Received: 2 August 2017 Revised: 3 January 2018 Accepted: 18 January 2018

DOI: 10.1002/oa.2642

RESEARCH ARTICLE

Functional relationship between dental macrowear and diet in

Late Pleistocene and recent modern human populations
Luca Fiorenza1,2 | Stefano Benazzi3,4 | Gregorio Oxilia3,5 | Ottmar Kullmer6,7

1
Department of Anatomy and Developmental
Biology, Monash University, Melbourne, VIC, Abstract
Australia Many aspects of diet and behaviour can be gleaned from dental wear, including environmental
2
Earth Sciences, University of New England, conditions, food‐fracture properties, food processing techniques, and cultural habits. Specifically,
Armidale, NSW, Australia the angulation (flat vs. steep) of molar wear has been used as an indicator of food toughness and
3
Department of Cultural Heritage, University has also been implicated in the use of grinding stones and pottery in food processing. In this work,
of Bologna, Bologna, Italy
4
we focus on the sequential phases of the power stroke of mastication in a functional way,
Department of Human Evolution, Max Planck
measuring the inclination of molar wear facets through the occlusal fingerprint analysis method.
Institute for Evolutionary Anthropology,
Leipzig, Germany Specifically, we have calculated the angulation of wear facets in upper and lower molars of
5
Department Biology, University of Florence, Palaeolithic humans, extant hunter‐gatherers, and proto‐farmers to discern differences between
Florence, Italy groups with different diets.
6
Senckenberg Research Institute, Frankfurt, Contrary to previous analyses, our study shows that the molars of Late Pleistocene specimens are
Germany
characterized by significantly steeper angles than those of modern hunter‐gatherers.
7
Department of Paleobiology and
The flat molar wear found in the latter group could be related to the excessive mixture of
Environment, Institute of Ecology, Evolution,
and Diversity, Johann Wolfgang Goethe exogenous materials accidentally introduced into their foods, as indicated by ethnographic
University, Frankfurt, Germany evidence. On the contrary, the steep wear angles characterizing the Palaeolithic group are
Correspondence probably associated with the consumption of a less abrasive diet, which could be ultimately
Dr Luca Fiorenza, Department of Anatomy and
due to food preparation techniques that incorporated less dust and grit into their diets.
Developmental Biology, Monash University,
Melbourne, VIC 3800 Australia.
Email: [email protected] KEY W ORDS

Funding information early farmers, food preparation, hunter‐gatherers, Neanderthals, wear facets
Faculty of Medicine, Nursing and Health
Sciences, Grant/Award Number: SGS16‐0344

1 | I N T RO D U CT I O N (Hinton, 1982; Kaidonis, Townsend, & Richards, 1993; Molnar, 1971,

1972). In particular, it is hypothesized that food preparation practices
The study of tooth wear has been an invaluable tool for the recon- can introduce a large quantity of abrasive exogenous materials, such
struction of dietary and cultural habits in prehistoric populations and as sand, dust, or ash, which may influence the wear patterns of these
extinct human species (e.g., El Zaatari, Grine, Ungar, & Hublin, 2011; human populations (Hinton, 1982; Kaidonis et al., 1993; Molnar,
Fiorenza, 2015; Fiorenza, Bennazi, Tausch, et al., 2011; Fiorenza 1971, 1972; Watson, 2008). Smith (1984) examined the patterns of
et al., 2015; Fiorenza, Benazzi, & Kullmer, 2011; Fiorenza & Kullmer, molar wear in hunter‐gatherers and agriculturalists and associated
2013, 2015; Hinton, 1982; Molnar, 1972; Smith, 1984). Dental wear changes in human subsistence by measuring their wear plane angles.
can be examined at two different scales: macrowear (visible to the For example, hunter‐gatherers and agriculturalists are characterized
naked eye or at low magnification) and microwear (visible only under by flatter and more oblique wear, respectively. This is related with dif-
microscope). Several different factors can contribute to the progres- ferent food preparation techniques that reduce food toughness or
sion of dental wear, including diet, food processing techniques, and fibrousness (Smith, 1984). More interestingly, wear plane angles
exogenous abrasives, and untangling the influence of each factor can between various hunter‐gatherer groups were similar despite regional
be difficult (Addy & Shellis, 2006). That said, several archaeological differences in diet. Similarly, Le Luyer, Rottier, and Bayle (2014)
and ethnographic studies have highlighted the importance of specific examined enamel thickness distribution and variation in Neolithic and
cultural and environmental factors in the formation of specific wear Medieval populations, and these researchers found a correlation
patterns in historic and modern hunter‐gatherer human populations between oblique tooth wear pattern and food preparation techniques.

Int J Osteoarchaeol. 2018;28:153–161. wileyonlinelibrary.com/journal/oa Copyright © 2018 John Wiley & Sons, Ltd. 153
154 FIORENZA ET AL.

Thus, it seems that the overall angulation of molar wear can distinguish We measure and compare the wear facet angulation of Paleolithic
between human populations that adopted significantly different human molars with those of recent modern hunter‐gatherers and early
subsistence strategies (e.g., hunter‐gatherers vs. agriculturalists) but is farmers (with known diets and known cultural habits). Because this
unable to detect smaller scale differences in diet in the hunter‐gatherer approach is limited to molars with a moderate degree of wear
groups. Additionally, analyses of surface microwear texture complexity (Fiorenza, 2009), we could only examine individuals at a young age,
among modern hunter‐gatherer and Paleolithic human teeth were ranging between young adults (with third molars erupted), adolescent
found to be correlated with the level of abrasiveness with food (with second molars erupted), and children (with the first molars
preparation methods associated with cultural changes in the human erupted and in full occlusion). Preliminary results have shown that at
diets (El Zaatari, 2008, 2010; El Zaatari & Hublin, 2014). This leaves more advanced wear stages, there are no significant differences in
open the question of what general wear plane angles can really tell wear facet angles between human groups with different diets and
us about changes in diet, culture, and technology. different subsistence strategies (Fiorenza, 2009). Primary tooth
In our study, we tested whether the angulations of wear facets can morphology becomes obliterated as wear increases; this causes wear
discriminate between human populations from different chronological facets to coalesce and renders their identification impossible. Although
times that relied on contrasting diets and preparation technologies, this limitation may prevent the use of large sample sizes (in particular
using the well‐established three‐dimensional approach known as for the modern hunter‐gatherer groups), it is important to note that
occlusal fingerprint analysis (OFA, Kullmer et al., 2009). Wear facets increased longevity in humans became common very late in our evolu-
are flat and shiny areas with well‐defined borders produced mainly tion. For example, Caspari and Lee (2004) estimated that only 33% of
by the attritional contacts between upper and lower teeth (e.g., Every, Neanderthal individuals of their sample reached adulthood, whereas
1972). The surface of the wear facets is covered with minute subpar- 67% died at a young age. Thus, our approach allows us to take into
allel striations mirroring the orientation of tooth movements (Kay, account a large proportion of Palaeolithic human populations, some-
1977). Every facet is matched by a corresponding facet on a tooth in thing that it was not possible, for instance, in Smith's study (1984),
the opposing arch, and when dentin is exposed, it remains flat with where only 14 Palaeolithic specimens were analyzed.
no “scooping” (Kaidonis, 2008). The OFA method is based on quantita- We test the null hypothesis of no correlation between the angula-
tive analyses that measure different wear facet variables and links tion of wear facets and diet (Hypothesis 0), and we investigated
them with dental function, masticatory movements, culture, and diets whether functional wear facet angles can be employed for larger scale
(Fiorenza, 2015; Fiorenza et al., 2015; Fiorenza, Benazzi, Tausch, subsistence divisions (e.g., hunter‐gatherer vs. farming) that are related
et al., 2011; Fiorenza, Benazzi, Tausch, Kullmer, & Schrenk, 2010; to major differences in food preparation methods and technology
Fiorenza, Benazzi, Viola, et al., 2011; Fiorenza & Kullmer, 2013, (Hypothesis 1). If the null hypothesis is rejected, hunter‐gatherer
2015, 2016; Harvati et al., 2013; Kullmer et al., 2009; Oxilia et al., groups that rely on different diets should be also characterized by
2015; Ulhaas, Kullmer, & Schrenk, 2007). different wear facet angles. If this hypothesis is instead accepted, we
Unlike microwear, dental macrowear is a cumulative process that can examine other possible causes that are beyond the determination
occurs throughout the individual's lifetime and is not influenced by of flat versus oblique wear facets. This information can help us to bet-
the “The Last Supper” effect (Grine, 1986; Janis, 1990). One of the ter understand the mechanisms and processes behind the formation of
major advantages of this approach is the use of high‐resolution 3D tooth wear in human populations and to evaluate whether occlusal
models of teeth, which allows detailed spatial measurements of the macrowear patterns contain information on culturally related changes
wear facet plane angles, thereby overcoming the limitations of 2D in the human diet.
analyses seen in Smith's work (1984). Moreover, our virtual approach
takes into account the functional aspects of tooth macrowear during
the sequential phases (power stroke) of the masticatory cycle (which 2 | MATERIAL AND METHODS
was not examined by Smith, 1984), providing more detailed
information on the attritional occlusal jaw movements and mecha- The sample used in this study consists of maxillary and mandibular
nisms behind the formation of each wear facet and thus permitting molars of prehistoric humans, modern hunter‐gatherers, and proto‐
the differentiation between dietary and nondietary movements and farmers. The specimens listed in Table 1 include fossil individuals from
wear (Fiorenza, 2015; Fiorenza, Benazzi, Kullmer, 2011; Fiorenza & the Middle and Upper Palaeolithic (PALAEO, n = 34), recent hunter‐
Kullmer, 2013, 2015, 2016). We have shown that relative facet areas gatherers (MHG, n = 15), and Natufians (FARM, n = 7) from the Levant
(obtained by dividing each facet area by the total occlusal wear area) who lived during the transition from hunting and gathering to early
of hunter‐gatherer molars differ between groups that consumed farming (Bar‐Yosef, 2011; Bowles & Choi, 2013; Table 1). The recent
different foods and are therefore a good indicator of diet (Fiorenza, sample consists of populations with known diets, including predomi-
Benazzi, Tausch, et al., 2011). Moreover, we have also demonstrated nantly meat‐eating groups such as Inuit (n = 4) and Vancouver islanders
that the formation and general wear facet pattern in Pleistocene and (n = 5), and also mixed‐diet hunter‐gatherers such as the Khoe‐San
modern humans are not influenced by tooth morphology (Fiorenza, (n = 6). The Inuit diet is almost entirely composed of animal proteins,
Benazzi, Viola, et al., 2011), corroborating previous works, which found including fish, sea mammals, terrestrial herbivores, birds, and, occa-
that differences in dental morphology do not generate differences in sionally, vegetable materials such as berries (Baarrengaard, 1949;
occlusal wear patterns if two mammalian taxa consume similar diets Gotfredsen, 1997; Pedersen, 1947; Pedersen & Scott, 1951). These
(Janis, 1990). populations often eat their food raw but sometimes frozen or dried
FIORENZA ET AL. 155

TABLE 1 List of Palaeolithic humans, modern hunter‐gatherers, and proto‐farmers used

Group N Specimen Locality Dating (ka)

PALAEO 34 Krapina 47, 48, 134, 135, 136, and 165 Croatia 130 ± 10
Krapina 166, 167, 169, 171, 175, and 177 Croatia 130 ± 10
Mountgaudier 5 France 130
Petit Puy 2 and 4b France 40–60
Combe Grenal 1 France 40–60
Kulna 1 Czech Republic 38–44
Vindija V206 and V259 Croatia 32–42
Le Moustier 1 France 41
Skhul 6 Israel 100–130
Qafzeh 3, 9, 11, 15, and 27 Israel 90–100
Oase 1 Romania 35
Les Rois France 32–35
Mladec 1 and 2 Czech Republic 31
Barma Grande 4 Italy 24,8
Le Placard France 22
Sungir 2 and 3 Russia 26.2–27.2
MHG 15 Inuit (n = 4) Greenland Recent
Vancouver Islanders (n = 5) Canada Recent
Khoe‐San (n = 6) South Africa Recent
FARM 7 Natufians Israel 11.5–15

Note. Groups: Palaeolithic individuals (PALAEO), modern hunter‐gatherers (MHG), and proto‐farmers (FARM). Numbers of MHG individuals are shown in
parentheses.

(Baarrengaard, 1949; Gotfredsen, 1997; Pedersen, 1947; Pedersen & phase (also called power stroke) where crushing and grinding of the
Scott, 1951). The subsistence economy of Vancouver islanders was food takes place (Kay & Hiiemae, 1974). During the first part of the
mainly based on the exploitation of various animal proteins but power stroke (Phase I), the opposing molar crests tend to slide past
dominated by various species of salmon, which were abundant and each other, and the trapped food is exposed to a shearing action
available at predictable times and places (Ames & Maschner, 1999). generated by forces parallel to the contact plane. At the end of Phase
They were consumed fresh with the remainder dried, smoked, and I, food is compressed (crushing) between basins and cusps of molars
stored. Finally, the Khoe‐San people were mostly gatherers rather than (the occlusal force is perpendicular to the contact plane) that are
hunters; their diets consisted predominantly of plant foods (60–80%), moving in centric occlusion (or maximum intercuspation). Phase II is
whereas meat was only of secondary dietary importance because usually an anterior‐medial movement, where the lower molars move
animals were scarce and their presence unpredictable (Lee, 1973, out of centric occlusion and the food is processed by grinding (which
1978; Lee & DeVore, 1976; O'Dea et al., 1991; Sahlins, 1968; is the resulting action of the combination of perpendicular and parallel
Silberbauer, 1981). forces to the contact plane). It is during the chewing cycle, after the
food is sufficiently softened and reduced in volume, that the surfaces
of the upper and lower teeth get into close occlusal contacts and wear
2.1 | Identification of wear facets facets start to develop.
The identification of occlusal wear facets is based on the terminology We further divided the Phase I facets into buccal and lingual pro-
proposed by Maier and Schneck (1981), who recognized 13 portions following the functional interpretations of Janis (1990) who
complementary pairs of facets in hominoid molars (Kullmer et al., analyzed the occlusal macrowear patterns in different mammalian
2009). We divided the facets into three groups based on their contri- groups. The distinction of wear facets into power stroke phases, and
bution to the power stroke phases of the chewing cycle: Buccal Phase the proportions of shearing, crushing, and grinding wear, is particularly
I facets (coloured in blue; Facets 1, 2, 3, and 4), Lingual Phase I facets important to understand the functional relationship between dental
(coloured in red; Facets 5, 6, 7, and 8), and Phase II facets (coloured wear and the physical properties of the foods consumed. This can be
in green; Facets 9, 10, 11, 12, and 13; Fiorenza, Benazzi, Tausch, useful for dietary interpretations as well (Fiorenza, Benazzi, Tausch,
et al., 2011; Janis, 1990; Kay & Hiiemae, 1974). All mammals chew et al., 2011). In addition, flat worn areas resulting from tip crushing
on one side of the jaw at a time and move the lower jaw in an orbital were also visible on the surface of the major cusp tips (Janis, 1990).
circuit (Janis, 1990). This orbital circuit consists of two distinct However, tip crushing facets were not considered here because their
functional steps: puncture‐crushing, where the food is initially pulped formation is still debated and probably cannot only be attributed to
and tooth‐to‐tooth contact rarely occurs, and a rhythmic chewing the normal power stroke of the chewing cycle.
156 FIORENZA ET AL.

Complementary occlusal wear facets tend to coalesce in advanced Finally, for each specimen, we calculated the average angle of the
wear stages, often preventing their clear identification and contouring. inclination of every phase of the power stroke.
Therefore, we selected only molars with a slight degree of wear based
on the amount of cusp removal and dentine exposure (Wear Stage 2;
2.3 | Statistical analyses
Smith, 1984). However, because most of our modern dental sample
is characterized by heavily worn teeth, we also included lower molars We calculated the median, standard deviation, and interquartile range

in an effort to increase the sample size. We initially tested whether of three variables (angles of Buccal Phase I, Lingual Phase I, and Phase

using lower and upper molars affects the obtained result. We also II wear facets) for each sample. Because the sample examined in this

compared the inclinations of first, second, and third molars to investi- study consists of few individuals, which prevent the assumption of a

gate whether there was any difference by dental element. To accom- normal distribution, we employed the multivariate and nonparametric

plish this, we used upper and lower molars from the same sample one‐way permutational multivariate analysis of variance test, which is

(Neanderthals from Krapina, Croatia). We also compared the inclina- a based on any distance measure between two or more groups. Statis-

tion of upper and lower molars in the modern hunter‐gatherer sample tical significance is computed with permutation test of group member-

(see Supporting Information). ship (n = 9,999). Finally, for the comparison of Facet 9 inclinations
between upper and lower molars, we used a Mann–Whitney univariate
test. The statistical analysis was accomplished with the software PAST
v.3.13 (PAlaeontological STatistics Hammer, Harper, & Ryan, 2001).
2.2 | Occlusal fingerprint analysis
Three‐dimensional computer models were produced by digitizing high‐
3 | RESULTS
quality dental casts (Fiorenza, Benazzi, & Kullmer, 2009) through a
white‐light scanning system with an x–y resolution of 45 μm
(smartSCAN 3D C‐5, Breuckmann GmbH). Postprocessing of the
3.1 | Analysis of the power stroke
polygonal models, together with the identification and analysis of wear Overall, Lingual Phase I facets show the greatest variation and steeper
facets, was carried out using Polyworks® V12 (InnovMetric software), inclination values than those of Buccal Phase I and Phase II areas
a 3D metrology software. The OFA method can be summarized as (Figure 2). This is further confirmed by the Mann–Whitney univariate
encompassing three sequential steps: use of a reference plane, facets test where a significant difference (p < .05) is found between Buccal
identification, and calculation of wear facet plane inclination and Lingual Phase I facets and between Phase II and Lingual Phase I
(Figure 1). facets (Table 2). When the wear patterns of the different groups are
The computation of a reference plane (Step 1) is based on the use considered, Palaeolithic humans are characterized by steep Buccal
of a reference Cartesian xy‐plane created along the cervical line of Phase I and Lingual Phase I wear facets and distinctively flat Phase II
each individual tooth by means of the least square best‐fit method wear facets (Table 3). On the other hand, the MHG sample displays
(Fiorenza et al., 2010; Kullmer et al., 2009; Ulhaas et al., 2007). Wear the lowest wear plane angles in both power stroke phases, whereas
facets were manually identified and segmented (Step 2) on the surface the Natufians have equivalent Buccal Phase I and Phase II wear facet
of the 3D models using the polyline tool of Polyworks® V12 inclinations, coupled with steep Lingual Phase I wear angles. The wear
(InnovMetric software). In order to create the plane of each facet, we plane angles of each group have been illustrated schematically using
selected all triangles contained in the facet perimeter and used the the transverse occlusal profile (Figure 3), where the power stroke
best‐fit plane function; the inclination (Step 3) is the angle measured phases are depicted by slopes that represent the average wear plane
between the facet and the reference plane (Kullmer et al., 2009). angles given in Table 3.

FIGURE 1 Occlusal fingerprint analysis method: Computation of a reference plane along the cervical line (Step 1), wear facet identification (Step 2)
based on Maier and Schneck (1981) labelling system, and inclination, measured between the reference and facet plane (Step 3). Buccal Phase I wear
facets (Facets 1, 1.1, 2, 2.1, 3, and 4); Lingual Phase I wear facets (Facets 5, 5.1, 6, 6.1, 7, and 8); Phase II wear facets (Facets 9, 10, 11, 12, and 13)
[Colour figure can be viewed at wileyonlinelibrary.com]
FIORENZA ET AL. 157

FIGURE 2 Boxplots illustrating the variation in the degree of wear

facet inclination within the different chewing phases in the entire
sample analyzed in this study (N = 56). Buccal Phase I wear facets
(Facets 1–4), Lingual Phase I wear facets (Facets 5–8), and Phase II
wear facets (Facets 9–13)

FIGURE 3 Occlusal profile. Digital 3D polygonal models of lower

TABLE 2 Between‐group comparisons of wear facet inclinations
molars in occlusal view showing the different wear facets grouped by
grouped according to chewing cycle phases (Kay & Hiiemae, 1974)
chewing cycle phases (Kay & Hiiemae, 1974): Buccal Phase I wear
Facets Buccal Phase I Lingual Phase I Phase II facets (Facets 1–4; blue), Phase II wear facets (Facets 9–13; green), and
Lingual Phase I wear facets (Facets 5–8; red). Occlusal profiles of each
Buccal Phase I ***
polygonal model is shown on the right; the slope (inclination) of the
Lingual Phase I 0.011 ***
different chewing cycle phases has been calculated using mean angle
Phase II 0.1989 0.0003 *** values (see Table 2). Groups: Palaeolithic individuals (PALAEO), modern
Note. Buccal Phase I facets (Facets 1–4), Lingual Phase I facets (Facets 5–8),
hunter‐gatherers (MHG), and Natufians (FARM). This figure is available
and Phase II facets (Facets 9–13). in colour online at wileyonlinelibrary.com/journal/oa [Colour figure
can be viewed at wileyonlinelibrary.com]
Mann–Whitney univariate test.
Significant p values (<.05) are highlighted in bold.
previous studies, the Palaeolithic group shows a significant more
oblique wear than those of MHG individuals (p = 0.0002), which it is
3.2 | Wear facet inclination and diet
comparable with those of early farmers (Table 6).
For testing the first null hypothesis, that there is no correlation
between the inclination of wear facets and diet, we analyzed each
MHG group individually (Table 4). Inuit molars are marked by
4 | DISCUSSION
extremely flat Buccal Phase I and Phase II facets, with relatively steep
Lingual Phase I facets. In contrast, Vancouver Islanders and Khoe‐San
are characterized by similar values in both chewing phase facets, with
4.1 | Dental macrowear in the Palaeolithic humans
the former possessing the flattest wear within the entire MHG sample It has been generally accepted that Palaeolithic humans had a much
analyzed in this study. A pairwise comparison between these three tougher and more fibrous diet than their modern counterparts (Molnar,
MHG groups did not show any statistical significance (Table 5). 1971, 1972). In particular, Smith (1984) found that Palaeolithic humans
As anticipated, if we compare the wear facet inclinations between tended to show flatter wear than modern hunter‐gatherers, displaying
modern hunter‐gatherers and early farmers, we find a statistical signif- the lowest plane angles of the entire sample analyzed in the study. This
icant difference between the two groups (p = .0314), with the former has suggested that the dentition of Palaeolithic humans underwent the
sample characterized by a flatter occlusal wear. However, contrary to heaviest demand on the masticatory apparatus. However, our study

TABLE 3 Explorative statistical analysis of wear facet inclinations with median, standard deviation (SD), and interquartile range (IQR)

Buccal Phase I Lingual Phase I Phase II

Group N Median SD IQR Median SD IQR Median SD IQR

PALAEO 34 27.1 5.6 8.8 31.4 6.9 11.1 22.5 7.3 9.6
MHG 15 20.1 5.7 7.5 21.1 7.4 15.0 22.3 4.2 5.9
FARM 7 26.5 6.6 9.3 31.2 10.8 14.0 26.7 8.4 14.2

Note. Group labels follow Table 1.

158 FIORENZA ET AL.

TABLE 4 Explorative statistical analysis of wear facet inclinations with median, standard deviation (SD), and interquartile range (IQR) within the
modern hunter‐gatherers (MHG) group
Buccal Phase I Lingual Phase I Phase II
Groups N Median SD IQR Median SD IQR Median SD IQR

INUIT 4 21.2 3.3 6.0 21.9 5.3 8.7 21.9 1.5 2.4
VANCOUV. 5 20.0 4.7 8.9 13.4 10.6 20.4 20.2 4.1 7.7
KHOE 6 19.8 3.2 5.2 22.1 5.3 8.2 24.6 5.6 6.8

Note. MHG Groups: INUIT (Inuit), VANCOUV. (Vancouver Islanders), and KHOE (Khoe‐San).

TABLE 5 Multivariate statistical analysis of wear facet inclinations in 2015). Moreover, although some scholars suggest that Neanderthals
the modern hunter‐gatherers (MHG) sample
extracted lipid sources directly from bone gnawing and chewing
Groups INUIT VANCOUV KHOE (Costamagno, 2013; Marean, 2005), we do not find microtrauma pat-
INUIT *** terns and large enamel chipping areas on Neanderthal postcanine
VANCOUV 0.3946 *** teeth that indicate this behaviour (Fiorenza et al., 2015). We therefore
KHOE 0.8329 0.5773 *** assume that the diets of Palaeolithic humans, as highlighted by others
(e.g., Kaifu, Kasai, Townsend, & Richards, 2003), probably were not as
Note. One‐way permutational multivariate analysis of variance test,
Permutation N = 9,999, P (same) = .7173. abrasive as those of Arctic hunter‐gatherers and Khoe‐San people.
Significantly p values (<.05) are highlighted in bold. This could be related to the food itself and/or to the use of less elab-
Group labels follow Table 4 orated food‐processing methods (such as desiccation, frozening, and
smoking) used by Palaeolithic humans, which probably introduced
lower amounts of dust and grit in their daily diet.
TABLE 6 Multivariate statistical analysis of wear facet inclinations Abrasion is the friction of exogenous materials forced over tooth
within the entire sample analyzed in this study
surfaces (Kaidonis, 2008; Kaifu et al., 2003). The action of food on a
Groups PALAEO MHG FARM tooth surface is not anatomically specific; the action generally occurs
PALAEO *** over the whole occlusal surface producing a wear area as opposed to
MHG 0.0002 *** a facet (Kaidonis, 2008). However, the abrasive action of food over
FARM 0.9187 0.0314 *** the enamel surface tends to remodel the primary tooth morphology,
reducing thus the slopes of the molar cusps, and therefore flattening
Note. One‐way permutational multivariate analysis of variance test,
Permutation N = 9,999, P (same) = .0002. the occlusal wear (Luke & Lucas, 1983).
Significantly p values (<.05) are highlighted in bold. The different results between our and Smith's (1984) study could
Group labels follow Table 1. be related to two main aspects: first, lower accuracy in measuring the
molar wear angles with 2D tools (which it becomes especially difficult
in heavily worn teeth), second, and more important, the terms used by
shows that the Palaeolithic individuals are characterized by the most
Smith (1984), such as “flat” versus “angled,” do not refer to the same
oblique wear, with significantly steeper wear facet planes than those
mechanisms analyzed in this study. Earlier works simply measured
of recent hunter‐gatherers. Several authors have argued that the
gross differences in cusp height between buccal and lingual sides of
severity of dental wear in Plio‐Pleistocene ancestors was not as exten-
the tooth as wear progressed to extreme, leaving a flattened tooth ver-
sive as in modern hunter‐gatherer people where tooth wear resulted
sus a tooth with markedly heavy wear on one side. The OFA approach
mainly from the excessive mixture of exogenous materials into their
takes place at a much finer scale than 1980s “wear plane angle” stud-
foods (Kaifu, 2000). In particular, Palaeolithic humans are characterized
ies. Here, we only considered wear facet angles associated to the
by steeper angles in Buccal and Lingual Phase I facets that are gener-
different phases of the masticatory cycle, independent from cusp
ally formed during the shearing phase of the chewing cycle (Kay &
morphology (e.g., Fiorenza, Benazzi, & Kullmer, 2011). The information
Hiiemae, 1974).
decoded in wear facet patterns is used to reconstruct major occlusal
Because we did not find any major differences between groups
movements and sequential tooth‐to‐tooth contacts between upper
that exploited different dietary sources, we need to take into account
and lower teeth, with the aim to show how contact areas are created
other possible factors, such as environmental abrasiveness and food
and what movements are responsible for their formation (e.g., Kullmer
processing methods, to explain why the Palaeolithic group is character-
et al., 2009).
ized by high wear angles. The analyses of cut marks, together with the
discovery of fractured animal bones, suggest intentional butchering
and marrow extraction (Mann, 2000). Moreover, although hearths
4.2 | Dental macrowear in modern hunter‐gatherers
and burned materials suggest controlled use of fire at many
Neanderthal sites (Lev, Kislev, & Bar‐Yosef, 2005), we have little
and proto‐farmers
archaeological evidence of other food processing techniques with the Smith (1984) hypothesized that a diet composed largely of ground
exception of possible evidence of heating of grains (Henry, Brooks, & grains and cooked (and therefore softer) foods had the effect of reduc-
Piperno, 2011; Mariotti Lippi, Foggi, Aranguren, Ronchitelli, & Revedin, ing food toughness (and hence leading to a reduction in perpendicular
FIORENZA ET AL. 159

masticatory forces) resulting in a more oblique wear. Furthermore, properties causing an increase in toughness (Molnar, 1972). All these
Smith (1984) noted that different hunter‐gatherer groups exhibited daily activities and food preparation methods employed by Inuit and
similar wear plane angles despite regional differences in diet. Our Vancouver Islander populations, besides changing food physical
results partially confirm these two main findings. For example, if we properties and increasing their toughness, seem to have also favoured
look at the wear facet plane angles on a broader scale, we can differen- the occasional ingestion of exogenous materials such as dust and grit,
tiate between traditional hunter‐gatherers (flat macrowear patterns) which may have significantly increased the amount of enamel loss.
and proto‐farmers such as the Natufians (more oblique wear). Although the Khoe‐San did not dry their meat nor cook it in earth
Although the Natufians are considered hunter‐gatherers who lived ovens (Lee, 1973, 1978; Lee & DeVore, 1976; O'Dea et al., 1991;
during the transitional period just before the Neolithic Revolution, they Sahlins, 1968; Silberbauer, 1981), their teeth still exhibit an extremely
were living in sedentary communities, cultivating wild plants, harvest- flat wear. Moreover, because they mostly inhabited an arid and sandy
ing cereals, and processing them with grinding stones. For this reason, environment, such as the Kalahari region, we should also take into
they have been considered farmers by many, with a lifestyle and tech- account the importance of grit and dust present in the environment.
nology significantly different from those of traditional hunter‐gatherer As highlighted by Kaidonis (2008), in hunter‐gatherers from desert,
communities (Bar‐Yosef, 2011; Bowles & Choi, 2013). The steeper the environment abrasion represents the most prevalent mechanism
wear facet planes of proto‐farmer lingual facets found in this study of wear. Thus, we can hypothesize that the flat wear patterns found
could indicate either a dietary difference with the modern hunter‐gath- in our Khoe‐San sample could be due to the combination of the masti-
erer group or the consumption of more refined and softened foods. cation of hard food items and foreign small particles unintentionally
The overall flat occlusal wear found in our MHG sample may have introduced in their mouth. Finally, Natufian people did not process
been caused by both intrinsic and extrinsic abrasiveness of their diet their foods as Arctic hunter‐gatherers did, and besides the introduction
and by different cultural habits. Inuit and Vancouver Islanders con- of grinding stones, their diet was significantly less abrasive than the
sumed foods low in abrasiveness such as meat and fish (Ames & diets consumed by the Inuit, Vancouver Islanders, and Khoe‐San
Maschner, 1999; Baarrengaard, 1949; El Zaatari, 2008, 2010; (Bar‐Yosef, 2011; Bowles & Choi, 2013).
Gotfredsen, 1997; Pedersen, 1947; Pedersen & Scott, 1951). Meat We need to stress here that, due to limitations in the method, we
itself, for instance, is not hard enough to scratch the enamel surface. could only use slightly worn teeth, preventing thus the analysis of
Differences in microwear signatures among populations with a diet larger comparative samples. Because hunter‐gatherers and prehistoric
mostly consisting of meat have been generally attributed to variations humans were characterized by extensive tooth wear, our approach can
in the amount of physical properties of exogenous abrasive particles, be limited to relatively young individuals and thus does not reflect the
either present in the environment or introduced through food process- overall pattern found in one specific population. Moreover, it would be
ing methods (El Zaatari, 2008; Kaidonis, 2008; Kaifu et al., 2003; Lucas interesting to investigate if there were any signals in dental wear pat-
et al., 2013; Luke & Lucas, 1983; Organ, Teaford, & Larsen, 2005; tern that could show a difference or a division of labour by age and
Teaford, Larsen, Pastor, & Noble, 2001). sex. Thus, future studies will necessitate the use of a larger sample size
Inuit populations, for example, consumed high amounts of raw, for better understanding the relationships between dental macrowear,
dried, or frozen meat (Baarrengaard, 1949; Gotfredsen, 1997; chewing behaviour, culture, and diet. Moreover, they should also focus
Pedersen, 1947; Pedersen & Scott, 1951). They also extracted marrow on potentially different patterns based on the stone‐tool technology
from crushing and chewing bones, a behaviour that put an enormous used and other functional aspects. For example, El Zaatari and Hublin
pressure on their teeth causing the formation of large enamel chipping (2014) found that changes in microwear texture signals of modern
areas (Turner II & Cadien, 1969). Raw meat requires a long chewing human teeth can be associated with major technological complexes
time before it can be reduced to smaller pieces and swallowed, overall of the Upper Paleolithic. More specifically, the analysis of the
increasing the masticatory effort (Zink & Lieberman, 2016; Zink, microwear texture showed significant differences between individuals
Lieberman, & Lucas, 2014). Because attritional wear facets are mostly from Aurignacian and Gravettian contexts relative to those of
produced by tooth‐to‐tooth contacts, prolonged mastication can Magdalenian culture, with the latter characterized by a more abrasive
increase the number of repetitive contact loadings between upper diet (El Zaatari & Hublin, 2014). Finally, it would be interesting as well,
and lower teeth, and thus, leading to an increased degree of wear to investigate how wear plane angles vary at different wear stages.
and flattening of the occlusal surface (Borrero‐Lopez, Pajares,
Constantino, & Lawn, 2014; Brace, 1977; Hylander, 1976; Lucas,
2004). This effect can be easily seen in modern human societies with ACKNOWLEDGEMENTS

the intake of soft foods in their diets, which significantly decrease The authors gratefully thank the curators and institutions which
the amount of time we chew. This is also reflected in our dentition, allowed us to study comparative and fossil specimens: Almut
which is characterized by very slightly worn teeth if compared with Hoffmann (Museum für Vor‐ und Frühgeschichte, Berlin, Germany),
hunter‐gatherer populations that relied on diets that required exten- Maria Teschler Nicola (Naturhistorisches Museum Wien, Vienna,
sive and powerful chewing (Dahl, Carlsson, & Ekfeldt, 1993; Kaifu, Austria), Marta Dočkalová (Moravské Zemské Muzeum, Anthropos
2000; Kaifu et al., 2003; Kullmer et al., 2009; Scott & Turner II, 1988). Institute, Brno, Czech Republic), Bence Viola (Department of
Similarly, Vancouver Islanders used to store large quantities of Anthropology, University of Vienna, Austria), Yoel Rak (Department
salmon in drying and smoking houses (Ames & Maschner, 1999). Meat of Anatomy and Anthropology, University of Tel Aviv, Israel), Fabio
(or fish) desiccation induces a significant change in its physical Parenti (Istituto Italiano di Paleontologia Umana, Rome, Italy), Chris
160 FIORENZA ET AL.

Stringer and Rob Kruszynski (Natural History Museum of London, Fiorenza, L. (2009). Occlusal wear pattern analysis of functional morphology in
England), National Museum of Wales (Cardiff, Wales), Angiolo del Neanderthals and early Homo sapiens dentition (Ph.D. dissertation). Frank-
furt: Johann Wolfgang Goethe University.
Lucchese (Museo Preistorico dei Balzi Rossi, Ventimiglia, Italy), and Erik
Fiorenza, L. (2015). Reconstructing diet and behaviour of Neanderthals
Trinkaus (Department of Physical Anthropology at Washington from Central Italy through dental macrowear analysis. Journal of Anthro-
University in Saint Louis, USA). We also would like to thank Christine pological Sciences, 93, 1–15.
Hemm for her help in surface scanning the original and cast dentition Fiorenza L, Benazzi S, Henry A, Salazar‐García DC, Blasco R, Picin A, Wroe
included in this sample and Iva Nikolic for copy‐editing our manuscript. S, Kullmer O. 2015. To meat or not to meat? New perspectives on
Neanderthal ecology. Yearbook of Physical Anthropology 156, Supple-
This study was supported by the Faculty of Medicine, Nursing and
ment S59: 43–71.
Health Sciences at Monash University through the Strategic Grant
Fiorenza, L., Benazzi, S., & Kullmer, O. (2009). Morphology, wear and 3D
Scheme 2016 (Grant SGS16‐0344) digital surface models: Materials and techniques to create high‐resolu-
tion replicas of teeth. Journal of Anthropological Sciences, 87, 211–218.
ORCID Fiorenza, L., Benazzi, S., & Kullmer, O. (2011). Para‐masticatory wear facets
and their functional significance in hunter‐gatherer maxillary molars.
Luca Fiorenza http://orcid.org/0000-0001-7110-3398 Journal of Archaeological Science, 38, 2182–2189.
Fiorenza, L., Benazzi, S., Tausch, J., Kullmer, O., Bromage, T. G., & Schrenk,
RE FE R ENC E S F. (2011). Molar macrowear reveals Neanderthal eco‐geographical die-
Addy, M., & Shellis, R. P. (2006). Interaction between attrition, abrasion and tary variation. PLoS One, 6, e14769.
erosion in tooth wear. Monographs in Oral Science, 20, 17–31. Fiorenza, L., Benazzi, S., Tausch, J., Kullmer, O., & Schrenk, F. (2010). Brief
Ames, K. M., & Maschner, D. G. (1999). Peoples of the Northwest Coast. Their communication: Identification reassessment of the isolated tooth
archaeology and prehistory. London: Thames and Hudson. Krapina D58 through occlusal fingerprint analysis. American Journal of
Physical Anthropology, 143, 306–312.
Baarrengaard, D. D. S. (1949). Dental conditions and nutrition among
natives in Greenland. Oral Surgery, Oral Medicine, Oral Pathology, 2, Fiorenza, L., Benazzi, S., Viola, B., Kullmer, O., & Schrenk, F. (2011). Rela-
995–1007. tionship between cusp size and occlusal wear pattern in Neanderthal
and Homo sapiens first maxillary molars. The Anatomical Record, 294,
Bar‐Yosef, O. (2011). The origins of agriculture: New data, new ideas. 453–461.
Current Anthropology, 52, S175–S193.
Fiorenza, L., & Kullmer, O. (2013). Dental wear and cultural behaviour in
Borrero‐Lopez, O., Pajares, A., Constantino, P. J., & Lawn, B. R. (2014). A middle Paleolithic humans from the near east. American Journal of
model for predicting wear rates in tooth enamel. Journal of the Physical Anthropology, 152, 107–117.
Mechanical Behavior of Biomedical Materials, 37, 226–234. Fiorenza, L., & Kullmer, O. (2015). Dental wear patterns in early modern
Bowles, S., & Choi, J. K. (2013). Coevolution of farming and private prop- humans from Skhul and Qafzeh: A response to Sarig and Tillier. Homo,
erty during the early Holocene. Proceedings of the National Academy of 66, 414–419.
Sciences of the United States of America, 110, 8830–8835. Fiorenza, L., & Kullmer, O. (2016). Occlusion in an adult male gorilla with a
Brace, C. L. (1977). Occlusion to the anthropological eye. In J. A. McNamara supernumerary maxillary premolar. International Journal of Primatology,
(Ed.), The biology of occlusal development ()Monograph No. 7, Craniofacial 37, 762–777.
Growth Series (pp. 179–209). Ann Arbor: Center for Human Growth and Gotfredsen, A. B. (1997). Sea bird exploitation on coastal Inuit sites, west
Development, the University of Michigan. and southeast Greenland. International Journal of Osteoarchaeology, 7,
Caspari, R., & Lee, S‐H. (2004). Older age becomes common late in human 271–286.
evolution. Proceedings of the National Academy of Sciences of the United Grine, F. E. (1986). Dental evidence for dietary differences in
States of America, 101, 10895–10900. Australopithecus and Paranthropus: A quantitative analysis of perma-
nent molar microwear. J. Hum. Evol, 15, 783–822.
Costamagno, S. (2013). Bone grease rendering in Mousterian contexts: the
case of Noisetier Cave (Fréchet‐Aure, Hautes‐Pyrénées, France). In J. L. Hammer, Ø., Harper, D. A. T., & Ryan, P. D. (2001). PAST: Paleontological
Clark, & J. D. Speth (Eds.), Zooarchaeology and modern human origins: Statistics software package for education and data analysis.
Human hunting behavior during the Later Pleistocene (pp. 209–225). Palaentologia Electronica, 4, 9.
New York: Springer. Harvati, K., Darlas, A., Bailey, S. E., Rein, T. R., el Zaatari, S., Fiorenza, L., …
Dahl, B., Carlsson, G. E., & Ekfeldt, A. (1993). Occlusal wear of teeth and Psathi, E. (2013). New Neanderthal remains from Mani peninsula, S.
restorative materials: A review of classification, etiology, mechanisms Greece: The Kalamakia Middle Palaeolithic cave site. J. Hum. Evol, 64,
of wear, and some aspects of restorative procedures. Acta Odontologica 486–499.
Scandinavica, 51, 299–311. Henry, A. G., Brooks, A. S., & Piperno, D. R. (2011). Microfossils in calculus
demonstrate consumption of plants and cooked foods in Neanderthal
El Zaatari, S. (2008). Occlusal molar microwear and the diets of the Ipiutak
diets (Shanidar III, Iraq; Spy I and II, Belgium). Proceedings of the National
and Tigara populations (Point Hope) with comparisons to the Aleut and
Academy of Sciences of the United States of America, 108, 486–491.
Arikara. Journal of Archaeological Science, 35, 2517–2522.
Hinton, R. (1982). Differences in interproximal and occlusal tooth wear
El Zaatari, S. (2010). Occlusal microwear texture analysis and the diets of
among prehistoric Tennessee Indians: Implications for masticatory func-
historical/prehistoric hunter‐gatherers. International Journal of
tion. American Journal of Physical Anthropology, 57, 103–115.
Osteoarchaeology, 20, 67–87.
Hylander, W. L. (1976). Morphological changes in human teeth and jaws in
El Zaatari, S., Grine, F. E., Ungar, P. S., & Hublin, J. J. (2011). Ecogeographis a high‐attrition environment. In A. A. Dahlberg, & T. M. Graber (Eds.),
variation in Neandertal dietary habits: Evidence from occlusal molar Orofacial growth and development (pp. 301–330). The Hague: Mouton.
microwear texture analysis. Journal of Human Evolution, 61, 411–424.
Janis, C. M. (1990). The correlation between diet and dental wear in herbiv-
El Zaatari, S., & Hublin, J. J. (2014). Diet of upper Paleolithic modern orous mammals, and its relationship to the determination of diets of
humans: Evidence from microwear texture analysis. American Journal extinct species. In A. J. Boucot (Ed.), Evolutionary Paleobiology of Behav-
of Physical Anthropology, 153, 570–581. ior and Coevolution (pp. 241–259). Amsterdam: Elsevier Science.
Every, R. G. (1972). A new terminology for mammalian teeth: Founded on the Kaidonis, J. A. (2008). Tooth wear: The view of the anthropologist. Clin.
phenomenon of thegosis. Parts 1 & 2. Pegasus: Christchurch. Oral. Invest. Supp. 1, 12, 21–26.
FIORENZA ET AL. 161

Kaidonis, J. A., Townsend, G. C., & Richards, L. C. (1993). Nature and fre- O'Dea, K., Jewell, P. A., Whiten, A., Altmann, S. A., Strickland, S. S., &
quency of dental wear facets in an Australian Aboriginal population. Oftedal, O. T. (1991). Traditional diet and food preferences of Austra-
Journal of Oral Rehabilitation, 20, 333–340. lian Aboriginal hunter‐gatherers. Phyl. Trans. Biol. Sci., 334, 233–241.
Kaifu, Y. (2000). Was extensive tooth wear normal in our ancestors?: A Organ, J. M., Teaford, M. F., & Larsen, C. S. (2005). Dietary inferences from
preliminary examination in the genus Homo. Anthropological Science, dental occlusal microwear at Mission San Luis de Apalachee. American
108, 471–385. Journal of Physical Anthropology, 128, 801–811.
Kaifu, Y., Kasai, K., Townsend, G. C., & Richards, L. C. (2003). Tooth wear Oxilia, G., Peresan, M., Romandini, M., Matteucci, C., Spiteri, C., Henry, A.,
and the “design” of the human dentition: A perspective from evolution- … Benazzi, S. (2015). Earliest evidence of dental treatment in the Late
ary medicine. Yearbook of Physical Anthropology, 46, 47–61. Upper Paleolithic. Scientific Reports, 5, 12150.
Kay, R. F. (1977). The evolution of molar occlusion in the Cercopithecidae Pedersen, P. O. (1947). Dental investigations of Greenland Eskimos.
and early catarrhines. American Journal of Physical Anthropology, 46, Proceedings of the Royal Society of Medicine, 40, 726–732.
327–352. Pedersen, P. O., & Scott, D. B. (1951). Replica studies of the surfaces of
Kay, R. F., & Hiiemae, K. M. (1974). Jaw movement and tooth use in recent teeth from Alaskan Eskimos. West Greenland natives and American
and fossil primates. American Journal of Physical Anthropology, 40, whites. Acta Odontologica Scandinavica, 9, 262–292.
227–256. Sahlins MD. 1968. Notes on the original affluent society. Man the hunter,
Kullmer, O., Benazzi, S., Fiorenza, L., Schulz, D., Bacso, S., & Winzen, O. RB Lee, I. De Vore (eds.). Aldine publishing company: Chicago, 85‐89.
(2009). Technical note: Occlusal fingerprint analysis: Quantification of Scott, R. G., & Turner, G. II (1988). Dental anthropology. Annual Review of
tooth wear pattern. American Journal of Physical Anthropology, 139, Anthropology, 17, 99–126.
600–605.
Silberbauer, G. (1981). Hunter and habitat in the Central Kalahari Desert.
Le Luyer, M., Rottier, S., & Bayle, P. (2014). Brief communication: Compar- Cambridge: Cambridge University Press.
ative patterns of enamel thickness topography and oblique molar wear
Smith, B. H. (1984). Patterns of molar wear in hunter‐gatherers and agricul-
in two Early Neolithic and medieval population samples. American
turists. American Journal of Physical Anthropology, 63, 39–56.
Journal of Physical Anthropology, 155, 162–172.
Teaford, M. F., Larsen, C. S., Pastor, R. F., & Noble, V. E. (2001). Pits and
Lee, R. B. (1973). Mongongo: Ethnography of a major wild food resource.
scratches. Microscopic evidence of tooth use and masticatory behav-
Ecology of Food and Nutrition, 2, 307–321.
iour in La Florida. In C. S. Larsen (Ed.), Bioarchaeology of La Florida:
Lee, R. B. (1978). Ecology of a contemporary san people. In P. Tobias (Ed.), Human biology in the northern frontier New Spain (Vol. 82) (pp. –112).
The bushmen: san hunters and herders of Southern Africa (pp. 98–114). Gainesville: University Press of Florida.
Cape town: Human and Rousseau.
Turner, C. G. II, & Cadien, J. D. (1969). Dental chipping in Aleuts, Eskimos
Lee, R. B., & DeVore, I. (1976). Kalahari Hunter‐Gatherers: Studies of the! and Indians. American Journal of Physical Anthropology, 31, 303–310.
Kung San and their neighbors. Cambridge: Harvard University Press.
Ulhaas, L., Kullmer, O., & Schrenk, F. (2007). Tooth wear diversity in early
Lev, E., Kislev, M. E., & Bar‐Yosef, O. (2005). Mousterian vegetal food in hominid molars: A case study. In S. E. Bailey, & J. J. Hublin (Eds.), Dental
Kebara Cave, Mt. Carmel. Journal of Archaeological Science, 32, perspectives on human evolution: State of the art research in dental paleo-
475–484. anthropology (pp. 369–390). The Netherlands: Springer.
Lucas, P. W. (2004). Dental functional morphology: How teeth work. Watson, J. T. (2008). Changes in food processing and occlusal dental wear
Cambridge, UK: Cambridge University Press. during the early agricultural period in northwest Mexico. American Jour-
Lucas, P. W., Omar, R., Al‐Fadhalah, K., Almusallam, A. S., Henry, A. G., nal of Physical Anthropology, 135, 92–99.
Michael, S., … Atkins, A. G. (2013). Mechanisms and causes of wear in Zink, K. D., & Lieberman, D. E. (2016). Impact of meat and Lower
tooth enamel: Implications for hominin diets. J. R. Soc. Interface, 10, Palaeolithic food processing techniques on chewing in humans. Nature,
20120923. 531, 500–505.
Luke, D. A., & Lucas, P. W. (1983). The significance of cusps. Journal of Oral Zink, K. D., Lieberman, D. E., & Lucas, P. W. (2014). Food material proper-
Rehabilitation, 10, 197–206. ties and early hominin processing techniques. Journal of Human
Maier, W., & Schneck, G. (1981). Konstruktionsmorphologische Evolution, 77, 155–166.
Untersuchungen am Gebiß der hominoiden Primaten. Z. Morph.
Anthrop., 72, 127–169.
Mann, N. (2000). Dietary lean red meat and human evolution. European SUPPOR TI NG INF ORMATI ON
Journal of Nutrition, 39, 71–79.
Additional Supporting Information may be found online in the
Marean, C. W. (2005). From the tropics to the colder climates: Contrasting
faunal exploitation adaptations of modern humans and Neanderthals. In supporting information tab for this article.
F. D'Errico, & L. Backwell (Eds.), From tools to symbols: From early hom-
inids to humans (pp. 333–371). Johannesburg: Witwatersrand
University Press.
Mariotti Lippi, M., Foggi, B., Aranguren, B., Ronchitelli, A., & Revedin, A. How to cite this article: Fiorenza L, Benazzi S, Oxilia G,
(2015). Multistep food plant processing at Grotta Paglicci (Southern Kullmer O. Functional relationship between dental macrowear
Italy) around 32,600 cal B.P. Proceedings of the National Academy of
and diet in Late Pleistocene and recent modern human popula-
Sciences of the United States of America, 112, 12075–12080.
tions. Int J Osteoarchaeol. 2018;28:153–161. https://doi.org/
Molnar, S. (1971). Human tooth wear, tooth function and cultural variabil-
ity. American Journal of Physical Anthropology, 34, 175–189. 10.1002/oa.2642
Molnar, S. (1972). Tooth wear and culture: A survey of tooth functions
among some prehistoric populations. Current Anthropology, 13,
511–526.