *121*(3), 230-240.

White, T. D. (2000). *Human osteology* (2nd ed.). Academic Press.

Whiteside, J. A., & Andrews, J. R. (1989). Common elbow problems in the recreational athlete. *The Journal of Musculoskeletal Medicine*, *6*, 17-34.

Willvonseder, K. (1937). Gräber der älteren Bronzezeit von Leopoldsdorf, Niederösterreich. *Germania: Anzeiger Der Römisch-Germanischen Kommission Des Deutschen Archäologischen Instituts*, *21*(2), 88-95.

Wiltschke-Schrotta, K., Berner, M., & Teschler-Nicola, M. (2001). Anthropologische Auswertung der menschlichen Überreste aus den Gräbern der Glockenbecherkultur von Zwingendorf (Alicenhof), VB Mistelbach, Niederösterreich. *Archaeologia Austriaca*, *84*, 241-256.

Wissler, A., Blevins, K. E., & Buikstra, J. E. (2022). Missing data in bioarchaeology II: A test of ordinal and continuous data imputation. *American Journal of Biological Anthropology*, *179*, 349-364.

Wolff, J. (1892). *Das Gesetz der Transformation der Knochen*. Verlag von August Hirschwald.

Zápotocký, M. (1962). Nové Mladoeneolitické Hroby z Litoměřicka. *Sborník Severočeského Musea-Historia*, *3*, 151-161.

Zhang, R., Ye, B., & Liu, P. (2019). Dimension reduction of high-dimensional dataset with missing values. *Journal of Algorithms & Computational Technology*, *13*, 1-8.

Zipp, P. (1979). Elektromyographie in der biomechanik des sports—Anwendungsbeispiel Bogenschießen. *Leistungssport*, *4*, 288-294.

## SUPPORTING INFORMATION

Additional supporting information can be found online in the Supporting Information section at the end of this article.

**How to cite this article:** Ryan-Despraz, J., Villotte, S., Desideri, J., & Besse, M. (2023). Multivariate assessments of activity-related skeletal changes: Interpreting Bell Beaker specialized male archery and social organization in Central Europe. *American Journal of Biological Anthropology*, *182*(2), 237-263. https://doi.org/10.1002/ajpa.24817