Article
Homo 72/4 (2021), 327–346
J. Comp. Hum. Biol.
Published in print December 2021
Inferences on mobility and subsistence patterns from
degenerative joint disease and entheseal changes. Trends in
the farmer/forager border (Central-Western Argentina)
Eva Ailén Peralta1,*, Leandro H. Luna2, Adolfo F. Gil1
1
Instituto de Evolución, Ecología Histórica y Ambiente, CONICET, Urquiza Ave. 314, San Rafael, 5600, Argentina.
Instituto Multidisciplinario de Historia y Ciencias Humanas, CONICET; Facultad de Filosofía y Letras, Universidad de
Buenos Aires, Ciudad Autónoma de Buenos Aires, Argentina.
* Corresponding author: evaailenperalta@gmail.com
2
With 3 figures and 8 tables
Abstract: This study tests the hypothesis that the incorporation of cultigens about ca. 2000 years BP substantially changed
hunter-gatherer subsistence and mobility in the Atuel River valley (Central-Western Argentina), where the frontier of preHispanic domesticated resource dispersion was defined. Degenerative joint disease and entheseal change markers were
analyzed on skeletal remains from Cañada Seca-1, a burial archaeological site with commingled skeletal remains dated
about ca. 1500 years BP (MNI = 24). The results show lower mobility in comparison with hunter-gatherer remains from the
neighboring Pampa region and quite different manual activities compared to low-level producers. These trends are explained
as a result of a mixed subsistence strategy and mobility in an area where the incorporation of domesticated plants was neither a linear nor a fast process, and a stereotypical view proves to be insufficient to understand it. Although further information is required for future discussions, the present research highlights the potential of commingled skeletal remains for this
kind of study.
Keywords: mechanic stress; maize; food production; activity patterns
Seca-1 (CS-1) site, an ossuary of maize consumers located
in the Atuel River valley dated ca. 1500 years BP. The aim
was to identify the patterns of body use and their connection
with mobility and manual tasks. The region was occupied
by hunter-gatherers before ca. 2000 years BP and a low proportion of cultigens was later incorporated, whose significance for human economy constitutes a main topic of debate.
Considering this context, the present research evaluates from
a bioarchaeological perspective whether human strategies
were modified after 2000 years BP in the Atuel valley, with
subsequent changes in daily activities and in the distribution
of mechanical stress indicators. Finally, the present study
evaluates the applicability of bone activity markers in contexts of subsistence shifts.
Introduction
The Atuel River valley (Central-Western Argentina) has
been seen as the Southern border of American pre-Hispanic agriculture, supported by the presence of macroremains of domestic plants such as Zea mays, Cucurbita sp.,
Chenopodium quinoa, Phaseolus vulgaris, and Lagenaria sp.
since ca. 2200 years 14C BP (Lagiglia 1980; Lagiglia 2001;
Neme & Gil 2012). Further research has proposed a flexible
subsistence strategy for groups that lived in the Atuel River
valley, alternating between hunter-gathering and low-level
food production. This assumption is supported by stable
isotope values that indicated an incipient consumption of
domesticated C4 plants such as maize (Gil et al. 2010; Gil
et al. 2011; Gil et al. 2018). Although the role of maize in
the diet has been strongly debated, and its consumption can
be supported (Gil et al. 2018; Peralta 2019), the problem of
its impact on hunter-gatherer subsistence and mobility is still
unsolved in the area. To address this issue, we approached
the study of degenerative joint disease (DJD) and entheseal
changes (EC) in a sample of 24 individuals from Cañada
Hunter-gatherers, agriculture and small-scale
food production: concepts and bioarchaeological
implications
It is generally assumed that domestication impacted multiple aspects of human life, including subsistence, technology, diet, mobility, demography, and human health
© 2021 E. Schweizerbart’sche Verlagsbuchhandlung, 70176 Stuttgart, Germany
DOI: 10.1127/homo/2021/1526
eschweizerbart_xxx
www.schweizerbart.de
0018-442X/2021/1526
$ 5.00
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E. A. Peralta, L. H. Luna, A. F. Gil
(Cohen 1977; Redman 1990; Colledge & Conolly 2007;
Vigne 2015). Traditionally, hunter-gathering and farming were built as polarized subsistence systems and later
re-discussed as extremes of a broad spectrum of adaptive
strategies (Winterhalder & Kennett 2006; Gepts et al. 2012;
Winterhalder & Kennett 2020). Some authors have characterized intermediate subsistence systems as low-level food
production in which groups highly dependent on huntergathering introduced cultigens and eventually domesticated fauna (Smith 2001; Winterhalder & Kennett 2006;
Winterhalder & Kennett 2020). This proposal was used in
southern Mendoza to characterize the archaeological contexts that include cultigens (Gil et al. 2018; Peralta 2019;
Gil et al. 2020). It is important to clarify that the lines of
inquiry driven in the region did not support intensive agriculture so far. Intensive agriculture and sedentarism took
place in Northwestern Argentina, where studies about subsistence shifts through entheseal changes and degenerative
joint disease are very scarce (Peralta 2019; Peralta 2020).
The present research explores the implications of lowlevel food production in terms of body use and physical
activities.
A double process of resource intensification and mobility reduction has been suggested for the Atuel valley for the
end of the late Holocene period (ca. 2000 yr BP) (Neme
2007; Gil et al. 2018). In highly mobile societies, physical demands in the lower extremities have been frequently
associated with daily human locomotion (Ruff & Hayes
1983; Cohen & Armelagos 1984; Quevedo 2000; Marchi
et al. 2006; Sparacello & Marchi 2008; Marchi 2008;
Lieverse et al. 2011). Residential and logistic movements
characteristic of hunter-gatherers have been linked to high
levels of degenerative joint disease of the knee, ankle-foot,
and lumbar vertebrae (Larsen 1997; Knüsel 2000; Quevedo
2000; Larsen & Ruff 2011; Lieverse et al. 2011). In similar contexts, a high prevalence of entheseal changes was
observed at the plantaris, soleus, gluteus, and adductor
muscles (Lieverse et al. 2011; Scabuzzo 2012; Lieverse
et al. 2013). By contrast, the opposite scenario, in which
lower extremities are exposed to less stress due to a reduction in mobility, is expected in the context of an intensification process (Bettinger & Baumhoff 1982; Larsen & Ruff
2011; Lieverse et al. 2011). At the same time, an increase in
diversity and intensity of manual tasks with food production
could be expected in some contexts and would be related
to higher physical demands in upper limbs evidenced by
higher prevalences of entheseal changes and degenerative
joint disease (Sparacello & Marchi 2008; Henderson 2013;
Sparacello et al. 2020; Varalli et al. 2020). Considering
this model, the present study hypothesizes that lower limbs
were less affected and upper limbs were more affected in
CS-1 than in other hunter-gatherer populations, while both
extremities were similarly affected compared to populations
that included low-level food production.
Late Holocene Central-Western Argentina and
human occupation of the Atuel valley
The Atuel River valley (34°48′55″LS, 68°22′07″LO) is
located in Central-Western Argentina, in Mendoza province,
an arid-semiarid environment occupied by hunter-gatherers
since ca. 10000 years BP. The Atuel valley shows drastic
changes in the archaeological record ca. 2200 years 14C BP.
These mainly consist of an increase in the number and size
of archaeological sites, the first occupation of marginal areas
(the high Andes and Payunia desert), the development of
new technologies (ceramic and stone grinding technology),
the increase in the use of obsidian as raw material (probably
indicating changes in social networks), and the exploitation
of new food resources, like small size fauna, and a wider
spectrum of wild animals. The earliest evidence of domestic
plants was recorded both in the valley and in the whole region
during this time (see Neme 2007; Neme & Gil 2008; Llano
& Andreoni 2012; Otaola et al. 2012; Salgán et al. 2012;
Neme et al. 2015; Sugrañes 2016). It has been proposed that
these changes were the result of the intensification process
in resource exploitation, consequence of the spatial saturation and a derived imbalance between human demography
and available resources ca. 2000 years BP (Neme 2007).
Intensification involved diet breadth expansion, including
low-ranking resources with higher procurement and processing costs, which required a reduction in mobility to achieve
these tasks (Bettinger & Baumhoff 1982).
The presence of cultigens was assumed to be associated
with a shift towards a farming/agricultural strategy. The
change from a hunter-gatherer economy to a food production lifestyle was firstly explored in the region as a historicalcultural phenomenon (Lagiglia 1980) and more recently as a
consequence of the above-mentioned intensification process
(Neme 2007). In terms of food consumption, stable isotope
analyses improved the knowledge about dietary patterns during the last 2200 years 14C BP. Based on δ13Ccol, δ13Capa,
and δ15N data, maize signal was detected in the human diet
of several groups in the Atuel River valley with high temporal and spatial variability in the incorporation of this cultigen (Gil et al. 2018; Peralta 2019). Dietary intakes in CS-1
showed a maize consumption of about 30% (Gil et al. 2018;
Peralta 2019; Peralta et al. 2021). Simultaneously, human
mobility was explored through δ18O values. In CS-1, δ18O
values had a low resolution to evaluate changes in mobility
associated with the incipient incorporation of cultigens and
low-level food production (Ugan et al. 2012; Gil et al. 2018).
As a consequence of the intensification process, domesticated resources (principally Zea mays) were partially incorporated as part of the human diet, but the precise impact of
this introduction in terms of mobility and subsistence patterns remains unclear. The hypotheses of this study state
that in the context of the introduction of domesticated
resources and low-level food production: 1) mobility tended
to decline, exposing lower extremities to less mechanical
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Inferences on mobility and subsistence patterns from degenerative joint disease
329
et al. 1984; Bridges 1989; Cheverko & Bartelink 2017;
Schrader 2019; Becker 2020). Specific changes have also
been recorded, such as a high level of DJD in the agriculturalist female left arm (Bridges 1992). Punctual activities have
also been proposed in particular hunter-gatherer contexts,
including spear-throwing, bow and arrow use, grinding, and
scraping of animal hides (Angel 1966; Merbs 1983; Schrader
2019; Becker 2020).
Entheseal changes (EC) are modifications on muscle
attachment sites (entheses) (Benjamin et al. 1986; Dutour
1986; Benjamin & Ralphs 1998; Benjamin et al. 2002;
Villotte & Knüsel 2013). They are the object of similar
controversies, and some authors have paid attention to the
multiple factors that may determine these changes, mainly
age, sex, body size, and pathological conditions (Wilczak
1998; Weiss 2003; Weiss 2004; Weiss 2007; Henderson
2008; Sparacello & Marchi 2008; Weiss et al. 2012; Villotte
et al. 2016). Previous studies attempted to rank muscles from
most to least used for human samples from different origins,
but some problems of equifinality arose (al-Oumaoui et al.
2004; Lieverse et al. 2009). Specific activities were proposed
in specific contexts, such as watercraft and spear throwing
(Lieverse et al. 2009; Sparacello et al. 2020), some authors
propose that agriculturalists have lower scores for entheseal changes than hunter-gatherers and industrial workers
(Henderson 2013), while others suggest increased upper
body demands with maize intensification (Shuler et al. 2012).
Lower extremity stress seems to be linked to both
high hunter-gatherer mobility and high agricultural workload (Eshed 2010; Lieverse 2013; Schlader 2015). Many
researchers have identified a trend showing a decrease in
DJD and EC in the lower limbs as a consequence of a decline
in workload derived from reduced mobility (Hoyme & Bass
1962; Larsen 1995; Formicola 1997; Ruff et al. 2006; Holt
& Formicola 2008; Sparacello & Marchi 2008; Larsen &
Ruff 2011; Marchi et al. 2011; Stock et al. 2011; Carlson
& Marchi 2014; Varalli et al. 2020). In turn, it was considered that changes in subsistence-related activities are better
reflected in the upper limbs in a transitional context between
hunting-gathering and low-level food production (Eshed
et al. 2004; Eshed 2010; Varalli et al. 2020). This is because
the arms and shoulder are seen as more ‘activity-specific’
than the legs and pelvic girdle in which changes may be
caused mainly by walking (Henderson 2008).
stress than in previous hunter-gatherer contexts, and that
2) daily tasks regarding food processing activities increased
in intensity and extent displaying a more similar pattern to
that observed among low-level food producers in the upper
limbs. Expectations are: 1) lower frequencies of DJD and
EC in lower extremities and higher frequencies in upper
extremities compared to the patterns observed by Scabuzzo
(2012) for typical hunter-gatherers of the Pampa Region
(Argentina) and 2) similar frequencies of DJD and EC in
both extremities compared to those observed by Salega
(2016) for hunter-gatherers that introduced low-level food
production in the Central Highlands Region (Argentina). A
greater frequency of DJD and EC in males than in females,
mainly in the lower limbs, is also expected. This entails that
males were commonly engaged in hunting risks and higher
mobility than females, in line with the ethnoarchaeological evidence available (Lee & DeVore 1968; Brown 1970;
Bridges 1989; Molleson 1994; Panter-Brick 2002; Eshed
et al. 2004). Additionally, it is assumed that these differences
would not be correlated with body size (Weiss 2003; Weiss
2004; Weiss 2007).
Stress markers of activity
DJD and EC have been suggested as useful proxies for past
activity patterns. Studies based on these indicators have
contributed to discussions about the characterization of lifestyles and subsistence for different types of societies (Cohen
& Armelagos 1984; Kennedy 1989; Bridges 1992; Bridges
1994; Capasso et al. 1999; Lieverse et al. 2009; Eshed et al.
2010; Villotte et al. 2010; Schrader 2012; Peralta 2017;
Peralta 2020). Several controversies have arisen related to
the multi causality of these bone markers, which may be not
affected by daily activities only but also by age, sex, body
size, and genetic conditions, among others (Weiss & Jurmain
2007; Santos et al. 2011; Jurmain et al. 2012). Many attempts
have also been made to establish standardized methods that
could be replicated by different researchers with low intra
and interobserver errors (Mariotti et al. 2004; Villotte 2006;
Mariotti et al. 2007; Weiss & Jurmain, 2007; Villotte et al.
2010; Jurmain et al. 2012; Henderson et al. 2013; Henderson
et al. 2016; Henderson et al. 2017a; Henderson et al. 2017b).
Degenerative Joint Disease (DJD), or osteoarthrosis, is
an alteration of synovial joints producing changes in both
periarticular and internal bone tissues (Rogers & Waldron
1995; Waldron 2009; Berenbaum 2013). Some researchers did not observe significant differences in prevalence
among groups with different types of subsistence (Bridges
1992; Knüsel 1993; Bridges 1994), although others identified major changes in articular loadings. Many studies have
proposed that workload declined with the introduction of
agriculture, resulting in a reduction of DJD (Larsen 1982;
Larsen 1984; Cohen & Armelagos 1984; Schrader 2019;
Becker 2020), while others identified a consistent trend of
DJD increasing with food production (Lallo 1973; Goodman
Material and methods
Foragers and farmers in the Atuel valley:
Cañada Seca-1 (CS-1) site
Many archaeological contexts with burials, including CS-1,
were identified in the Atuel valley, most of them dated during the end of the Late Holocene (Neme & Gil 2010; Peralta
2019). CS-1 is an archaeological site with an exclusive
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E. A. Peralta, L. H. Luna, A. F. Gil
ses studied are presented in Table 1. Bones were considered
observable when more than 75% of the surface/margin of
auricular surface or entheses was present and in a good condition of preservation. This criterion was independent of the
complete or fragmented bone condition.
funerary function. Since it is an ossuary with commingled
remains, it was not possible to individualize skeletons
(Peralta et al. 2021). The elements were recovered without
systematic excavations during the 1990s, and the burial
context was unfortunately lost (Lagiglia 1991), a common
aspect of field activities in those times. The site is located
in the Atuel River middle valley (34°74’ LS, 68°23’ LO;
610 m.a.s.l.) (Fig. 1) and is dated in 1398 years cal. BP
(Peralta et al. 2021). Isotopic analyses show δ13Ccol, δ13Capa,
and δ15N mean values of –15‰, –9.7‰, and 11‰, respectively (Gil et al. 2018; Peralta et al. 2021). These data were
framed within the isotopic ecology of the Atuel valley and
interpreted as a result of C4 energy and protein intakes
(Harrison & Katzenberg 2003; Kellner & Schoeninger
2007; Schoeninger 2009). In the Atuel valley, C4 resources
are mainly defined as Zea mays (Gil et al. 2010; Gil et al.
2011; Gil et al. 2018; Gil et al. 2020). A maize consumption
close to 30% was proposed for CS-1 employing FRUITS,
a Bayesian model for diet reconstruction (Gil et al. 2018;
Peralta et al. 2021). In consequence, the chronological and
spatial location of CS-1 and its isotopic evidence become
relevant for a discussion about bioarchaeological concerns
in low-level food production societies.
The Minimum Number of Individuals (MNI) for CS-1 is
24, including 4 subadults, 2 juveniles, and 18 adults (Peralta
2019; Peralta et al. 2021). We identified one young adult
female, three middle adult males, three middle adult females,
and two old adult males. Since CS-1 only included commingled skeletal remains, os coxae and skulls were not associated with appendicular bones. A total of 155 appendicular
bones from both sides (humerus, radius, ulna, femur, tibia,
and calcaneus) were analyzed. Frequencies of elements,
articular surfaces, and fibrous and fibrocartilaginous enthe-
Sex and age estimations
Sex was estimated in previous research (Peralta 2019;
Peralta et al. 2021); the procedure included both traditional
methodologies for the os coxae (Phenice 1969; Bruzek
2002) and skulls (Buikstra & Ubelaker 1994) and alternative ones based on bone metrics of long bones and the calcaneus (Steele 1976; Berrizbeitia 1989; Holland 1991; Bruzek
1995; Silva 1995; Introna et al. 1997; Trancho et al. 1997;
Seidemann et al. 1998; Wilbur 1998; Alemán Aguilera et al.
2000; López-Bueis et al. 2000; Robledo et al. 2000; Trancho
et al. 2000; Murphy 2002; Frutos 2003; Luna 2008). Age
estimations were only obtained following the recording of
the pubic symphysis and auricular surface (Todd 1920; Todd
1921; Meindl et al. 1980; Lovejoy et al. 1985; Meindl &
Lovejoy 1989). Unfortunately, the commingled condition of
the burial precluded the evaluation of age-at-death influence
on long bone EC and DJD (Peralta 2017; Peralta 2020).
Degenerative joint disease and entheseal changes
recording methods
The recording of DJD was performed analyzing articulation
surfaces and margins separately (Luna et al. 2017). Each
articulation was divided into four quadrants, each of which
was rated on a scale from 0 to 3. Results were added and
divided by the number of quadrants effectively recorded,
which allowed including articulations with different degrees
of preservation. Intensity margin scores were defined as fol-
Fig. 1. Location of Cañada Seca-1 (CS-1) in the Atuel valley.
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Inferences on mobility and subsistence patterns from degenerative joint disease
Table 1. Recorded articular surfaces, fibrous entheses (F), and fibrocartilaginous entheses (FC). Px: proximal articulation; Dx: distal
articulation; R: right; L: left.
Element
Surface/entheses
Laterality
N
N_F
N_M
N_IN
Humerus
Px
R
10
11
9
3
L
9
5
13
19
7
10
9
Dx
Pectoralis major (F)
Deltoid (F)
12
R
9
L
13
9
L
13
Brachioradialis (F)
R
9
L
13
Extensor (FC)
R
9
L
11
Flexor (FC)
R
8
L
11
Subscapularis (FC)
R
8
L
7
Teres minor (FC)
R
5
L
7
R
5
L
6
R
6
L
7
Infraspinatus (FC)
Ulna
11
L
R
Supraspinatus (FC)
Radius
R
Px
R
18
L
19
Dx
R
8
L
8
Biceps (FC)
R
12
L
10
Px
R
13
L
13
Dx
R
9
L
11
Supinator (F)
R
12
L
15
R
12
L
15
Pronator quadratus (F)
Triceps (FC)
Brachialis (FC)
R
11
L
13
R
12
L
13
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332
E. A. Peralta, L. H. Luna, A. F. Gil
Table 1. continued.
Element
Surface/entheses
Laterality
N
N_F
N_M
N_IN
Femur
Px
R
14
13
10
8
L
15
Dx
R
11
L
10
Gluteal tuberosity (F)
R
15
L
16
Linea aspera (F)
R
15
L
16
Iliopsoas (FC)
R
8
L
12
11
9
3
5
8
2
Gluteus minimus (FC)
Gluteus medius (FC)
Tibia
Calcaneus
R
8
L
11
R
9
L
10
R
11
L
8
Dx
R
12
L
11
Quadriceps (F)
R
10
L
12
Soleal line (F)
R
10
L
12
Plantaris (FC)
R
8
L
7
Px
activity markers in human skeletal remains (Scabuzzo 2010;
Scabuzzo 2012; Salega 2016). To make possible the comparison between CS-1 results and previously published data,
we decided to replicate the methodological procedures.
Robusticity (rugosity or hypertrophy), stress lesions (pitting or furrowing), and ossifications (bony spurs or projections) were recorded. Robusticity and stress lesions were
considered as a continuum of increasing muscle and/or
ligament strain and were combined in the recording procedure (Hawkey & Merbs 1995). The resulting scores ranged
from 0 to 6: 0 = absence of expression; 1 = cortex slightly
rounded, palpable elevation, although without distinct crests
or ridges; 2 = uneven cortical surface with a mound-shaped
elevation; no sharp ridges or crests formed; 3 = sharp crests
or ridges formed; 4 = pitting (< 1 mm depth) into the cortex with lytic appearance; 5 = deeper pitting (> 1 mm but
< 3 mm depth) covering more surface area; may vary in
length but never > 5 mm; 6 = marked pitting > 3 mm depth
and > 5 mm length. Since ossifications are considered the
result of abrupt macro trauma, rather than continuous muscle
use, they were analyzed and interpreted separately using the
following score: 0 = absence; 1 = slight exostosis, usually
lows: 0 = no changes associated with DJD; 1 = slight irregularities (< 1 mm); 2 = osteophytes clearly visible (1–5 mm
long); 3 = osteophytes > 5 mm. Cases of ankylosis were
included in the last category. In turn, surface scores were
recorded using the following categories: 0 = no porosity;
1 = < 1/3 of surface affected by porosity; 2 = 1/3 to 2/3 of
surface affected; 3) > 2/3 of surface affected. Eburnation was
included at this stage. The percentage of affectation was also
relieved; in this case, four categories were considered: 0 =
not affected; 1 = less than 25% of periarticular alteration and
surface porosity; 2 = 25 to 50% affected; 3 = more than 50%
affected. Results were presented by articulation and anatomical portion (upper or lower limbs). Each articulation was
composed of both joint surfaces (three for elbow), and each
surface was recorded separately and later grouped by joint.
Carpals, metacarpals, tarsals (excluding the calcaneus) and
metatarsals were excluded from the recording of the wrist
and the ankle.
The methodology developed by Hawkey & Merbs (1995)
was applied to fibrous entheses. Although the method have
some weaknesses, the election was conditioned by previous regional research that employed this method to explore
eschweizerbart_xxx
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Inferences on mobility and subsistence patterns from degenerative joint disease
Table 2. Frequency of degenerative joint disease in each joint considering margin and surface.
Margin
Joint
Shoulder
0
1
Surface
2
3
0
1
2
3
%
n
%
n
%
n
%
n
%
n
%
n
%
n
%
n
68.4
13
26.3
5
5.3
1
0
0
78.9
15
21.1
4
0
0
0
0
F
50
4
50
4
0
0
0
0
50
4
50
4
0
0
0
0
M
100
6
0
0
0
0
0
0
100
6
0
0
0
0
0
0
IN
60
3
20
1
20
1
0
0
100
5
0
0
0
0
0
0
Elbow
50
33
40.9
27
7.6
5
1.5
1
84.8
56
13.6
9
1.5
1
0
0
F
52.6
10
47.4
9
0
0
0
0
89.5
17
10.5
2
0
0
0
0
M
46.7
14
40
12
13.3
4
0
0
86.7
26
13.3
4
0
0
0
0
IN
52.9
9
35.3
6
5.9
1
5.9
1
76.5
35
17.6
1
5.9
1
0
0
Wrist
69.4
25
25
9
5.6
2
0
0
97.2
35
2.8
1
0
0
0
0
F
80
8
20
2
0
0
0
0
90
9
10
1
0
0
0
0
M
62.5
10
31.3
5
6.3
1
0
0
100
16
0
0
0
0
0
0
IN
70
7
20
2
10
1
0
0
100
10
0
0
0
0
0
0
Hip
100
29
0
0
0
0
0
0
96.6
28
3.4
1
0
0
0
0
F
100
13
0
0
0
0
0
0
92.3
12
7.7
1
0
0
0
0
M
100
8
0
0
0
0
0
0
100
8
0
0
0
0
0
0
IN
100
8
0
0
0
0
0
0
100
8
0
0
0
0
0
0
Knee
55
22
42.5
17
2.5
1
0
0
92.5
37
7.5
3
0
0
0
0
F
43.8
7
50
8
6.3
1
0
0
93.8
15
6.3
1
0
0
0
0
M
62.5
10
37.5
6
0
0
0
0
87.5
14
12.5
2
0
0
0
0
IN
62.5
5
37.5
3
0
0
0
0
100
8
0
0
0
0
0
0
Ankle
43.5
10
52.2
12
4.3
1
0
0
100
23
0
0
0
0
0
0
F
62.5
5
37.5
3
0
0
0
0
100
8
0
0
0
0
0
0
M
18.2
2
72.7
8
9.1
1
0
0
100
11
0
0
0
0
0
0
IN
75
3
25
1
0
0
0
0
100
4
0
0
0
0
0
0
rounded in appearance, extending < 2 mm from the cortical
surface; 2 = distinct exostosis extending > 2 mm but < 5 mm
from the cortex surface; 3 = exostosis extending > 5 mm
from the bone surface or covering an extensive area of the
cortical surface.
The methodology proposed by Henderson et al. (2016;
2017a) was posteriorly applied for the fibrocartilaginous
entheses. For the recording process, each enthesis was
divided into two zones: Zone 1 is the entheseal margin at
which fibers attach most obliquely to the bone, while Zone
2 encompasses the remaining fibrocartilaginous entheseal
footprint and the remaining margin. In most entheses, Zone
2 is the closest to the joint surface (Henderson et al. 2016;
Henderson et al. 2017a). Eight features were recorded:
bone formation (BF1) and erosion in Zone 1 (ERO1), textural change (TC), bone formation (BF), erosion (ERO),
fine porosity (FPO), macroporosity (MPO), and cavitation
(CAV) in Zone 2. Each feature was characterized from 0 to
2, except for TC that was recorded as absent (0) or present
(1) (see Table S1 for score characterization of each feature).
Fibrocartilaginous entheseal changes were analyzed with
Principal Components Analyses (PCA) using the software
PAST 3.15 (Hammer et al. 2001) to determine which of
them best explain the variance identified. The results of both
fibrous and fibrocartilaginous entheses were presented separately showing the frequencies of changes by entheses and
anatomical portion (upper and lower limbs).
Finally, since Weiss (2003; 2004) demonstrated that
muscle markers may correlate with body size, it is necessary
to discard the influence of body size in EC variation before
trends of activity patterns can be inferred and interpreted.
Therefore, Weiss’ methodology to infer body size (2003;
2004; 2007) was used. Z-scores were calculated to create
two composite (or aggregate) variables: upper/lower limb
muscle marker and upper/lower limb size. The upper/lower
limb muscle marker composite was generated by averaging
z-scores for 77 component variables: 38 insertion sites from
humeri (including pectoralis major, deltoid, and brachioradialis) and 39 femoral insertion sites (including linea aspera
and gluteal tuberosity). Each variable was scored in the cat-
eschweizerbart_xxx
334
E. A. Peralta, L. H. Luna, A. F. Gil
Table 3. Frequency of entheseal changes in fibrous enthuses.
Ossifications
Robusticity-stress lesions
Enthesis
0
1
2
3
4
0
1
2
3
%
n
%
n
%
n
%
n
%
n
%
n
%
n
%
n
%
n
Pectoralis major
57.1
8
35.7
5
0
0
0
0
7.1
1
100
0
0
0
0
0
0
0
F
57.1
4
28.6
2
0
0
0
0
14.3
1
100
0
0
0
0
0
0
0
M
66.7
2
33.3
1
0
0
0
0
0
0
100
0
0
0
0
0
0
0
IN
50.0
2
50
2
0
0
0
0
0
0
100
0
0
0
0
0
0
0
Deltoid
84.6
11
15.4
2
0
0
0
0
0
0
100
0
0
0
0
0
0
0
F
83.3
5
16.7
1
0
0
0
0
0
0
100
0
0
0
0
0
0
0
M
100
2
0
0
0
0
0
0
0
0
100
0
0
0
0
0
0
0
IN
80
4
20
1
0
0
0
0
0
0
100
0
0
0
0
0
0
0
Brachioradialis
94.1
16
5.9
1
0
0
0
0
0
0
100
0
0
0
0
0
0
0
F
100
10
0
0
0
0
0
0
0
0
100
0
0
0
0
0
0
0
M
75
3
25
1
0
0
0
0
0
0
100
0
0
0
0
0
0
0
IN
100
3
0
0
0
0
0
0
0
0
100
0
0
0
0
0
0
0
Supinator
69.6
16
17.4
4
13
3
0
0
0
0
100
0
0
0
0
0
0
0
F
83.3
5
16.7
1
0
0
0
0
0
0
100
0
0
0
0
0
0
0
M
72.7
8
9.1
1
18.2
2
0
0
0
0
100
0
0
0
0
0
0
0
IN
50
3
33.3
2
16.7
1
0
0
0
0
100
0
0
0
0
0
0
0
Pronator quadratus
80
16
20
4
0
0
0
0
0
0
100
0
0
0
0
0
0
0
F
100
4
0
0
0
0
0
0
0
0
100
0
0
0
0
0
0
0
M
57.1
4
42.9
3
0
0
0
0
0
0
100
0
0
0
0
0
0
0
IN
88.9
8
11.1
1
0
0
0
0
0
0
100
0
0
0
0
0
0
0
Gluteal tuberosity
42.1
8
31.6
6
15.8
3
10.5
2
0
0
100
0
0
0
0
0
0
0
F
33.3
3
44.4
4
11.1
1
11.1
1
0
0
100
0
0
0
0
0
0
0
M
33.3
2
33.3
2
33.3
2
0
0
0
0
100
0
0
0
0
0
0
0
IN
75
3
0
0
0
0
25
1
0
0
100
0
0
0
0
0
0
0
Linea aspera
54.5
12
27.3
6
18.2
4
0
0
0
0
86.4
19
4.5
1
9.1
2
0
0
F
50
4
37.5
3
12.5
1
0
0
0
0
87.5
7
12.5
1
0
0
0
0
M
50
4
25
2
25
2
0
0
0
0
87.5
7
0
0
12.5
1
0
0
IN
66.7
4
16.7
1
16.7
1
0
0
0
0
83.3
5
0
0
16.7
1
0
0
Quadriceps
62.5
10
31.3
5
6.3
1
0
0
0
0
100
0
0
0
0
0
0
0
F
87.5
7
12.5
1
0
0
0
0
0
0
100
0
0
0
0
0
0
0
M
28.6
2
57.1
4
14.3
1
0
0
0
0
100
0
0
0
0
0
0
0
IN
100
1
0
0
0
0
0
0
0
0
100
0
0
0
0
0
0
0
Soleal line
55
11
30
6
15
3
0
0
0
0
90
18
10
2
0
0
0
0
F
60
6
30
3
10
1
0
0
0
0
90
9
10
1
0
0
0
0
M
55.6
5
22.2
2
22.2
2
0
0
0
0
88.9
8
11.1
1
0
0
0
0
IN
0
0
100
1
0
0
0
0
0
0
100
1
0
0
0
0
0
0
eschweizerbart_xxx
335
Inferences on mobility and subsistence patterns from degenerative joint disease
Table 4. Mean score (x) for each enthesis and rank ordering (r) of mean score from high to low.
Robusticity-Stress lesions
Enthesis
Pectoralis major
Females
Males
n
x
r
n
x
r
7
0.86
1
3
0.33
3
Deltoid
6
0.17
2
2
0
5
Brachioradialis
10
0
4
4
0.25
4
Supinator
6
0.17
3
11
0.45
1
Pronator quadratus
4
0
5
7
0.43
2
Gluteal tuberosity
9
1
1
6
1
1
Linea aspera
8
0.62
2
8
0.75
3
Quadriceps
8
0.12
4
7
0.86
2
Soleal line
10
0.5
3
9
0.67
4
while ECs were recorded using both Henderson et al. (2016;
2017a) and Hawkey & Merb’s (1995) proposals. Data available in publications for the Pampa and Central Highlands
are uneven compared with data obtained for CS-1, so comparisons were conditioned by how results were structured by
each author. Concerning DJD, the frequencies between the
CS-1 and Pampa samples were compared by sex accounting
for degrees of severity. The frequencies between the CS-1
and Central Highland samples were only compared considering the presence of articular defects in the margin and
surface without distinguishing by sex or severity. Regarding
EC, comparisons at the fibrous entheses were conducted
between the CS-1 and Pampa samples; sex and degrees of
severity of the individuals were considered. Change frequencies at fibrocartilaginous entheses were compared between
the CS-1 and Central Highland samples. Differences were
tested with the Mann-Whitney test using PAST 3.15.
egories of robusticity and stress lesions defined by Hawkey
& Merbs (1995). A composite variable of upper limb size
was created by averaging z-scores for two humeral size variables (humeral vertical head diameter and humeral epicondylar breadth). A composite variable of lower limb size was
created by averaging z-scores for two femoral size variables
(maximum head diameter and epicondylar breadth). Weiss’
measures of maximum length were not considered due to the
large number of fractured humeri and femora. All humeral
and femoral measurements were made according to Buikstra
& Ubelaker (1994). Ulna, radial or tibial measurements in
the size aggregate variable for upper and lower limbs were
not included due to the impossibility of matching isolated
bones. The composite variable upper/lower limb muscle
marker was correlated using two-tailed Spearman tests with
the composite upper/lower limb variable (Weiss 2003; Weiss
2004). To correlate the composite variable upper/lower limb
muscle marker with sex, the eta coefficient was employed.
Both tests were run with SPSS software.
All the recording was fulfilled by the first author.
Intraobserver error in the recording of DJD and EC was
calculated after two observations made two weeks after the
other. Differences between both observations were tested
applying the Intraclass Correlation Coefficient (ICC) for DJD
variables (Landis & Koch 1977) and the Gamma Coefficient
(GC) for EC variables (Flom 2021).
Results
Intraobserver error
ICC values for osteophytes and porosity were 0.92 and 0.89,
respectively (“almost perfect” following Landis & Koch
1977). GC values for fibrocartilaginous EC variables ranged
from 0.89 to 1, while for fibrous EC, variables were between
0.76–0.79 (evidence of strong association following Flom
2021).
Samples for comparison
To interpret the trends obtained about body use in CS-1, published DJD and EC data from two neighboring bioarchaeological samples were used for comparison. Scabuzzo (2010;
2012) recorded DJD and fibrous EC following Merbs (1983)
and Hawkey & Merbs (1995) methodology, respectively, in
hunter-gatherers’ skeletons from the Pampean region. In turn,
Salega (2016) obtained information about a low-level food
production population from the Central Highlands. The last
author recorded DJD using a method developed by herself,
Degenerative joint disease
Regardless of intensity, the elbow was the most affected joint
(50%) in the upper limbs. This joint reached low percentages
of moderate (7.6%) and high scores (1.5%) for periarticular
osteophytes. Intraarticular porosity had a DJD prevalence
of 15.1% at the elbow (Table 2). The shoulder followed the
elbow with 31.6% of DJD prevalence at margins and showed
the highest percentage of DJD for intraarticular porosity
eschweizerbart_xxx
336
E. A. Peralta, L. H. Luna, A. F. Gil
Fig. 2. Feature loadings at PC1. PC1 explains 33% of the variance.
Fig. 3. PCA comparing fibrocartilaginous entheseal changes between males (circles) and females (triangles).
(21.1%). The wrist was less affected at both the joint margin
and surface (30.6% and 2.8%, respectively) compared with
the elbow and shoulder. Moderate scores were observed at
margins in the shoulder, elbow, and hand-wrist joints. Severe
scores only occurred at margins in the elbow.
Considering the whole sample, the ankle was the most
affected joint at margin level (56.5%, Table 2). The knee
showed a high percentage of DJD at margins (45%) and a
lower percentage on the joint surface (7.5%). Moderate
scores were recorded only at margins in the knee and anklefoot joints. Severe scores were not observed, and the hip
was almost unaffected by DJD. Females showed higher
percentages of DJD at margins in the shoulder (50%) and
knee (56.3%) and on the surface in the shoulder (50%),
wrist (10%), and hip joints (7.7%). Males showed higher
percentages at margins in the elbow (53.3%), wrist (37.5%),
eschweizerbart_xxx
337
Inferences on mobility and subsistence patterns from degenerative joint disease
Table 5. Frequency of entheseal changes in fibrocartilaginous entheses.
Enthesis
%
Biceps
72.7
F
80.0
M
76.9
IN
50
Brachialis
84
F
83.3
M
91.7
IN
71.4
Extensor
85
F
83.3
M
100
IN
0
Flexor
84.2
F
81.8
M
100
IN
50
Infraspinatus 100
F
100
M
100
IN
100
Subscapularis 73.3
F
66.7
M
100
IN
33.3
Supraspinatus 90.9
F
80
M
100
IN
100
Teres minor 100
F
100
M
100
IN
100
Triceps
95.8
F
100
M
91.7
IN
100
Iliopsoas
50
F
55.6
M
28.6
IN
75
Gluteus
84.2
minimus
F
88.9
M
66.7
IN
100
Gluteus medius 84.2
F
100
M
71.4
IN
75
Plantaris
46.7
F
80
M
25
IN
50
0
n
16
4
10
2
21
5
11
5
17
10
7
0
16
9
6
1
13
6
6
1
11
4
6
1
10
4
5
1
12
6
5
1
23
5
11
7
10
5
2
3
BFO1
1
%
n
27.3 6
20
1
23.1 3
50
2
16
4
16.7 1
8.3 1
28.6 2
10
2
16.7 2
0
0
0
0
15.8 3
18.2 2
0
0
50
1
0
0
0
0
0
0
0
0
20
3
33.3 2
0
0
33.3 1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
4.2 1
0
0
8.3 1
0
0
45
9
33.3 3
71.4 5
25.0 1
2
0
%
0
0
0
0
0
0
0
0
5
0
0
100
0
0
0
0
0
0
0
0
6.7
0
0
33.3
9.1
20
0
0
0
0
0
0
0
0
0
0
5
11.1
0
0
n
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
1
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
1
1
0
0
%
54.5
60
53.8
50
56
83.3
50
42.9
95
100
100
0
100
100
100
100
84.6
83.3
83.3
100
73.3
83.3
50
100
90.9
80
100
100
91.7
83.3
100
100
83.3
100
75
85.7
80
77.8
71.4
100
n
12
3
7
2
14
5
6
3
19
12
7
0
19
11
6
2
11
5
5
1
11
5
3
3
10
4
5
1
11
5
5
1
20
5
9
6
16
7
5
4
ERO1
1
%
n
45.5 10
40
2
46.2 6
50
2
44 11
16.7 1
50
6
57.1 4
5
1
0
0
0
0
100 1
0
0
0
0
0
0
0
0
15.4 2
16.7 1
16.7 1
0
0
26.7 4
16.7 1
50
3
0
0
9.1 1
20
1
0
0
0
0
8.3 1
16.7 1
0
0
0
0
16.7 4
0
0
25
3
14.3 1
20
4
22.2 2
28.6 2
0
0
TCH
2
0
%
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
n
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
%
54.5
60
53.8
50
56
83.3
50
42.9
95
100
100
0
100
100
100
100
84.6
83.3
83.3
100
73.3
83.3
50
100
90.9
80
100
100
91.7
83.3
100
100
83.3
100
75
85.7
80
77.8
71.4
100
1
n
12
3
7
2
14
5
6
3
19
12
7
0
19
11
6
2
11
5
5
1
11
5
3
3
10
4
5
1
11
5
5
1
20
5
9
6
16
7
5
4
0
%
n
%
n
45.5 10 95.5 21
40
2 100 5
46.2 6 92.3 12
50
2 100 4
44 11 84 21
16.7 1 100 6
50
6 83.3 10
57.1 4 71.4 5
5
1 100 20
0
0 100 12
0
0 100 7
100 1 100 1
0
0 89.5 17
0
0 90.9 10
0
0 100 6
0
0
50
1
15.4 2 100 13
16.7 1 100 6
16.7 1 100 6
0
0 100 1
26.7 4 93.3 14
16.7 1 100 6
50
3 100 6
0
0 66.7 2
9.1 1 100 11
20
1 100 5
0
0 100 5
0
0 100 1
8.3 1 100 12
16.7 1 100 6
0
0 100 5
0
0 100 1
16.7 4 83.3 20
0
0 80.0 4
25
3 83.3 10
14.3 1 85.7 6
20
4
90 18
22.2 2 100 9
28.6 2 100 7
0
0
50
2
16 15.8
3
0
0
68.4 13 31.6
6
0
0
68.4 13 31.6
8
4
4
16
8
5
3
7
4
2
1
1
2
0
3
0
2
1
6
0
5
1
0
0
0
0
0
0
0
13.3
20
12.5
0
0
0
0
0
0
0
0
2
1
1
0
88.9 8 11.1 1
33.3 2 66.7 4
75
3
25
1
47.4 9 52.6 10
37.5 3 62.5 5
71.4 5 28.6 2
25.0 1
75
3
93.3 14 6.7 1
100 5
0
0
87.5 7 12.5 1
100 2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
88.9 8 11.1 1 100 9
33.3 2 66.7 4 83.3 5
75
3
25
1 100 4
47.4 9 52.6 10 84.2 16
37.5 3 62.5 5 87.5 7
71.4 5 28.6 2 71.4 5
25
1
75
3 100 4
93.3 14 6.7 1 93.3 14
100 5
0
0 100 5
87.5 7 12.5 1 87.5 7
100 2
0
0 100 2
11.1
33.3
0
15.8
0
28.6
25
40
0
62.5
50
eschweizerbart_xxx
6
94.7 18
BFO
1
%
n
4.5 1
0
0
7.7 1
0
0
16
4
0
0
16.7 2
28.6 2
0
0
0
0
0
0
0
0
5.3 1
9.1 1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
16.7 4
20
1
16.7 2
14.3 1
10
2
0
0
0
0
50
2
2
%
0
0
0
0
0
0
0
0
0
0
0
0
5.3
0
0
50
0
0
0
0
6.7
0
0
33.3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
n
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
5.3
1
0
0
0
16.7
0
15.8
12.5
28.6
0
6.7
0
12.5
0
0
1
0
3
1
2
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
338
E. A. Peralta, L. H. Luna, A. F. Gil
Table 5. continued.
Enthesis
0
%
Biceps
81.8
F
100
M
76.9
IN
75
Brachialis
100
F
100
M
100
IN
100
Extensor
100
F
100
M
100
IN
100
Flexor
100
F
100
M
100
IN
100
Infraspinatus 92.3
F
83.3
M
100
IN
100
Subscapularis 73.3
F
66.7
M
83.3
IN
66.7
Supraspinatus 100
F
100
M
100
IN
100
Teres minor
100
F
100
M
100
IN
100
Triceps
100
F
100
M
100
IN
100
Iliopsoas
100
F
100
M
100
IN
100
Gluteus
minimus
F
M
IN
Gluteus medius
F
M
IN
Plantaris
F
M
IN
n
18
5
10
3
25
6
12
7
20
12
7
1
19
11
6
2
12
5
6
1
11
4
5
2
11
5
5
1
12
6
5
1
24
5
12
7
20
9
7
4
94.7 18
ERO
1
%
n
18.2 4
0
0
23.1 3
25
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
7.7 1
16.7 1
0
0
0
0
20
3
33.3 2
16.7 1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
%
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
6.7
0
0
33.3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
n
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
%
90.9
60
100
100
100
100
100
100
95
100
100
0
100
100
100
100
84.6
83.3
83.3
100
86.7
100
83.3
66.7
90.9
80
100
100
100
100
100
100
100
100
100
100
100
100
100
100
n
20
3
13
4
25
6
12
7
19
12
7
0
19
11
6
2
19
8
7
4
18
9
5
4
20
9
7
4
11
5
5
1
13
6
5
2
10
4
5
1
FPO
1
%
n
9.1 2
40
2
0
0
0
0
0
0
0
0
0
0
0
0
5
1
0
0
0
0
100 1
0
0
0
0
0
0
0
0
15.4 2
16.7 1
16.7 1
0
0
13.3 2
0
0
16.7 1
33.3 1
9.1 1
20
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
%
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
n
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
%
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100.
100
100
100
100
100
100
100
100
81.8
80
80
100
100
100
100
100
100
100
100
100
100
100
100
100
n
22
5
13
4
25
6
12
7
20
12
7
1
19
11
6
2
13
6
6
1
15
6
6
3
9
4
4
1
12
6
5
1
24
5
12
7
20
9
7
4
2
0
MPO
1
%
n
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
9.1 1
20
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
CAV
2
0
1
%
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
9.1
0
20
0
0
0
0
0
0
0
0
0
0
0
0
0
n
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
%
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
100
n
22
5
13
4
25
6
12
7
20
12
7
1
19
11
6
2
13
6
6
1
15
6
6
3
11
5
5
1
12
6
5
1
24
5
12
7
20
9
7
4
%
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
n
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
5.3
1
0
0
94.7 12
5.3
1
0
0
100
19
0
0
0
0
94.7 18
5.3
1
88.9 8 11.1
100 6
0
100 4
0
84.2 16 15.8
75
6
25
85.7 6 14.3
100 4
0
86.7 13 6.7
100 5
0
87.5 7
0
50
1
50
1
0
0
3
2
1
0
1
0
0
1
0
0
0
0
0
0
0
6.7
0
12.5
0
0
0
0
0
0
0
0
1
0
1
0
100 6
0
83.3 5 16.7
100 1
0
100 24
0
100 5
0
100 12
0
100 7
0
86.7 13 6.7
100 5
0
87.5 7
0
50
1
50
0
1
0
0
0
0
0
1
0
0
1
0
0
0
0
0
0
0
6.7
0
12.5
0
0
0
0
0
0
0
0
1
0
1
0
100 9
100 6
100 4
100 19
100 8
100 7
100 4
100 15
100 5
100 8
100 2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
100 9
0
100 6
0
75
3
25
89.5 17 10.5
100 8
0
71.4 5 28.6
100 4
0
100 15
0
100 5
0
100 8
0
100 2
0
0
0
1
2
0
2
0
0
0
0
0
eschweizerbart_xxx
Inferences on mobility and subsistence patterns from degenerative joint disease
and ankle joints (81.8%), and on the surface in the elbow
(13.3%) and knee (12.5%). Sex differences were not statistically significant.
339
Analysis (PCA) showed that BF1 was the variable with the
highest loading at PC1 followed by TC (Figs 2 and 3). Males
had higher frequencies of entheseal changes in the lower
limbs than females, and females showed higher frequencies of EC in some upper limb entheses in comparison with
males (Table 5). The Mann-Whitney test showed statistically
significant differences between males and females only for
the lower limbs considering BF1 (p = 0.01). Humeral and
femoral entheseal changes in terms of robusticity did not
significantly correlate with body size (composite variable
for humerus and femur) or sex: Humeral Entheseal Changes/
Humeral Size = 0.015; Humeral Entheseal Changes/Sex =
0.046; Femoral Entheseal Changes/Femoral Size = 0.094;
Femoral Entheseal Changes/Sex = 0.042.
Entheseal changes
Regarding fibrous entheses in the upper limbs, the pectoralis major and the supinator showed the highest percentages
in terms of robusticity (35.7% and 30.4%, respectively)
(Table 3). The pronator quadratus (20%), deltoid (15.4%),
and brachioradialis (5.9%) had lower frequencies of changes
associated with robusticity. The lower limb entheses showed
percentages ranging from 30 to 40% for score 1 and from 6
to 18% for scores 2 and 3. The most affected entheses were
the gluteal tuberosity (57.9%) and the linea aspera (45.5%).
Ossifications were observed only in the lower limbs in the
soleal line of the tibia (10%) and linea aspera of the femur
(13.6%).
Males and females showed similar percentages of
robusticity in the lower limbs at the linea aspera and gluteal tuberosity entheses. Nevertheless, females were more
affected at the pectoralis major and deltoid insertions in
comparison with males, while males were more affected
in the remaining entheses. Considering the total scores for
each enthesis (Table 4), females exhibit remarkably higher
means in the upper extremities, while males display higher
means in the lower extremities. Statistically significant differences between males and females were corroborated at
the quadriceps.
Regarding fibrocartilaginous entheses, the large number
of features hinders variability analysis. Principal Components
CS-1 and regional contexts
Comparing DJD mean scores for the upper limb joints, the
prevalence of females from CS-1 is higher than in Pampa in
the shoulder and elbow, while males have a higher prevalence in Pampa than in CS-1 in all joints (Table 6). The differences are not statistically significant between the upper
extremities of CS-1 and Pampa. Comparing lower limb
joints of CS-1 and Pampa samples, the prevalence of DJD
in the Pampa group is higher than in CS-1. The differences
are statistically significant between the lower extremities of
males for both samples. Comparing EC for fibrous entheses,
the Pampa sample shows a higher prevalence of robusticity/
stress lesions in both upper and lower extremities than the
CS-1 sample, and the difference is significant for both males
and females (Table 6).
Table 6. Entheseal changes and degenerative joint disease mean scores and rank ordering for CS-1 and hunter-gatherer group of
Pampa comparing lower limbs. p-values of the Mann-Whitney test are shown (values ≤ 0.05 are indicated in italic). n = number of
elements, x = mean score.
CS-1
Joint
Pampa
Females
n
Males
x
n
Females
x
n
Males
p (CS-1 vs. Pampa)
x
n
x
Shoulder
8
0.5
6
0
9
0.11
9
0.33
Elbow
19
0.47
30
0.66
8
0.12
8
1.12
Wrist
10
0.2
16
0.43
8
0.75
9
0.55
Hip
13
0
8
0
8
0.38
6
0.83
Knee
16
0.56
16
0.38
8
0
9
1.11
Ankle-foot
8
0.38
11
0.82
9
0.78
10
1.6
Pectoralis major
7
0.85
3
0.33
6
0.66
4
1.25
Deltoides
6
0.16
2
0
6
1.33
7
2
0
4
0.25
6
0.83
8
2
Linea aspera
8
0.63
8
0.75
14
1
19
2.57
Quadriceps
8
0.125
7
0.86
14
1.25
19
2
Soleal line
10
0.5
9
0.67
14
0.6
19
2.14
0.16 (Females)
0.55 (Males)
0.46 (Females)
0.00 (Males)
Enthesis
Brachioradialis
eschweizerbart_xxx
0.00 (Females)
0.00 (Males)
0.03 (Females)
0.00 (Males)
340
E. A. Peralta, L. H. Luna, A. F. Gil
Table 7. Comparisons of prevalence of degenerative joint disease between CS-1 and Córdoba samples. p-values of the MannWhitney test are shown (values ≤ 0.05 are indicated in italic).
Joints
Margin
Surface
n
%
shoulder_cba
6
10.2
shoulder_cs1
6
31.6
elbow_cba
10
10.6
elbow_cs1
33
50
wrist_cba
3
3.6
wrist_cba1
11
30.6
0.00
hip_cba
0
0
hip_cs1
0
0
1
knee_cba
4
5.5
knee_cs1
18
45
ankle_cba
4
4
ankle_cs1
13
56.5
p
0.001
0.00
0.00
0.00
n
%
5
5
4
21.1
17
10.4
10
15.2
6
4.8
1
2.8
8
8.2
1
3.4
5
3.9
3
7.5
2
1.2
0
0
p
0.01
0.21
0.64
0.44
0.32
0.60
in CS-1 than in the Córdoba sample. An explanation would
be that the CS-1 sample is composed of a greater percentage of middle-aged adults than the Córdoba sample, which
mostly includes young adults (Salega 2016; Peralta 2019),
and it is known that DJD is strongly correlated with age
(Jurmain 1977; Masoro & Austad 2006; Seibel et al. 2006).
Interestingly, the upper extremities showed a higher prevalence of DJD and EC in CS-1 with statistically significant
differences, indicating a distinctive spectrum of manual tasks
between both samples. The lower extremities did not show
any statistically significant differences concerning EC, suggesting that the pattern of mobility in CS-1 would be similar
to that of low-level producers from Córdoba.
CS-1 males were significantly more affected than females
at the fibrocartilaginous entheses of the lower limb, which
could be interpreted as an indicator of a division of labor in
which males would have been mainly involved in hunting
trips, while females would have stayed at the basecamp doing
a broad range of manual tasks. In this sense, although statistical differences are not significant, females were remarkably
more affected than males in some joints and fibrous entheses
of the upper extremities, which would be related to the incorporation of new technologies (such as ceramics and grinding
tools) and demands inherent to a wide spectrum of resource
exploitation (Neme 2007; Gil et al. 2018). These sexual differences could be a consequence of body size, as noted by
Weiss (2003; 2004; 2007). In this regard, the correlation
tests did not show high concordance between body size and
entheseal changes in terms of robusticity, supporting that the
difference in entheseal changes could be explained by the
distinctive patterns of body use by males and females.
Occupational stress markers help understand past human
subsistence strategies; however, their use requires some cau-
In CS-1, DJD frequencies are higher than those for lowlevel producers, and periarticular differences are statistically
significant for the shoulder, elbow, wrist, knee, and ankle
(Table 7). Concerning fibrocartilaginous entheses, ECs are
more frequent in the CS-1 sample than in the Córdoba sample. Subscapularis, extensor, flexor, triceps, gluteus medius,
and plantaris insertions show a higher prevalence of BF1,
BF, and ERO in the Córdoba sample, but differences are not
statistically significant. Statistically significant differences
were observed at the insertions of the infraspinatus, in TC
and FPO; subscapularis, in TC and FPO; biceps in ERO1,
TC and ERO, and triceps, in TC (Table 8).
Discussion and conclusions
The goal of this paper was to discuss patterns of mobility
and potential manual activities in an area where hunter-gatherers and low-level food producers interacted. Concerning
the expectations raised at the beginning of this study, lower
frequencies of DJD and EC at fibrous entheses in lower
extremities and higher frequencies in upper extremities
were expected in CS-1 compared to hunter-gatherer populations; at the same time, similar frequencies of DJD and
EC at fibrocartilaginous entheses in both extremities were
expected in CS-1 compared to groups with low-level food
production. As regards terrestrial hunter-gatherer organization, CS-1 trends can be explained as a result of the lower
physical demand associated with low mobility and a reduced
workload (Scabuzzo 2012). With regard to low-level food
producers from Córdoba, physical activities in CS-1 would
be quite different and more intense. Notably, both upper and
lower extremities were significantly more affected by DJD
eschweizerbart_xxx
341
Inferences on mobility and subsistence patterns from degenerative joint disease
Table 8. Comparisons of prevalence of EC between CS-1 and Córdoba samples (considering only fibrocartilaginous entheses).
p-values of the Mann-Whitney test are present (values ≤ 0.05 are indicated in italic).
Entheses
BF1
n
%
supraspinatus_cba
0
0
supraspinatus_cs1
1
9.1
infraspinatus_cba
0
0
infraspinatus_cs1
0
0
subscapularis_cba
6 16.7
subscapularis_cs1
4 26.7
extensor_cba
12 32.4
extensor_cs1
3
15
flexor_cba
3
8.6
flexor_cs1
3 15.8
biceps_cba
6 14.3
biceps_cs1
6 27.3
triceps_cba
4 11.4
triceps_cs1
1
4.2
gluteus medius_cba
5 22.7
gluteus medius_cs1
3 15.8
plantaris_cba
14 58.3
plantaris_cs1
8 53.3
ERO1
p
0.09
1
0.43
0.16
0.04
0.21
0.34
0.07
0.77
n
%
1
2.9
1
9.1
0
0
2 15.4
1
2.8
4 26.7
0
0
1
5
0
0
0
0
0
0
10 45.5
2
5.7
4 16.7
0
0
10 52.6
1
4.2
1
6.7
TC
p
0.41
1
0.54
1
1
0.004
0.25
1
0.46
n %
0
0
1 9.1
0
0
2 15.4
0
0
4 26.7
0
0
1
5
0
0
0
0
0
0
10 45.5
0
0
4 16.7
1 4.3
10 52.6
0
0
1 6.7
BF
p
0.07
0.02
0.00
n
%
0
0
0
0
0
0
0
0
11 29.7
0.17
1
0.00
0.01
0.18
0.52
1
6.7
0
0
0
0
2
5.4
2
5.3
1
2.4
1
4.5
5
14.3
4
16.7
2
8.7
3
15.8
0
0
1
6.7
ERO
p
1
1
0.08
1
0.50
0.67
0.81
0.43
0.21
n
%
2
5.4
0
0
1
2.9
1
7.7
3
8.1
4 26.7
0
0
0
0
1
2.7
0
0
0
0
4 18.2
3
8.6
0
0
2
8.7
3 15.8
1
4
2 13.3
FPO
p
0.46
0.48
0.08
1
0.50
0.00
0.15
0.15
0.30
n
%
0
0
1
9.1
0
0
2 15.4
0
0
2 13.3
0
0
1
5
0
0
0
0
0
0
2
9.1
0
0
0
0
0
0
0
0
0
0
2 13.3
MPO
p
0.07
0.02
0.03
0.17
1
0.05
1
1
0.07
n
%
1
2.7
2 18.2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
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Manuscript received: 03 June 2021
Revisions requested: 11 August 2021
Revised version received: 11 November 2021
Accepted: 11 November 2021
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