Journal of Archaeological Science 124 (2020) 105266

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Journal of Archaeological Science

journal homepage: http://www.elsevier.com/locate/jas

Factors affecting molar size in Sus scrofa

Melinda A. Zeder a, *, Ximena Lemoine b
a
Department of Anthropology, National Museum of Natural History, Smithsonian Institution, USA
b
Department of Anthropology, Washington University in St. Louis, USA

A R T I C L E I N F O A B S T R A C T

Keywords: Teeth are thought to be less affected than post-cranial elements by factors such as age, region, sex, and post-
Sus scrofa Pleistocene climate change, making changes in tooth size a better gage of domestication-induced morpholog­
Domestication ical change in Sus scrofa. Rigorous empirical evaluation of this assumption, however, has been lacking. Here we
Molars
examine the impact of multiple factors on molar size in an assemblage of modern wild boar from Southwest Asia,
Sexual dimorphism
Logarithm size index scaling
as well as a large assemblage of ancient S. scrofa from an archaeological site in southeastern Turkey dated to
11,600 cal. BP. We find that age related interstitial wear has a profound impact on molar lengths. Comparisons of
length measurements of S. scrofa molars should only be made if one strictly controls for the state of wear, as well
as the stage of crown development and eruption in third molars. Breadth measurements, on the other hand, are
little affected by progressive wear and, as a result, are preferable to length measurements in the analysis of molar
size in S. scrofa. We also evaluate the impact of logarithm size index (LSI) scaling on molar dimensions. Molar
lengths are subject to a number of factors that distort meaningful patterning rendering LSI values of length di­
mensions of little utility in the evaluation of molar size in S. scrofa. With the exception of the upper first molar,
LSI values of molar breadths appear to both capture and amplify patterns seen among unconverted measure­
ments of individual teeth. Our evaluation also confirms the assumption that the size of post-cranial elements is
more profoundly affected than molars by factors such as region and Post-Pleistocene climate change. While
sexual dimorphism also has a greater impact on the size of post-crania than on molars, the degree of sexual
dimorphism in molar breadths, as well as in M3 lengths, is still substantial and must be taken into account in any
study of S. scrofa molar size. Before interpreting downward shifts in molar size as an indicator of domestic status,
archaeozoologists need to first to consider the possible impact of differences in the proportions of males and
females in archaeological assemblages. We conclude with a number of recommendations for new protocols for
the analysis of S. scrofa dental metrics. We also offer observations about the impacts of factors like region, sex,
and change over time on the size of the molars of this species that may open up new areas of research.

1. Introduction
There is a long history of archaeozoologists using linear measure­
ments of teeth to distinguish wild from domestic forms of ancient Sus
scrofa. In the 1860s Ludwig Rütemeyer’s (1861) study of skulls and jaws
recovered from Swiss Lake Dweller sites found that ancient S. scrofa
from had demonstrably shorter snouts than modern European wild boar
and were more similar to modern domestic pigs. He interpreted these
changes in skull morphology as an artifact of domestication. The
reduction in snout length in these animals was particularly apparent in a
shortening of the diastema between the canines and the premolars, a
crowding of pre-molars resulting in an overall reduction in the length of
the cheek-tooth row, and a reduction in the size of canines and in the
length molar teeth. This pioneering work was followed by that of
Swedish zoologist Adolf Pira who monitored the progressive changes in
the snout length of S. scrofa remains from archaeological sites in Sweden
dating from the Paleolithic through the Iron Age (Pira 1909). Pira’s
research also documented a progressive shortening in both upper and
lower jaws through time that seemed to be particularly evident in a
reduction of the length of the upper third molars overtime.
Since this early work, a reduction in molar size has been widely
viewed to be an effective marker of domestication in S. scrofa across the
wide distribution of this species, from the Near East (e.g., Ervynck et al.,
2001; Flannery, 1961; 1983; Hongo and Meadow 1998; Price and
Arbuckle, 2015; Stampfli 1983), to Europe (e.g., Albarella et al., 2006a;
Boessneck et al., 1963; Bökönyi 1974; Rowley-Conwy 2003, 2011), and

* Corresponding author.
E-mail addresses: zederm@si.edu (M.A. Zeder), ximena.lemoine@wustl.edu (X. Lemoine).

https://doi.org/10.1016/j.jas.2020.105266
Received 20 August 2020; Received in revised form 21 October 2020; Accepted 21 October 2020
Available online 1 November 2020
0305-4403/© 2020 Elsevier Ltd. All rights reserved.
M.A. Zeder and X. Lemoine
Asia (e.g., Dai et al., 2019; Luo 2012; Luo and Zhang 2008; Ma 2005;
Pike-Tay et al., 2016; Wang et al., 2015; Yuan and Flad 2002). While size
reduction in post-cranial elements has also been used as a marker of
domestication in S. scrofa (e.g., Hongo and Meadow 1998), reduction in
tooth size is generally believed to be a more reliable marker in this
species principally because teeth are thought to be less affected than
post-cranial dimensions by sex-linked size differences (Albarella et al.,
2006a; Price and Arbuckle, 2015; Mayer et al., 1998). Recently,
archaeozoologists have begun to advocate for the use of breadth over
length measurements of teeth as a more effective measure of size
changes with domestication (Albarella and Payne 2005; Pike-Tay et al.,
2016; Rowley-Conwy et al., 2012). Breadth measurements are argued to
be less affected by age-related interstitial wear than length measure­
ments which are thought undergo progressive reduction over the life­
time of the animal. Increasingly, archaeozoologists are also applying the
logarithm size indexing (LSI) method (Meadow 1999) to dental metrics,
a technique that allows measurements of multiple different teeth to be
combined and compared over time and space (e.g., Albarella et al.,
2006b; Price and Arbuckle, 2015; Price and Hongo 2019).
Given the ubiquity of this domestication marker, it is surprising how
little is known about the impact of a wide range of different factors on
tooth size in S. scrofa. While the degree of interstitial wear isreasonably
thought to affect the occlusal length measurements of S. scrofa molars,
(e.g., Price and Arbuckle, 2015), we know of no actual published
empirical data that demonstrates this. Similarly, sexual dimorphism is
assumed to have only a negligible effect on molar size compared to
post-crania (Payne and Bull 1988; Price and Evin 2017) even though we
know of only one comparative study (Mayer et al., 1998) that tests this
assumption. While regional body-size variation in S. scrofa is well
documented (Groves 1981, 2008), there is little data on regional vari­
ation in tooth size or the degree to which dental size and the size of
post-crania covary by region, sex, or over time.
This article addresses these gaps in our understanding of the different
factors affecting molar size in S. scrofa. As in earlier articles dedicated to
developing new methods for demographic profiling in this species
Fig. 1. Map showing locations of modern and ancient boar.
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Journal of Archaeological Science 124 (2020) 105266
(Lemoine et al., 2014; Zeder et al., 2015; Zeder and Lemoine 2020), we
focus on a large and geographically constricted assemblage of modern
wild boar from Southwest Asia of known sex and region of origin, whose
age at death has been estimated using new methods for dental and
post-cranial age determination (Lemoine et al., 2014; Zeder et al., 2015).
Also as in other earlier studies, we include data from the large Hallan
Çemi (c. 11,600 cal BP) assemblage of S. scrofa to provide a window into
variation over time.
We address six questions: 1) What is the impact of progressive age-
related wear on length and breadth measurements of molar teeth?; 2)
What is the degree of regional variation in dental measurements?; 3)
What is the impact of sex on dental measurements?; 4) How does molar
size vary over time; 5) Are there scaling differences between different
molars and different dimensions when converted into LSI values that
might distort meaningful patterning in LSI size profiles, and how well do
the combined LSI values of multiple molars mirror size differences in
unconverted measurements of individual molars; and 6) What is the
correlation in the size of molars and post-crania by region, sex, and over
time. We do not directly address the impact of domestication on molar
size because we lack a comparative assemblage of domestic pig. But
before dental measurements can be confidently used as a marker of
domestication, these as yet unanswered questions need to be resolved. It
is our hope that a controlled assessment of the impact of these different
factors on linear measurements of S. scrofa molars will assist archae­
ozoologists in the application and interpretation of dental metrics in the
context of on-going domestication of this species.
2. The modern and ancient samples
The modern sample considered here consists of the upper and lower
molars of 34 wild boar from various collecting localities within South­
west Asia (Fig. 1). These specimens include 20 females and 14 males
ranging in age from a few months to very old animals at the end of their
natural life span (Table 1, Tables S1 and S2). A full description of this
sample can be found in Zeder and Lemoine (2020). Metric data was
M.A. Zeder and X. Lemoine Journal of Archaeological Science 124 (2020) 105266

recorded for 258 permanent molars of these modern wild boar

(Table S3).
The archaeological S. scrofa are from the site of Hallan Çemi in
southeastern Turkey (Fig. 1). Occupied for a brief time around 11,600
BP, Hallan Çemi is one of the earliest sedentary communities established
in the region at the beginning of the Early Holocene (Rosenberg 1994;
Zeder and Spitzer 2016). The nearly two metric tons of animal remains
recovered during salvage excavations in the 1990s included a large
number of S. scrofa bones and teeth (8000 NISP, 6000 MNE), making
this among the largest archaeological assemblage of S. scrofa in South­
west Asia. Metric data was collected on 236 upper and lower S. scrofa
molars from Hallan Çemi following the same protocols used in the Fig. 2. Occlusal length measurement guide. Inferior view of the left side of a
measurement of the modern assemblage (Table S4). S. scrofa maxilla with dotted black lines showing occlusal length measurements
Dimensions measured in this study include: 1) Greatest overall length, for the permanent M1, M2, and M3. The interstitial margins between teeth are
taken mesio-distally and equivalent to “L” following von den Driesch indicated with solid white lines and the distalmost margin of the M3 is marked
1976; 2) Occlusal length, taken mesio-distally from each interstitial by a solid black line. Measurements for permanent M1 and M2 are taken mesio-
margin for the M1 and M2, and for the M3 from the M2-M3 interstitial distally as the greatest length between the interstitial margins of its adjacent
margin to the distalmost portion of the third cusp visible above the teeth. Occlusal measurement for the permanent M3 is taken medio-distally as
alveolar margin (Fig. 2); and 3) Greatest breadth, taken lingual-labially the greatest length between the M2/M3 interstitial margin and the distalmost
point of the third cusp of the M3 exposed above the alveolar surface.
from the widest point of the crown and equivalent to “B” in von den
Driesch 1976. Measurement of greatest overall length was not possible
for teeth within alveolar bone without damaging the specimen—as was the problems introduced if measurements from both occlusal and overall
the case with the majority of our modern specimens and many of the lengths are used without distinction by comparing overall and occlusal
ancient ones. As a result, greatest occlusal lengths were the only length length measurements of the same upper and lower M2s—the teeth for
measurement recorded for most of these teeth and are, thus, the primary which we had the most robust sample sizes for these two measurements.
focus here. When we compare these two length measurements, we see that occlusal
We note that while we generally use occlusal lengths as a proxy for length measurements are systematically and significantly smaller than
molar length, occlusal and overall lengths should not be used inter­ overall length measurements (Fig. 3, Table S5). Archaeozoologists
changeably in archaeozoological analyses because they measure two should thus be explicit about which length measurements are being used
different things and are not directly comparable. We can get an idea of and never mix the two to prevent invalid identification of patterns of
molar size change that may instead actually be the result of the differ­
ence between occlusal and overall length measurements.
Table 1 Statistical tests for significance include a combination of one-way
Modern Sus scrofa specimens by region, sex, and age (see Table S1). ANOVAs to assess variation across multiple variables, and Wilcoxon
Province Long Bone Age Class Female Male Total signed-rank tests to assess the significance of two way comparisons be­
Kızılcahamam C 0 1 1 tween variables. All statistical tests were performed using JMP® Version
Kızılcahamam D 1 1 2 14 (SAS Institute Inc., 2019). Descriptive statistics for all molar mea­
Kızılcahamam E 4 2 6 surements discussed in this paper, along with the results of these sta­
Kızılcahamam F 2 1 3 tistical tests, are reported in the supplemental tables.
Kızılcahamam G 2 1 3
Kızılcahamam H 1 0 1
Kızılcahamam I 1 1 2
Kızılcahamam Total 11 7 18
Khuzistan J 0 1 1
Khuzistan K 0 1 1
Khuzistan Total 0 2 2
Highland Zagros C 1 0 1
Highland Zagros F 1 0 1
Highland Zagros H 2 0 1
Highland Zagros I 1 1 1
Highland Zagros Total 5 1 4
Caspian B 0 1 1
Caspian E 0 1 1
Caspian I 1 0 1
Caspian K 1 0 1
Caspian Total 2 2 4
Sistan B 1 0 1
Sistan C 0 1 1
Sistan J 0 1 1
Sistan K 1 0 1
Sistan Total 2 2 4
All Regions B 1 1 2
All Regions C 1 2 3
All Regions D 1 1 2
All Regions E 4 3 7
All Regions F 3 1 4
All Regions G 2 1 3
All Regions H 3 0 3
All Regions I 3 2 5
All Regions J 0 2 2
All Regions K 2 1 3
All Regions Total 20 14 34 Fig. 3. Comparison of length and occlusal length measurements of upper and
lower M2s. Sample sizes in Table S5.

3
M.A. Zeder and X. Lemoine
Fig. 4. Wear stages and wear groups for S. scrofa teeth (adapted from Grant 1982).
3. Impact of wear on molar size
The impact of progressive age-related wear on molar dimensions as a
potentially confounding factor in the evaluation of these measurements
as proxies for phenotypic change was recognized early on (e.g. Payne
and Bull 1988), and dental attrition is often mentioned in more recent
attempts at refining (e.g. Mayer et al., 1998) and applying (e.g. Wang
et al., 2015) linear molar measurements in the evaluation of archaeo­
logical assemblages. Interstitial wear is suspected of having a particu­
larly pronounced effect on molar lengths—which are thought to
decrease with age due to attrition, making length measurements a less
reliable indicator of phylogenetic changes (such as those that may come
about through domestication) than breadth measurements (e.g. Kusat­
man 1991; Price 2016; Price and Evin 2017, see also Ervynck et al.,
2001). Until now, however, there has been no controlled study of the
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Journal of Archaeological Science 124 (2020) 105266
impact of age-related wear on the different linear dimensions of molars.
In addition to metric data, eruption and wear stages were recorded
for individual teeth following the system outlined in Lemoine et al.,
(2014) (Fig. 4). In order to assess the impact of progressive wear on the
length and breadth of S. scrofa molars, we combined the 19 different
eruption and wear stages into four wear groups: 1) Wear Group 1: wear
stages 1–9, including unworn teeth in the process of eruption and teeth
just beginning to come into wear, 2) Wear Group 2: wear stages 10–12,
corresponding to light wear, 3) Wear Group 3: wear stages 13–15,
denoting moderate wear, and 4) Wear Group 4: wear stages 16–19,
including molars in heavy wear (see Tables S3 and S4 for eruption and
wear stage assignments of teeth used in this study).
Given the limited number of overall length measurements of molars,
we can only look at the impact of wear on occlusal length and breadth in
the modern sample. As expected, the impact of progressive wear on
M.A. Zeder and X. Lemoine Journal of Archaeological Science 124 (2020) 105266

Fig. 5. Modern S. scrofa occlusal lengths and breadths by wear group. Sample sizes in Table S6.

occlusal length measurements is quite profound. Both the upper and
lower M1s and M2s show dramatic decreases in occlusal length with
each progressive wear stage (Fig. 5, Table S6a). Wilcoxon signed rank
tests for differences between wear groups (Table S6c) find statistically
significant differences (α = 0.05) between all four groups for the upper
and lower M1s. For the upper and lower M2s, decreases in occlusal
length are most evident between groups 2 and 3, with specimens in wear
groups 1 and 2 similar in length to one another. We should note that
sample sizes are very small for teeth in groups 3 and 4 for both the upper
(n = 4) and lower (n = 4) M2s, likely due to the relatively later eruption

5
M.A. Zeder and X. Lemoine
Fig. 6. Hallan Çemi S. scrofa LM1 overall and occlusal lengths and breadths by wear group. Sample sizes in Table S7.
(and thus less time subject to attrition) of this tooth compared to the
M1s.
This problem of late eruption resulting in smaller sample sizes in the
later wear groups is especially pronounced for the upper and lower M3s.
Thus, the insignificant results from our one-way ANOVAs for the
occlusal lengths of the M3s (Table S6a) must be very conservatively
interpreted. Although small sample size impedes our ability to fully
assess the effect of progressive dental wear on M3 length, it seems likely
that because these teeth are subject to only unidirectional interstitial
wear forces from the adjacent M2, their lengths are not as affected by
progressive wear—at least in contrast to the M1 and M2. The M3’s
prolonged and progressive cusp-by-cusp crown development and erup­
tion, however, presents special challenges that make measurements of
this tooth especially prone to observer-introduced error (Ide et al., 2013;
Tonge and McCance 1973). If the stage of development is not precisely
accounted for, measuring loose M3s collected from archaeological sites
that have not yet completed ossification, but may already show signs of
attrition on the mesial cusps, risks including teeth with under-developed
distal aspects that can erroneously drive down M3 occlusal and overall
length measurements of a given assemblage—especially if that assem­
blage contains a large number of S. scrofa between 8 and 24 months of
age when this is most likely to occur.
In contrast to length, progressive, age-associated dental attrition
seems to have little impact on molar breadth measurements (Fig. 5,
Table S6b). The distribution of breadth measurements for all upper and
lower molars is quite constant across the four different wear groups,
with no significant changes detected by ANOVAs (Table S6c). This
constancy is particularly apparent in both upper and lower M1s for
which we have the largest sample of teeth across all four wear groups.
Again, small sample sizes in the latter two wear groups for the upper and
lower M3s and M2s necessitate some caution in interpreting these
results.
Although limited, the number of overall length measurements for the
Hallan Çemi molars is somewhat larger than it is in the modern sample,
and, while the sample is too small for reliable statistical tests, there are
enough overall length measurements of lower first molars in the Hallan
Çemi sample to demonstrate that overall length measurements are
similarly affected by interstitial wear (Fig. 6, Table S7). Both overall and
occlusal length measurements of the lower M1 show a similar downward
progression in each successive wear group, while breadth measurements
remain generally steady. Moreover, this comparison also highlights the
difference between overall and occlusal length measurements with
overall length measurements larger than occlusal lengths in each wear
group, further underscoring the dangers of combining overall and
occlusal length measurements.
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Journal of Archaeological Science 124 (2020) 105266
Examination of the impact of age related wear on molar size, then,
confirms that progressive wear has a significant impact on the both
occlusal and overall length dimensions of molar teeth in S. scrofa. Molar
length comparisons should only proceed by controlling for wear –
comparing molars in one or perhaps two continuous wear groups. As a
rule, molar teeth in heavy wear should not be used for these purposes
and M3 lengths should be treated with extreme caution, or be excluded
altogether, due to the problems the development of this tooth presents to
accurate and reliable measurement. Our results also confirm, however,
that breadth measurements are little affected by wear and thus likely
serve as more reliable and valid measures differences in S. scrofa molar
size than length measurements. In addition, our results reinforce the
importance of recording and considering the status eruption and wear
on all molar teeth in any evaluation of molar size.
4. Impact of region on molar size
Our earlier study of post-cranial elements of the modern wild boar
assemblage found significant variability between the Turkish specimens
in our sample and those from Iraq and Iran (Zeder and Lemoine 2020).
Here we examine whether these differences are also reflected in the
molar size of wild boar in this same modern assemblage. Ideally, eval­
uation of the impact of region on size should be based on well-defined
populations from discrete collecting localities. The Turkish wild boar
in our sample meet this criterion being from a single population in the
Kızılcahamam preserve collected by Sabastian Payne and Gail Bull in the
1970s (Payne and Bull 1988). In contrast, modern boar from Iraq and
Iran are from multiple populations over this broad territory that were
collected by different expeditions over more than 50 years (Table S1).
Even when combined into four general regions (Table 1), there are too
few specimens to allow for a valid region-by-region comparison. In our
earlier study (Zeder and Lemoine 2020) we determined that post-cranial
dimensions of boar from different collecting localities in Iraq and Iran
were similar enough to one another that they could be combined into a
single Iraq/Iran group that could then be compared to the post-crania of
the Turkish wild boar. The same is true for molar dimensions (Table S8)
which we have again combined into a single Iraq/Iran group for com­
parison with the Turkish boar. We also restrict our comparison of molar
size in these two regional samples to females (for which we have the
most specimens) to control for any sex related variability in size.
Moreover, given the clear impact of age-related wear on occlusal lengths
discussed above, we further limit the comparison of occlusal lengths to
the teeth of female boar in wear groups 1 and 2.
Molar dimensions of wild boar from these two regions are very
similar (Fig. 7, Table S9). In almost all cases, there is no statistical
M.A. Zeder and X. Lemoine
Fig. 7. Occlusal lengths and breadths by region. Sample sizes in Table S9.
difference between either the occlusal lengths (controlled for wear) or
the breadths of upper and lower molars of the Turkish and the Iraq/Iran
female boar. The only statistically significant differences are between
the occlusal lengths of the lower M1 (that are slightly larger in the Iraq/
Iran specimens than in the Turkish boar) (p-value = 0.0384, α = 0.05)
(Table S9a) and in the breadths of the upper M1 (that are slightly smaller
in the Iraq/Iran specimens than in the Turkish ones) (p-value = 0.0403,
α = 0.05) (Table S9b).
5. Impact of sex on molar size
In contrast to post-cranial elements which are highly dimorphic in
S. scrofa (see Zeder and Lemoine 2020), it is generally thought that
molars do not to display a strong degree of sexual dimorphism (Ervynck
et al., 2001; Mayer et al., 1998; Payne and Bull, 1988; Price and
Arbuckle, 2015; Price and Evin 2017). This opinion is not universal, with
an early application of geometric morphometric analysis to S. scrofa
molar size and shape maintaining that there is, in fact, “clear dimor­
phism in molar size in wild boars” (Evin et al., 2013:742). Empirical
7
Journal of Archaeological Science 124 (2020) 105266
evaluation of the impact of sex on molar size in S. scrofa, however, is
limited to one study that documented some sex-based variation in the
size upper and lower M2 and M3 lengths and breadths among diverse
sample of wild boar from Europe, North Africa, and Asia that also
included feral swine and wild boarxferal swine hybrids (Mayer et al.,
1998). We consider this question with a less geographically and
phylogenetically diverse sample of modern wild boar. Given the simi­
larity in molar sizes of the Turkish and the Iraq/Iran wild boar, our
examination of sex-linked differences in molar size are based on a
combined sample of specimens from both regions. As before, however,
we limit our evaluation of occlusal length measurements to specimens in
wear groups 1 and 2 to control for age-related wear.
With the exception of the upper and lower M3s, sex-linked differ­
ences in occlusal lengths are either only weakly significant (for upper
M2s) (p-value = 0.0976) or not significant (Fig. 8, Table S10a). Occlusal
lengths of both upper and lower M3s, however, are significantly larger
than females at a high degree of confidence (p < 0.05). Sex-linked dif­
ferences in molar breadths, in contrast, appear to be much stronger
(Table S10b). With the exception of the lower M1s and, to a lesser
M.A. Zeder and X. Lemoine
degree, the Upper M3s, the molar breadths of males are significantly
larger than those of females showing highly significant differences (p-
value ≤ 0.05).
Thus, it would appear that, sexual dimorphism plays a role in molar
size in wild boar, especially in molar breadths, but also in the occlusal
lengths of upper and lower M3s when controlled for wear. The degree of
sexual dimorphism detected in M3 occlusal lengths is especially
important given the ubiquity of early studies that focus on M3 lengths in
attempting to distinguish between wild and domestic S. scrofa. Before
interpreting downward sifts in molar breadths or M3 lengths as an in­
dicator of on-going domestication, one needs first to consider the pos­
sibility that these changes are instead attributable to demographic
differences in the proportions of males and females in these
assemblages.
6. Impact of time on molar size
Our earlier study also revealed a marked reduction in the size of post-
cranial elements in modern wild boar when compared to ancient boar
from Hallan Çemi, raising once again the question of whether the molar
Fig. 8. Occlusal lengths and breadths by sex. Sample sizes in Table S10.
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Journal of Archaeological Science 124 (2020) 105266
sizes show similar variation. When we compare modern and ancient
boar occlusal lengths and breadths (Fig. 9, Table S11), however, we find
little or no change over the nearly 12,000 years that separate these two
assemblages. Molar breadths do show some indication of decrease over
time in upper and lower M1s and M2s, and lower M3s Table S11b) that
are especially apparent in the box plots in Fig. 9, but these differences
are only statistically significant for the upper M1s and then only weakly
so (p = 0.072).
7. The application of logarithm size index scaling to dental
measurements
Logarithm size index (LSI) scaling of metric data is one of the most
widely used techniques in the metrical analyses of archaeological animal
bones. Normalizing measurements of different dimensions to a single
standard allows metric data from different elements to be combined into
a single size profile making it possible to draw comparisons between
assemblages despite limited sample size and/or differential element
representation. While this technique is most commonly used to examine
variation in the size of post-cranial elements, LSI scaling is also
M.A. Zeder and X. Lemoine
increasingly being applied to dental metric data (Albarella et al., 2006b;
Price and Arbuckle, 2015; Price and Hongo 2019). The generally more
restricted number of teeth recovered from archaeological sites makes
the application of LSI scaling to dental metrics a particularly attractive
way of obtaining a picture of size variation in dentition across time and
space.
There is, however, little standardization in how this method is
applied to dental metrics. Some applications combine breadth mea­
surements of anterior and posterior cusps of both upper and lower mo­
lars, along with breadths of deciduous fourth pre-molars (e.g. Albarella
et al. 2006a, 2006b; Hamilton and Thomas, 2012; Price and Arbuckle,
2015; Price and Evin 2017; Price et al., 2017); some construct separate
profiles for dental length and breadth measurements (Albarella et al.,
2006a); still others combine length and breadth measurements into
single size profiles (Albarella et al., 2006b; Hamilton and Thomas 2012).
Fig. 9. Occlusal lengths and breadths over time. Sample sizes in Table S11.
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Journal of Archaeological Science 124 (2020) 105266
It is often not clear whether upper and lower teeth are combined in these
size profiles, or if length measurements are of the overall length of the
tooth, its occlusal surface, or some combination of the two. Sometimes it
is unclear precisely which teeth are included in these profiles, or in what
proportions they contribute to the total sample. In all cases the degree of
wear of the individual teeth are not precisely specified nor controlled for
in the aggregate LSI analyses.
The underlying assumption of size index scaling is that all di­
mensions of a single specimen will vary isometrically so that individual
measurements of different elements of the same specimen will all scale
the same way when compared to the standard specimen (i.e., the dif­
ference between each dimension of the elements of a single specimen
and those of the chosen standard will be identical, or at least very
similar, to one another). This basic assumption is violated, however, if
there is substantial variation in how different dimensions of a single
M.A. Zeder and X. Lemoine Journal of Archaeological Science 124 (2020) 105266

specimen scale relative to the standard—a potential source of bias that

Table 2
might mask any real patterning in the size profile of an assemblage based
Dental metrics for standard Sus scrofa (FMNH 97887, see Table S3).
on scaled values. Our earlier application of LSI scaling to S. scrofa post-
cranial metric data from the same modern and ancient samples used in FMNH 97887 UM1 UM2 UM3 LM1 LM2 LM3

the present study found that this assumption does not hold for all post- Breadth 19.59 22.23 24.54 14.20 18.21 21.03
crania (Zeder and Lemoine 2020). LSI values of breadth measurements Occlusal Length 16.56 23.93 44.90 14.86 21.04 49.92
of the glenoid process of the scapula, for example, scaled so much
smaller than those of all other post-cranial elements that they were
similar, as expected under the assumption of isometric scaling, there is a
excluded from the study. Less marked differences were also found for
steady downward progression of LSI values of occlusal length values of
other post-cranial dimensions, resulting in the division of the elements
each individual tooth (Fig. 10, Table S12a). Moreover, there is little
in the modern and ancient S. scrofa assemblages into two groups of
correspondence between the same molar in upper and lower tooth rows;
larger and smaller scaling dimensions.
upper molars uniformly scale higher relative to the standard than the
So, it is important to ask whether there are similar scaling issues in
corresponding lower molars. With the exception of the upper and lower
LSI values of dental metric data. Specifically, is there enough similarity
M3s, there is virtually no overlap in the means, medians, quartile values,
in the ways that individual molar dimensions scale relative to a standard
minimum and maximum of the LSI values for any of these teeth, which
to permit combing them by tooth row or as a combination of both upper
are all significantly different from one another at a high degree of
and lower tooth rows? We also need to consider whether size profiles
confidence (p-value < 0.0001) (Table S12c).
based on scaled values reflect patterning seen in unaltered metrics of
The extreme variation in the scaled values of occlusal length mea­
individual molar dimensions. In particular, do we see the same
surements can in part be attributed to the animal chosen for use as a
patterning in dental size by region, sex, and over time that are evident in
standard. Specimen FMNH 97887 is a very large older male between
the unconverted metrics of individual molars?
about 5 to 8 years of age which was chosen as the standard animal in our
We use the same adult male from Sistan province Iran as the standard
earlier study because all of its post-cranial elements were fused (see
for computing LSI values of molar measurements as used in our earlier
Zeder and Lemoine 2020). The advanced age of this specimen, however,
study of post-cranial dimensions (FMNH 97887, Table S1) to allow for
makes it a less than ideal standard for the conversion of occlusal length
direct comparisons of post-cranial and molar size variation (see below).
measurements into LSI values as this specimen had highly worn first and
We also use the same formula, d = log(x/y) — where x is the dimension
second molars (both in wear group 4), but third molars in light wear
of the modern specimens, y the dimension of the standard, and d the
(wear group 2) (Table S3). The occlusal measurements for this indi­
resulting scaled dimension. Due to the lack of overall length measure­
vidual reflect these differential wear states; upper and lower M1 occlusal
ments for this specimen (and indeed for the majority of specimens in our
lengths are considerably smaller than those of other modern wild boar in
sample of modern wild boar), we can only standardize occlusal lengths
the sample (male and female), the M2s are about the same size as those
and maximum breadth measurements. LSI values for occlusal lengths
of other specimens, and the M3 occlusal lengths are considerably larger
and breadth measurements of for this specimen are included in Table S3.
than those of other specimens. So it is no surprise that the LSI values of
The measurements of this specimen used to compute LSI values for both
the combined occlusal length of M1s and M2s in wear groups 1 and 2 are
the modern and ancient samples are presented in Table 2. Once again we
as a whole greater than 0 on the log normalized scale (meaning they are
restrict the evaluation of occlusal lengths to specimens in wear groups 1
larger than the standard), while the M3s are less than 0 (meaning they
and 2.
are smaller than the standard). But while the differential impact of
progressive wear of the molars of this individual likely contributes to the
7.1. Scaling downward progression of the occlusal length LSI values of the other
modern specimens relative to this standard, it cannot account for the
There is no agreement between LSI values of occlusal lengths of in­ marked offset in the LSI values of the same molar in the upper and lower
dividual teeth within the same tooth row. Instead of being roughly

Fig. 10. LSI values of individual molars of modern S. scrofa. Sample sizes in Table S12.

10
M.A. Zeder and X. Lemoine
jaws. It would seem, then, a number of different factors affect LSI values
for molar length measurements — differential impacts of wear on molar
length, the choice of standard used in logarithmic scaling, and the failure
of length measurements to scale isometrically — that raise concerns
about utility of the LSI scaling technique for molar length
measurements.
The choice of standard has no impact on the LSI values of molar
breadths for the modern specimens. Despite the advanced state of wear
of the first and second molars of this individual, the breadths of all its
molars (upper and lower) are consistently larger than those of most
other modern S. scrofa in the assemblage (Table S3). This is why the LSI
values of molar breadths of the other modern boar computed using this
standard mostly fall below 0 on the log normalized scale (Fig. 10,
Table S12b). Moreover, with the exception of the upper M1s, there is a
much closer correspondence between the LSI values of molar breadths
than of occlusal lengths. LSI values of upper M1 breadths scale signifi­
cantly smaller than those of other molars with a high degree of statistical
significance (p-values < 0.0001) (Table S12c). In contrast, there is no
statistical difference between the LSI values of the breadths of upper M2s
compared to upper M3s, nor is there a statistically significant difference
between the LSI values of the lower M1 breadths and those of lower M3s.
And while there are significant differences between the LSI breadth
values of lower M1s and M2s, as well as between lower M2s and M3s,
these differences are not as highly significant as those between the LSI
values for upper M1 breadths and other molars. So, while breadth
measurements of individual teeth do not strictly conform to the
assumption of isometric scaling, the general similarities of these values
for molar breadths (excepting the upper M1s) are likely enough to
permit them to be combined into size profiles of upper and lower tooth
rows.
As to the question of whether LSI values of individual teeth from
upper and lower teeth can be combined (Fig. 11, Table S13), clearly this
is not advisable for upper and lower occlusal length measurements
which Wilcoxon signed-rank tests indicate differ from one another with
a high degree of significance (p-values < 0.0001). The sharp difference
between occlusal length LSI values and those for breadth measurements
also rules out the possibility of combining occlusal length and breadth
LSI values (Fig. 11). Combined LSI values of the breadths of upper and
lower molars also differ significantly from one another when LSI values
for the upper M1s are included (p = 0.0055). Excluding the upper M1
Fig. 11. LSI values of upper and lower molars of modern S. scrofa. Occlusal
lengths = wear groups 1&2. Sample sizes in Table S13.
11
Journal of Archaeological Science 124 (2020) 105266
breadths decreases the difference between upper and lower tooth rows,
but the difference is still statistically significant at a relatively weak level
(p = 0.0934).
LSI values of occlusal lengths, then, are not a reliable way to monitor
variations in molar size, either as size profiles of combined occlusal
length values (by jaw or for both jaws combined), and certainly not as
part of size profiles based on a combination of length and breadth LSI
values. LSI values of molar breadth measurements are less problematic
but are likely best considered separately for upper and lower tooth rows.
Accordingly, we limit our further consideration of log normalized metric
data to LSI values of breadth measurements by tooth row, excluding the
M1s, in comparisons of LSI values of breadths in upper tooth rows.
7.2. LSI values by locality, sex, and time
The examination of unaltered measurements of individual teeth
shows little difference between Turkish and the Iraq/Iran S. scrofa in our
modern assemblage. Nor does there appear to be much change in molar
size over time. In contrast, we found considerable, and in many cases
highly significant, differences in breadth measurements by sex. The
question then arises whether these patterns are also seen in size profiles
based on combined LSI values of different molars.
Turning first to locality, LSI values of molar breadths of upper and
lower molar rows seem to show a tendency for the molars of Turkish
boar to be slightly larger than those of the wild boar from Iraq and Iran
(Fig. 12). These differences are not, however, statistically significant
(Table S14). Thus, the LSI values of molar breadths confirm and
strengthen the impression gained from the evaluation of unconverted
metrics of individual teeth that there is no significant difference in molar
breadths of wild boar across this broad territory from southeastern
Turkey to southeastern Iran.
On the other hand, LSI values of upper and lower molar breadths
amplify the small, but statistically insignificant, tendency toward size
reduction seen in the unconverted measurements of individual teeth of
modern wild boar when compared to those from Hallan Çemi (Fig. 13,
Table S15). The difference between the LSI values from the ancient and
Fig. 12. LSI values of breadth measurements by region. Sample sizes
in Table S14.
M.A. Zeder and X. Lemoine Journal of Archaeological Science 124 (2020) 105266

in molar size.

8. Size variation in molars and post-crania

In addition to making it possible to combine metric data from mul­

tiple elements into robust size profiles, logarithm scaling techniques also
provide a means for comparing variability in body size between different
components of an animal’s skeleton. In particular, they allow us to ask if
all skeletal components experience the same responses to forces
affecting body size in animals, or whether, instead, different parts of the
skeleton respond to different factors in different ways. Given the ubiq­
uity of LSI-based analyses of archaeological animal bone it is surprising
that this topic has not been pursued in any depth. The metric data
compiled here for the large assemblages modern and ancient wild boar
make such a comparison possible. Specifically we ask four questions: 1)
Are there differences in how post-cranial and dental dimension scale
relative to one another; 2) How do the sizes of molars and post-crania
vary by region; 3) How do the sizes of these respective skeletal ele­
ments vary by sex; and 4) How do they vary over time?

8.1. Scaling of molar and post-cranial metric data

Fig. 13. LSI values of breadth measurements over time. Sample sizes
in Table S15. Scaling issues are addressed by comparing the distribution of LSI
values for upper (excluding M1s) and lower molars of the modern
modern boar molar breadth is now significant at a high degree of con­ S. scrofa to those of post-crania in the two different larger and smaller
fidence (p ≤ 0.05) for both the upper and lower jaws scaling groups mentioned above and discussed at length in Zeder and
Sex-linked differences in molar breadths are especially amplified Lemoine (2020) (Fig. 15, Table S17). If the assumption of isometric
when measurements of individual teeth are converted into LSI values scaling applies to all skeletal elements (dental and post-cranial) then the
and combined into tooth rows (Fig. 14). Wilcoxon signed-rank tests for
LSI values of the breadths upper and lower molar teeth find a highly
significant difference between the molar breadths of upper and lower
teeth (p < 0.05) (Table S16).
LSI profiles of molar breadth measurements (excluding the upper
M1s), then, both capture and amplify patterns seen among unconverted
measurements of individual teeth making the application of logarithm
scaling to molar breadths an effective technique for evaluating variation

Fig. 15. LSI values of upper and lower molars compared to post-cranial group 1
and group 2 elements for modern S. scrofa. Sample sizes in Table S17.

Fig. 16. LSI values of upper and lower molars compared to post-cranial group 1
Fig. 14. LSI values of breadth measurements by sex. Sample sizes in Table S16. and group 2 elements for modern S. scrofa by region. Sample sizes in Table S18.

12
M.A. Zeder and X. Lemoine
Fig. 17. LSI values of upper and lower molars compared to post-cranial group 1
and group 2 elements for modern S. scrofa by sex. Sample sizes in Table S19.
distribution of the LSI values for these elements should be roughly the
same. Instead, we see a clear downward progression in how these
different groups of elements scale relative to one another. While, as
mentioned above, the distributions of LSI values of upper and lower
molars are roughly similar (though statistically different at a weaker
level of significance), the LSI values of both groups of post-cranial ele­
ments differ from those of molars and each other at a high degree of
significance (p < 0.0001) (Table S17b). Thus, it seems that LSI values of
the dimensions of dental and post-cranial elements within the same
individuals scale differently from one another and should be examined
individually.
8.2. Comparison of LSI values of molars and post-crania by region
We can evaluate the impact of region on the dental and post-cranial
dimensions by comparing LSI values of molars and post-crania of wild
boar from the Turkish population to those of the combined Iraq/Iran
assemblage of S. scrofa. Once again, we limit this comparison to females
to control for any sex-linked differences that might arise from the un­
even representation males and females in the Turkish and Iraq/Iran
assemblages. We further limit the post-cranial assemblage to wild boar
older than 8 months (age class D in Zeder et al., 2015, Table S2) to
eliminate specimens of very young animals represented in the
post-cranial assemblage but not in the assemblage of dental dimensions
of permanent molars. Although LSI values of molar breadths showed a
statistically insignificant tendency for Turkish wild boar to be slightly
larger than those of wild boar from Iraq and Iran, there are major dif­
ferences in the size of the post-cranial elements of boar from these two
regions (Fig. 16, Table S18). In both post-cranial scaling groups wild
boar from Iraq and Iran are significantly larger than those of the Turkish
boar at a high degree of confidence (p < 0.05). Regional differences,
then, have a much greater impact on the size of post-cranial elements
than on molar dimensions.
8.3. Comparison of LSI values of molars and post-crania by sex
In evaluating the impact of sex on the size of molars and post crania
we once again limit our sample of post-crania to wild boar older than 8
months of age (Fig. 17, Table S19). As we saw earlier, there is a clear,
and statistically significant, tendency for male molar breadths to be
somewhat larger than those from females. There is, however, a good
deal of overlap in the interquartile ranges of males and females in the LSI
values of molar breadth measurements in upper and lower molars (see
Fig. 17). In contrast, there is a much greater separation between males
and females in the LSI values of post-crania in both scaling groups which
are statistically different from one another at a high degree of confidence
13
Journal of Archaeological Science 124 (2020) 105266
(p < 0.0001). These sharp differences are seen in all descriptive statistics
for post-cranial LSI values in means, medians, and in the wide separation
between the interquartile ranges of male and female post-cranial LSI
values.
Our earlier study used these marked differences in the interquartile
ranges of post-cranial LSI values to develop a method for identifying
male and female S. scrofa in archaeological assemblages (Zeder and
Lemoine 2020). Following this method, specimens with LSI values
smaller than the third quartile value of females (the top of the female
box in a box plot) are classified as female, while those greater than the
first quartile value of males (the bottom of the box for the males) are
classified as male. The range of values between them is considered an
overlap zone where sex cannot be determined. The marked separation
between the interquartile ranges of females and males in post-cranial LSI
values means that these cut-off points capture the majority of male and
specimens in the sample, with only a handful of specimens falling into
the overlap range of values that cannot be assigned to sex. The large
overlap between the interquartile ranges of female and male molars LSI
values, however, speaks against applying this technique to dental met­
rics. The number of specimens that would fall into the overlap range
constitute more than half of an assemblage of molars, making any age
profile based on the small sample of sexable molars highly unreliable.
Although there does not appear to be a way to cleanly separate dentition
by sex that would make it possible to construct sex-specific age profiles
based on dental eruption and wear in archaeological assemblages, sex is
nonetheless a significant factor in molar size that must be taken into
consideration in the interpretation of dental metrics.
8.4. Comparison of LSI values of molars and post-crania over time
As noted above, the LSI values for upper and lower molars amplify a
small, but significant, downward trend in molar size over the roughly
12,000 years that separate these two assemblages. The reduction in the
LSI values of post-crania over time, however, is much more dramatic,
with the Hallan Çemi post-cranial LSI values differing from modern wild
boar with a high degree of statistical confidence (p < 0.0001)
(Table S20). Box-plots of LSI values for ancient and modern S. scrofa
clearly demonstrate the reduction on the size of post-cranial elements
(and thus in overall body size) over time (Fig. 18). While there is
considerable overlap in the ranges of LSI values for upper and lower
molars in the ancient and modern assemblages, there is no overlap in the
third to first interquartile ranges of the post-cranial LSI values of the
ancient and modern assemblages (the boxes of the box plots), and little
overlap between the outlying first and fourth quartile ranges (the tails of
Fig. 18. Comparison of LSI values of upper and lower molars compared to post-
cranial group 1 and group 2 elements for modern and Hallan Çemi S. scrofa
(post-crania include immature). Sample sizes in Table S20.
M.A. Zeder and X. Lemoine
the boxes).
The differential impact of time on molar breadths post-cranial ele­
ments is further underscored by a comparison of the relationships be­
tween the LSI values of molars and post-cranial elements within the
modern and ancient assemblage. As noted above, the LSI values of post-
cranial elements in the modern wild boar are significantly smaller than
the values for molar breadths from the same animals at a high degree of
statistical confidence. Post-cranial LSI values of Hallan Çemi S. scrofa
are also significantly different from Hallan Çemi molar breadths, but in
this case the post-cranial LSI values are substantially larger (more pos­
itive) than the molar breadths, highlighting the degree to which post-
cranial elements have reduced in size over time when compared to teeth.
This pattern is unlikely to be an artifact of differential sex compo­
sition of the two assemblages (i.e., that the smaller size of the modern
S. scrofa is attributable to a higher proportion of females in the assem­
blage compared to Hallan Çemi). Based on the method for identifying
the sex of archaeological post-crania developed in Zeder and Lemoine
(2020), we estimate that females make up almost identical proportions
of both the modern and the Hallan Çemi assemblage (65% and 63%
respectively). Given the relative absence of domestic pigs across the
Near East today, the substantial difference between the size of the Hallan
Çemi and the modern boar is also likely not attributable to recent
introgression between wild and domestic S. scrofa in the modern sample.
Instead the marked reduction in the size of post-cranial elements evident
in modern wild boar when compared to those from Hallan Çemi is likely
an artifact of climatic shifts toward warmer temperatures that began in
the Late Pleistocene and have been held to account for similar changes in
the size of a number of species since the Early Holocene (caprines, ga­
zelle, foxes, and rabbits) (Albarella et al., 2009; Davis 1977, 2019; Zeder
2008).
9. Recommendations and observations
Based on this analysis, we can make a number of recommendations
about of the impact of various factors on molar size in S. scrofa. Spe­
cifically, the data presented here point to a number of protocols that
might be considered “best practices” in the analysis of S. scrofa dental
metrics. We can also offer several concrete observations about the im­
pacts of factors such as region, sex, and change over time on the size of
the molars of this species that may open up new areas of research, as well
as help better inform the traditional use of tooth size as a marker of
domestication.
9.1. Controlling for the impact of wear on dental metrics
Our data confirm the hypothesis that interstitial wear has a dramatic
impact on molar size, particularly on molar length measurements in
upper and lower M1 and M2s. As a rule, comparisons of length mea­
surements of S. scrofa should only be made if one strictly controls for
state of wear. Age-related wear is likely to have a smaller impact on
length measurements of the M3, but here the prolonged sequence of
crown development and eruption of this tooth (both upper and lower)
present special challenges to obtaining accurate length measurements
which should only be used with care for analyses of molar size. Given the
marked differences we document between overall length and occlusal
length measurements, care must also be taken to specify which of these
two different length measurements are being used and to never combine
the two. It would also be useful to note whether length measurements
are taken on loose teeth or teeth still situated within alveolar bone with
adjacent teeth still present, as this may affect molar length measure­
ments. We realize that these restrictions will limit the size of datasets of
length measurements suitable for comparative studies. Failing to do so,
however, renders such comparisons essentially meaningless.
Breadth measurements, on the other hand, are little affected by
progressive wear and, as a result, are preferable to length measurements
in the study molar size in S. scrofa. It is also easier to measure the
14
Journal of Archaeological Science 124 (2020) 105266
breadth of fully erupted molars that are within alveolar bone in a way
that makes the measurements directly comparable to those of loose
teeth. As a general rule, however, recording of eruption and wear of all
teeth, preferably using the system based on Grant, 1982, should be
standard practice in any study of S. scrofa dental metrics, whether using
length or breadth measurements. Where this information is not available
(i.e., in older literature), it may be possible to refer to other information
on age structure (i.e., dental or long bone age profiles) to assess whether
there are any potential age-related differentials in wear that might be
affecting metric data.
9.2. Protocols for the application of log normalizing techniques on dental
metrics
Size profiles composed of LSI values of molar lengths amplify biases
introduced by differential wear. Even when controlling for wear, scaling
differences in molar length measurements introduces another source of
bias that would likely either blur or override any “real” patterning in
normalized length measurements that might be of interest. Application
of logarithm size scaling techniques to length measurements should,
then, be avoided. Size profiles composed of combined length and
breadth molar measurements are particularly unreliable.
LSI values of most molar breadth measurements, on the other hand,
are relatively unaffected by either differential wear or scaling issues.
Moreover, since combined LSI values of multiple teeth amplify
patterning in molar size variation, log normalized size profiles based on
combined LSI values of molar breadths are preferable to raw measure­
ments of individual molars as a way of monitoring meaningful vari­
ability in molar size. The offset of LSI values of upper M1s from other
molar breadth values means, however, that these teeth should be
excluded from size profiles based on normalized data. While LSI values
for upper and lower tooth rows are generally similar to one another and
might in the case of very small assemblages be combined, doing so
would mask differences in how teeth in the upper and lower jaws
respond to factors such as locality, sex, time, and, possibly, domestic
status. Therefore, we recommend against combining LSI values of upper
and lower molars in size profiles of normalized data.
9.3. Impacts of region, sex, and time on molar size
The impacts of region, sex, and time on molar size are variable in
degree, teeth, and dimension affected. Unconverted occlusal lengths and
breadth measurements individual molars, as well as LSI values of molar
breadths, indicate that there is essentially no difference in molar sizes
between the Turkish wild boars and the combined sample of boar from
Iraq and Iran. LSI values for upper and lower molar breadths, on the
other hand, amplify the small, but statistically insignificant, tendency
for unconverted measurements of individual molar breadths of modern
boar to be reduced in size compared to those of ancient S. scrofa.
In contrast to region and time which do not seem to have a major
impact on molar size, sex seems to have a much more significant, and
consistent, impact on the breadth of most molars as well as the occlusal
lengths of both upper and lower M3s. Sex-linked differences in molar
breadths are especially strong in LSI values of upper and lower tooth
rows. It would be interesting to see if domestic S. scrofa show the same
degree of sexual dimorphism in molar, and post-crania, size. Domesti­
cation is likely to result in a relaxation of the selective advantage of large
male body size in mate competition and attraction leading to a reduction
in the degree of sexual-dimorphism in size. Large male body size with its
extra energetic requirements might also be actively selected against in
domestic pigs. The differential impact of the relaxation of the selective
advantage of the larger size in post-crania and teeth (and on different
teeth, i.e., canines and molars) in domesticates is another promising area
of inquiry.
The impact of all of these factors on molar size is much more muted,
and less consistent, than it is on post-cranial size. Post-cranial body size
M.A. Zeder and X. Lemoine
of wild boar from Iraq and Iran is substantially larger than that of the
Turkish S. scrofa across both scaling groups and all the elements within
them. There is also a significant reduction in post-cranial body size in
modern wild boar compared to the Early Holocene S. scrofa from Hallan
Çemi. Moreover, the difference between the size of male and female
post-cranial body size is much greater than in molar size, making it
possible to classify individual specimens within an archaeological
assemblage as either male or female with some confidence (Zeder and
Lemoine 2020). In contrast, although there are marked differences in
male and female molar breadths, dental metrics can only give a general
idea of the distribution of males and females within assemblages and
cannot be used to construct sex-specific demographic profiles as is
possible with post-cranial elements.
9.4. Molar size and domestication
There is, then, some truth to the hypothesis that molars are less
affected by factors such as regional variation, sexual dimorphism, and
Fig. 19. Histograms of LSI values of breadth measurements of modern S. scrofa by sex and tooth row. x-axis = NISP, y-axis = LSI value.
15
Journal of Archaeological Science 124 (2020) 105266
time than post-crania, which is often cited as a reason why teeth are
more useful marker of domestication in S. scrofa (Ervynck et al., 2001;
Mayer et al., 1998; Payne and Bull, 1988; Price and Arbuckle, 2015). But
precisely how (and why) domestication might be expected to affect
molar size and how the impact of domestication differs from other fac­
tors such as sex on molar size is not entirely clear. A useful study of
modern wild boar from across Eurasia and an assortment of domestic
breeds of pigs found evidence of considerable differences in linear
measurements of both lengths and breadths of upper and lower M2s and
M3s (Evin et al., 2014). And it would be interesting if a follow-on study
took factors such as age-related wear and sex into consideration in
evaluating size differences within this data set of wild and domestic
S. scrofa. Similarly, Mayer et al.’s (1998) earlier study of molar size in
wild, domestic, and feralized boar argued that domestication-induced
size reduction was greater than that seen between female and male
S. scrofa; but the broad geographical spread of specimens examined and
the failure to take age-related wear to account in the evaluation of length
measurement makes it hard to interpret these results. While we cannot
M.A. Zeder and X. Lemoine
directly address the question of the degree and distinctiveness of the
impact of domestication on molar size here, our results introduce an
important cautionary note to the use of molar size as a marker of
domestication in S. scrofa.
As we have seen, the profound impact of age related wear severely
restricts the utility of length measurements of most molars in evaluating
changes in S. scrofa molar size. Without strictly accounting for wear,
length measurements, especially those of occlusal surfaces, are of little
utility whether as unconverted measurements of individual teeth or as
LSI values. Moreover, though less than in post-crania, the degree of
sexual dimorphism noted in molar breadths, as well as in the length of
M3s, is still substantial and must be taken into account in any study of
S. scrofa molar size. Before interpreting downward sifts in molar
breadths, or M3 lengths, as indicators of on-going domestication, one
needs first to consider the possibility that these changes are instead
attributable to demographic differences in the proportions of males and
females in these assemblages. Given the consistency in the way the
molars of ancient S. scrofa respond to factors such as interstitial wear
and LSI scaling despite the large chronological gap that separates them,
it seems likely that ancient S. scrofa molars display similar sex-based
differences in molar breadth and M3 length measurements as found
for the modern wild boar.
Histograms of molar dimensions of male and female modern wild
boar — a traditional way such data are displayed in studies of S. scrofa
domestication (e.g., Price and Arbuckle, 2015) — underscore the need to
Fig. 20. Histograms of M3 occlusal length measurements of modern S. scrofa. . x-axis = NISP, y-axis = measurement in mm.
16
Journal of Archaeological Science 124 (2020) 105266
take sex into consideration in any evaluation of S. scrofa molar size.
Fig. 19 contrasts histograms of LSI values for upper and lower molars of
males and females, while Fig. 20 presents histograms of occlusal length
measurements of male and female M3s. The differences seen here be­
tween male and female molar size are at least as marked as differences in
molar size that earlier studies have interpreted as markers of domesti­
cation (e.g. Ervynck et al., 2001; Price and Arbuckle, 2015). This is
especially the case for occlusal length measurements of lower M3s
(Fig. 20) — a measurement that has been used in earlier studies as a
marker of domestication in S. scrofa (e.g. Flannery, 1983; Stampfli
1983).
This is not to say conclusions drawn in these studies about the do­
mestic status of S. scrofa with reduced molar size are incorrect. But given
the degree of sexual dimorphism evidenced in molar size documented
here, sex must be taken into consideration before drawing conclusions
about domestication status. Until this is done, archaeozoologists must
consider the strong possibility that any size differences observed in the
molars of different S. scrofa assemblages are at least as likely to be
attributable to differences in the sex composition of the assemblages as
they are to domestic status — if not more so. Similar caution has been
urged in drawing conclusions about domestication on the basis of
reduction in the size of post-cranial elements caprines (Zeder 2001)
—caution that we now know extends to both post-crania and teeth in
S. scrofa.
Sex should not be seen, however, as a confounding factor in research
M.A. Zeder and X. Lemoine
seeking to track the transition from wild to domestic in S. scrofa. Being
able to monitor shifts in the sex, and age, of S. scrofa, and other species
is, in fact, central to any such effort. Indeed, alterations in demographic
profiles of targeted animals are likely to precede any archaeologically
detectable change in the size of skeletal elements (Zeder 2008), making
such demographic shifts likely leading-edge markers of the process of
domestication. The complex demographic structure of wild boar pop­
ulations and the variable ways in which hunters might target them,
combined with the range of possible management practices documented
for pigs (Lemoine 2020; Price 2016; Price and Hongo 2019), makes
accounting for the sex, and age, of archaological S. scrofa assemblages
even more critical. The discovery that both post-crania and teeth reflect
sex-linked dimorphism in body size in S. scrofa will enhance our ability
to reconstruct the strategies employed by transitional hunter-herders
that helped drive the process of domestication in this species.
10. Concluding remarks
This study has shown that a number of enduring and widely held
assumptions about the utility of molar size for tracing domestication in
S. scrofa either need adjustment or in some cases should be abandoned
altogether. The more stringent protocols called for here will require
significant changes in how archaeozoologists go about collecting and
analyzing dental metrics in this species. These include restricting com­
parisons of molar lengths to teeth in similar states of wear, specifying
whether one is dealing with occlusal or overall lengths and not
combining the two, restricting log normalizing techniques to breadth
measurements and not combining LSI values of upper and lower molars,
and, above all, considering the possibility that sex, and not domestica­
tion, is the primary factor affecting molar size in archaeological as­
semblages. These results also call for extra scrutiny of the conclusions
drawn in earlier studies that do not control for such factors. We hope,
however, that a better understanding of the range of factors affecting
molar size in S. scrofa will improve the reliability and power of future
research on this species, as well as help in evaluating the validity of
previous work.
Declaration of competing interest
None.
Acknowledgments
This research was supported by grants from the Wenner-Gren
Foundation [Gr. 8619], the Committee for Research and Exploration of
the National Geographic Society [9113-13], and the Scholarly Studies
Humanities Fund of the Smithsonian Institution to Zeder and by Wen­
ner-Gren Grant [Gr. 9416] and National Science Foundation Graduate
Research Fellowship [GR.DGE-1143954] to Lemoine. Once again, we
would like to express our very great appreciation to Sebastian Payne
who provided us access to the remarkable collection of Turkish wild
boar he collected. We would also like to thank the Zoology Department
of the Field Museum of Natural History and the Division of Mammals of
the National Museum of Natural History for providing access to the
modern museum collections analyzed in this study.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.jas.2020.105266.
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