International Journal of Osteoarchaeology

Int. J. Osteoarchaeol. 20: 67–87 (2010)

Published online 23 December 2008 in Wiley InterScience
(www.interscience.wiley.com) DOI: 10.1002/oa.1027

Occlusal Microwear Texture

Analysis and the Diets of Historical/
Prehistoric Hunter-Gatherers
S. EL-ZAATARI*
Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology,
Deutscher platz 6, D-04103 Leipzig, Germany

ABSTRACT With the exception of few studies, occlusal microwear of pre-agricultural modern humans has
not been documented. In this study, microwear fabrics of samples from seven historic/
prehistoric hunter-gatherer populations with known and diverse dietary habits, representing
mostly meat-eaters from different environments, arctic/tundra (Tigara from Point Hope), cold-
steppe (Fuegians) and Mediterranean (Chumash), and mixed-diet hunter-gatherers from
tropical climates (Andamanese and Khoe-San from Matjes River, Riet River, and Oakhurst
Shelter), were analysed to better understand how dietary differences affect microwear in these
groups and to establish a reasonable comparative database for interpreting fossil hominins
microwear.
Significant microwear differences, related to diet and food preparation techniques, between
the meat-eaters and mixed-diet hunter-gatherers were detected. Finer scale differences
within each of these dietary categories were also observed. Ethnographic accounts indicate
that the Tigara and Andamanese ingested hard particles attached to their food as a result of
their food preparation techniques; their microwear fabrics also reflect highly abrasive diets.
On the other hand, as expected, the microwear signatures of the Chumash and Fuegians
indicate a diet low in abrasives, reflecting their almost exclusive reliance on marine meat for
subsistence and the low amounts of extraneous particles attached to this meat. The mixed-
diet Khoe-San occupy an intermediate position between the Tigara and Andamanese on the
one hand, and the Chumash and Fuegians on the other, with regard to the level of abrasives
ingested. The Khoe-San ate large amounts of hard plants, most likely responsible for abrading
their enamel surface. Copyright ß 2008 John Wiley & Sons, Ltd.

Key words: microwear; diet; hunter-gatherers; Tigara; Fuegians; Andamanese; Khoe-San;

Chumash

Introduction Teaford & Robinson, 1989; Teaford & Glander,

Numerous studies have demonstrated the poten-
tial of occlusal dental microwear to yield direct
evidence of tooth use and diet in a variety of
animal taxa (e.g. Walker, 1976; Ryan, 1981;
Gordon, 1982; Teaford & Walker, 1984; Teaford,
1985, 1986, 1993; Taylor & Hannam, 1987;
* Correspondence to: Department of Human Evolution, Max Planck
Institute for Evolutionary Anthropology, Deutscher platz 6,
D-04103 Leipzig, Germany.
e-mail: sireen.elzaatari@eva.mpg.de
1991; Solounias & Moelleken, 1992; Teaford &
O’Leary, 1992; Ungar, 1994, 1996; Walker et al.,
1994; Daegling & Grine, 1999; King et al., 1999a;
MacFadden et al., 1999; Lewisa et al., 2000;
Godfrey et al., 2004; Merceron et al., 2004, 2005;
Merceron & Ungar, 2005; El Zaatari et al., 2005;
Green et al., 2005; Hopley et al., 2006). Occlusal
dental microwear has also proven to be an
important tool for inference of the diets of fossil
hominins (e.g. Grine, 1981, 1986, 1987; Walker,
1981; Puech & Albertini, 1983, 1984; Puech et al.,
1983; Grine & Kay, 1988; Ryan & Johanson, 1989;

Copyright # 2008 John Wiley & Sons, Ltd. Received 7 January 2008
Revised 21 August 2008
Accepted 27 August 2008
68
Ungar & Grine, 1991; Teaford et al., 2002; Scott
et al., 2005; Grine et al., 2006; Ungar et al., 2006,
2008), as well as recent human groups from
archaeological deposits (e.g. Harman & Rose,
1988; Bullington, 1991; Molleson & Jones, 1991;
Teaford, 1991; Pastor, 1992; Molleson et al., 1993;
Hojo, 2001; Schmidt, 2001; Teaford et al., 2001;
Organ et al., 2005; Mahoney, 2006a,b; El Zaatari,
2008; Hogue & Melsheimer, 2008). However,
only a few studies have documented occlusal
microwear of pre-agricultural modern humans. In
addition, all these studies employed conventional
techniques of occlusal molar microwear analysis,
and they were aimed at either examining dietary
changes resulting from the transition to garden-
ing/farming (Molleson & Jones, 1991; Pastor,
1992; Schmidt, 2001; Mahoney, 2006b) and the
effects of the European contact (Teaford, 1991;
Teaford et al., 2001), or at documenting dietary
variability amongst distinct populations living in
the same geographical area (El Zaatari, 2008).
Thus, no study to date has compared the occlusal
molar microwear of various hunter-gatherer groups
with different dietary habits and living in a wide
range of environmental conditions. Also, no study
has done so using microwear texture analysis.
Microwear texture analysis is a new, automated
and repeatable approach to the study of dental
microwear and it has been presented recently by
Ungar et al. (2003). This new technique replaces
the scanning electron microscopy with scanning
confocal profilometry, and introduces scale-
sensitive fractal analysis as a tool for the analysis
of microwear features in three dimensions (Ungar
et al., 2003; Scott et al., 2005, 2006). Microwear
texture analysis has proven to be as effective as
conventional methods in separating extant and
fossil species with different dietary habits (Scott
et al., 2005, 2006; Ungar et al., 2007a,b).
Therefore, in this paper, this new technique is
used to compare the occlusal molar microwear
signatures of archaeological, historical and pre-
historic hunter-gatherer populations that were
living in different geographical locations and
environmental conditions, and had diverse
dietary habits for which good ethnographical
or historical data are available. The aim of this
study is to establish how differences in the diets of
these groups are reflected on the occlusal surfaces
of their molars. This study will also contribute to
Copyright # 2008 John Wiley & Sons, Ltd.
S. El-Zaatari
a comparative and interpretative database for the
future interpretation of fossil hominins’ occlusal
microwear signatures.
Microwear and diet
Studies of microwear analysis of extant animals
have shown that although variations in microwear
patterns can be attributed to various factors [e.g.
the characteristics of the abrasives ingested with
food (Teaford & Glander, 1991; Daegling &
Grine, 1999), enamel microstructure (Maas,
1991) and properties of jaw biomechanics
(Gordon, 1982)], the mechanical properties of
the diet itself are the primary cause of these
variations. For extant primates with known diets,
studies of molar microwear have shown that a diet
of harder objects, such as hard seeds and nuts,
results in heavily pitted, complex surfaces, a diet
of grasses or leaves results in heavily scratched,
more anisotropic surfaces, and a mixed diet, or a
diet consisting of foods with intermediate
properties, leads to intermediate microwear
patterns (Teaford & Walker, 1984; Scott et al.,
2006). Yet, studies of microwear in non-human
primates have also shown that the characteristics
of the abrasives ingested with the food can affect
the microwear signatures (e.g. Teaford & Glan-
der, 1991; Daegling & Grine, 1999). This is
similar to the case in carnivores where the
microwear pattern was found to be affected by
hard, non-meat items, such as bone, sometimes
chewed with the food (Van Valkenburgh et al.,
1990).
Examinations of molar microwear using con-
ventional microwear analysis techniques in
modern human populations, whose diet included
– or consisted mostly of – meat, have shown that
abrasives ingested with the food can play an
important role in shaping their microwear
patterns (e.g. Teaford et al., 2001; Organ et al.,
2005; El Zaatari, 2008). Since meat by itself is not
hard enough to markedly scratch the enamel
surface (Puech et al., 1981; Gordon, 1986; Teaford
& Runestad, 1992), it has been argued that, with
the lack of extraneous abrasive particles, the
microwear signature of meat-eating generally
results from the effects of tooth-on-tooth
abrasion during mastication, which is expected
Int. J. Osteoarchaeol. 20: 67–87 (2010)
Microwear Texture Analysis of Hunter-Gatherer Groups
to create large numbers of small pits and fine
scratches (Puech et al., 1981; Teaford & Runestad,
1992; Molleson et al., 1993; Teaford et al., 2001).
Therefore, differences in the microwear signa-
tures among modern human populations that ate
mostly meat have generally been linked to
variations in the amount and physical properties
of the extraneous abrasive particles usually
present in the environment and are attached to
the food as part of the food preparation
techniques (e.g. Teaford et al., 2001; Organ
et al., 2005).
This study compares the microwear signatures
of several hunter-gatherer human populations
from different environments with various levels of
meat-eating and different food preparation
techniques. Since the effects of meat-eating on
occlusal molar microwear are still not well
understood, this study is expected to shed more
light on the occlusal molar microwear signature
caused by meat-eating in the presence and
absence of extraneous abrasive particles.
Material
A total of 117 individuals representing seven
recent hunter-gatherer groups were analysed for
the purposes of this study. These groups
consisted of Andamanese (n ¼ 30), Chumash
(n ¼ 13), Fuegians (n ¼ 6), three groups of pre-
historic and historical (Khoe-San) hunter-gath-
erers from South Africa (Oakhurst Shelter, n ¼ 9;
Matjes River, n ¼ 14; and Riet River, n ¼ 20), and
Tigara from Point Hope, Alaska (n ¼ 25).
Andamanese
Study sample
The Great and Little Andamans, along with other
neighbouring islands, form a volcanic chain that
stretches between the province of Pegu and the
northernmost point of Sumatra in the Bay of
Bengal (Man, 1885). The sample of Andamanese
included in this study consists of 30 protohisto-
rical specimens dating to the period immediately
after the first permanent European settlement of
the islands in AD 1858 (Man, 1883, 1885). They
belonged to individuals who lived in the area of
Copyright # 2008 John Wiley & Sons, Ltd.
69
Port Blair, South Andaman. These dental remains
are currently housed at the British Museum of
Natural History, London.
Climate
The Andaman Islands have a tropical climate
where the temperature is very uniform through-
out the year (Man, 1885).
Dietary habits
Historical records indicate that the Andamanese
relied on marine hunting to provide fish, dugong
and sea turtles (Myka, 1993). A third of their daily
food consumption consisted of edible roots, fruits
and honey. They also consumed meat from pig
(most frequently hunted in the rainy season),
paradoxurus, iguana, turtle and shellfish (Man,
1883). Food was almost always cooked, but eaten
half-roasted, such that it remained relatively
tough (Pal, 1984). The food was usually cooked
uncovered, which would have enabled contami-
nation by windborne sand (Pal, 1984).
Chumash
Study sample
The skeletons comprising this collection (n ¼ 13)
were excavated in 1927 from the site of SCrI-3 on
the island of Santa Cruz, one of the four Northern
Channel Islands 20 miles off the coast of southern
California. This collection dates to between 4000
and 5000 years BP (Walker, 1986) and is currently
housed at the British Museum of Natural History,
London.
Climate
The climate of this area is predictable most of the
year. The island chain forms a sheltered channel
where moderate winds and air temperatures rarely
fluctuate (Fagan, 1995). The area has a Medi-
terranean climate characterised by cool and wet
winters, and warm and dry summers (Kenneth,
2005).
Dietary habits
The Chumash were fishers and hunters. Their
diet was based on fish, marine mammals and other
marine resources (e.g. shellfish). They also ate
Int. J. Osteoarchaeol. 20: 67–87 (2010)
70
small terrestrial animals available on the island
(Fagan, 1995). Isotopic studies show that the
Chumash relied heavily on marine protein
resources (Walker, 1996). Ethnohistorical
accounts also indicate that these people relied
on several kinds of plants including islay,
manzanite, mangle berries, tarweed seeds, tubers
and sage (Timbrook, 1993; Walker, 1996).
Fuegians
Study sample
Tierra del Fuego lies in the southernmost part of
South America (Garson, 1886). The area has an
Antarctic landscape with high mountains (Gar-
son, 1886). The skeletons included in this study
(n ¼ 6) date to AD 1880 and come from the
Yamana tribe (Manzi, 1986). This tribe occupied
the Beagle Channel islands and the islands of the
Chilean archipelago in the southwestern part of
Tierra del Fuego (Yesner et al., 2003). These
remains are currently housed in the Museum of
Anthropology at the University of Rome.
Climate
The Beagle Channel has an average summer
temperature of 108C. Average winter temperature
can reach
128C, but the constant winds
dramatically increase the cold. The southern
area of Tierra del Fuego is generally forested;
however, the temperate forests gradually give
way to shrub and tundra further south, the area
where the individuals sampled for this study lived.
Dietary habits
The ethnohistorical record of the late 1800s
indicates that the Yamana tribe of the Chilean
archipelago relied on fishing and seal-hunting,
although seals rank lower in importance (Snow,
1861; Bridges, 1885). Guanaco (Lama guanicoe)
may also have formed a terrestrial source of food
(Yesner et al., 2003). Stable isotope analysis of
Yamana skeletons indicates that their diet was
based on marine resources, but that terrestrial
animals also formed an important part of their
diet (Yesner et al., 2003).
Copyright # 2008 John Wiley & Sons, Ltd.
S. El-Zaatari
Prehistoric/historical hunter-gatherers
from Africa
Study sample
The molars included in this study are of hunter-
gatherers from South Africa. These specimens
derive from two prehistoric sites (Oakhurst
Shelter n ¼ 9 and Matjes River n ¼ 14) and one
historical site (Riet River n ¼ 20). The Oakhurst
Rock Shelter lies 13 miles east of George. It is
2 miles from the lakes that border the sea, and
4 miles away from the ocean. Excavations in the
site were undertaken in 1932–1935 by A.J.H.
Goodwin of the University of Cape Town. The
site is dated to between 9000 and 5000 BP (Morris,
1992a). Matjes River Rock Shelter is located on the
western side of the mouth of the Matjes River, east
of Robberg. The majority of the skeletal remains
from this site were excavated in the 1920s by
Dreyer (Sealy, 2006). Additional remains were
recovered in the 1950s by Hoffman and Meiring
(Sealy, 2006). The individuals from Matjes River
date to between 8000 and 2000 BP (Sealy &
Pfeiffer, 2000). The Riet River skeletons come
from the Orange River Valley, Northern Cape
province, and date to between the 11th and 19th
centuries AD (Morris, 1992b). The remains from
Oakhurst Shelter are currently housed at the
University of Cape Town, those from Matjes River
are housed at the National Museum of Bloemfon-
tein, and those from Riet River are housed at the
McGregor Museum in Kimberley.
Climate
The Oakhurst Shelter and Matjes River sites are
in a montane forest environment (Goodwin,
1938). This area is characterised by heavy rainfall
in a warm temperate climate. The terrestrial fauna
in the area consists mainly of small browsers,
although some medium-sized and large species
are also present. Riet River falls in an arid to semi-
arid climate characterised by hot summers and
cold and frosty winters.
Dietary habits
In these three sites shellfish were found indicating
some possibility of fishing activity (Goodwin,
1938; Sealy, 2006). Carbon-nitrogen isotope
analyses of specimens from Oakhurst Shelter
Int. J. Osteoarchaeol. 20: 67–87 (2010)
Microwear Texture Analysis of Hunter-Gatherer Groups
and Matjes River indicate that these individuals
had a mixed diet, but included a large amount of
marine resources (Sealy & van der Merwe, 1986,
1988; Sealy et al., 1992; Sealy & Pfeiffer, 2000;
Sealy, 2006). The Riet River inhabitants had a
mixed subsistence strategy; they were hunter-
gatherers and herders (Burchell, 1967). Stable
isotope analyses support this subsistence strategy
(Lee Thorp et al., 1993). Ethnographic studies
show that 80% or more of the diet of modern
Kalahari hunter-gatherers consists of plant food
(Silberbauer, 1981).
Tigara
Study sample
Point Hope, Alaska, is a peninsula formed by a
narrow ribbon of gravel and sand protruding
approximately 32 km into the Arctic Ocean some
200 km north of the Arctic Circle. It is the
westernmost point of land in Alaska north of the
Bering Strait. For the purposes of this study,
occlusal molar microwear data were collected
from a total of 25 individuals belonging to the
pre-contact Tigara population from Point Hope
dating to AD 1200–1700 and uncovered by
excavations between 1939 and 1941 (Larsen &
Rainey, 1948). The specimens are currently
housed at the American Museum of Natural
History.
Climate
The Arctic tundra vegetation of Point Hope is
characterised by the lack of trees and the
presence of moss, lichens, grass and small
flowering plants. In winter, the peninsula is
covered by hard-packed snow, although the
ocean currents leave an area of water close to the
shore free of pack ice, making Point Hope famous
for its great variety of sea mammals (Larsen &
Rainey, 1948).
Dietary habits
The material culture of the Tigara suggests that
their dietary habits were similar to those of the
Point Hope Eskimos who inhabited the area after
the European contact (Costa, 1980). The diets of
the Eskimos living at Point Hope in the 20th
Copyright # 2008 John Wiley & Sons, Ltd.
71
century consisted of 35–60% fat, 35–65% protein
and very little carbohydrate (Waugh, 1930). The
meat component of their diet came mainly from
fish, eaten fresh or dried on open racks; whale,
eaten fresh or kept frozen underground as a year-
round staple; and seal, often eaten raw at the site
of the catch or kept frozen (de Poncins, 1941;
Giddings, 1967). It should be mentioned that
Point Hope consists of sandy beaches, and the
freezing of meat on open racks or its storage
underground would have most likely caused sand
particles to be integrated with the meat.
Uncooked seal skin together with its attached
subcutaneous fat was often chewed for prolonged
periods of time (Balikci, 1970).
Methods
Specimen preparation
For the purposes of this study, moulds of
permanent molar tooth crowns of the fossil and
modern human groups were prepared (by SEZ)
following established procedures (Teaford &
Oyen, 1989b) at the institutions where the
specimens are housed. Specimens were first
cleaned with cotton swabs soaked with distilled
water. Acetone and/or ethyl alcohol were used
only when the occlusal surface was covered with
preservatives (e.g. Glyptal) that could not be
removed with water. Impressions were then made
using Coltene President MicroSystemTM (Col-
tène1) (polysiloxane vinyl) impression material,
and casts were made with Epo-Tek 301 epoxy
resin and hardener (Epoxy Technologies). This
procedure allows for a reproduction of features
with a resolution of a fraction of a micron
(Teaford & Oyen, 1989b).
Specimen examination
Dental microwear texture analysis, a combination
of confocal microscopy and scale-sensitive fractal
analysis, was used in this study for the analysis of
dental microwear. Specimens were examined
using a Sensofar Plm Confocal Imaging Profiler
(Solarius Development Inc., Sunnyvale, CA).
Low magnification (10) was first used to scan
Int. J. Osteoarchaeol. 20: 67–87 (2010)
72
Figure 1. Examples of artefacts and actual wear surfaces.
(A) and (B) Surfaces covered with glue taken at 10 and
100 magnification respectively. (C) Surface covered
with bubbles as a result of poor casting (taken at 10).
(D) Artefacts resulting from a sharp tool having scratched
the surface during specimen preparation (20). (E) Sur-
face covered with dirt (100). (F) and (G) ‘Good’ surfaces
(100).
the entire occlusal surface and to eliminate teeth
that showed obvious post mortem damage and
which lacked good microwear preservation. Post
mortem damage, which can be caused either by
natural causes or during collection and prep-
aration of the specimens, is generally easily
detected. Naturally caused post mortem wear (e.g.
sediment abrasion) will be represented on all
surfaces of the tooth and not only the occlusal
surface (Grine, 1986; Teaford, 1988; King et al.,
1999b), while preparation damage is fairly
distinctive (Teaford, 1988). Artefact fields caused
by glue, dirt and bubbles stuck to the tooth
surface during the moulding or casting processes
are easy to identify at low magnification.
Copyright # 2008 John Wiley & Sons, Ltd.
S. El-Zaatari
Examples of post mortem damage identified on
some teeth are presented in Figure 1.
Each individual was represented by one molar
(M1 or M2) with the best microwear preser-
vation. Both upper and lower molars were
included. Using conventional techniques of
dental microwear, some studies have suggested
that the molar microwear signature might differ
along the tooth row in the same individual
(Gordon, 1982; Mahoney, 2006a). However, the
results of these studies do not show consistent
differences in microwear along the tooth row, and
thus are not conclusive. Yet, because this current
study includes both first and second molars,
possible differences in the microwear signature
between the two molars for each population were
tested before comparisons among groups were
conducted.
Analysis was restricted to crushing/grinding
facets (i.e. facets 9, 10n, or x) to control for
interfacet variation (Gordon & Walker, 1983).
Following Scott et al. (2005, 2006), scans of the
crushing/grinding facets of specimens with good
wear surfaces were taken at a magnification of
100 with a lateral sampling interval of 0.18 mm
and a vertical resolution of 0.005 mm. At this
magnification, each scan covers an area of
138  102 mm. Four adjoining scans were taken
for each surface, resulting in a total area of
276  204 mm. Microscopy was conducted at the
University of Arkansas at Fayetteville.
Using the Solarmap Universal software (Solar-
ius Development Inc., Sunnyvale, CA), scans
were then levelled, modified (to erase defects),
and photostimulations and 3D images were
generated. This program also allows for the
elimination of some identifiable small defects
from the analysis, if necessary, either by first
tracing their boundaries or by applying the
thresholding option and then the ‘erase defect
operator’ in Solarmap. Such defects can be dust
particles or casting bubbles that are present on
otherwise ‘good’ surfaces.
The resulting data were analysed in Toothfrax
and SFrax (Surfract, www.surfract.com) using
scale-sensitive fractal analysis. Scale-sensitive
fractal analysis follows the principles of fractal
geometry, which has demonstrated that the
appearances of surfaces differ with the scale of
observation. Following Scott et al. (2006), five
Int. J. Osteoarchaeol. 20: 67–87 (2010)
Microwear Texture Analysis of Hunter-Gatherer Groups
variables were considered: surface complexity,
scale of maximum complexity, anisotropy,
heterogeneity, and texture fill volume.
Complexity
Based on the principles of scale-sensitive fractal
analysis, the area of a rough surface changes with
the scale of observation. Thus, at sufficiently
coarse scales, surface textures would appear
smooth, but these same surfaces would appear
rough with increasing resolution at increasingly
finer scales. Each surface has a fixed planometric
area, which is the two-dimensional area that it
occupies. In addition, each surface has a relative
area for each scale of observation. Relative areas
are calculated by the area-scale tiling algorithm,
that is, by adding all the areas of triangular tiles of
a particular scale and dividing this sum by the
planometric area of the surface. For each scan,
relative areas were calculated for scales ranging
from 7200 to 0.02 mm2. The area-scale fractal
complexity (Asfc) is the slope of the steepest part
of the curve fit to the log-log plot of relative area
over the range of scales multiplied by
1000.
Complexity is characterised by changes in
relative area with the scale of measurement.
More complex surfaces have microwear features
of different sizes overlying each other (Figure 2).
Figure 2. Examples of surfaces with relatively high
(A) and low (B) complexity. Highly complex surfaces
(A) are surfaces that have many microwear features over-
lying each other. The opposite is true for surfaces of low
complexity (B). This figure is available in colour online at
www.interscience.wiley.com/journal/oa.
Copyright # 2008 John Wiley & Sons, Ltd.
73
Figure 3. Examples of microwear surfaces with relatively
high (A) and low (B) anisotropy. Surfaces with high aniso-
tropy (A) are surfaces with many parallel scratches com-
pared with low anisotropic surfaces (B). This figure is
available in colour online at www.interscience.wiley.
com/journal/oa.
Median Asfc values were calculated from the four
scans of each individual.
Scale of maximum complexity
Scale of maximum complexity (Smc) is the scale at
which the surface is most complex. Median Smc
values from the four scans of each specimen were
calculated.
Anisotropy
Surfaces dominated by many parallel striations
tend to be more anisotropic (Figure 3). Each
profile has a fixed projected length, but it also has
relative lengths for different scales of observation
and different orientations. Relative lengths are
calculated by the length-scale tiling algorithm,
which is the sum of line segments of a given scale
fitted to a profile divided by the projected length
of the same profile. Relative length measures are
taken at different orientations for a given scale of
observation. These lengths are then defined as
vectors and calculated at 58 intervals for a total of
36 measurements at each scale. Vectors are then
normalised using the exact proportion method by
dividing them by the sum of relative lengths from
all orientations. This is done to eliminate the
Int. J. Osteoarchaeol. 20: 67–87 (2010)
74
effect of pits and scratch depth on the anisotropy
graph. The normalised vectors are displayed
graphically in a rosette diagram. The exact
proportion Length-scale anisotropy of relief
(epLsar) is the length of the mean vector at the
scale of 1.8 mm. This is the finest scale for which
epLsar could be determined, given the need to
balance scale with number of pixels available at all
orientations to calculate robust relative length
values. Median epLsar values were calculated from
the four scans for each specimen.
Heterogeneity
Heterogeneous surfaces will differ in texture from
place to place across the surface. Heterogeneity
of Area-scale fractal complexity (HAsfc) is calcu-
lated by dividing the median absolute deviation
(the absolute distance between each observation
and the median) of Asfc by the median Asfc for the
four scans for each specimen.
Texture fill volume
The volume-filling versus scale algorithm is used
to calculate the volume of a surface by filling it
with cuboid prisms at different scales. Different
scales can be obtained by changing the dimen-
sions of the ‘prism’ faces. The distance from the
base, where it rests on surface relief, to the
highest vertical elevation on the surface forms
the rectangular dimension of each cube. With
reduction in scale, more cubes fit onto and into
the surface, thus filling a greater volume.
The total volume that is filled changes as a result
of variations in two components: the shape of the
surface and the texture of the surface. Structural fill
volume results from the gross shape of a surface (i.e.
it is the fill volume at a sufficiently coarse scale).
Texture fill volume, however, is a result of the
microwear surface. The textural fill volume was
calculated as the total fill volume (generated at a
scale of 2 mm) minus the structural fill volume (at a
scale of 10 mm). Median textural fill volume of the
four scans of each individual was calculated.
Statistical analyses
Differences in surface complexity, scale of
maximum complexity, anisotropy, heterogeneity
Copyright # 2008 John Wiley & Sons, Ltd.
S. El-Zaatari
and textural fill volume within each group (i.e.
between the M1s and M2s in each group) and
among the groups were assessed statistically.
Data were rank-transformed before analysis to
reduce the possible effects of violating assump-
tions associated with parametric statistical tests
(Conover & Iman, 1981). Data for the five
variables were compared between the M1 and M2
of each group and among the different groups
using a multivariate analysis of variance model
(MANOVA) (Neff & Marcus, 1980). Single
classifications of variance ANOVAs on each
variable along with the multiple comparisons
tests were used to determine the sources of
significant variation (Sokal & Rohlf, 1995). Both
Fisher’s least significant difference (LSD) and
Tukey’s honestly significant difference (HSD) post
hoc tests were used to balance Type I and Type II
errors (Cook & Farewell, 1996).
Results
Comparisons between M1 and M2 of
each group
Figure 4 and Table 1 present the means of the five
microwear variables for both M1 and M2 in each
group. The microwear signatures of the M1s and
M2s in each group were first compared using
MANOVAs conducted on the five variables for
each group separately. These statistical compari-
sons indicate no significant differences in the
microwear signatures between these two molars
in any of the groups analysed. Thus, it is possible
to combine data from both teeth in each
sample for the purposes of among-groups
comparisons.
Comparisons among groups
Comparative surface scans of the groups are
presented in Figure 5. Five parameters were
considered in this analysis: Asfc, Smc, epLsar, HAsfc
and Tfv. The results of the MANOVA statistical
tests indicate significant differences among the
groups considered. Individual ANOVAs show
significant differences in all variables except
HAsfc. Therefore, HAsfc will not be considered
Int. J. Osteoarchaeol. 20: 67–87 (2010)
 hunter-gatherers
www.interscience.wiley.com/journal/oa.

Comparisons among the mixed diet

Copyright # 2008 John Wiley & Sons, Ltd.

both Fisher’s LSD and Tukey’s HSD tests.
Figure 4. Means for M1 and M2 for the five variables for

individual. This figure is available in colour online at

Oakhurst Shelter M1 sample also represents only one

Shelter and the historic site of Riet River. The

sampled for African (Khoe-San) aborigines and
further here. In the majority of cases, significant
sample represents only one individual, and that for the
each group: (A) Asfc, (B) Smc, (C) epLsar, (D) HAsfc, and
(E) Tfv. Note that the value presented for the Fuegian M2

The Khoe-San samples included individuals from

Molars of mixed diet hunter-gatherers were
differences amongst the groups are detected by

the prehistoric sites of Matjes River and Oakhurst

individuals who lived in the Andaman Islands.
Microwear Texture Analysis of Hunter-Gatherer Groups

Table 1. Descriptive statistics for Asfc, Smc, epLsar, Tfv and HAsfc for the M1 and M2 of each group

Tigara Andamanese Matjes River Oakhurst Shelter Riet River Chumash Fuegians

M1 M2 M1 M2 M1 M2 M1 M2 M1 M2 M1 M2 M1 M2

n 11 14 24 6 10 4 1 8 7 13 8 5 5 1
Asfc
Mean 5.656 7.286 6.383 6.193 3.258 4.354 3.529 3.550 3.609 3.715 2.703 2.920 0.980 0.788
SD 5.312 6.268 4.450 3.477 2.104 4.077 — 1.711 1.995 2.830 2.387 2.543 0.314 —
Smc
Mean 0.193 0.229 0.214 0.210 0.197 0.208 0.150 0.179 0.192 0.202 0.216 0.150 0.426 0.266
SD 0.053 0.082 0.240 0.146 0.053 0.048 — 0.054 0.044 0.062 0.058 0.001 0.132 —
epLsar
Mean 0.0032 0.0029 0.0025 0.0027 0.0030 0.0030 0.0011 0.0021 0.0025 0.0025 0.0024 0.0021 0.0048 0.0025
SD 0.0019 0.0011 0.0011 0.0014 0.0013 0.0015 — 0.0010 0.0013 0.0014 0.0007 0.0009 0.0011 —
HAsfc
Mean 0.277 0.284 0.222 0.155 0.211 0.160 0.327 0.325 0.192 0.206 0.150 0.257 0.100 0.148
SD 0.179 0.223 0.144 0.079 0.120 0.050 — 0.222 0.131 0.089 0.078 0.157 0.021 —
Tfv
Mean 11479.1 12252.5 10168.5 10789.9 9776.2 12569.1 4132.5 9914.4 11528.1 11938.7 6725.6 6492.4 5129.7 5700.4
SD 4797.1 4695.0 4470.1 6194.2 4070.7 2570.3 — 6132.1 5202.7 4167.5 2659.1 4260.1 3929.7 —

Int. J. Osteoarchaeol. 20: 67–87 (2010)

75
76 S. El-Zaatari

Table 2. Descriptive statistics for Asfc, Smc, epLsar,

HAsfc and Tfv for the Khoe-San groups

Matjes River Oakhurst Shelter Riet River

n 14 9 20
Asfc
Mean 3.571 3.548 3.678
SD 2.677 1.601 2.514
Smc
Mean 0.200 0.176 0.198
SD 0.050 0.051 0.056
epLsar
Mean 0.0030 0.0020 0.0025
SD 0.0013 0.0010 0.001
HAsfc
Mean 0.198 0.325 0.201
SD 0.105 0.208 0.103
Tfv
Mean 10574.2 9272.0 11795.0
SD 3835.5 6051.2 4422.4

resulted from a high degree of variability in their

diets. Therefore, specimens from these four sites
were treated as a single sample for the purposes of
further comparisons.

Andamanese and Khoe-San

Figure 5. Representative micrographs of the groups ana-
lysed. (A) Fuegians; (B) Chumash; (C) Khoe-San;
(D) Tigara; and (E) Andamanese. This figure is available
in colour online at www.interscience.wiley.com/journal/oa.
microwear signatures of these three Khoe-San
samples were first compared. Following this,
differences and similarities in the microwear
signatures between the Khoe-San and the
Andamanese were assessed.
The microwear signature of the Andaman
Islanders differs from those of the Khoe-San
groups in that the former has significantly higher
Asfc (Figure 6; Tables 4–5). Fisher’s LSD also
detected significant difference in Smc between the
two groups, where the Andamanese were found
to have a significantly higher mean value
compared with the Khoe-San. However, because
of the large overlap in the Smc ranges of variation
of these two groups, this significant difference
could be an artefact of the wide range of variation
in the Andamanese sample and the narrow range
of variation in the Khoe-San sample for this
variable. No significant differences were found in
Tfv and epLsar values between the two mixed diet
hunter-gatherer groups (Figure 6; Tables 4–5).

The different Khoe-San groups Comparisons among mainly meat-eating

No significant differences in the microwear
signatures of individuals from Matjes River, Riet
River and Oakhurst Shelter were recorded,
although these samples came from different
environments (Tables 2–3). This probably
hunter-gatherers
Dental samples of three groups of mostly meat-
eating hunter-gatherers are included in this
analysis: Tigara, Chumash and Fuegians.

Copyright # 2008 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. 20: 67–87 (2010)
Microwear Texture Analysis of Hunter-Gatherer Groups 77

Table 3. Results of statistical analysis for the Khoe-San samples

A. MANOVA results

F Hypothesis df Error df P

Wilks’ Lambda 0.814 0.782 10 72.000 0.646

Pillai’s Trace 0.191 0.782 10 74.000 0.646
Hotelling’s Trace 0.223 0.781 10 70.000 0.647

B. Individual ANOVAS

Sum of squares df Mean square F P

Asfc
Effect 99.971 2 49.985 0.547 0.653
Error 6521.529 40 163.038
Smc
Effect 335.983 2 167.992 1.510 0.226
Error 5742.517 40 143.563
epLsar
Effect 518.611 2 259.305 1.510 0.226
Error 6077.389 40 151.935
HAsfc
Effect 428.561 2 214.281 2.101 0.115
Error 6193.439 40 154.836
Tfv
Effect 137.716 2 68.858 0.521 0.670
Error 6484.284 40 162.107

The Fuegians had significantly higher epLsar
and Smc values compared with those of the
Chumash. Also, Fisher’s LSD suggests that the
Fuegians had significantly lower Asfc values
compared with those of the Chumash
(Figure 6; Tables 4–5). Both the Fuegian and
Chumash values for Asfc and Tfv were signifi-
cantly lower than that of the Tigara. Compared
with the Smc value for the Tigara, that of the
Fuegians was significantly higher. The Chumash
had Smc values not significantly different from
those of the Tigara. The mean epLsar values of the
Chumash did not differ significantly from those
of the Tigara, but Fisher’s LSD suggests that the
Fuegians had significantly higher epLsar values
compared with those of the Tigara (Figure 6;
Tables 4–5).
in having significantly lower epLsar and Smc values.
Fisher’s LSD suggests that the Andamanese had
significantly higher Tfv compared with both the
Fuegians and Chumash. The Andamanese dif-
fered from the Tigara in having significantly
lower mean Smc. This, however, might be an
artefact of the wide range of variation in the
Andamanese sample and the narrow range of
variation in the Tigara sample for this variable.
The Khoe-San had significantly higher Asfc and
Tfv, but lower Smc and epLsar values compared
with the Fuegians. They also had a significantly
higher mean Tfv value compared with the
Chumash. Compared with the Tigara, the
Khoe-San probably had a significantly lower Asfc
as detected by Fisher’s LSD test but not Tukey’s
HSD test (Figure 6; Tables 4–5).

Mainly meat-eating vs. mixed diet Discussion

hunter-gatherers
The microwear signature of the Andamanese
differed from that of the mostly meat-eating
Fuegians and Chumash in having a significantly
higher mean Asfc value (Figure 6; Tables 4–5).
The Andamanese also differed from the Fuegians
Few studies have documented the occlusal molar
microwear of pre-agricultural modern humans
using conventional techniques (Teaford, 1991;
Molleson & Jones, 1991; Pastor, 1992; Schmidt,
2001; Teaford et al., 2001; Mahoney, 2006b; El
Zaatari, 2008). Yet, no study has examined the

Copyright # 2008 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. 20: 67–87 (2010)
78 S. El-Zaatari

Figure 6. Summary statistics for the groups analysed. (A) Asfc; (B) Smc; (C) epLsar; and (D) Tfv. Circles ¼ sample
means; vertical lines indicate 1 SD of the mean.

Table 4. Descriptive statistics for Asfc, Smc, epLsar, HAsfc and Tfv for modern groups

Tigara Andamanese Khoe-San Chumash Fuegians

n 25 30 43 13 6
Asfc
Mean 6.569 6.345 3.616 2.787 0.948
SD 5.807 4.219 2.360 2.344 0.291
Smc
Mean 0.213 0.213 0.194 0.190 0.400
SD 0.072 0.222 0.053 0.055 0.135
epLsar
Mean 0.0029 0.0026 0.0026 0.0023 0.0044
SD 0.0015 0.0012 0.0013 0.0007 0.0014
HAsfc
Mean 0.267 0.208 0.226 0.191 0.109
SD 0.201 0.135 0.138 0.121 0.027
Tfv
Mean 11912.2 10292.8 10869.5 6635.9 5224.8
SD 4656.7 4746.3 4621.6 3191.8 3522.5

Copyright # 2008 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. 20: 67–87 (2010)
Microwear Texture Analysis of Hunter-Gatherer Groups 79

Table 5. Results of statistical analysis for the modern samples

A. MANOVA results

F Hypothesis df Error df P

Wilks’ Lambda 0.518 3.941 20 359.145 0.00

Pillai’s Trace 0.586 3.807 20 444.000 0.00
Hotelling’s Trace 0.745 3.968 20 426.000 0.00
B. Individual ANOVAS

Sum of squares df Mean square F P

Asfc
Effect 35161.358 4 8790.34 10.02 0.000
Error 98295.642 112 877.64
Smc
Effect 23892.032 4 5973.008 6.98 0.000
Error 95794.968 112 855.312
epLsar
Effect 11706.268 4 2926.567 2.7 0.034
Error 121415.732 112 1084.069
HAsfc
Effect 7533.863 4 1883.466 1.67 0.161
Error 125917.637 112 1124.265
Tfv
Effect 22826.077 4 5706.519 5.78 0.000
Error 110631.923 112 987.785

C. Multiple comparisons tests (matrices of pairwise mean differences)

Tigara Andamanese Khoe-San Chumash

Andamanese
Asfc 4.7
Smc 25.6z
epLsar 10.0
Tfv 14.3
Khoe-San
Asfc 17.6y 22.3z
Smc 9.7 15.9y
epLsar 8.7 1.3
Tfv 9.2 5.1
Chumash
Asfc 32.2z 36.8z 14.5
Smc 11.1 14.5 1.4
epLsar 14.3 4.3 5.6
Tfv 41.3z 27.0y 32.1z
Fuegians
Asfc 64.6z 69.3z 47.0z 32.4y
Smc 38.5z 64.1z 48.2z 49.6z
epLsar 33.0y 43.0z 41.7z 47.3z
Tfv 48.8z 34.5y 39.6z 7.5
y
Fisher’s LSD test, p  0.05.
z
Fisher’s LSD and Tukey’s HSD test, p  0.05.

occlusal microwear of a wide range of recent
hunter-gatherer groups. Thus, the interpretation
of occlusal microwear patterns in these groups is
not well understood.
Using microwear texture analysis, this study
examines the occlusal molar microwear of a
combination of first and second molars in seven
(including three Khoe-San groups) different
hunter-gatherer populations. Studies looking at
possible variations in microwear as a result of
molar position in the tooth row are limited
(Gordon, 1982; Mahoney, 2006a). Also, these

Copyright # 2008 John Wiley & Sons, Ltd. Int. J. Osteoarchaeol. 20: 67–87 (2010)
80
studies do not provide consistent results. In a
sample of chimpanzees, Gordon (1982) showed
that, although the microwear signature of the
third molars differs significantly from those of the
first and second molars, those of the first and
second molars do not differ significantly from one
another. Based on preliminary results, Mahoney
(2006a) suggested that there are significant
differences in some microwear variables between
M1 and M2 in a Bronze–Iron Age human sample
from the Jordan Valley.
The analysis of the microwear patterns of both
M1 and M2 from the groups sampled in this
current study reveals no significant differences in
the microwear signatures between these teeth.
Since this study examines samples from seven
different hunter-gatherer populations, it can be
argued that first and second molars provide
similar microwear patterns, at least in hunter-
gatherers. This result is valuable for the analysis
of microwear in fossil hominins where combining
both molars is standard practice in an effort to
increase sample sizes for the fossil groups (e.g.
Grine et al., 2006; Ungar et al., 2006).
The five hunter-gatherer groups included in
this study had different dietary habits, came from
different geographical regions, and lived under a
variety of climatic conditions, ranging from
Arctic to tropical environs. Based on their dietary
habits, these groups can be divided into two
major categories: (1) mainly meat eaters (Tigara,
Chumash and Fuegians); and (2) mixed diet
hunter-gatherers (Andamanese and Khoe-San).
The group that consumed mostly meat can be
further divided into three categories based on
habitat: (1) those from Arctic environments
(Tigara); (2) those from cold-steppe environ-
ments (Fuegians); and (3) those from a Medi-
terranean climate (Chumash). Differences in the
dietary habits of these groups were clearly
reflected in their dental microwear patterns.
Significant microwear differences between the
principally meat-eating and the mixed diet
hunter-gatherers were detected, and finer-scale
divisions within each of these dietary categories
were also observed.
The mostly meat-eating hunter-gatherer group
from the Arctic environments of Point Hope, the
Tigara, has the most complex surfaces among the
groups analysed. High levels of surface complex-
Copyright # 2008 John Wiley & Sons, Ltd.
S. El-Zaatari
ity have been found to be a result of high numbers
of microwear features overlying each other,
usually resulting from highly abrasive diets (Scott
et al., 2006). Archaeological and ethnological
accounts indicate that the Tigara had a mainly
marine diet, relying on whale meat, fish and seal
(Rainey, 1941, 1971; Larsen & Rainey, 1948;
Lester & Shapiro, 1968). Meat by itself is not a
highly abrasive food item. Yet ethnological
evidence suggests that the Tigara dried their
meat on open racks or kept it frozen underground
as a year-long staple (de Poncins, 1941; Giddings,
1967). Point Hope has numerous sandy beaches,
and freezing meat on open racks or in under-
ground storage would have provided ample
opportunity for sand to be integrated into it.
Thus, the food preparation techniques of the
Tigara probably led to high levels of extraneous
abrasive particles being ingested. Microwear
analysis, using conventional techniques, of the
same sample of Tigara specimens included in this
current study also shows a highly abrasive diet for
the Tigara (El Zaatari, 2008).
Concerning the two mostly meat-eating
hunter-gatherer samples from climates warmer
than the Arctic, the Chumash and the Fuegians
have the lowest texture fill volume and surface
complexity among the groups considered, thus
indicating a microwear signature characterised by
few pits and few overlying features overall. This
reflects a relatively weakly abrasive diet for both
the Chumash and Fuegians. The Chumash were
fishers and hunters whose diet was based on fish,
marine mammals and other marine resources
(e.g. shellfish) (Fagan, 1995). They also hunted
small terrestrial animals available on the
island. The Fuegians relied on fishing and
seal-hunting, although seals rank lower in
importance (Snow, 1861; Bridges, 1885). Gua-
naco (Lama guanicoe) might also have formed a
terrestrial food source for the Fuegians (Yesner
et al., 2003). Unlike the Tigara, the Chumash and
Fuegians did not dry their meat or store it
underground. Thus, the low levels of surface
complexity and texture fill volume for the
Chumash and Fuegians reflect a diet low in
abrasives. This is consistent with a highly meat-
eating diet with low amounts of extraneous
abrasive particles ingested as a result of food
preparation techniques.
Int. J. Osteoarchaeol. 20: 67–87 (2010)
Microwear Texture Analysis of Hunter-Gatherer Groups
The differences between the microwear sig-
natures of the Chumash and Fuegians probably
suggest slight differences in the diets of these two
groups. The significantly higher surface complex-
ity and lower scale of maximum complexity and
anisotropy levels of the Chumash, compared with
those of the Fuegians, suggest that the Chumash
ingested more abrasive particles that were
relatively smaller than those ingested by the
Fuegians. Ethnohistorical accounts indicate that
although both groups relied on marine animals
and some terrestrial ones for food, the Chumash
also relied on several kinds of plants including
islay, manzanite, mangle berries, tarweed seeds,
tubers and sage (Timbrook, 1993; Walker, 1996).
The differences in the microwear signature of
these two groups might be linked to the
Chumash’s inclusion of such plants in their diet.
However, this remains only a suggestion, as there
is no mention in the ethnohistorical records as to
whether the Fuegians included any plants in their
diets. These records only mention that the
Fuegians, especially the Yamana tribe, to which
the specimens included in this study belong,
relied almost exclusively on marine animals for
their subsistence (Snow, 1861; Bridges, 1885).
The carbon-nitrogen signatures of the Fuegians
and Chumash suggest that these two groups
relied mainly on marine animals for their
subsistence (Walker & DeNiro, 1986; Yesner
et al., 2003). The low levels of occlusal microwear
surface complexity and texture fill volume of
these groups reflect this mainly meat-eating diet,
which did not include a large amount of abrasive
particles. Thus, the stable isotopes and microwear
data are in agreement here. Unfortunately in this
case, since ethnographic accounts state that the
Fuegian diet consisted of large amounts of
shellfish (Snow, 1861; Bridges, 1885), the stable
isotope signature cannot clarify whether the
Chumash included slightly more terrestrial
resources in their diet compared with the
Fuegians, since a marine diet with high amounts
of low trophic-level food (such as shellfish) gives
a very similar carbon-nitrogen signature to one
which is predominantly based on marine foods,
but included some terrestrial resources (Schoe-
ninger et al., 1983; Katzenberg & Weber, 1999).
The Khoe-San mixed diet hunter-gatherers
exhibit intermediate levels of surface complexity,
Copyright # 2008 John Wiley & Sons, Ltd.
81
lower than those of the meat-eaters from Point
Hope and higher than those of the meat-eaters
from warmer climates. The Khoe-San had a diet
consisting of a wide range of marine, estuarine
and terrestrial (plants and animal) foods (Good-
win, 1938; Burchell, 1967; Sealy, 2006). Ethno-
graphic studies show that 80% or more of the diet
of modern Kalahari hunter-gatherers consists of
plant food (Silberbauer, 1981). Even though the
Kalahari is quite sandy, it should be mentioned
that the San do not dry meat or store it
underground. It is reasonable to suggest that
the abrasives ingested by the Khoe-San were not
due to dust or sand particles attached to their
dietary staples, but rather due to the ingestion of
hard dietary items such as nuts, seeds, tubers, and
so on, that were widely available and exploited by
these hunter-gatherers.
The Khoe-San sample included in this study
comprises specimens from the sites of Oakhurst
Shelter, Matjes River and Riet River. The results
of stable isotope analyses indicate that the diets of
the inhabitants of these three sites were similar.
The carbon and nitrogen signatures of these
groups show that they all had a mixed diet
consisting both of marine and terrestrial (plant
and animal) resources (Burchell, 1967; Sealy &
van der Merwe, 1986, 1988; Sealy et al., 1992;
Sealy & Pfeiffer, 2000; Sealy, 2006). In accord-
ance with the stable isotope results, microwear
texture analysis did not detect any significant
differences in the occlusal microwear signatures
of these three Khoe-San groups. Also, the dental
microwear results are in agreement with the stable
isotope data in classifying the Khoe-San as
having a mixed diet.
The Andaman Islanders have relatively com-
plex surfaces, which place them with the Point
Hope population. Also, the surfaces of the
Andamanese attain maximum complexity at
relatively fine scales, indicating the presence of
numerous small microwear features that overly
one another. Although the mixed-diet Andama-
nese came from a tropical environment, historical
accounts indicate that these people relied more
on hunting and fishing than gathering plants for
their subsistence (Myka, 1993). Unlike the Khoe-
San, whose diet consisted more of plants than
meat, meat comprised some 66% of the
Andamanese diet (Man, 1883). However, this
Int. J. Osteoarchaeol. 20: 67–87 (2010)
82
does not explain why the Andamanese have such
complex surfaces, placing them with Arctic
hunter-gatherers rather than with meat-eating
hunter-gatherers from warmer climates, such as
the Chumash and the Fuegians. Historical
accounts also note that the Andamanese diet
consisted mostly of meat that was eaten half
roasted, such that it remained relatively tough
(Pal, 1984). The food was usually cooked
uncovered, which would have enabled contami-
nation by windborne sand (Man, 1883; Pal,
1984). The high complexity of the surfaces of the
Andamanese molars might thus reflect their food
preparation techniques which allowed for very
small abrasive particles to be attached and
ingested with their food.
The results of the occlusal molar microwear
analysis show that dental microwear texture
analysis is able to differentiate between modern
humans with different dietary habits. Five
variables were gleaned from occlusal microwear
texture analysis. The modern groups are best
differentiated on the basis of the level of complex-
ity of their wear surfaces. To a lesser extent, the
scale of maximum complexity, anisotropy, and
texture fill volume were also differentiating factors.
The heterogeneity of surface texture was not found
to be a good differentiating factor among the
modern groups analysed in this study.
The results of this study show that surface
complexity clearly distinguishes the groups based
on the amount of hard objects included in their
diets. Based on complexity of the surface texture,
the samples can be divided into three groups in
decreasing order: (1) Tigara and Andamanese;
(2) Khoe-San and Chumash; and (3) Fuegians.
Thus the increase in surface complexity was
found to correlate with the increased ingestion of
hard items. Scott et al. (2006) also found that
surface complexity was consistent with a high
bite force to fracture hard objects. The scale of
maximum complexity appears to be linked to the
scale of wear-causing particles (Scott et al., 2006).
Small microwear features caused by small abrasive
particles are detected at fine-scale measurements,
and large features which are caused by large
abrasive particles are detected at coarse scales
(Scott et al., 2006). This variable differentiates the
Fuegians as having the highest Smc compared
with the other modern human samples, and the
Copyright # 2008 John Wiley & Sons, Ltd.
S. El-Zaatari
Andamanese as having the lowest value compared
with all the modern human groups, with the
exception of the Chumash. Anisotropy is linked
to the directionality of the jaw movements during
mastication. It serves to differentiate non-human
primates with different diets since highly
anisotropic surfaces are found among primates
that consume more tough pliant items such as
grasses and leaves (Scott et al., 2005, 2006). The
results of this study indicate that the Fuegians
exhibit the most anisotropic surfaces of all the
modern groups. The Fuegian diet consisted
almost exclusively of meat, and lacked hard
abrasive items. Since meat is a tough, but not
hard, dietary item, its chewing would have
required repetitive directional jaw movements,
thus the higher level of anisotropy for the
Fuegians compared with the rest of the groups
analysed. Scott et al. (2006) show that texture fill
volume is greater in surfaces with larger, deeper
and more symmetrical areas of wear (i.e. heavily
pitted surfaces). Texture fill volume differentiates
the Fuegians and Chumash from the rest of the
groups as having the lowest value. This is
expected, since individuals that do not eat a
large number of hard objects have been shown
not to have highly pitted occlusal surfaces
compared with those who have a mainly hard-
object diet (e.g. Teaford & Walker, 1984; Harman
& Rose, 1988; Teaford, 1988; Teaford & Oyen,
1989a).
Finally, this study sheds light on the microwear
signature of meat-eating through the analysis of
the microwear pattern of mostly meat-eating
hunter-gatherers with various degrees of
inclusion of abrasive particles. Of these groups,
the Fuegians included the least amount of
abrasives in their diet, whereas the Tigara
included the highest amount. The microwear
signature of the Fuegians is characterised by low
levels of complexity and texture fill volume,
reflecting the presence of few microwear features
overlying each other and few pits, thus indicating
low amounts of abrasives. This is in accord with
meat-eating, since meat on its own does not form
a hard, abrasive dietary item. The Fuegian
microwear signature is also associated with high
anisotropy, which is expected since meat is
considered a tough item and requires some level
of directionality in the jaw movement during
Int. J. Osteoarchaeol. 20: 67–87 (2010)
Microwear Texture Analysis of Hunter-Gatherer Groups
mastication. The high scale of maximum com-
plexity for this group indicates the absence of
small abrasive particles. On the other hand, the
mostly meat-eating Tigara show the opposite
microwear pattern compared with the Fuegians.
Ethnographic evidence suggests that the Tigara
ingested high amounts of abrasive particles
attached to their food as a consequence of their
food preparation techniques. Thus, the results of
this study show that the microwear signature
of meat-eating is highly affected by the amount of
extraneous abrasive particles.
Conclusion
In this study, the microwear fabrics of five modern
hunter-gatherer populations with varied dietary
habits were documented to better understand
how dietary differences affect microwear in these
groups. The modern hunter-gatherer samples
were selected from historical and prehistoric
populations that represent: (1) mixed diet hunter-
gatherers from tropical climates (Khoe-San and
Andamanese); (2) meat-eating hunter-gatherers
from Arctic/tundra environments (Tigara); and
(3) meat-eating hunter-gatherers from climates
warmer than the Arctic (Fuegians and Chumash).
Microwear differences between the meat-eating
hunter-gatherers and the mixed diet hunter-
gatherers were detected, and significant differ-
ences were also found among the meat-eating
hunter-gatherers that inhabited different environ-
ments. To this end, the microwear fabrics of the
Tigara and Andaman Islanders reflect the highly
abrasive diet of these individuals, which is most
likely related to the ingestion of hard particles
clinging to their foods as a result of their food
preparation techniques (i.e. drying the meat on
open racks on the sandy beaches of Point Hope,
Alaska, for the Tigara; and cooking the food
uncovered, allowing windborne sand particles to
be included with it, for the Andamanese). The
microwear signatures of the Chumash and
Fuegians, on the other hand, classify them as
having the least abrasive diets among the modern
groups. This is expected, as the dietary habits of
the Chumash and Fuegians consisted mostly of
meat from marine animals, which by themselves
are too soft to indent the enamel surface. Also,
Copyright # 2008 John Wiley & Sons, Ltd.
83
there is no indication in the ethnohistorical
accounts that these two groups ingested any kind
of extraneous abrasive particles. Significant
differences in the microwear signatures between
the Fuegians and Chumash were also detected
reflecting subtle dietary differences, probably
related to a somewhat greater amount of hard
plant foods eaten by the Chumash. The mixed-
diet Khoe-San occupy an intermediate position
between the Tigara and the Andamanese on
the one hand, and the Chumash and Fuegians on
the other, with regard to the level of abra-
sives ingested. The mixed diet of the Khoe-
San included large amounts of plant foods, such
as nuts, seeds and tubers, which were pro-
bably responsible for abrading their enamel
surfaces.
Acknowledgements
I would like to thank James Brink, Zoe Hender-
son, Louise Humphrey, Rob Kruszynski, Giorgio
Manzi, Alan Morris, David Morris, Ken Mow-
bray, Chris Stringer and Ian Tattersall for allow-
ing access to the specimens under their care.
I would also like to thank Frederick Grine, Peter
Ungar and Anthony Olejniczak for helpful dis-
cussions and two anonymous reviewers for their
comments on an earlier version of this paper. This
study was supported by the National Science
Foundation, the Leakey Foundation and the Rug-
gles-Gates Fund for Biological Anthropology (the
Royal Anthropological Institute of Great Britain
and Ireland).
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