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Contents lists available at ScienceDirect
HOMO - Journal of Comparative
Human Biology
journal homepage: www.elsevier.com/locate/jchb
Assessing change in diet and biological affinity
between the 4th and 3rd millennia cal BCE in
the Portuguese Estremadura: A preliminary
dental comparison of Feteira II and Bolores
B.C. Horwath a,b,c, A.J. Waterman b,c, K.T. Lillios b,∗, J.D. Irish d
a
Department of Anthropology, University of Alaska, Fairbanks, Fairbanks, AK, USA
Department of Anthropology, University of Iowa, Iowa City, IA 52242, USA
Department of Natural and Applied Sciences, Mount Mercy University, Cedar Rapids, IA, USA
d
School of Natural Sciences and Psychology, Liverpool John Moores University, Liverpool L3 3AF,
United Kingdom
b
c
a r t i c l e
i n f o
Article history:
Received 29 June 2013
Accepted 17 November 2013
Available online xxx
a b s t r a c t
Although the social and political changes accompanying the transition from the Neolithic through Copper Age, between the 4th
and 3rd millennia cal BCE, in southwestern Iberia are reasonably
well understood, much less is known about whether population
movements and dietary changes accompanied these transformations. To address this question, human dental remains from the
Middle through Late Neolithic site of Feteira II (3600–2900 cal BCE)
and the Late Neolithic site of Bolores (2800–2600 cal BCE) in the
Portuguese Estremadura were used to examine diet (microwear)
and affinity (dental non-metrics). Microwear features were not
found to be significantly different between Feteira II and Bolores,
suggesting that the emergence of social complexity during this
period did not result in large-scale changes in subsistence practices during the period of use at these sites. Using the Arizona State
University Dental Anthropology System and supporting statistics,
no significant difference between the samples from Feteira II and
Bolores was observed, suggesting that no population replacement
occurred between the Middle Neolithic and Late Neolithic/Copper
Age. However, at Bolores there is some indication that there may
have been demographic exchanges between southern Iberian and
North African populations during the Late Neolithic/Copper Age.
© 2013 Elsevier GmbH. All rights reserved.
∗ Corresponding author. Tel.: +1 319 335 3023; fax: +1 319 335 0653.
E-mail address: [email protected] (K.T. Lillios).
0018-442X/$ – see front matter © 2013 Elsevier GmbH. All rights reserved.
http://dx.doi.org/10.1016/j.jchb.2013.11.003
Please cite this article in press as: Horwath, B.C., et al., Assessing change in diet and biological
affinity between the 4th and 3rd millennia cal BCE in the Portuguese Estremadura. . . . HOMO - J.
Comp. Hum. Biol. (2014), http://dx.doi.org/10.1016/j.jchb.2013.11.003
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Introduction
During the 4th and 3rd millennia cal BCE in the Iberian Peninsula (the Late Neolithic to Copper
Age), as throughout much of Europe, profound social and economic changes occurred. These include
the development of hierarchical societies that supported craft specialization and engaged in intensive
agricultural production (Berrocal et al., 2013; Chapman, 1990, 2003; Gilman, 1987). Since the late
1800s, archeologists have documented this cultural transformation in Portugal and Spain through the
study of hundreds of settlements and thousands of burials (in caves, rockshelters, and megaliths).
Many of the settlements established during this period, such as Zambujal and Leceia in Portugal
(Cardoso, 2000; Sangmeister and Schubart, 1981) and Los Millares in Spain (Molina González and
Cámara Serrano, 2005), were fortified and situated strategically on hilltops, suggestive of a new social
environment with heightened competition and militarism. Long distance trade also expanded. Objects
made from ostrich eggshells and elephant ivory from North Africa were acquired by populations in
southern Iberia (García Sanjuán et al., 2013; Harrison and Gilman, 1977; Schuhmacher et al., 2009), and
valued stones, including amphibolite and variscite, were trafficked throughout the Peninsula (Lillios,
1997; Odriozola et al., 2010).
Despite these social and economic changes in the Iberian Peninsula during the Late
Neolithic/Copper Age, their relationship to human biological dynamics is poorly understood. Bioanthropological studies of prehistoric diet and biological affinity have primarily addressed the Mesolithic
to Neolithic transition (Cunha and Cardoso, 2001; Fox and Martin, 1999; Jackes, 2009; Jackes et al.,
2001; Lubell and Jackes, 1994). While such studies are critical to understanding the biological and
cultural impacts of the Neolithic Revolution, they do not address the biological and cultural ramifications of post-Neolithic economic and social developments in the region. Thus, it is still unclear how
these changes impacted the bodies of people living at the time. Specifically, it is unknown whether
demographic or dietary shifts accompanied the emergence of complex societies in the Iberian Peninsula (but see preliminary research by Silva, 2003; Waterman, 2006, 2012). In order to contribute to
a more nuanced understanding of the dynamics of the post-Neolithic social landscape, this study
seeks to assess the biological nature of social changes in the Neolithic and Copper Age of the Iberian
Peninsula by investigating diet and demographics in human dental remains from two burial sites in
the Estremadura region of Portugal: the Middle to Late Neolithic site of Feteira II (Waterman, 2006;
Zilhão, 1984) and the Late Neolithic site of Bolores (Lillios et al., 2010). Dietary changes were evaluated
through microwear analysis, and demographic shifts were evaluated through dental morphological
traits.
Materials
In this study, dental morphology and dental microwear patterns were compared from selected
individuals from Feteira II and Bolores. These sites were chosen because they represent burial samples
dated to different time spans but located in comparable geographic landscapes only about 20 km apart
(Figs. 1 and 2).
Feteira is a burial cave located near Lourinhã. The site was discovered in 1981 when a cave gallery
(Feteira I) was exposed during a construction project (Zilhão, 1984). Years later, another portion of the
cave along with a large deposit of commingled human remains was discovered (Feteira II). Feteira II
was excavated between 1995 and 1997. AMS radiocarbon dates on two human bones returned dates
between 3600 and 2900 cal BCE, corresponding to the Middle to Late Neolithic (Waterman, 2006).
Based on dental analyses, Feteira II housed a minimum of 68 individuals, 26 of which are subadults. A
total of approximately 1500 deciduous and permanent teeth and tooth fragments were recovered.
Bolores is a rock-cut tomb carved out of Jurassic sandstone and shale located 16 km southeast of
Feteira II near Torres Vedras. The site was discovered in 1986, and test excavations recovered human
remains and Late Neolithic material culture (Kunst and Trindade, 1990). In 2007, 2008, 2010 and 2012,
intensive excavations were undertaken (Lillios et al., 2010). Ten AMS dates taken from human bones
found throughout the mortuary deposit, including the basal level, cluster between 2800 and 2600 cal
BCE, placing the site in the Late Neolithic and at a slightly later date than Feteira II. In this study, the
remains from the 1986, 2007, 2008, and 2010 seasons were analyzed (analyses of the 2012 remains
Please cite this article in press as: Horwath, B.C., et al., Assessing change in diet and biological
affinity between the 4th and 3rd millennia cal BCE in the Portuguese Estremadura. . . . HOMO - J.
Comp. Hum. Biol. (2014), http://dx.doi.org/10.1016/j.jchb.2013.11.003
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3
Fig. 1. Location of Feteira II and Bolores.
are in progress). Based on these dental remains, the minimal number of individuals is 22, representing
11 adults and 11 subadults. A total of approximately 345 deciduous and permanent teeth and tooth
fragments were recovered.
Methods
For adult teeth, all second left mandibular molars without post-mortem damage or casting defects
on wear facet x or 10n (phase II wear facets located on the protoconid halfway between the central groove and the cusp tip) were observed for microwear. Deciduous second molars were used in
subadults. Molds were taken using Coltène® President Jet Plus Light Body, a polyvinylsiloxane impression material (Benyon, 1987; Fiorenza et al., 2009; Galbany et al., 2004; Schmidt, 1999). The teeth were
not treated with any preservative and were cleaned with only dry techniques. Casts were made using
Specifix-20, a cold cure Epoxy Resin produced by Streurs. Once the mold was dried, it was mounted
to an aluminum stub and placed inside the chamber on the Ladd sputter coater.
Scanning electron microscopy (SEM) micrographs were taken of the phase II wear facets on the
protoconid (Gordon, 1984; Kay, 1977; Kay and Hiiemae, 1974; Ungar, 1996). The SEM was set to
secondary emission mode with a magnification of 500× approximately 0.02 m2 , a working distance
of 25, and KV of 25 (Greene, 2006).
Images of the phase II wear facets were analyzed by Microwear 4.02 (Ungar, 2002). Summary statistics generated by Microwear 4.02 were analyzed in PASW® 18.0 using an independent samples t-test.
In this case, multiple t-test statistics were computed to compare the following variables: percentage of
pits, scratch width in microns, scratch breadth in microns, pit width in microns, percentage of scratch
occurrence, percentage of pit occurrence, scratch tally and pit tally (Ungar, 1995; Ungar and Teaford,
Please cite this article in press as: Horwath, B.C., et al., Assessing change in diet and biological
affinity between the 4th and 3rd millennia cal BCE in the Portuguese Estremadura. . . . HOMO - J.
Comp. Hum. Biol. (2014), http://dx.doi.org/10.1016/j.jchb.2013.11.003
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Fig. 2. Calibrated dates for Feteira II and Bolores (calibrated using OxCal v4.2.2 Bronk Ramsey, 2013)
2002). Adult and subadult dentitions were pooled because no significant differences were revealed
between microwear formation (also see Gamza, 2010).
Fifteen comparative samples were used to characterize the morphology of the Feteira II and Bolores
dentitions (Table 1 and Fig. 3). They derive from sites in Ancient Palestine, Italy, Greece, Turkey, and
North Africa. The latter are included, in part, to assess whether North African cultural items found
in Portugal during the Late Neolithic and Copper Age were accompanied by any gene flow. Origin
locations include Tunisia, Algeria, Morocco, Libya, Egypt (Abydos, Kharga, Tarkan, Hierakonpolis, and
Saqqara) and Nubia (Kerma). The frequency of dental traits can distinguish groups from one another,
for example Sub-Saharan African samples are characterized by low frequency of double shoveling in
UI1 and high frequency of traits such as the Bushman canine, two rooted UP1, LM2 Y-groove pattern,
and LM1 cusp seven, among other traits (Irish, 1997).
Thirty-six non-metric dental and osseous traits were recorded in the permanent dentition using 24
reference plaques from the Arizona State University Dental Anthropology System (ASUDAS). Because
of the fragmentary nature of the Bolores and Feteira II samples, composite individuals were created
to limit the probability of counting an individual multiple times for a given trait. In cases where
both antimeres are present, only the highest expression was used for statistical analysis; if only one
tooth was present, it was automatically included (Turner et al., 1991). Males and females were pooled
following standard ASUDAS protocol (Irish, 1997).
All data were entered into a PASW 18.0 database and the Mean Measure of Divergence distance
statistic (MMD) was used to determine similarity among all samples. The Freeman and Tukey (1950)
transformation was used in conjunction with the MMD statistic to adjust for variance in small samples
(Green and Suchey, 1976; Irish, 2010).
Please cite this article in press as: Horwath, B.C., et al., Assessing change in diet and biological
affinity between the 4th and 3rd millennia cal BCE in the Portuguese Estremadura. . . . HOMO - J.
Comp. Hum. Biol. (2014), http://dx.doi.org/10.1016/j.jchb.2013.11.003
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5
Table 1
Comparative dental morphology samples used in study.
Samples
Dates
Abydos (ABY)
Bedouin (BED)
3000–2686 BCE
1800–1900 CE
Bolores (BOL)
Capsian (CAP)
2800–2600 BCE
6500–3000 BCE
Feteira II (FET)
Greek (GRK)
3600–2900 BCE
475–300 BCE
historic period
3500–3200 BCE
30 BCE–CE
395/modern
period
1800–1900 CE
1750–1500 BCE
3300–1098 BCE
2613–2494 BCE
1200–520 BCE
1800s CE
3000–2890 BCE
2055–1773 BCE
Classic period
∼300 BCE
Hierakonpolis (HRK)
Italy (ITY)
Kabyle/Berber (KAB)
Kerma (KER)
Lachish- Bronze (LCB)
Saqqara (SAQ)
Lachish- Iron (LCI)
Shawia/Berber (SHA)
Tarkhan (TAR)
Thebes (THE)
Turkey (TRK)
Geographical
location
Egypt
Morocco,
Tunisia, Libya
Portugal
Tunisia,
Algeria
Portugal
Greece
MNI
Source
Egypt
Italy
247
90
Irish (2006) and Irish and Friedman (2010)
Irish (2012, personal communication)
Algeria
Nubia
Palestine
Egypt
Palestine
Algeria
Egypt
Egypt
Turkey
32
63
34
41
365
26
51
54
40
Guatelli-Steinberg et al. (2001) and Irish (2000)
Irish (2005) and Irish and Friedman (2010)
Dicke-Toupin (2012)
Irish (2006) and Irish and Friedman (2010)
Dicke-Toupin (2012)
Guatelli-Steinberg et al. (2001) and Irish (2000)
Irish (2006) and Irish and Friedman (2010)
Irish (2006) and Irish and Friedman (2010)
Irish (2012, personal communication)
54
49
Irish (2006) and Irish and Friedman (2010)
Guatelli-Steinberg et al. (2001) and Irish (2000)
22
22
Current study
Irish (2000)
68
77
Current study
Irish (2012, personal communication)
MNI – minimum number of individuals.
The MMD and geographic distances (km) were compared using the Pearson’s R correlation coefficient. The geographic location of each sample was plotted in decimal degrees on a Robinson Projection
world map. Straight-line distances were calculated between samples using the measure tool in
ArcGIS® 9.3.1 (ESRI, 2011). Correlations were then examined using Wright’s (1938, 1940, 1943)
Fig. 3. Location of comparative dental morphology samples.
Please cite this article in press as: Horwath, B.C., et al., Assessing change in diet and biological
affinity between the 4th and 3rd millennia cal BCE in the Portuguese Estremadura. . . . HOMO - J.
Comp. Hum. Biol. (2014), http://dx.doi.org/10.1016/j.jchb.2013.11.003
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Table 2
Comparison of summary statistics for microwear features.
Features
Sites
n
Mean
SD
Feature tally
Feteira II
Bolores
Feteira II
Bolores
Feteira II
Bolores
Feteira II
Bolores
Feteira II
Bolores
Feteira II
Bolores
17
5
17
5
17
5
17
5
17
5
17
5
74.71
81.60
34.64
33.10
2.49
2.75
66.94%
63.00%
4.85
4.84
33.06%
37.00%
29.52
30.05
4.92
7.61
0.63
0.98
11.99%
13.75%
0.95
1.07
11.99%
13.75%
Scratch length
Scratch breadth
Percentage of scratches
Pit width
Percentage of pits
p-Value
0.666
0.593
0.597
0.538
0.982
0.538
n – sample size.
SD – standard deviation.
isolation-by-distance stepping stone model to determine if a correlation exists (Irish, 2010; Kimura
and Weiss, 1964; Konigsberg, 1990).
Results
Results from the independent samples t-test between Feteira II and Bolores reveal no significant
differences in microwear (Table 2). For this comparison, adults and subadults were pooled (see Gamza,
2010). Sample size was significantly reduced because of post mortem wear or other damage.
Table 3 shows the percentage of teeth that present a trait and the total number of teeth scored for
biological affinity. Results from Bolores, Lachish Bronze Age, and Capsian should be interpreted with
caution as they may not be representative of the populations from which they derive (Irish, 2005,
2006). Traits were compared pairwise using Kendall’s tau b to determine if any correlations existed.
High correlations (e.g., 0.6 and higher) can skew the results of the MMD statistic (Harris and Sjøvold,
2004; Irish, 1998a; Sjøvold, 1977). Four traits were found to be highly correlated at 0.6 or greater:
labial curvature, deflecting wrinkle, C1–C2 crest, and protostylid. As such, only MMD comparisons
with the four traits removed will be discussed further.
The MMD is a measure of dissimilarity; therefore, if a sample pair exhibits a high value, then greater
phenetic dissimilarity is identified, and vice versa (Irish, 2005, 2010). To determine if the MMD value
represents a significant difference it must be greater than two times the standard deviation; in such a
case the null hypothesis is rejected at the 0.025 level (Irish, 2005, 2010; Sjøvold, 1977).
Feteira II (0.037) and Bolores (0.037) are not significantly different. Bolores is also similar to the
Capsian (0.06) and Lachish Bronze Age (0.051) samples; Feteira II is similar to the Lachish (0.141)
sample. The remainder of the comparisons, as seen in Table 4, yielded significant differences.
A significant positive correlation was found between phenetic and geographic distance for Bolores
(0.628) (p = 0.007) and Feteira II (0.514) (p = 0.035). This conclusion fits the stepping stone isolationby-distance model. Positive relationships are represented in both scatter plots (Figs. 4 and 5). When
contrasting Bolores with the Capsian, Greek, Thebes, Saqqara, Kerma, and Tarkhan samples, all fall
with the 95% confidence interval. Italy, Turkey, and Feteira II are close to the 95% confidence interval.
These results fit the isolation-by-distance stepping stone model, which states that as populations
increase in geographic distance from each other, the more different they should be phenetically. In
this model, limited gene flow occurs along a continuum with neighboring groups (Kimura and Weiss,
1964; Konigsberg, 1990; Wright, 1938, 1940, 1943).
A number of groups do not fit the stepping stone model relative to the two Portuguese samples:
Bedouin, Kabyle, Shawia, Lachish Iron Age, Hierakonpolis, and Abydos, and Lachish Bronze Age. The
Lachish (n = 34) sample is small, with many missing data; thus, it is more likely to look like other samples. The Northwest African samples – Capsian, Bedouin, Kabyle, and Shawia – appear closer to Bolores
than the Egyptian samples in phenetic and geographic distances. This relationship could provide
Please cite this article in press as: Horwath, B.C., et al., Assessing change in diet and biological
affinity between the 4th and 3rd millennia cal BCE in the Portuguese Estremadura. . . . HOMO - J.
Comp. Hum. Biol. (2014), http://dx.doi.org/10.1016/j.jchb.2013.11.003
ABP
BED
CAP
%
n
%
n
%
n
%
n
%
n
%
0
0
0
11
0
0
0
10
0
11
61.5
0
0
0
31
0
0
5.1
39
0
43
16.7
2.5
40
0
21
0
39
5.9
17
0
17
27.8
5.4
37
8.3
24
2.4
41
8
25
12.5
24
37.5
0
5
0
4
0
10
0
5
0
5
60
1.5
68
0
5
4.3
70
0
5
0
5
35
5.4
167
8.3
109
0
125
17.6
102
4.5
110
25.2
3.9
76
0
29
10.4
77
26.9
26
0
27
13.3
n
%
13
46.2
42
22
18
25
24
43.5
5
60
20
5.3
115
36.7
n
%
n
%
13
0
16
54.5
41
0
41
84.6
16
0
17
20
23
0
29
12
5
22.2
9
42.7
19
8.7
23
8.3
n
%
n
%
n
%
n
%
n
%
11
77.8
9
44.4
9
45.5
11
0
9
10
39
73.7
38
31.4
35
59.5
37
0
36
8.6
15
66.7
33
10
20
90.5
21
0
28
11.8
25
58.8
34
8.8
34
54.5
33
0
20
5.6
n
%
n
%
n
%
10
12.5
8
75
4
0
35
33.3
30
84.4
32
0
34
69.4
49
92.9
28
1.9
36
50
32
69
29
0
7
100
10
30
10
100
6
0
9
0
13
33.3
12
85.7
7
0
GRK
HRK
ITY
KAB
KER
LCI
LCB
SAQ
0
29
12.5
8
3.4
29
14.3
7
12.5
8
21.4
5.4
56
7.7
13
1.8
55
22.2
9
0
7
9.1
0
21
0
15
0
355
6.3
16
0
16
3.1
0
7
14.3
7
0
32
0
7
0
7
0
2.8
36
0
11
0
39
0
7
0
8
33.3
30
36.7
14
50
11
8.3
32
29.4
7
30
109
5
120
12.6
30
2.6
39
19.2
12
0
16
27.3
12
16.7
18
18.2
34
0
77
8.3
12
50
54
5.7
53
8.5
48
0
33
16.7
103
86.6
157
15.5
97
80.8
104
0
142
19.5
26
59.7
72
17.5
63
61.3
62
0
41
5.8
11
63.6
22
11.8
17
57.9
19
0
22
0
11
91.7
48
24.1
29
51.6
31
5.4
37
4
54
61.9
63
58.3
36
0
164
59.8
164
75.6
119
3.6
69
5.9
59
76.9
39
9.6
23
52.2
23
68.4
19
6.2
50
80.4
51
90.2
41
1.6
SHA
TAR
THE
TRK
0
26
14.3
7
0
25
0
7
25
8
46.2
6.8
44
6.7
30
0
44
7.1
28
0
26
8.8
5.6
54
4.8
21
0
51
15.8
19
0
18
20.8
0
36
0
10
0
35
0
10
0
10
15.4
9
66.7
13
25
34
28.1
24
30
13
15.4
10
0
14
0
6
0
10
0
12
0
4
22.2
32
5.4
37
3.8
20
3
33
10.5
13
0
19
6.3
36
77
217
29.5
251
71.3
164
1.6
128
5.5
4
85.7
21
24
25
89.5
19
0
9
0
6
95.7
23
0
9
1
16
0
15
0
9
68.4
19
10
20
55.6
18
7.7
13
4.8
26
75
40
0
23
67.9
28
2.6
38
0
19
85.7
42
14.3
28
90.3
31
0
37
4.8
16
60
25
4.5
22
85.7
21
0
13
25
255
56.8
199
85.9
64
4.9
8
54.5
11
100
5
0
18
89.7
29
82.6
23
6.1
21
52.2
23
72.2
18
0
45
75
32
72.2
18
4
42
85.3
34
81.3
32
0
24
69
29
62.1
29
5.7
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Traits
Winging UI 1
(+ = ASU 1)
Labial Curvature UI1
(+ = ASU 2–4)
Palatine Torus
(+ = ASU 2–3)
Shoveling UI 1
(+ = ASU 2–6)
Double Shoveling UI 1
(+ = ASU 2–6)
Interruption Groove
UI2
(+ = ASU +)
Tuburculum Dentale
UI2
(+ = ASU 2–6)
Bushman Canine UC
(+ = ASU 1–3)
Distal Accessory Ridge
UC
(+ = ASU 2–5)
Hypocone UM2
(+ = ASU 3–5)
Cusp 5 UM1
(+ = ASU 3–5)
Carabelli’s Trait UM1
(+ = ASU 1–5)
Parastyle UM 3
(+ = ASU 1–5)
Enamel Extension UM
1
(+ = ASU 1–3)
Root Number UP 1
(+ = ASU +)
Root Number UM 2
(+ = ASU 3 + )
Peg/Reduced Incisor UI
2
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Please cite this article in press as: Horwath, B.C., et al., Assessing change in diet and biological
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Table 3
Dental trait frequencies and percentages for dental morphology samples.
7
44
10
40
0
52
2.5
40
6.7
27
0
40
21.1
10
0
12
16.7
73
0
44
17.6
197
1.3
156
8.1
83
2.7
74
23.5
16
0
22
3.4
63
0
41
16.7
61
0.5
218
16.1
10
0
21
15
n
%
n
%
n
%
n
%
n
%
n
%
n
%
n
%
n
%
8
0
0
66.7
3
37.5
8
0
0
9.1
11
0
0
8.3
12
18.2
11
12.5
42
0
0
69.2
26
33.3
24
0
0
0
39
0
0
14.3
35
48.6
35
4.6
45
5.4
37
65.4
26
22.2
18
0
53
15.2
46
17
47
3.5
29
37.8
37
21.7
38
8.8
34
64.3
28
37.5
24
2.9
35
46.9
32
9.4
32
12.5
32
42.9
28
15.6
12
0
5
84.6
13
45.5
11
0
19
41.2
17
17.7
17
17.7
17
38.9
18
20
68
3
66
60
10
36.4
11
0
34
43.5
23
30.3
33
0
19
47.6
21
17.6
184
2.8
168
81.4
129
21.6
74
0.5
178
37.1
175
22.2
144
9.5
137
36.5
137
20.5
81
4.9
82
34.9
43
51.4
35
0
73
26.2
61
12.5
72
2
51
35.6
45
12.5
29
12
25
69.2
13
60
10
0
19
27.8
18
10.5
19
31.3
16
33.3
18
6.7
60
3.3
60
86.4
22
43.8
16
0
60
41.3
46
5.3
57
0
28
41.2
34
11.1
310
4.8
21
71.4
28
20.8
24
3.8
78
34
50
0
81
3.9
51
37.7
69
0
22
0
21
0
27
0
33
3.6
28
0
32
3.3
30
93.1
29
0
0
23
0
28
0
30
5.1
39
17.4
46
2
49
5.6
36
84.4
32
13.3
45
32
3
33
0
33
5.9
34
6.3
32
0
26
0
33
88.9
27
20
25
10
0
9
0
15
16.7
18
0
15
0
12
5.9
17
85.7
14
23.1
13
17
5.9
17
0
19
5.6
18
7.1
28
3.4
29
0
22
91.3
23
13
23
112
48
3.9
6.4
102
47
1.4
0
139
51
6.2
5.4
177
56
14.3 10.5
175
57
6.1
3.3
179
60
2.2
0
136
43
89.1 100
128
57
7.4 16.3
148
43
15
0
14
0
16
5.9
17
5.3
19
20
10
0
17
88.9
18
21.4
14
27
0
27
0
26
17.1
35
25
52
1.9
52
2
49
94
50
15.7
51
14
0
18
1.5
67
6.1
33
3
33
0
69
0
6
100
5
14.3
49
n
%
n
%
n
%
n
%
n
%
n
%
n
%
n
%
n
8
0
8
0
10
10
10
0
9
0
8
0
7
100
9
0
0
33
0
12
20
13
0
23
23.1
50
0
43
4.1
54
5.1
39
19.6
35
0
30
21.9
20
0
2
100
3
100
1
0
8
40
5
0
7
0
2
37.5
8
0
35
0
33
66.7
12
14.3
7
0
37
22.7
22
24.3
37
0
10
25
12
0
26
0
23
92.3
13
29.4
17
4.2
24
36.8
19
8.3
24
9.5
21
31.6
19
5
49
4.2
48
77.8
18
0
2
0
49
30.6
36
16.3
43
5
20
50
28
12.5
51
1.9
53
70.3
37
42.9
14
0
52
25
48
22.6
53
2.8
36
26.3
38
13.3
32
0
37
82.4
17
40
10
0
30
5.9
17
13.8
29
0
19
41.2
17
6.7
2
0
2
0
5
0
2
0
2
0
9
0
2
100
1
0
6
8
0
5
0
14
0
20
6.7
30
6.1
33
0
26
86.7
30
0
23
20
0
20
0
21
4.8
21
10.5
19
0
16
0
22
95.5
22
23.5
17
16
0
16
0
20
3.7
27
13.6
44
4.4
45
0
33
85
40
5.6
36
30
0
26
0
41
6.8
44
11.1
36
0
35
0
39
91.7
36
22.5
40
15
0
13
0
17
0
19
0
25
0
22
5.3
19
89.5
19
31.3
16
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18
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(+ = ASU P or R)
Odontome P1-P2
(+ = ASU +)
Congenital Absence
UM 3
(+ = ASU −)
Midline Diastema UI 1
(+ = 0.05 mm)
Lingual Cusp LP 2
(+ = ASU 2–3)
Anterior Fovea LM 1
(+ = ASU 2–4)
Mandibular Torus
(+ = ASU 2–3)
Grove Pattern LM 2
(+ = ASU Y)
Rocker Jaw
(+ = ASU 6+)
Cusp Number LM 1
(+ = ASU 6+)
Cusp Number LM 2
(+ = ASU 5+)
Deflecting Wrinkle LM
1
(+ = ASU 2–3)
C1-C2 Crest LM 1
(+ = ASU +)
Protostylid LM1
(+ = AUS 1–6)
Cusp 7 LM 1
(+ = ASU 2–4)
Tomes Root LC
(+ = ASU 2+)
Root Number LC
(+ = ASU 2+)
Root Number LM 1
(+ = ASU 3+)
Root Number LM 2
(+ = ASU 2+)
Torso Angle LM 3
(+ = ASU +)
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Table 3 (Continued)
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Fig. 4. Scatterplot of Bolores distance (km) vs. mean measure of divergence values.
Fig. 5. Scatterplot of Feteira II distance (km) vs. mean measure of divergence values.
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Table 4
Mean measure of divergence values.
Site
Bolores MMD with
4 traits
Bolores MMD
without 4 traits
Feteira II MMD
with 4 traits
Feteira II MMD
without 4 traits
ABY
BED
BOL
CAP
FET
GRK
HRK
ITY
KAB
KER
LCB
LCI
SAQ
SHA
TAR
THE
TRK
0.059
0.023
0
0
0
0.084
0.069*
0.059
0.001
0.135*
0.005
0.066
0.167*
0
0.149*
0.092*
0.085
0.159*
0.110*
0
0.060
0.037
0.147*
0.149*
0.179*
0.119*
0.327*
0.051
0.107*
0.293*
0.160*
0.329*
0.242*
0.189*
0.132*
0.180*
0
0.140*
0
0.200*
0.182*
0.162*
0.116*
0.207*
0.086
0.134*
0.270*
0.147*
0.201*
0.174*
0.137*
0.189*
0.263*
0.037
0.260*
0
0.234*
0.245*
0.268*
0.189*
0.366*
0.141
0.163*
0.393*
0.284*
0.322*
0.296*
0.240*
MMD – mean measure of divergence.
*
Statistically significant difference between sites.
Table 5
Summary of dietary studies for Feteira II and Bolores.
Dietary indicator
Microwear
Macrowear
Caries
Isotope
Difference between
Feteira II and
Bolores
No significant
difference, except
between adults and
sub-adults for
scratch breadth
No significant
difference
No significant
difference
Some differences
Conclusion
Source
Soft foods and high
abrasives
Present study
Soft foods and high
abrasive
No singular reliance on
high sugar carbohydrates
Bolores-C3 plants and
terrestrial proteins
Feteira II-C3 and C4
plants and terrestrial and
marine proteins
Waterman and Horwath (2009)
Waterman (2006) and Lillios et al. (2010)
Lillios et al. (2010) and Waterman (2012)
evidence for a connection between Northwest Africa and Portugal during the Late Neolithic/Copper
Age. When correlations for Feteira II are compared, the general groupings stay the same; however,
phenetic distance increases.
Discussion
This study provides important insights into dietary patterns of the people from Feteira II and Bolores
and builds on previous studies (Table 5). As noted above, independent samples t-tests revealed no significant differences between Feteira II and Bolores in the type and occurrence of microwear features.
This result, however, must be considered in light of the small sample size. Various events influence the
formation of microwear features and these events can be used to reconstruct general dietary information. In general, pits and scratches differ between living groups based on dietary differences (El-Zaatari,
2008; Harmon and Rose, 1988; Ma and Teaford, 2010; Mahoney, 2006; Molleson et al., 1993; Organ
et al., 2005; Pastor, 1993; Scott et al., 2012; Teaford, 1991). When observing microwear features in
this study, scratches, as opposed to pits, dominate. No significant difference was determined between
Please cite this article in press as: Horwath, B.C., et al., Assessing change in diet and biological
affinity between the 4th and 3rd millennia cal BCE in the Portuguese Estremadura. . . . HOMO - J.
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11
groups for the number of scratches; however, each group had a higher percentage of scratches than
pits. Scratches are more common when the dietary texture of food is soft, as well as when sand or grit
created by stone tools are present (Grine, 1987; Schmidt, 1998, 2001, 2010; Teaford, 1986, 1988a,b;
Teaford and Lytle, 1996; Teaford and Runestad, 1992; Ungar and Spencer, 1999; Ungar and Teaford,
1996). Previous research on the aspects and degree of occlusal attrition in the samples from Feteira
II and Bolores burial sites determined that soft foods and highly abrasive dietary elements characterized the diets of these populations, with cupped wear and “reverse curve of Monson” patterns
suggesting the consumption of soft foods with a substantial amount of grit, such as stone-ground
grains (Waterman and Horwath, 2009). Additionally, lingual surface attrition of the maxillary anterior teeth (Irish and Turner, 1987, 1997; Turner and Machado, 1983) was observed on the maxillary
central incisors, and unusually angled wear is present on some mandibular molars – which suggest
paramasticatory activities (Waterman and Horwath, 2009). Because occlusal wear patterns and scores
were similar in the Feteira II and Bolores populations, it was concluded that diet was generally similar
through the Middle and Late Neolithic/Copper Age. Thus, qualitatively, the microwear data reported in
this paper are consistent with previous findings. Additionally, analyses of dental pathologies (caries,
enamel hypoplasia) for the two sites show no significant difference (Waterman, 2006) suggesting that
health status as well as diet remained consistent over the 4th and 3rd millennia cal BCE.
However, recently completed isotopic analyses on bone samples from Feteira II and Bolores suggest
that there are some slight variations between the diets of the Feteira II and Bolores populations (Lillios
et al., 2010; Waterman, 2012). In particular, nitrogen and carbon isotope analyses indicate that, while
individuals from Feteira II and Bolores ate primarily C3 plants and terrestrial proteins, some individuals
from Feteira II consumed marine proteins and C4 plants, most likely millet or seaweed (Waterman,
2012). Because some marine proteins, such as shellfish, can also contribute grit it may be that the
high percentage of scratches at Feteira II indicates sand from marine food sources, while at Bolores
the high percentage of scratches could be the result of phytolith-rich plant foods or grit from stone
processing techniques (Grine, 1987; Schmidt, 2001; Teaford, 1988a,b; Ungar and Spencer, 1999; Ungar
and Teaford, 1996; Teaford and Lytle, 1996). Thus, we need to consider the possibility of equifinality
when interpreting the microwear at the two sites.
A comparison of non-metric dental traits using the MMD indicates that individuals at Feteira II
are not significantly different from Bolores. This conclusion is in agreement with the research of
Jackes et al. (2001), which argues for demographic continuity between the Mesolithic through the
Late Neolithic/Copper Age in Portugal.
When the biological affinity of the individuals interred at Feteira II and Bolores are compared
with other Mediterranean groups, however, there are some intriguing differences. The relationship
between the Feteira II sample and the northwest African samples is different from that of Bolores and
the northwest African samples. When comparing genes (as expressed through dental morphology) and
geography between Feteira II and northwest Africa, all the northwest African samples are phenetically
more distant than they are geographically distant. The opposite is true for Bolores. That is to say, while
the input of genes from northwest Africa was not significant enough to render Bolores and Feteira II
to be biologically divergent, there does seem to be a demographic relationship between Bolores burial
sample and northwest African groups during the third millennium cal BCE. The MMD values indicate that only Capsian and Bolores are similar when comparing North African groups to Bolores and
Feteira II (Table 4). There are a number of traits that are expressed at similar frequencies between
Bolores and Capsian that are different from Feteira II: distal accessory ridge (BOL = 54.5%, CAP = 42.7%,
FET = 84.6%), interruption groove UI2 (BOL = 61.5%, CAP = 60.0%, FET = 16.7%), and deflecting wrinkle
LM1 (BOL = 12.5%, CAP = 20.0%, FET = 4.6%). North African groups in Irish (1997, 1998b) trend toward
the simplification of dental traits, which makes them somewhat similar to Europeans. North African
groups have been found to possess similar frequencies of four-cusp LM2, two-rooted lower canine,
incidence of shoveling, and UC distal accessory ridge to European groups. They differ by exhibiting
higher frequencies of two-rooted UM2, LM cusp 7, three-rooted UM2, Bushman canine, Carabelli’s
cusp, LM2 Y groove and lower frequencies of UM1 enamel extension, and absence or peg/reduced
UM3. North African groups have traits frequencies in common with Sub-Saharan groups and European
groups with potential influence from both directions (Irish, 1997, 1998b). Evidence for interactions
between North Africa and the Iberian Peninsula during this time has previously been only known
Please cite this article in press as: Horwath, B.C., et al., Assessing change in diet and biological
affinity between the 4th and 3rd millennia cal BCE in the Portuguese Estremadura. . . . HOMO - J.
Comp. Hum. Biol. (2014), http://dx.doi.org/10.1016/j.jchb.2013.11.003
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through artifactual remains. Objects made from North African elephant ivory and ostrich eggshells
have been found in Late Neolithic/Copper Age sites in southern Portugal and Spain (Harrison and
Gilman, 1977; Schuhmacher et al., 2009). Interestingly, current genomic data suggests that gene flow
from North Africa to the Iberian Peninsula was more recent (240–300 years ago; Botigué et al., 2013).
Thus, it would appear that the process of population contact between the Iberian Peninsula and northern Africa is more complex than presented by material culture or genetic evidence alone. The sample
size at Bolores is small, and work at additional burial sites will be necessary to determine the validity
of this conclusion.
Conclusions
In this study, dental microwear and non-metric dental traits were compared in individuals from
the burial sites of Feteira II (Middle to Late Neolithic, 3600–2900 cal BCE) and Bolores (Late Neolithic,
2800–2600 cal BCE) in order to better understand the dynamics of social change in the Portuguese
Estremadura. Microwear analysis suggests that no dietary change accompanied the Middle to Late
Neolithic/Copper Age transition in the Estremadura. This finding is consistent with previous studies of
macrowear (occlusal attrition) and dental pathology for the Feteira II and Bolores populations. However, isotopic evidence suggests that diet became more homogenous over time and that at Feteira II
some individuals ingested more seafood and different plant foods (Lillios et al., 2010; Waterman, 2012).
Because of this discrepancy, more research investigating microwear and equifinality in combination
with isotopic studies of diet at other related sites is warranted.
This study also investigated whether the increase in long-distance trade goods during the Late
Neolithic/Copper Age, which included objects from North Africa, was accompanied by an influx of
new genes and people. Using the Arizona State University Dental Anthropology System and supporting statistics, no significant difference between the samples from Feteira II and Bolores was
observed, suggesting that no population replacement occurred between the Middle Neolithic and
Late Neolithic/Copper Age. However, at Bolores there is some indication that there were demographic
as well as cultural exchanges between southern Iberian and North African populations during the Late
Neolithic/Copper Age. Future research to assess this hypothesis, by including other burial sites in the
Portuguese Estremadura, is being planned.
Acknowledgements
The authors would like to thank the following for their support of this research: the National Science
Foundation (#1153568), Sigma Xi, the Geist Fund (University of Alaska Fairbanks), the University of
Iowa Social Science Funding Program, and the University of Iowa Office of International Programs.
Cidália Duarte kindly permitted us to study the Feteira collection.
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