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Biomechanical approach to the reconstruction of activity patterns in Neolithic Western Liguria Italy.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 131:447–455 (2006)
Biomechanical Approach to the Reconstruction of
Activity Patterns in Neolithic Western Liguria, Italy
Damiano Marchi,1* Vitale S. Sparacello,2 Brigitte M. Holt,3 and Vincenzo Formicola4
1
Department of Biological Anthropology and Anatomy, Duke University, Durham, North Carolina 27708
Dipartimento di Scienze Antropologiche, University of Genoa, Genoa, Italy 16159
3
Department of Anthropology, University of Massachusetts, Amherst, Massachusetts 01003
4
Dipartimento di Biologia, University of Pisa, Pisa, Italy 56126
2
KEY WORDS
cross-sectional geometry; bioarchaeology; femur; humerus; mobility
ABSTRACT
This paper investigates the changes in
upper and lower limb robusticity and activity patterns
that accompanied the transition to a Neolithic subsistence
in western Liguria (Italy). Diaphyseal robusticity measures were obtained from cross-sectional geometric properties of the humerus and femur in a sample of 16 individuals (eight males and eight females) dated to about 6,000–
5,500 BP. Comparisons with European Late Upper Paleolithics (LUP) indicate increased humeral robusticity in
Neolithic Ligurian (NEOL) males, but not in females, with
a significant reduction in right-left differences in both
sexes. Sexual dimorphism in robusticity increases in
upper and lower limb bones. Regarding the femur, while
all female indicators of bending strength decrease steadily
through time, values for NEOL males approach those of
LUP. This suggests high, and unexpected, levels of me-
chanical stress for NEOL males, probably reflecting the
effects of the mountainous terrain on lower limb remodeling. Comparisons between NEOL males and a small sample of LUP hunter-gatherers from the same area support
this interpretation. In conclusion, cross-sectional geometry data indicate that the transition to Neolithic economies in western Liguria did not reduce functional requirements in males, and suggest a marked sexual division of
labor involving a more symmetrical use of the upper limb,
and different male-female levels of locomotory stress.
When articulated with archaeological, faunal, paleopathological, and ethnographic evidence, these results support
the hypothesis of repetitive, bimanual use of axes tied to
pastoral activities in males, and of more sedentary tasks
linked to agriculture in females. Am J Phys Anthropol
131:447–455, 2006. V 2006 Wiley-Liss, Inc.
Western Liguria represents one of the most important
areas for understanding the Upper Paleolithic and Neolithic peopling of Southern Europe. This region is extraordinarily rich in paleoanthropological material, including
Early Upper Paleolithic (EUP) and Late Upper Paleolithic
(LUP) skeletons from the Grimaldi and Arene Candide
caves, as well as several Neolithic (NEOL) and Bronze Age
burials found in Arene Candide and other nearby caves.
Such material provides the rare opportunity to study regional diachronic variations in biological parameters such
as health status, body size, and robusticity in the framework of changing economic and lifestyle contexts.
The spread of the Neolithic in Liguria dates back to
the beginning of the seventh millennium BP (uncalibrated chronology), and is characterized by a dramatic
drop in hunting activities and by a well-established production economy. Faunal and archaeological evidence
suggests that, from the beginning, Neolithic subsistence
focused on terrestrial resources, and that pastoralism
played an important economic role (Macphail et al.,
1997; Rowley-Conwy, 1997). The importance of pastoralism was probably due both to the hilly nature of the terrain (less suitable for agriculture) and to the presence of
caves and rock shelters providing natural ways to control the movements of animals (Maggi and Nisbet, 2000).
The increase in the number of sheep and the appearance
of goats during the Middle Neolithic (Rowley-Conwy,
1997) imply an increased reliance on animal domestication through time.
While agriculture was probably not the focus of Neolithic subsistence in Liguria, evidence of agricultural
practices can be drawn from the presence of querns1
(Starnini and Voytek, 1997; Tiné, 1999) and from the
carbonized remains of cereals (barley and wheat), as well
as their impressions on clay (Ewett and Renfrew, 1971;
Arobba et al., 1987, 1997). There is no evidence of
legumes, sometimes accompanying the cultivation of barley and wheat in Neolithic Italy, until later, at the beginning of the Copper Age (Arobba et al., 1997).
Comparisons between LUP and NEOL western Ligurian samples show a diachronic reduction in stature and
a decline in health conditions, in keeping with trends
known to accompany the transition to agriculture elsewhere (Cohen and Armelagos, 1984). Osteometric analy-
C 2006
V
WILEY-LISS, INC.
C
1
Quern stones are a pair of stone tools for hand-grinding a wide
variety of materials. The lower (stationary) stone is called a quern,
while the upper (mobile) stone is called a handstone.
Grant sponsor: Ministero dell’Istruzione, dell’Universı̀tà e della
Ricerca Cofinanziamento (MIUR COFIN) 2003; Grant numbers:
PRIN 2003054059-002.
*Correspondence to: Damiano Marchi, Department of Biological
Anthropology and Anatomy, 05 Biological Sciences Bldg., Science
Dr., Box 90383, Durham, NC 27708-0383.
E-mail: dmarchi1@duke.edu
Received 5 November 2005; accepted 22 February 2006.
DOI 10.1002/ajpa.20449
Published online 9 May 2006 in Wiley InterScience
(www.interscience.wiley.com).
448
D. MARCHI ET AL.
sis, however, shows that, while lower limb robusticity
decreases, there is an increase in upper limb robusticity
in the NEOL sample (Formicola, 1983, 1986, 1997; Formicola and Canci, 2003). The observed changes in robusticity suggest that the Neolithic transition in Liguria did
not result in a wholesale decline in bone strength, as
documented at many other sites, and underscores the
need to use measures of robusticity that reflect specific
mechanical loads and physical activity.
Traditional osteometric parameters provide only a
baseline measure of robusticity (Trinkaus and Ruff,
2000). By taking into account the internal architecture
of the diaphysis, we can better analyze robusticity and
its possible links with specific behaviors (Biewener,
1982; Jungers et al., 1998; Ruff, 2000). Numerous studies showed that, due to the plasticity of cortical bone, diaphyseal shape and size vary according to activity levels
and types (Jones et al., 1977; Woo et al., 1981; Lanyon
et al., 1982; Burr et al., 1989; Ruff, 1992; Trinkaus
et al., 1994). The cross-sectional geometry (CSG) of long
bone diaphyses was used extensively to assess the
effects of subsistence pattern changes on past populations (Ruff, 1987; Bridges, 1989; Ledger et al., 2000;
Stock and Pfeiffer, 2001; Holt, 2003; Weiss, 2005). In
particular, it was demonstrated that bone cross-sectional
shape more accurately reflects types of mechanical
loads, while bone cross-sectional size more accurately
reflects general levels of mechanical loads (Ruff, 1995;
Trinkaus and Ruff, 1999; Stock and Pfeiffer, 2001). For
instance, there is strong evidence that running and
long-distance travel over uneven surfaces increase
antero-posterior bending loads in the femur and tibia
(Ruff and Hayes, 1983; Ruff, 1987). This relationship
was used extensively to evaluate changing patterns of
mobility in prehistoric populations (Bridges, 1989;
Larsen, 1995; Trinkaus et al., 1998; Trinkaus and Ruff,
1999; Bridges et al., 2000; Holt, 2003).
We examine here the patterns of physical activity in a
Ligurian sample across the Paleolithic-Neolithic transition. Evidence from the archaeological record associated
with the western Ligurian Neolithic skeletal samples
indicates an economy based on herding and agriculture.
Given the known ethnographic documentation of sexual
division of labor, we expect a differing involvement of
the sexes in those activities, with males more involved
in pastoral activities, and females in more sedentary agricultural tasks. If this hypothesis is correct, we should
find: 1) increased sexual dimorphism in lower limb
robusticity, 2) maintenance of a high level of locomotory
stress in males, and 3) decreased upper limb asymmetry
in robusticity in males, resulting from the decreased use
of hunting-related throwing technologies.
MATERIALS
The NEOL sample includes 16 individuals (eight
males and eight females; see Appendix A for sample
composition) from three caves (Arene Candide, Arma
dell’Aquila, and Pollera) located near Finale Ligure
(Savona, Italy). The skeletons belong to the Middle Neolithic (Square-Mouthed Pottery culture), radiometrically
dated between 6,000–5,500 BP (uncalibrated chronology; Maggi, 1997a). The bones studied include the humerus and femur. To evaluate the degree of upper limb
asymmetry, both right and left humeri were studied,
while based on the minor degree of bilateral asymmetry
in lower limb bones (Ruff and Jones, 1981; Auerbach
and Ruff, 2006), for the femur, only one side was analyzed (the right when it was the best preserved; the
left, otherwise). Due to differences in the proportion of
cortical to total area between subadults and adults
(Ruff et al., 1994), only full adults (epiphyses completely sealed) not showing signs of senescence (osteoarthritis) were sampled. Moreover, only complete skeletons in a good state of preservation were taken into
account, allowing a reliable sex diagnosis. Data on
upper and lower limb bones from LUP and Mesolithic
(only lower limb bones) samples (Appendices B and C)
provided comparative material (Churchill, 1994; Holt,
1999, 2003). The major part of the LUP and the totality
of the Mesolithic sample are pan-European, while the
Neolithic sample is only from Liguria. Thus, many of
the differences between NEOL and comparative samples might result from terrain and other localized
effects, rather than (or in combination with) differences
in subsistence strategy. The effect of the composition of
the comparative sample is considered in the subsequent
interpretation of results.
METHODS
Reconstruction of cross sections
Subperiosteal and estimates of endosteal contours
were reconstructed, using polysiloxane molds and measurements of biplanar radiographs of the diaphysis. This
method proved to be accurate and economical for reconstructing cross sections without damaging important fossil material (Trinkaus and Ruff, 1989; Churchill, 1994;
O’Neill and Ruff, 2004). Cross sections were reconstructed at the mid-distal humerus (35% bone length)
and midshaft femur (50% bone length). The 35% level in
the humerus falls below the deltoid tuberosity. Reconstructed cross sections were analyzed with a version of
the program SLICE (Nagurka and Hayes, 1980), adapted
as a macroroutine for use with NIH Image 1.52. Statistical analyses were carried out with Statistica 5.1 (Statsoft, 1997).
Cross-sectional geometric dimension
standardization
The robusticity of the humerus and femur is influenced by the combined effects of body mass and physical activity (Ruff et al., 1991, 1993). To isolate the
effects of activity, estimates of strength must be scaled
to some measure of body mass. Given strong evidence
that body breadth affects long bone diaphsyseal structure (Ruff et al., 1991; Ruff, 2000), we standardized
femoral cross-sectional properties by body mass, calculated using a multiple regression formula based on stature and bi-iliac breadth (Bib) (Ruff et al., 2005). Four
male and four female skeletons were not sufficiently
complete to measure Bib. In these cases, Bib was estimated from maximum femoral length (Trinkaus, 1996).
Further details can be found in Holt (1999). Stature
was estimated using femoral lengths and equations calibrated on European Neolithic skeletal samples (Formicola and Franceschi, 1996), while body breadth was
derived directly from bi-iliac breadth (M-2 in Martin
and Saller, 1957).
Cross-sectional areas (cortical area, CA; total area, TA)
are related to cross-sectional rigidity in axial loading
(compression or tension) (Ruff et al., 1993), and are di-
American Journal of Physical Anthropology—DOI 10.1002/ajpa
449
NEOLITHIC LIGURIAN ACTIVITY PATTERNS
TABLE 1. Temporal differences in diaphyseal geometric
properties of humerus: males
LUP1
Mean (n)
Right
TA4,5
CA
Ix6
Iy
Imax
Imin
J
Imax/Imin
Ix/Iy
Left
TA
CA
Ix
Iy
Imax
Imin
J
Imax/Imin
Ix/Iy
497.11
403.73
129.94
115.57
NEOL2
SD
(6)
(6)
(5)
(5)
89.30
78.73
48.95
50.36
245.52 (5)
98.78
1.15 (8)
0.13
452.85
337.73
104.34
93.70
(6)
(6)
(5)
(5)
63.73
28.38
24.95
29.18
199.44 (5)
47.36
1.12 (7)
0.19
Mean (n)
SD
525.43
409.54
158.02
144.71
161.86
140.23
302.73
1.18
1.12
(7)
(7)
(6)
(6)
(6)
(6)
(6)
(7)
(7)
85.99
74.79
47.43
45.00
45.50
47.35
91.98
0.08
0.10
488.92
378.07
126.81
119.22
134.63
111.40
246.46
1.24
1.09
(7)
(7)
(6)
(6)
(6)
(6)
(6)
(7)
(7)
56.87
61.03
38.08
36.35
36.89
37.42
63.43
0.10
0.10
TABLE 2. Temporal differences in diaphyseal geometric
properties of humerus: females
LUP1
Percent
difference3
Mean (n)
Right
TA4,5
CA
Ix6
Iy
Imax
Imin
J
Imax/Imin
Ix/Iy
5.70
1.44
21.61
25.21
23.30
2.31
Left
TA
CA
Ix
Iy
Imax
Imin
J
Imax/Imin
Ix/Iy
7.97
11.94
21.53
27.23
23.57
2.55
1
Comparative Late Upper Paleolithic sample (from Churchill,
1994).
2
Ligurian Neolithic sample.
3
Percent difference ¼ ((NEOL value LUP value)/LUP value))
3 100).
4
All variables are explained in text.
5
CA and TA are divided by body mass.
6
Ix, Iy, Imax, Imin, and J are divided by product of body mass
and square of bone length.
vided by body mass. Second moments of area are used to
evaluate resistance to bending. These include the area
moments of inertia about the M-L axis (Ix) and about the
A-P axis (Iy), proportional to bending rigidity in the A-P
and M-L planes, respectively; the maximum (Imax) and
the minimum Imin area moments of inertia, proportional
to maximum and minimum bending rigidity, respectively; and the polar moment of inertia (J), polar
moment of inertia, proportional to average bending torsional rigidity. Bending and torsional second moments of
area are divided by the product of body mass and the
second power of bone length (Ruff, 2000, and personal
communication).
The upper limb of hominids, whose primary function
is not locomotor and therefore not weight-bearing, may
not scale strictly to body mass. For this reason, in some
previous works (Churchill, 1994), cross-sectional areas
(cortical and total areas) were standardized by dividing
by humeral articular length squared, and area moments of inertia (Ix, Iy, Imax, Imin, and J) by dividing by
humeral articular length raised to the power of four.
More recently, however, it was empirically demonstrated that the humerus follows the same scaling relationships to body size as the femur (Ruff, 2000). Humeral length alone does not take into account differences
in body proportions, which appear to affect upper as
well as lower limb bones. For a particularly broad-bodied sample such as the Ligurian Neolithic, the use of
humeral length alone as a body-size measure would
tend to overestimate relative humeral strength. Thus,
following Ruff (2000), we used the same method of
standardization for the humerus and femur.
513.66
394.94
135.53
101.81
NEOL2
SD
(3)
(3)
(3)
(3)
84.92
116.45
43.39
39.70
237.34 (3)
83.08
1.36 (4)
0.07
443.39
305.72
112.84
86.01
(5)
(5)
(5)
(5)
48.24
72.46
17.71
15.12
198.86 (5)
31.56
1.32 (5)
0.11
Mean (n)
SD
426.38
320.68
92.64
82.61
96.25
79.00
175.25
1.22
1.12
(8)
(8)
(8)
(8)
(8)
(8)
(8)
(8)
(8)
39.02
20.76
18.52
11.38
16.93
11.70
27.51
0.11
0.17
417.37
317.43
92.79
82.62
97.49
77.17
175.41
1.25
1.12
(8)
(8)
(8)
(8)
(8)
(8)
(8)
(8)
(8)
31.70
24.39
16.87
8.51
14.55
9.64
23.41
0.09
0.15
Percent
difference3
16.99*
18.80
31.65*
18.86
26.16
17.22*
5.87
3.83
17.77**
3.40
11.79
15.06*
1
Comparative Late Upper Paleolithic sample (from Churchill,
1994).
2
Ligurian Neolithic sample.
3
Percent difference ¼ ((NEOL value LUP value)/LUP value))
3 100).
4
All variables are explained in the text.
5
CA and TA are divided by body mass.
6
Ix, Iy, Imax, Imin, and J are divided by product of body mass
and square of bone length.
*
P < 0.05. All significance levels are from Mann-Whitney Utest between temporal groups within sex.
**
0.05 < P < 0.1.
Ratios of area moments of inertia (Ix/Iy and Imax/Imin)
were calculated to evaluate variability in diaphyseal
shape (Ruff and Hayes, 1983). Given that humeral Imax
and Imin were not available for the LUP comparative
sample (Churchill, 1994), only Ix/Iy was included for the
humerus.
Due to the small sample size and possible non-normal
sample distribution, statistical evaluation of humeral differences between NEOL and the comparative LUP sample was carried out using a nonparametric Mann-Whitney U-test between temporal groups within sex. The
same test was used for humeral bilateral asymmetry
analyses. Statistical evaluation of femoral differences
among NEOL and the comparative LUP and Mesolithic
samples was carried out using a Kruskal-Wallis analysis
of ranks between temporal groups within sex, followed
by post hoc multiple comparisons of mean ranks for all
group tests. Sexual differences for the humerus and femur within a time period were analyzed using the
Mann-Whitney U-test.
RESULTS
Humerus
Cross-sectional geometric properties, temporal differences, and changes in asymmetry and in sexual dimorphism are reported in Tables 1–4. Male humeral crosssectional indicators of robusticity (Ix, Iy, CA, TA, and J)
consistently increase relative to the LUP sample, for
both the left and right sides, although the change is
American Journal of Physical Anthropology—DOI 10.1002/ajpa
450
D. MARCHI ET AL.
TABLE 3. Bilateral asymmetry of humerus of Ligurian
Neolithics and of comparative Late Upper Paleolithic sample
LUP1 (n)
Males
TA3,4
CA
Ix
Iy
Imax
Imin
J
Imax/Imin
Ix/Iy
Females
TA
CA
Ix
Iy
Imax
Imin
J
Imax/Imin
Ix/Iy
17.91
25.25
28.57
25.33
(5)
(5)
(3)
(3)
26.93 (3)
0.54 (6)
16.45
22.85
11.93
11.97
(3)
(3)
(3)
(3)
11.91 (3)
0.16 (3)
NEOL2 (n)
8.84
8.03
21.15
17.57
18.20
20.90
15.80
3.26
2.56
(6)
(6)
(4)
(4)
(4)
(4)
(4)
(6)
(6)
2.03
1.09
0.36
0.41
1.60
1.10
0.35
2.99
0.07
(8)
(8)
(8)
(8)
(8)
(8)
(8)
(8)
(8)
TABLE 4. Sexual dimorphism of humerus of Ligurian
Neolithics and of comparative Late Upper Paleolithic sample
LUP-NEOL5
8
**
NS
NS
NS
NS
*
*
NS
NS
NS
NS
1
Comparative Late Upper Paleolithic sample (from Churchill,
1994).
2
Ligurian Neolithic sample.
3
All variables are explained in text.
4
Bilateral asymmetry calculated as: ((right left)/(mean of
right and left)) 3 100.
5
Probabilities of between-sample comparison. NS, nonsignificant difference. Significance levels are from Mann-Whitney Utest between temporal groups within sex.
*
P < 0.05.
**
P < 0.01.
8
0.05 < P < 0.1.
never statistically significant (Table 1). Female cross-sectional parameters decrease relative to the LUP sample,
with the exception of left CA. The change is statistically
significant only for right TA and Ix (Table 2).
The diaphyseal shape ratio (Ix/Iy) decreases in both
NEOL males and females compared to the LUP sample.
In males, the reduction is slight (about 2% for both the
right and left sides). In females, the reduction is more
than 15%. Changes are statistically significant only for
females (Table 2).
Asymmetry in humeral robusticity decreases in both
NEOL males and females (Table 3; significant results
only for female CA and TA and male CA) compared to
LUP. This is a consequence of the relatively greater
increase in robusticity in the left humerus in males,
while in females, the decreased asymmetry results from
the greater decline in robusticity on the right side. Bilateral asymmetry in the NEOL sample is never statistically significant. Only CA is significant for the LUP sample (P < 0.05, result not reported in Table 3).
The NEOL sample shows significant sexual dimorphism (males larger than females) for all cross-sectional
indicators of robusticity (for both the right and left humerus). Differences in the indicators of diaphyseal shape
(Imax/Imin and Ix/Iy) (Table 4), however, are not significant.
Femur
Temporal differences, sexual dimorphism, and midshaft femoral CSG properties are listed in Tables 5 and
Right
TA3,4,5
CA
Ix6
Iy
Imax
Imin
J
Imax/Imin
Ix/Iy
Left
TA
CA
Ix
Iy
Imax
Imin
J
Imax/Imin
Ix/Iy
LUP1
NEOL2
3.22
2.23
4.12
13.52
23.23**
27.71**
70.58**
75.17**
66.17**
77.51**
72.74**
3.18
0.46
3.44
15.65*
2.13
10.47
7.53
8.94
0.29
15.15***
17.14**
19.10*
36.67*
44.298
38.09*
44.36**
40.51**
0.87
2.66
1
Comparative Late Upper Paleolithic sample (from Churchill,
1994).
2
Ligurian Neolithic sample.
3
All variables are explained in the text.
4
Sexual dimorphism calculated as: ((males females)/females)
3 100.
5
CA and TA are divided by body mass.
6
Ix, Iy, Imax, Imin, and J are divided by product of body mass
and square of bone length.
*
P < 0.05. All significance levels are from Mann-Whitney Utest between sexes within temporal group.
**
P < 0.01.
***
P < 0.001.
8
0.05 < P < 0.1.
6. NEOL males show lower (nonsignificant) cortical area,
total area, Iy, Imin, and J values than Mesolithic and
LUP males. However, NEOL males exhibit higher (nonsignificant) Ix and Imax values than Mesolithic males,
resulting in significantly higher diaphyseal shape ratios
(Table 5). All indicators of cross-sectional robusticity
decrease constantly through time in females, with the
exception of Iy, although none of these changes reach
statistical significance (Table 5). The NEOL sample
shows greater values of sexual dimorphism for Ix, Imax,
J, Imax/Imin, and Ix/Iy than both the LUP and Mesolithic
samples taken as comparisons. However, NEOL sexual
dimorphism is statistically significant only for indicators
of diaphyseal shape (Table 6).
DISCUSSION
Humerus
Cross-sectional geometric properties of LUP and
NEOL humeri indicate a pattern of increased robusticity
in NEOL males and decreased robusticity in females,
suggesting an increased manipulatory activity in males
but not females. The different trajectories affecting
males and females result in significantly increased sexual dimorphism for the NEOL sample, and indicate a
different degree of mechanical loading between sexes in
European hunter-gatherers and early farmers from western Liguria. Evidence for change in loading patterns is
American Journal of Physical Anthropology—DOI 10.1002/ajpa
451
NEOLITHIC LIGURIAN ACTIVITY PATTERNS
TABLE 5. Temporal differences in diaphyseal geometric properties of femur
1
MESO2
LUP
Mean (n)
SD
Mean (n)
NEOL3
SD
Mean (n)
Males
TA5,6
CA
Ix7
Iy
Imax
Imin
J
Imax/Imin
Ix/Iy
953.79
768.83
308.65
231.24
320.73
218.82
539.55
1.44
1.33
(12)
(12)
(12)
(12)
(12)
(12)
(12)
(12)
(12)
111.85
121.04
72.07
60.1
80.02
50.47
122.85
0.23
0.25
967.79
741.31
275.03
246.25
275.84
223.93
521.26
1.24
1.13
(22)
(22)
(22)
(22)
(16)
(16)
(22)
(18)
(24)
113.16
104.4
61.24
55.75
69.30
53.80
110.18
0.14
0.16
904.03
690.02
289.99
206.40
298.89
197.51
496.40
1.51
1.4
Females
TA
CA
Ix
Iy
Imax
Imin
J
Imax/Imin
Ix/Iy
951.46
751.83
280.16
221.41
294.16
207.43
501.60
1.40
1.25
(6)
(6)
(7)
(7)
(7)
(7)
(7)
(7)
(7)
143.67
183.81
94.78
45.50
88.91
48.62
133.27
0.19
0.27
940.17
749.55
254.26
228.61
265.94
205.72
482.98
1.31
1.12
(11)
(11)
(11)
(11)
(6)
(6)
(11)
(10)
(15)
81.29
86.38
33.70
38.74
38.46
26.77
60.99
0.14
0.2
875.99
680.10
238.49
203.66
243.73
198.43
442.15
1.23
1.17
Percent difference4
SD
NEOL-LUP
NEOL-MESO
(8)
(8)
(8)
(8)
(8)
(8)
(8)
(8)
(8)
148.06
141.84
96.84
63.47
99.69
62.29
158.13
0.22
0.17
5.22
10.25
6.04
10.74
6.81
9.74
8.00
8.63
5.26
6.85
7.85
4.96
17.32
6.50
13.26
5.61
21.77*
23.89**
(8)
(8)
(8)
(8)
(8)
(8)
(8)
(8)
(8)
62.3
28.67
55.18
26.48
53.02
26.35
75.12
0.17
0.2
7.93
9.54
14.87
8.01
17.15
4.34
11.85
12.14
6.40
6.83
9.26
6.20
10.91
8.35
3.55
8.45
6.11
4.46
1
Comparative Late Upper Paleolithic sample (from Holt, 1999).
Comparative Mesolithic sample (from Holt, 1999).
Ligurian Neolithic sample.
4
Percent difference ¼ ((Group 1 Group 2)/Group 2)) 3 100).
5
All variables are explained in text.
6
CA and TA are divided by body mass.
7
Ix, Iy, Imax, Imin, and J are divided by product of body mass and square of bone length.
*
P < 0.05. Significance levels are from Kruskal-Wallis analysis of ranks between temporal groups within sex.
**
P < 0.01.
2
3
also provided by the decreased lateralization exhibited
particularly by males that points to decreased unimanual work, such as throwing-based hunting activities,
likely modeling the dominant arm in LUP hunter-gatherers (Churchill et al., 1996; Holt et al., 2000; Schmitt
and Churchill, 2003)
A recurring tool found in Ligurian Middle Neolithic
archaeological deposits is the ax, mostly made from
high-quality stone (eclogite). Microwear traces indicate
that these tools were used in woodworking (Starnini and
Voytek, 1997), possibly for deforestation practices linked
both to agriculture and to provide grassland for domesticated animals (Garibaldi et al, 1996; Canci et al., 1998).
It was hypothesized that pollarding (recurrently cropping branches) of trees represented an alternative strategy to provide an additional source of animal fodder
(Maggi and Nisbet, 2000). Indications for this practice in
the Neolithic come from botanical analyses carried out
on sediments of Arene Candide, showing that a significant part of animal fodder was derived from branches
and leaves (Maggi, 1997b).
Studies performed by Canci et al. (1998), Canci and
Dini (2003), and Canci and Marini (2003) on the same
NEOL sample documented in males a high incidence of
bilateral enthesopathies at the insertions of the costoclavicular ligament, pectoralis major, latissimus dorsi,
and biceps and triceps brachii, as well as frequent cases
of dorsal and bisulcate patterns of the axillary border of
the scapula. Those changes are likely the effect of repetitive and intense movements of adduction and abduction
of the upper limb. These results, when articulated with
faunal and archaeological data, support the hypothesis
of a repetitive, bimanual use of axes tied to deforestation
and pollarding practices. The decreased lateralization of
TABLE 6. Sexual dimorphism of femur of Ligurian Neolithic
and of comparative Late Upper Paleolithic and Mesolithic
samples
TA4,5,6
CA
Ix7
Iy
Imax
Imin
J
Imax/Imin
Ix/Iy
LUP1
MESO2
NEOL3
0.25
2.26
10.17
4.44
9.03
5.49
7.57
5.71
9.60
3.22
0.10
8.67
9.20
5.53
10.69
8.89
5.34
0.89
3.20
1.46
21.60
1.34
22.63
0.46
12.27
22.76*
19.66*
1
Comparative Late Upper Paleolithic sample (from Holt, 1999).
Comparative Mesolithic sample (from Holt, 1999).
3
Ligurian Neolithic sample.
4
All variables are explained in text.
5
Sexual dimorphism calculated as: ((males females)/females)
3 100.
6
CA and TA are divided by body mass.
7
Ix, Iy, Imax, Imin, and J are divided by product of body mass
and square of bone length.
*
P < 0.05. Significance levels are from Mann-Whitney U-test
between sexes within temporal group.
2
NEOL male humeri would be in agreement with this
interpretation. Processing cereals by grinding stones is,
however, a further possible explanation for the decreased
lateralization in Neolithic populations. In this activity,
the right and left sides are involved in the process, and
the forces operating on the left side are equivalent to
those of the right. The processing of food (and hence the
operation of grinding cereals in primitive cultures) is
American Journal of Physical Anthropology—DOI 10.1002/ajpa
452
D. MARCHI ET AL.
usually performed by females (Murdock and Provost,
1973). Thus, the decline of humeral lateralization of
NEOL females, rather than that observed in males, is
more likely affected by this activity.
Considering the different functional requirements of
the hypothesized male and female activities, this division
of labor might well explain the high sexual dimorphism in
relative humeral strength exhibited by the NEOL sample.
Femur
Measures of femoral cross-sectional areas, second moments of area proportional to the bending rigidity in the
M-L planes (Iy and Imin), and average bending torsional
rigidity (J) for the NEOL sample decrease relative to
those of the LUP and Mesolithic samples. This result is
in agreement with analyses of external measurements
taken on LUP and NEOL male Ligurian samples (Formicola, 1986).
A different picture arises from consideration of second
moments of area proportional to bending rigidity in the
A-P planes (Ix and Imax) and diaphyseal shape ratios
(Imax/Imin and Ix/Iy). The indicators of diaphyseal shape
increase significantly in the male NEOL sample compared to the Mesolithic sample, and slightly exceed the
values for the LUP sample. These results are surprising,
given the patterns observed in skeletal populations and
theoretical expectations for decreased mobility with the
advent of food-producing economies (Larsen, 1997).
Comparisons among the European EUP, LUP, and
Mesolithic (Holt et al., 2000; Holt, 2003) pointed out a
significant increase in femoral circularity over time, possibly indicating reduced mobility among hunter-gatherer
populations following the Last Glacial Maximum. Therefore, the high antero-posterior ellipticity of midshaft femoral cross sections in NEOL males might reflect a high
level of mobility.
From the above discussion, it appears that the NEOL
population (males) was more mobile than the Mesolithic
population and, similar to the LUP population, committed to a hunting-gathering economy. However, an important role in the robusticity of the femur may be played
by the mountainous territory (Ruff, 1999). The Mesolithic comparative sample is composed of specimens from
around Europe, mainly from nonmountainous regions
(Holt, 1999). In order to assess the role of terrain in
modeling femoral robusticity in Ligurian Neolithics, we
compared a male LUP subsample from Liguria (Arene
Candide 4, 10, and 12; see Appendix B) and the rest of
the European LUP male sample. Unfortunately, no LUP
Ligurian females are available for comparisons with
their European contemporaries. Given the small sample
size of the Ligurian LUP male subsample, none of the
changes are statistically significant. However, some preliminary trends may be noted. Figure 1a shows that for
the three indicators of robusticity analyzed (Ix, Iy, and
J), the Ligurian LUP subsample exhibits greater values
than the rest of the LUP, while Figure 1b emphasizes
the more antero-posteriorly oriented shape of Ligurian
femora (both in LUP and NEOL). Given the hilly nature
of the Ligurian terrain and archaeological evidence of a
pastoral-based mobile subsistence, it is reasonable to
hypothesize that the antero-posteriorly buttressed shape
of the NEOL femora results from the combined effect of
lifestyle and terrain.
In an extensive analysis of sexual division of labor in
185 societies, Murdock and Provost (1973) found that
Fig. 1. Comparisons between Ligurian Late Upper Paleolithic (LUP Lig), non-Ligurian Late Upper Paleolithic (LUP
non-Lig), and Ligurian Neolithic (NEOL) males. a: Midshaft femur polar moment of inertia (J) and second moments of area
about medio-lateral (Ix) and antero-posterior axis (Iy). b: Midshaft femur shape ratios (Ix/Iy and Imax/Imin). Ix, Iy, and J are divided by product of body mass and square of bone length.
activities which are exclusively or usually carried out
by males involve more mobility than those carried out
by females. Accordingly, NEOL females exhibit significantly less elliptical midshaft femora than males, with
differences in most cases exceeding those found in European hunter-gatherers. On the whole, the results of
the CSG analysis of the femur provide evidence of high
sexual dimorphism in the Neolithic Ligurian population, and point to a marked sexual division of labor,
with females probably involved in more sedentary agricultural tasks.
CONCLUSIONS
Understanding the relationships between limb bone
robusticity and Neolithic subsistence in western Liguria
was the main goal of this paper. Although the results
were predominantly nonsignificant, comparisons with
LUP hunters-gatherers are suggestive of increased humeral robusticity in NEOL males, with a significant reduction of bilateral asymmetry. Upper limb asymmetry also
declines in NEOL females, despite a general reduction in
robusticity. The different trends affecting male and
female humeral robusticity result in significantly increased sexual dimorphism.
American Journal of Physical Anthropology—DOI 10.1002/ajpa
453
NEOLITHIC LIGURIAN ACTIVITY PATTERNS
These results suggest that the activities performed by
Ligurian Neolithics involved a marked sexual division of
labor and more symmetrical use of the upper limb by
both males and females than is evident from the pan-European LUP comparative samples. When articulated
with archaeological, faunal, paleopathological, and ethnographic information, these findings support the hypothesis that males engaged in the repetitive, bimanual
use of axes tied to pastoral activities such as procurement of fodder, while females attended to agricultural
tasks.
The hypothesis of such a division of labor between
genders is emphasized by femoral CSG data. While all
female indicators of bending strength decrease steadily
through time, NEOL male diaphyseal shape ratios approach those of the LUP and even show an increase relative to the Mesolithic group. This indicates a level of mechanical stress for NEOL males that is surprising, given
theoretical expectations for decreased mobility with the
advent of food-producing economies and results from
previous studies of skeletal robusticity correlated with
these changes. Interestingly, however, expectations are
fulfilled at a regional level (Fig. 1a), i.e., when comparing NEOL males and LUP hunter-gatherers from the
same area. Liguria is a mountainous region, and this
finding provides further evidence for the role of the terrain for lower limb remodeling. Comparative pan-European samples living in a similar environment are needed
in order to better understand the connection of subsistence economy and nature of the terrain to cross-sectional
geometric properties of long bones.
ACKNOWLEDGMENTS
We express our gratitude to the Soprintendenza Archeologica della Liguria for permission to examine the
material. Thanks are also due to G. Rossi and P. Garibaldi of the Museo di Archeologia Ligure in Genova
Pegli, to G. Vicino of the Museo Civico di Archeologia in
Finale Ligure for active collaboration and assistance
when working in the museums, and to S.E. Churchill for
comments and suggestions during preparation of the
manuscript.
APPENDIX A. Ligurian Neolithic sample composition
1
2
Specimen
Sex
Arene Candide 7
Arene Candide 8
Arene Candide EIV
Arene Candide EVI
Arene Candide IX1
Arene Candide XII
Arma dell’Aquila I
Arma dell’Aquila V
Pollera 1 (Tiné)
Pollera 10
Pollera 12
Pollera 13
Pollera 14
Pollera 302
Pollera 33
Pollera 6246
Male
Male
Female
Male
Male
Female
Female
Female
Female
Male
Female
Male
Female
Male
Female
Male
Only left humerus present.
Only right humerus present.
APPENDIX B. Late Upper Paleolithic comparative
sample composition
Specimen
Sex
Reference
Arene Candide 4
Arene Candide 10
Arene Candide 12
Bruniquel 24
Cap Blanc 1
Chancelade 1
Grotte des Enfants 3
Laugerie-Basse 4
Neuessing 2
Oberkassel 1
Oberkassel 2
Riparo Continenza
Riparo Tagliente
Rochereil 1
Romanelli 1
Romito 3
Romito 4
San Teodoro 4
St. Germain-la-Rivière 4
Veyrier 1
Male
Male
Male
Female
Female
Male
Female
Male
Male
Male
Female
Male
Male
Male
Male
Male
Female
Female
Female
Male
Holt, 1999
Holt, 1999
Churchill, 1994;
Churchill, 1994;
Churchill, 1994;
Churchill, 1994;
Holt, 1999
Churchill, 1994
Churchill, 1994;
Churchill, 1994;
Churchill, 1994;
Holt, 1999
Holt, 1999
Holt, 1999
Churchill, 1994;
Churchill, 1994;
Churchill, 1994;
Holt, 1999
Churchill, 1994;
Churchill, 1994;
Holt,
Holt,
Holt,
Holt,
1999
1999
1999
1999
Holt, 1999
Holt, 1999
Holt, 1999
Holt, 1999
Holt, 1999
Holt, 1999
Holt, 1999
Holt, 1999
APPENDIX C. Mesolithic comparative sample composition
Specimen
Birsmatten
Bottendorf
Culoz 1
Culoz 2
Dragsholm A
Dragsholm B
Gramat 1
Hoedic 1
Hoedic 2
Hoedic 4
Hoedic 5
Hoedic 6
Hoedic 8
Hoedic 10
Kolbjerg
Korsor Glasvaerek
Le Rastel
Loschbour
Molara 2
Mondeval
Moita de Sebastiao
Moita de Sebastiao
Moita de Sebastiao
Moita de Sebastiao
Moita de Sebastiao
Moita de Sebastiao
Muge Arruda 3
Sejrø
Taviec 3
Taviec 4
Taviec 7
Taviec 8
Taviec 9
Taviec 11
Taviec 16
Unseburg
Uzzo 2
Uzzo 5
Uzzo 7
Vaegensø
Vatte di Zambana
Sex
1
2
7
9
18
31
Female
Female
Male
Male
Female
Female
Male
Female
Male
Male
Male
Male
Female
Female
Female
Male
Male
Male
Male
Male
Female
Female
Male
Male
Male
Male
Male
Male
Female
Male
Male
Male
Female
Male
Male
Female
Male
Male
Male
Male
Female
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Holt,
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