Journal of Human Evolution 63 (2012) 781e795

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Journal of Human Evolution

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Modeling Neanderthal clothing using ethnographic analogues

Nathan Wales
Department of Anthropology, University of Connecticut, 354 Mansfield Road, Storrs, CT 06269-2176, USA

a r t i c l e i n f o a b s t r a c t

Article history: Although direct evidence for Neanderthal clothing is essentially nonexistent, information about Paleo-
Received 16 June 2011 lithic clothing could provide insights into the biological, technological, and behavioral capabilities of
Accepted 15 August 2012 Neanderthals. This paper takes a new approach to understanding Neanderthal clothing through the
Available online 17 October 2012
collection and analysis of clothing data for 245 recent hunter-gatherer groups. These data are tested
against environmental factors to infer what clothing humans tend to wear under different conditions.
Keywords:
Beta regression is used to predict the proportion of the body covered by clothing according to a location’s
Homo neanderthalensis
mean temperature of the coldest month, average wind speed, and annual rainfall. In addition, logistic
Hunter-gatherer
OIS-3
regression equations predict clothing use on specific parts of the body. Neanderthal clothing patterns are
Environmental conditions modeled across Europe and over a range of Pleistocene environmental conditions, thereby providing
Beta regression a new appreciation of Paleolithic behavioral variability. After accounting for higher tolerances to cold
temperatures, it is predicted that some Neanderthals would have covered up to 80% of their bodies
during the winter, probably with non-tailored clothing. It is also likely that some populations covered the
hands and feet. In comparison with Neanderthals, Upper Paleolithic modern humans are found to have
worn more sophisticated clothing. Importantly, these predictions shed new light on the relationship
between Neanderthal extinction and their simple clothing.
Ó 2012 Elsevier Ltd. All rights reserved.

Introduction
Humans are the only extant species habituated to wearing
clothing, but it is yet to be determined whether any other hominin
species ever engaged in the same behavior. In particular, archae-
ologists and paleoanthropologists have contemplated whether
Neanderthals periodically wore garments. Over the course of the
past century, conventional wisdom about Neanderthal clothing has
changed, often in accordance to changing perceptions of the
Neanderthals in general (Drell, 2000). During the early part of the
twentieth century, Neanderthals were assumed to have lacked
clothing, as depicted in Franti
sek Kupka’s famous 1909 drawing
(Trinkaus and Shipman, 1993: fig. 70). By the middle of the twen-
tieth century, Neanderthals were often thought of utilizing very
basic clothing, such as scraps of fur around the waist, as seen in
Zdeuck Burian’s 1950 drawing of a Neanderthal clan (Trinkaus and
Shipman, 1993: fig. 75). More recently, some researchers have
suggested that Neanderthals wore advanced forms of tailored
clothing. Nevertheless, most of these revisions are based more
upon speculation than scientific data. Compared to recent advances
in understanding Neanderthal subsistence strategies (Kuhn and
E-mail address: nathan.wales@uconn.edu.
Stiner, 2006; Hardy et al., 2012), symbolic behavior (Caron et al.,
2011), mobility (Richards et al., 2008; Lalueza-Fox et al., 2011),
and evolutionary history (Endicott et al., 2010; Wills, 2011), the
paucity of data on Neanderthal clothing is remarkable. This is
largely due to the fact that most Paleolithic clothing decayed
millennia ago, along with other untold organic artifacts. Indirect
evidence, such as lithics used to process clothing, is ambiguous at
best. There is no single lithic type that is unequivocally associated
with the production of clothing, and while microwear analysis of
stone tools can document the processing of hides, the issue of
equifinality complicates interpretations (Keeley, 1980).
The lack of clothing-specific tools from Neanderthal sites is
made more apparent when compared with sites attributed to the
earliest anatomically modern humans in Europe. Soffer (2004) cites
multiple lines of evidence that anatomically modern humans in
Eurasia engaged in weaving and sewing, including the presence of
weaving battens and textile impressions at Dolní Vĕstonice, dating
at least 26,000 years before present (BP) (Adovasio et al., 2001;
Soffer et al., 2001). Other sites, including Vogelherd (Germany) and
Kostenki 15 (Russia) have sewing needles between 35,000 and
30,000 BP (Soffer, 2004; Hoffecker, 2005). In addition, Kvavadze
et al. (2009) claim to have identified dye-colored flax fibers from
Dzuduana Cave, Georgia, dating to 32,000e26,000 14C BP. While
Paleolithic textiles could have been used exclusively for cordage

0047-2484/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.jhevol.2012.08.006
782 N. Wales / Journal of Human Evolution 63 (2012) 781e795
and basketry, research on Upper Paleolithic Venus figurines
suggests otherwise. Many figurines have designs that researchers
interpret as clothing (Soffer et al., 2000; Conard, 2009).
Based on genetic analyses of the human body louse (Pediculus
humanus corporis) and head louse (P. humanus capitis), Kittler et al.
(2004) argue that anatomically modern humans were the first
species to wear clothing. The body louse could have evolved from
the head louse only after clothing created a unique ecological niche.
The researchers calculate that the body louse arose in Africa
between 114,000 and 30,000 years ago, long after Neanderthals
were well established in Eurasia. Toups et al. (2011) have since
conducted more robust analyses using multiple genetic markers
and estimate that body and head lice diverged at least 83,000 years
ago, with a median estimate of 170,000 years ago. It is important to
note that while these studies suggest that modern humans began
habitually wearing clothing at this time, it does not preclude
independent inventions of clothing by other hominins. If Nean-
derthals developed clothing, head lice presumably would invade
the niche and evolve into a new form of body louse. Given Green
et al.’s (2010) findings that Neanderthals and modern humans
interbred, it would be expected that body lice would be transferred.
In fact, a similar transmission of head lice from Homo erectus to
modern humans has been argued using genetic data (Reed et al.,
2004). So far, there are no indications that body lice were passed
between Neanderthals and modern humans. Various factors could
explain the lack of evidence for Neanderthal body lice. For example,
Neanderthal body lice may not have been well adapted to modern
human clothing and went extinct along with their host. Alterna-
tively, if interbreeding only took place in the Middle East as indi-
cated by the genetic data (Green et al., 2010), Neanderthals or
modern humans may have been unclad and not harbored the
parasites.
Previous research on Neanderthal clothing
Despite the seeming dearth of evidence for Neanderthal
clothing, many researchers have tackled the issue from various
angles. Using estimates of metabolic rates and thermal conduc-
tance, Aiello and Wheeler (2003) argue that Neanderthals needed
a significant amount of ‘cultural insulation,’ i.e., clothing, to survive
the harsh climates of Oxygen Isotope Stage 3 (OIS-3: 59,000e
27,000 years ago; van Andel et al., 2003). Although many
researchers have argued that Neanderthals were cold-adapted (e.g.,
Holliday, 1997; but see; Stewart, 2007), the authors calculate that
Neanderthals could tolerate only marginally colder ambient
temperatures than anatomically modern humans. However, if
Neanderthals wore a layer of clothing equivalent to a business suit,
they could withstand temperatures as frigid as
23.9  C. Likewise,
Sørensen (2009) predicts Neanderthal clothing using rates of
metabolic energy production and heat loss. He finds that Nean-
derthals living near Bispingen, Germany, during the Eemian Inter-
glacial (OIS-5e) would require tailored clothing and advanced
footwear to survive an average winter.
Gilligan (2007) provides a different perspective on Neanderthal
clothing, based partially on the lack of an archaeological signature
of sophisticated clothing-related artifacts. He hypothesizes that
Neanderthal adaptations predisposed them to delay the invention
of complex clothing. Modern humans, on the other hand, lack such
cold adaptations, and were obligated to invent complex multilay-
ered clothing with tight fitting sleeves and leggings (for discussion
and depictions of tailored and non-tailored clothing, see Gilligan,
2010a). Gilligan (2010a) further speculates that during rapid
climatic oscillations in OIS-3, possibly associated with extreme
wind chill levels, the lack of complex clothing in Neanderthals led
to hypothermia and ultimately caused their extinction. Froehle and
Churchill (2009) present a more mathematical analysis of Nean-
derthal energy requirements, but like Gilligan (2010a), suggest that
cold adaptations in Neanderthals discouraged the development of
complex clothing. The authors calculate that clad modern humans
would have had a competitive advantage over the Neanderthals,
eventually out-reproducing them and leading to Neanderthal
extinction.
The above studies provide plausible interpretations of Nean-
derthal clothing and interesting explanations for the population
turnover event of the European Paleolithic (Shea, 2008). However,
a more detailed picture of Neanderthal clothing has not been
attempted, nor has any researcher satisfactorily analyzed clothing
variability across geographic regions and temporal periods. Simply
put, it is naïve to assume that Neanderthals from Spain, northern
Germany, the Levant, and Siberia wore identical clothes. This paper
reconciles this inadequacy by using a new strategy to predict
Neanderthal clothing across Europe and over different climatic
regimes. Not only can the amount of clothing be modeled, but it is
also possible to predict whether specific body parts would have
been covered. For the first time, we have an empirical method to
infer continental-scale clothing patterns for a long-extinct hominin.
Most importantly, these projections allow archaeologists and
paleoanthropologists to more fully understand how Neanderthals
differed from modern humans behaviorally, technologically, and
biologically.
Methods
To predict Neanderthal clothing, this study attempts to first
understand modern hunter-gatherer clothing. Following Binford’s
(2001) Constructing Frames of Reference, analytical methods are
used to identify core variables that account for variability in forager
clothing. As such, this study can be seen as part of a growing set of
literature building upon Binford’s approach to better understand
prehistory (e.g., Johnson, 2008; Johnson and Hard, 2008; Gabler,
2011). While recent hunter-gatherers are not relicts from the
past, when used with appropriate restraint, their lifeways can
provide helpful insights for many anthropological questions (Kelly,
1995).
Clothing data are examined here in the context of environ-
mental variables. It is assumed that the primary function of clothing
is to protect the body from the environment, as it is known that
clothing insulates the body with a layer of air (Stenton, 1991; Buijs,
1997). The amount of clothing must be appropriate to local envi-
ronmental conditions; excessive clothing leads to overheating and
loss of thermal insulation through sweat accumulation in garments
(Gilligan, 2010b). Of course, clothing plays a role in cultural and
personal identity (Hansen, 2004), so it is to be expected that people
living in identical environmental conditions will not necessarily
have identical clothing. Rather, this research seeks to find over-
arching patterns in the use of clothing based on environmental
conditions. Detailed methods, including relevant equations and in-
depth explanations, are provided in Appendix A.
Clothing data
As discussed by Gilligan (2008), the collection of high-quality
clothing data from anthropological literature is quite challenging.
Early reports are often incomplete, inconsistent, or overtly ethno-
centric. Therefore, it was necessary to consult detailed ethnographic
descriptions, illustrations, and photographs of hunter-gatherer
clothing. From these sources, clothing data were recorded for the
339 hunter-gatherer groups in Binford’s (2001) database. When
possible, indigenous clothing information was recorded for males
and females in multiple seasons. For sources that describe several
 N. Wales / Journal of Human Evolution 63 (2012) 781e795
individuals of the same sex in the same season, the clothing of the
most representative individual was recorded. In total, data were
collected for 595 individuals, representing 245 hunter-gatherer
groups (72% of groups). Source information for the groups is
found in Appendix B. The global distribution of hunter-gatherer
groups with clothing data is depicted in Fig. 1.
For each individual, the following data were systematically
recorded: number of clothing articles, material of each garment,
percentage of the body covered by clothing, and body adornments,
including paint, jewelry, tattooing, and scarification. The
percentage of the body covered by clothing was determined using
Wallace’s (1951) ‘rule of nines’: hands, 1%; lower arms, 8.5%; upper
arms, 8.5%; torso, 36%; genitals, 1%; upper legs, 17.5%; lower legs,
17.5%; feet, 1%; and head, 9%. This method is commonly used by
doctors to assess the extent of injuries of burn victims, but it also
provides a standardized estimate of body surface area for this study.
To simplify computations and prevent recording biases, individual
body parts were recorded as being either covered or uncovered.
The dataset was modified to omit some cases and to identify
seasonal variability in clothing patterns. Cases with ambiguous
descriptions or European dress were omitted from all analyses.
Seasonal clothing requirements were investigated using the
maximum and minimum amount of clothing worn by each
hunter-gatherer group. To determine the maximum amount of
clothing worn by each hunter-gatherer group, the individual from
that group with the most clothing was kept as the value for the
entire group. In the same manner, the minimal amount of clothing
was determined using the individual wearing the least amount of
clothing in the group. Entire groups were also not considered when
clothing data was only available for the opposite season. For
example, groups with only winter clothing data were omitted for
the minimal clothing analysis. Maximal and minimal clothing
projections are based on 212 and 213 hunter-gatherer groups,
respectively. Naturally, individuals from a given population often
wear different clothes according to cultural expectations or
personal preference. However, this approach arguably still provides
a reasonable assessment of the clothing commonly worn by
foragers throughout the year, especially when using statistical
methods resistant to exceptional cases.
Figure 1. Geographic distribution of hunter-gatherer groups in clothing database.
783
To investigate the use of clothing on specific body parts, the
thermal properties of clothing articles were considered. Sandals
were not able to be consistently predicted using the method
described below and were omitted from predictions of footwear.
Likewise, small hats or visors were not counted as clothing on the
head.
Environmental data
Binford’s database of hunter-gatherers provides environmental
and geographic variables for each forging group, and serves as the
primary source of independent variables against which the clothing
data are tested. Because researchers have identified wind chill as an
important factor in clothing, supplementary wind data for each
hunter-gatherer group were collected from 3TIER, a company that
assesses renewable energy prospects (3TIER, 2009). Four
hunter-gatherer groups reside in polar areas outside the supported
geographic regions in 3TIER’s FirstLook software. These groups
were omitted from further analyses, leaving a total of 422 indi-
viduals in the study.
There are a number of competing wind chill equations, all of
which attempt to quantify the cooling effect of wind on the human
body. As discussed by Quayle and Steadman (1998), the first wind
chill equation was developed in 1945 and popularized by the
National Oceanic and Atmospheric Administration (NOAA), here
referred to as the NWS wind chill equation. Inadequacies in the
NWS formula led Quayle and Steadman to propose the more
accurate Steadman wind chill. In 2003, NOAA also established
a new wind chill temperature index (WCTI) to address criticisms
(Williamson, 2003). The WCTI equation was used by Aiello and
Wheeler (2003) in their analysis of Neanderthal thermoregula-
tion, however, Sørensen (2009) argues that their use is potentially
invalid. Despite the numerous debates on wind chill equations,
most concerns are related to extreme wind velocities and frigid
temperatures. As the data examined in this study deals with long-
term averages rather than wind gusts or abnormally cold days, the
issue is less controversial here. Furthermore, if an equation fails to
accurately reflect human thermal requirements, it should be
automatically ignored when using methods that identity the best
784 N. Wales / Journal of Human Evolution 63 (2012) 781e795
models. Wind chills based on the NWS, Steadman, and WCTI
equations were calculated for all hunter-gatherer groups. Wind
speed is usually computed using instantaneous wind velocity.
However, only long-term averages are available in the dataset, so
variables such as the mean temperature of the coldest month and
mean annual temperature were hesitantly used. Although this
approach is unconventional, it was determined that these average
wind chills, though modest, provide a better understanding of
clothing variability.
Beta regression
Relationships between the amount of the body covered by
clothing and environmental variables were explored using SPSS
14.0 (2005). It was found that the percentage of the body covered
by clothing in hunter-gatherer groups is correlated with a number
of environmental variables. For example, the mean temperature of
the coldest month is inversely correlated with the maximum
amount of clothing worn in hunter-gatherer groups (Fig. 2).
Although there is a clear relationship between clothing coverage
and temperature, it is inappropriate to use linear regression for
percentages and proportions. One fault lies in the fact that most
percentage data are heteroskedastic (Ferrari and Cribari-Neto,
2004). In addition, percentage data invalidate the assumption
that the response variables can have any value. Rather, percentage
data are restricted to the 0e1 interval.
The percentage of the body covered by clothing was examined
using beta regression by way of the Betareg package created by
Cribari-Neto and Zeileis (2009) for R statistical software (R
Development Core Team, 2009). The percentage of the body
covered by clothing was used as the dependent variable while
environmental factors served as independent variables, entered
individually and in combination with other variables. For each set of
independent variables, the software yields a pseudo R-squared (R2p )
value and Akaike information criterion (AIC). The R2p is analogous to
the R2 of linear regression; it describes how much of the variability
in the independent variable is accounted for by the model. Higher
Figure 2. Maximum percentage of the body covered by clothing in hunter-gatherer
groups versus the mean temperature of the coldest month (MCM). Fit line from beta
regression is depicted using the single dependent variable, however, equations best
able to predict clothing are multivariate and cannot be visualized in two-dimensional
space.
R2p values indicate that the model is better at predicting the inde-
pendent variable. AIC is a measure that can be used to find
a balance between the complexity of a model and its interpretive
ability (Akaike, 1974). In other words, adding more variables to
a model increases its predictive ability, but it also increases the
likelihood that correlations occur by chance. Different predictive
models were compared to find the one with the lowest AIC value.
This model has the highest predictive ability without including
unnecessary independent variables. The cauchit link function was
used for all beta regressions because it handles outliers relatively
well (Morgan and Smith, 1992). See Appendix A for further
discussion and relevant equations.
Logistic regression
Predictions for the use of clothing on specific body parts were
conducted using logistic regression in SPSS 14.0. Logistic regression
was selected over discriminant function analysis because it better
accommodates non-normally distributed data (Press and Wilson,
1978) and yields a value for the confidence of each prediction
rather than only group assignments (Menard, 2010). Environmental
variables were used to predict the likelihood that clothing is used to
cover the head, hands, and feet. Independent variables were
entered in a stepwise fashion using likelihood ratios. The amount of
variability accounted for by each model is described by the R2L
statistic, a measurement proportional to the reduction in the log
likelihood measure (Menard, 2010).
Logistic regression yields a function with values from 0 to 1, with
1 indicating 100% chance of success. In this study, ‘success’ means
than the body part is covered. For example, a value of 0.6 for a given
body part means that there is a 60% chance the body part is covered
by clothing. The most conclusive predictions have values less than
0.05 and above 0.95. One should note that the predicted coverage of
a body part does not necessarily indicate a specific garment. For
example, it is possible to keep hands covered with a large cloak
rather than fabricating tailored gloves.
Clothing predictions for Neanderthals
Predictions of Neanderthal clothing are based on paleoenvir-
onmental data from the Stage 3 Project (van Andel, 2003), an
interdisciplinary study that reconstructed European environments
during OIS-3 (60,000e24,000 years ago). Four paleoclimatic
reconstructions for different phases of the late Pleistocene and
Holocene have been published: OIS-3 warm phase, OIS-3 cold
phase, the Last Glacial Maximum (LGM), and modern values. The
LGM occurred 24,000 years ago, following the Neanderthal
extinction, however, this climatic model can be used as a proxy for
the OIS-4 glacial, as done by Aiello and Wheeler (2003). Modern
environmental conditions are used as an analog for OIS-5e, the last
interglacial period, dating to 130,000e117,000 BP (van Andel and
Tzedakis, 1996). It should be noted that some researchers are crit-
ical of these paleoclimatic projections because they are averages of
many millennia. For example, Gilligan (2007, 2010b) argues that
abrupt climatic fluctuations are not observed in these projections,
but such events would have had a dramatic effect on past species,
including Neanderthals. While short-term cold periods could pose
severe challenges to past species, regional-scale projections of
these events are currently unavailable and therefore are not
explicitly examined in this study.
When extrapolating a predictive model to another organism, as
done here, it is critical to evaluate whether the model can be
applied directly to the second population or if a correction factor is
required. For equations calculating the percentage of the body
covered by clothing, it was assumed that Neanderthals have a slight
 N. Wales / Journal of Human Evolution 63 (2012) 781e795
overall thermal advantage compared with modern humans. This
assumption is based on research demonstrating that compared
with anatomically modern humans, Neanderthals have a lower
ratio of body surface area to mass (Helmuth, 1998) and greater
muscle mass (Aiello and Wheeler, 2003). As discussed by Ruff
(1993), organisms with lower ratios of surface area to mass retain
heat better than those with proportionally more surface area, and
consequently, stout Neanderthals have a thermal advantage over
tropically-proportioned modern humans. Rugose muscle attach-
ment sites on Neanderthal thoracic and appendicular skeletal
elements indicate that they were more heavily muscled than
modern humans (e.g., Franciscus and Churchill, 2002; Mariotti and
Belcastro, 2011). Since muscle insulates better than fat (Veicsteinas
et al., 1982), Aiello and Wheeler (2003) suggest that Neanderthals
would have a 5% reduction in heat loss compared with less
muscular modern humans, a value from experimental research by
Glickman-Weiss et al. (1993). Whatever the sources of these
morphological differences, cold adaptation (Steegmann et al.,
2002), genetic drift (Weaver et al., 2007), or otherwise, it is
necessary to make the hominin populations more comparable by
altering the paleoenvironmental dataset to make Neanderthal
environments artificially warmer. All temperatures for Neanderthal
climate models were increased by 4  C, based on Aiello and
Wheeler’s (2003) calculation that heavily-muscled, Neanderthal-
shaped bodies can withstand sustained minimum temperatures of
6.5  C, in contrast 10.5  C for modern humans.
Other hypothesized thermal and metabolic differences between
Neanderthals and moderns humans were not included in the
models. For example, Aiello and Wheeler (2003) suggest a sparse
layer of 3.9 cm body hair would have provided Neanderthals with
the thermal equivalent of a light layer of clothing. Likewise,
Steegmann et al. (2002) hypothesize that Neanderthal adults had
deposits of brown adipose tissue, a fat that generates heat in infants
and some adult primates. Clear evidence for Neanderthal body hair
and brown adipose fat are lacking, so these factors were not
considered for the models. Of course, advances in understanding
phenotypic traits of Neanderthals through their DNA may allow
such questions to be answered one day (see Lalueza-Fox et al.,
2007).
Along the same lines, the analyses do not include any correction
for hypothesized elevated basal metabolic rates (BMR) in Nean-
derthals. Many researchers argue that formulas to predict BMR
from stature and mass underestimate humans who live in cold
climates and eat protein-rich diets, including the Neanderthals
(Aiello and Wheeler, 2003; Churchill, 2006; Snodgrass and Leonard,
2009). Furthermore, Froehle and Churchill (2009) calculate that
Neanderthals would have had a higher BMR than the earliest
modern humans in Europe. This would cause Neanderthals to have
to generate more internal heat at the expense of higher caloric
requirements. However, the clothing models found in the article are
based upon hunter-gatherers who are adapted to a range of
climates and also do not conform to BMR equations derived
primarily from European and Euro-American subjects. Measure-
ments of BMR demonstrate that there is wide variability among
human populations, and many groups have rates that are much
higher or lower than predicted by common methods (Wilson,
1945). To determine if Neanderthals fall within the modern range
of variability, BMR was calculated using average stature and weight
data for 35 human populations (Ruff, 1994) and 26 Neanderthal
individuals (Churchill, 2006) using the Kleiber (1961) BMR formula
for placental mammals and a method tested by Churchill (2006).
Although these formulae may underestimate cold-adapted
humans, they should bias the estimates for modern
hunter-gatherers and Neanderthals to a similar extent. Even with
this limited dataset, male and female Neanderthals are found to be
785
within the upper range of modern human variability for BMR. Both
equations predict female Neanderthals to have a lower BMR than
one human population (Czech), with a value nearly identical to that
of Inuit. Male Neanderthals are found to have a BMR just below Lau
Pacific Islanders when using Churchill’s method, however, the
Neanderthals are found to have a slightly higher value with the
Kleiber equation (1829 versus 1802 kcal/day). Overall, these values
suggest that Neanderthals generally fall within the range of modern
human BMR, and therefore no correction for BMR was included in
the models.
Although there is substantial reason to artificially elevate
temperatures used to predict Neanderthal clothing over the whole
body, it is a separate issue whether clothing use on individual body
parts needs a correction value. The crux of the problem is surface
area. The surface area of the head, hands, and feet determines how
quickly heat is transferred to the environment. To determine if
Neanderthal crania have a statistically different surface area than
those of modern humans, cranial measurements from five Nean-
derthals and 11 Upper Paleolithic modern humans compiled by
Kidder (1996) were analyzed. Surface area of the cranium was
estimated using three standardized measurements defined by
Howell (1973): glabella to opisthocranion (GOL), euryon to euryon
(XCB), and basion to bregma (BBH). These measurements effectively
represent the cranium’s maximum length, breadth, and height, and
were used to calculate the surface area of the ellipsoid demarcated
by these axes. Mean surface area of the sample of Neanderthal and
modern human crania are 78,317 mm2 and 74,696 mm2, respec-
tively. An independent sample T-test found no evidence that the
two groups are significantly different (equal variances not assumed,
t ¼
0.655, df ¼ 5.074, p ¼ 0.541). Hence, no correction value was
used when predicting the use of clothing on the head.
Predicting the surface area of hands and feet is possible using
equations developed by Liao et al. (2008) and Yu and Tu (2009),
respectively. However, parameters of these equations require data
on the articulated hand/foot, and are generally meant for
measurement on living humans. This creates a problem for fossil
specimens because reliable data for muscle and tissue volumes are
not available. Data on the skeletal remains for Neanderthal hands
(Niewoehner, 2000) and feet (Trinkaus, 1975) have been published,
and certain skeletal elements like the apical tuft of the thumb are
known to exhibit some differences between the populations
(Mittra et al., 2007). However, many skeletal elements of the hands
and feet have a great amount of overlap in Neanderthals and
modern humans. For example, metacarpal articular lengths for
Neanderthals and Upper Paleolithic modern humans overlap to
a great extent (Niewoehner, 2000). Until reliable data are made
available on relative hand and foot sizes of Neanderthals, it would
be speculative to assume that Neanderthal hands and feet had
different surface areas and thermal properties than those of
modern humans. Thus, like with the head, no correction was
utilized when modeling clothing on the hands or feet.
Table 1
Climatic phases as described by van Andel et al. (2003) with chosen simulation.
SPECMAP Climate phase Simulation model used Age
for projections (cal ka BP)
OIS-5ea Eemian Interglaciala Modern climate 130e117
OIS-5de5b Multiple phases Not modeled in this study 117e85
OIS-5a Early Glacial Warm Phase Warm 85e74
OIS-4 Transitional phase Warm 74e66
OIS-4 First Glacial Maximum Last Glacial Maximum 66e59
OIS-3 Stable Warm Phase Warm 59e44
OIS-3 Transitional phase Warm 44e37
OIS-3 Early Cold Phase Cold 37e27
OIS-2 Last Glacial Maximum Last Glacial Maximum 27e16
a
Added to van Andel et al.’s (2003) list for this study.
786 N. Wales / Journal of Human Evolution 63 (2012) 781e795

Table 2
Beta regression models to predict amount of body covered by clothing.

Model Beta regression equationa R2p Variable p-value

Maximum amount of body covered by clothing h ¼ 0.1798387
0.1947009*STD_MCM þ 0.0006376*CRR 0.6249 STD_MCM <0.001
CRR <0.001
Minimum amount of body covered by clothing h ¼
3.07270
0.08506*MCM þ 0.05124*LAT 0.4288 MCM <0.001
LAT <0.01
a
As discussed in Appendix A, predicted proportion of body covered m ¼ 0:5 þ arctanðhÞ=p. CRR: average annual rainfall (mm); LAT: latitude (nonnegative decimal degrees);
MCM: mean temperature of the coldest month ( C); STD_MCM: Steadman wind chill calculated using the mean temperature of the coldest month ( C) and mean annual wind
speed (m/s) at 10 m above the ground.

Neanderthal clothing patterns were predicted at each coordi-
nate of the Stage 3 Project grid using equations derived from the
hunter-gatherer clothing database. Results were mapped using
ArcMap 9.3 (ERSI, 2008) and values between the points were
interpolated using kriging. For each phase of OIS-3, the landmass of
Europe was adjusted to the appropriate sea level.
Archaeological site database
The locations of archaeological sites provide a geographical
context in which to interpret clothing predictions. Corresponding
sets of Middle Paleolithic and Upper Paleolithic sites are included for
all maps. Mousterian sites, including Châtelperronian sites, are listed
when clothing predictions are projected using Neanderthal-
morphology temperature corrections; it is assumed that these
techno-complexes were made by Neanderthals (but see Bar-Yosef
and Bordes, 2010). Early Upper Paleolithic, Aurignacian, Gravet-
tian, and Upper Paleolithic sites are mapped with clothing projec-
tions based on modern human morphology. The database is
primarily based on the collection of sites published by the Stage 3
Project (van Andel et al., 2003). OIS-5e Mousterian sites were
compiled from sites cited in Roebroeks et al. (1992), Baryshnikov
and Hoffecker (1994), Gamble (1999), Speleers (2000), Adler et al.
(2003), Richter (2005), Svoboda (2005), Wenzel (2007), and
Slimak et al. (2010). While there are likely dating problems with
some sites, and site-specific chronologies are always being revised

Figure 3. Predicted maximum amount of body covered by clothing for Neanderthals. Mousterian sites displayed according to climatic regime: a) OIS-5e Interglacial [130e117 ka BP]
(modern climate model), b) Glacial phase of OIS-4 [66e59 ka BP] (Last Glacial Maximum climate model), c) Warm phases of OIS-4 [74e66 ka BP] and OIS-3 [59e37 ka BP], and d)
cold phase of OIS-3 [37e27 ka BP].
 N. Wales / Journal of Human Evolution 63 (2012) 781e795 787

and improved, it should be possible to see broad regional patterns.
Sites are associated with different climatic regimes according to
their age as listed in Appendix C.
It is challenging to associate an archaeological site with specific
warm or cold fluctuations documented within the oxygen isotope
record. van Andel (2003) remedy this problem by dividing OIS-3
into broad subunits, reproduced in Table 1. These subunits serve
as the basis for the climatic regime used to interpret archaeological
sites. Many phases can be unambiguously associated with one of
the four simulations: warm phase, cold phase, glacial maximum,
and modern conditions. The transitional phases of OIS-4 and OIS-3
are more uncertain, but the warm climate model was chosen for
these projections. This choice reflects a more skeptical perspective
on Neanderthal technological capabilities. In other words, Nean-
derthals would have needed less clothing to live at a given location
during warm times than cold times. Given the uncertainty in some
site dates, this approach is a more conservative assessment of
Neanderthal behavior.
Results
Percent body covered
Beta regressions for the maximum and minimum percentage of
the body covered by clothing were performed using numerous
iterations of environmental variables. The set of variables with the
lowest AIC value was selected as the model that best predicts the
maximum/minimum percentage of body covered by clothing.
A comparison of the different models that were investigated is
listed in the Supplementary Online Material (Table S1). The equa-
tions representing the best models for predicting the percentage of
the body covered by clothing are found in Table 2.
As the R2p values of 0.6259 and 0.4288 for the maximum and
minimum clothing models may not be intuitively revealing, an
additional analysis was conducted to clarify how well these
equations predict clothing use within the existing dataset. The
approach used was based on the Prediction Sum of Squares
statistic (PRESS), which calculates a predicted value for the i-th
observation using all other observations (Allen, 1974). In a similar
fashion, the clothing of each hunter-gatherer group was predicted
using a beta regression equation based on the other cases.
However, rather than summing the squares of the errors as in
PRESS, the mean absolute error was calculated. This assessment
yields an error value on the same scale as the original data and can
be interpreted more straightforwardly. The mean absolute error
for the maximum and minimum clothing models is 0.135 and
0.152, respectively. In other words, when the maximum clothing
model predicts that a hunter-gatherer group will cover 30.0% of
their bodies with clothing, the observed value would fall between
16.5% and 43.5% (0.300  0.135) on average. A second method, the

Figure 4. Predicted maximum amount of body covered by clothing for anatomically modern humans. Aurignacian and Gravettian sites displayed according to climatic regime: a)
Warm phase of OIS-3 [59e37 ka BP], b) cold phase of OIS-3 [37e27 ka BP], c) Last Glacial Maximum [27e16 ka BP], and d) Holocene [12 ka BP to present] (modern climate model).
788 N. Wales / Journal of Human Evolution 63 (2012) 781e795
median absolute error rate provides a further way of interpreting
the amount of error and is not affected by outliers like MAE
(Swanson et al., 2011). The median absolute errors for the
maximum and minimum clothing modes were 0.092 and 0.113,
indicating, for example, that half of the predicted values for the
maximum amount of clothing worn by hunter-gatherer groups are
less than 9.2% away from the observed values. These error values
demonstrate that while the models are not perfect, they work
relatively well, especially considering the fact that the observed
data comes from different anthropological sources with multiple
sources of variability.
Figs. 3e6 depict clothing predictions for Neanderthals and
modern humans across Europe and temporal periods.
Head, hands, and feet covered
Numerous logistic regression equations were tested for their
ability to predict the use of clothing to cover individual body parts.
The best models to predict the use of clothing on individual body
parts are listed in Tables 3 and 4, accompanied by the necessary
statistics (Peng et al., 2002). While the goodness-of-fit p-values
and the R2L statistics indicate the models are robust, one can also
investigate how well the models correctly predict cases using
Table 5. The successfulness of a logistic regression model can be
intuitively understood by comparing its predictions against a naïve
baseline model based on the more common value (Hair et al., 1987).
Figure 5. Predicted minimum amount of body covered by clothing for Neanderthals.
For instance, more hunter-gatherer groups were recorded as leaving
the feet uncovered than covered: 129 versus 103 groups. By pre-
dicting that no groups will use clothing on the feet, one correctly
guesses 55.6% of the cases. In contrast, the logistic equation
correctly predicts 84.5% of the groups that cover the feet and 93%
that keep the feet uncovered, scoring 89.2% overall. All three models
improve over the chance baseline, however, in order to be consid-
ered a ‘good’ model, Hair et al. (1987) suggest that models should
improve over the baseline by a factor of 1.25. Multiplying the feet
model baseline 55.6% by 1.25 gives a required score of 69.5%. With
an overall score of 89.2%, the logistic equation for clothing on the
feet can be considered a ‘good’ model. Likewise, the logistic equa-
tion for clothing on the hands has an overall score of 89.6%,
exceeding 1.25 times the naïve model score of 65.2%. The baseline
score for the head model, however, is abnormally high because only
16.8% of hunter-gatherer groups cover their heads in the dataset (39
out of 232 groups). To handle very unequal group sizes, the
proportion chance criterion is used (Hair et al., 1987). Squaring and
summing the proportion of each category gives a baseline for the
head model of 0.720 (0.1682 þ 0.8322). The logistic regression model
for the use of clothing on the head correctly predicts 95.3% of
hunter-gather groups, surpassing the portion chance criterion by
a factor of 1.25 and demonstrates that it, like the other two models,
is a satisfactory predictive model.
Predicted maps for clothing use on individual body parts are
found in Figs. 7e9.
 N. Wales / Journal of Human Evolution 63 (2012) 781e795 789

Figure 6. Predicted minimum amount of body covered by clothing for anatomically modern humans.

Discussion Predictions for the maximum amount of body covered by

One of the most productive ways to explore these newly
developed clothing models is by comparing projected patterns
against archaeological site distributions. In other words, although
clothing patterns have been predicted for all of Europe, Neander-
thals and modern humans inhabited only specific regions of the
continent during different temporal periods. Based on site distri-
butions, one can infer prehistoric clothing variability and changes
in human behavior.
clothing provide insights into how Neanderthals coped over the
winter. Fig. 3 shows that many Mousterian sites are found in
locations with environmental conditions that require 70e80% of
the body to be covered by clothing. This high percentage indicates
that it was essential for Neanderthals to fabricate clothing. To gain
perspective on how much and what kind of clothing this estimate
may represent, consider the following hunter-gatherer examples
that fall into this interval. The clothing of Plains Cree men of Alberta
and Saskatchewan, as described by Mandelbaum (1940), commonly

Table 3
Logistic equations to predict the use of clothing on specific body parts.

Model Equation R2L Test c2 df p-value

a
Head covered e
1:076
0:419*STD CMAT 0.725 Omnibus 152.417 1 < 0.001
1 þ e
1:076
0:419*STD CMAT HosmereLemeshowb 6.314 8 0.612
Hands covered e2:722
0:451*STD CMAT 0.681 Omnibusa 202.418 1 < 0.001
1 þ e2:722
0:451*STD CMAT HosmereLemeshowb 3.949 8 0.862
Feet covered e0:982
0:330*MCM 0.662 Omnibusa 211.043 1 < 0.001
1 þ e0:982
0:330*MCM HosmereLemeshowb 5.279 8 0.727

MCM: mean temperature of the coldest month ( C); STD_CMAT: Steadman wind chill calculated using the mean annual temperature ( C) and average annual wind speed (m/s)
at 10 m above the ground.
a
Omnibus test evaluates whether the final model fits the data better than the null model. The null hypothesis that the overall model does not improve over the baseline
model is rejected when p < 0.05.
b
HosmereLemeshow statistic tests the goodness-of-fit of actual observations to the final model. The null hypothesis of a well-fitting model cannot be rejected when
p > 0.05.
790 N. Wales / Journal of Human Evolution 63 (2012) 781e795

Table 4
Variables statistics for logistic regression models.

Model Variable Coefficient (S.E) Wald p-value Odds ratio (95% CI)
Head covered Constant
1.076 (0.361) 8.882 0.003 0.341
STD_CMAT
0.419 (0.074) 32.310 < 0.001 0.658 (0.569, 0.760)
Hands covered Constant 2.722 (0.576) 22.326 < 0.001 15.204
STD_CMAT
0.451 (0.076) 35.171 < 0.001 0.637 (0.549, 0.739)
Feet covered Constant 0.982 (0.311) 9.975 0.002 2.670
MCM
0.330 (0.050) 43.409 < 0.001 0.719 (0.652, 0.793)

consisted of a leather breechcloth, hide leggings, moccasins with
grass insulation, a hat partially covering the head, and a buffalo
robe over the torso and one shoulder. According to the methods
described above, this attire corresponds to 77.5% of the body. In
a similar fashion, Boas (1890) records that Tlingit women from the
Pacific Northwest Coast wore cedar-bark petticoats, shirts,
mountain-sheep wool blankets, and waterproof hats made of roots.
They frequently had bare lower legs and feet, equivalent to 77% of
the body covered. The common dress of men and women from
southern Australia also covered between 70 and 80% of the body.
Groups such as the Jaralde wore possum skin cloaks over the torso
and legs, but kept the feet, one arm, and head unprotected
(Beveridge, 1883). While there are many ways to cover three-
quarters of the body with clothing, the significance of these
examples is that a simple fur draped around the waist would have
been insufficient for Neanderthals in the coldest parts of their
range. Rather, it would have been necessary to cover the torso
and part of the appendages or head. Even though most
hunter-gatherers rely on tailored clothing to cover 70% of the body,
Neanderthals arguably could have survived without complex (i.e.,
non-tailored) clothing, as was done by the Jaralde and other
Australian groups. By wearing an animal skin fashioned much like
a toga, Neanderthals would have covered a suitable amount of their
bodies during the winter. The lack of clothing-specific artifacts at
Neanderthal sites further suggests that no attempts were made to
sew animal skins into tight-fitting garments. Instead, Neanderthals
likely donned the hides of large mammalian prey species, including
aurochs, horses, and mammoths (Patou-Mathis, 2000; Bocherens
et al., 2005). If necessary, simple thongs of leather or sinew could
have been used to secure furs to a Neanderthal’s torso, although
this falls outside the traditional definition of tailored clothing.
It is also valuable to determine just how little clothing was
necessary for some Neanderthals to survive the winter. In the most
southwestern parts of the Neanderthal range, the maximum
amount of body covered by clothing ranges between 20 and 40% for
OIS-3 and the glacial phase of OIS-4. For Neanderthals living in
southwestern Iberia, winter clothing could have been quite simple,
covering just the torso or the genitals and upper legs.
The central conclusion from the maximum clothing model that
it was essential for many Neanderthals to cover a significant
portion of their bodies is partially consistent with Sørensen’s
(2009) calculations that Neanderthals needed clothing and blan-
kets to survive an average winter night. However, while he argues
that tailored clothing was essential, clothing patterns from recent
hunter-gatherers suggest that this is not necessarily the case. The
projections from this study build upon Sørensen’s (2009) findings
by providing a much broader understanding of Neanderthal
clothing across geographical regions and in different time periods.
It is now possible to see that Neanderthal clothing requirements in
northern Germany during OIS-5e are not an isolated case. Rather,
much of the Neanderthal range during OIS-5e and later periods
required significant protection against the elements.
Projections of the minimum amount of the body covered by
clothing are indicative of warm weather attire. The distribution of
Mousterian sites in OIS-5e and OIS-3 indicates that Neanderthals
needed to cover at most 40% of their body with clothing during the
summer. Again, there are many ways to reach this degree of body
coverage, but in general most hunter-gatherers do so by covering
their torso and genitals. For example, the summer attire of Mattole
men of northern California consists of a sleeveless buckskin shirt
covering the torso and reaching just past the genitals, approxi-
mately 37% of the body (Nomland, 1938). For Neanderthals living in
milder climates, such as along the Mediterranean, less than 20% of
the body would be expected to be covered during warm months.
Such a percentage can be reached by wearing a garment over the
genitals and upper legs, as is done by the Semang men of Thailand,

Table 5
Number of hunter-gatherer groups that use clothing on specific body parts.

Body part Model Observed Predicted Correct in category Overall correct in model

Covered Uncovered
Head Naïvea Covered 0 39 0.0% 83.2%
Uncovered 0 193 100.0%
b
Logistic regression Covered 33 6 84.6% 95.3%
Uncovered 5 188 97.4%
a
Hands Naïve Covered 0 80 0.0% 65.2%
Uncovered 0 150 100.0%
Logistic regressionc Covered 67 13 83.8% 89.6%
Uncovered 11 139 92.7%
Feet Naïvea Covered 0 103 0.0% 55.6%
Uncovered 0 129 100.0%
d
Logistic regression Covered 87 16 84.5% 89.2%
Uncovered 9 120 93.0%
a
The naïve model predicts group membership based on the more common value and provides the baseline against which the logistic regression model is tested.
b
Head model: Sensitivity ¼ 33/(6 þ 33)*100% ¼ 84.6%. Specificity ¼ 188/(188 þ 5)*100% ¼ 97.4%. False positive ¼ 5/(33 þ 5)*100% ¼ 13.2%. False negative ¼ 6/(6 þ 188)
*100% ¼ 3.1%.
c
Hands model: Sensitivity ¼ 83.8%. Specificity ¼ 92.7%. False positive ¼ 14.1%. False negative ¼ 8.6%.
d
Feet model: Sensitivity ¼ 84.5%. Specificity ¼ 93.0%. False positive ¼ 9.4%. False negative ¼ 11.8%.
 N. Wales / Journal of Human Evolution 63 (2012) 781e795
Figure 7. Probability that Neanderthals covered their heads with clothing.
who traditionally wear a skirt made of pounded bark (Brandt,
1961). Some Neanderthal populations may well have lived
completely nude in the summer.
Predictions for whether Neanderthals covered specific body
parts also yield interesting results. The models predict that no
Neanderthals would have covered their heads using large hats or
hoods. According to the models, it is also possible that Neanderthal
populations covered their hands with clothing, with some sites
having nearly a 90% likelihood. The use of clothing on the feet
follows a very similar pattern. The vast majority of Mousterian sites
fall in regions with a greater than 50% chance of covering the feet,
with some sites at a 95% likelihood. The use of footwear is another
point in which this model agrees with that of Sørensen (2009).
However, footwear does not have to be particularly advanced to
protect the feet from the elements. Sørensen (2009) notes that
small holes in a shoe permit wind to pass through and lead to heat
loss. While this is true with modern shoes that use stitched textiles,
simpler protective footwear, such as fur wrapped around a foot and
tied with a leather thong, can provide insulation while being
impervious to wind. This interpretation is consistent with
Trinkaus’s (2005) hypothesis that Neanderthals may have used
insulating footwear that did not provide traction or protection
against rough terrain. In general, these results suggest that some
Neanderthals continually lived in places where most modern
humans would cover their hands and feet. Thus it is reasonable to
expect that some populations relied upon such clothing articles,
791
however, many Neanderthals in Southern Europe would have lived
comfortably without them.
Predictive models for modern humans during the late Pleisto-
cene shed new light on clothing requirements during the Upper
Paleolithic. As seen in Figs. 4 and 6, early Upper Paleolithic modern
humans occupied harsher environments than their Neanderthal
predecessors. Additionally, the modern human morphology leads
to greater heat loss that must be supplemented by using more
clothing. The sites of OIS-3 modern humans are shown to
frequently be located in places that require 80e90% of the body to
be covered by clothing during the winter. By the LGM, the pattern
becomes more exaggerated and modern humans in Eastern Europe
would have covered more than 93% of their bodies. Among recent
hunter-gatherers, this degree of coverage is only found in high
latitudes, often requiring heavy parkas and tailored leggings. Only
a few body parts such as part of the head or hands would be
exposed to the elements. Arguably, this indicates that modern
humans had advanced technological skills in clothing manufacture.
The findings from this study are significant on a number of
levels. For one, the models provide an objective method to infer
prehistoric clothing. In addition to using these equations for
Neanderthals and Upper Paleolithic modern humans, the same
models can be applied to other hunter-gatherers in different loca-
tions and time periods. For instance, it would be possible to infer
the clothing patterns of foragers who disappeared before being
visited by ethnographers. When clothing is understood as an
792 N. Wales / Journal of Human Evolution 63 (2012) 781e795
Figure 8. Probability that Neanderthals covered their hands with clothing.
integral part of hominin survival strategies, these models provide
a novel method to infer hominin behavioral patterns. Interpreta-
tions of Paleolithic behaviors frequently rely upon stone tools,
butchered fauna, human fossils, items of personal adornment, and
artifacts related to ritual activities. The clothing models presented
here can be used in conjunction with such evidence to provide
a more comprehensive understanding of hominin behaviors. This
study shows that Neanderthals and Upper Paleolithic modern
humans effectively engineered their personal space in order to
retain heat and keep out the elements. A further strength of these
clothing models is their ability to evaluate behavioral variability at
a continental scale. As highlighted by Shea (2011), the concept of
behavioral variability is a theoretically sound way to investigate
behavioral changes over the course of human evolution. It provides
an alternative option to the flawed concept of ‘behavioral moder-
nity’ (see McBrearty and Brooks, 2000). In addition, these data can
be used to determine how Neanderthals and modern humans
differentially responded to their environments. Compared with
Neanderthals, modern humans in OIS-3 frequently occupied
regions where tailored clothing was required. This indicates that
modern humans spent more time preparing and maintaining
clothing articles than Neanderthals.
The differences in clothing patterns between Neanderthals and
modern humans may have played an important cultural role.
Tailored clothing often includes elements of personal adornment
and group affiliation, while non-tailored clothing generally does
not (Gilligan, 2010b). Thus, if a Neanderthal draped in furs
encountered a group of modern humans wearing tight-fitting
garments with beads and symbolic paint, it is possible that
clothing would have acted as a cultural boundary, more impervious
than a biological one. While speculative, this idea is consistent with
genetic research that finds evidence for gene flow between Nean-
derthals and modern humans in the Near East but not Eurasia
(Green et al., 2010). In a warm environment, like the Levant, both
populations would have worn simple clothing, if anything at all,
and the cultural divide might have been less pronounced.
Considering these clothing differences, it is also tempting to
wonder what role clothing might have played in the extinction of
the Neanderthals. Gilligan (2007) hypothesizes that the Neander-
thals’ lack of tailored clothing led to their downfall as the climate
deteriorated toward the LGM, due to hypothermia. The models
produced in this project provide a new way of looking at this
question. The fact that Neanderthals frequently inhabited locations
where they needed little protection from the elements makes it
unlikely that the entire Neanderthal species went extinct due to
insufficient thermal insulation. In other words, based on our
current understanding of OIS-3, some Neanderthals are expected to
have survived in glacial refugia despite the harsh conditions of
the LGM.
The true cause(s) of Neanderthal extinction may never be
known, but they were likely multifaceted and varied over time,
perhaps analogous to the multicausal extinction of the Beringian
 N. Wales / Journal of Human Evolution 63 (2012) 781e795
Figure 9. Probability that Neanderthals covered their feet with clothing.
woolly mammoth (MacDonald et al., 2012). Many argue that
modern humans are to blame in the extinction of Neanderthals
(Banks et al., 2008), but researchers point to different driving
factors, including dietary breadth (Hockett and Haws, 2005),
mobility patterns (Barton et al., 2011), and caloric requirements
(Snodgrass and Leonard, 2009). In reality, such factors could be
working in unison. Differences in clothing patterns can be added
to the list too, although in a less direct manner than suggested by
Gilligan (2007). Instead, clothing may be indirectly linked to
Neanderthal extinction by limiting the species’ range, thereby
keeping population levels low. The use of more sophisticated
clothing by modern humans would have provided marginal
evolutionary advantages by conserving heat and permitting
expansion into colder climates (Froehle and Churchill, 2009).
Such differences in population sizes and reproductive and
mortality rates can ultimately lead to population turnover events,
often in surprisingly short spans of time (Zubrow, 1989). In this
way, although the non-tailored clothing of the Neanderthals was
satisfactory for tens of thousands of years, it proved suboptimal
when confronted by another hominin occupying a similar niche.
Based upon current paleoenvironmental reconstructions, the
simple clothing of the Neanderthals did not lead to mass
extinction via hypothermia. However, it can be considered to be
part of a larger cultural and behavioral package that gave Nean-
derthals an evolutionary disadvantage and ultimately led to their
demise.
793
Conclusion
Attempting to infer the clothing of extinct hominins is
immensely challenging. Like all modeling exercises, this research is
a simplification of complex issues and relies on assumptions that
may not precisely correspond with reality. However, such obvious
limitations do not invalidate the overall findings. This study
corroborates many earlier conclusions about Neanderthal clothing,
but it also signifies a great advancement in the understanding of
prehistoric clothing variability. Rather than relying solely on esti-
mates of Neanderthal metabolic rates and body surface area, the
models presented here are grounded upon empirical clothing data
from recent hunter-gatherers. It is now possible to predict Nean-
derthal clothing patterns for Europe during OIS-3, including
whether clothing was used on specific body parts. Indeed, it is clear
that Neanderthals would have required clothing to survive from
70,000 years before present until their extinction. Yet the exact
amount of clothing an individual would have needed depended on
their location and the climatic regime. For Neanderthals in the
coldest regions, it was necessary to cover 70e80% of the body
during the winter. In one configuration this amount is reached by
covering the torso, genitals, and a portion of the appendages. Other
Neanderthals could have survived the cold season with far less
clothing, covering only a quarter of their bodies. Contrary to some
hypotheses, these models suggest that effectively all Neanderthal
populations could survive without advanced tailored clothing.
794 N. Wales / Journal of Human Evolution 63 (2012) 781e795
These findings aid archaeologists and paleoanthropologists in
understanding Neanderthal biological limitations, technological
capabilities, and behavioral variability. It is also possible to see that
modern humans inhabited areas that required more clothing than
Neanderthals, indicating they relied upon tailored clothing. These
differences in clothing patterns would have provided marginal
advantages for modern human survival, and indirectly may have
been one of the factors in the extinction of the Neanderthals. In the
end, we are left with an unparalleled view into Paleolithic clothing,
which may otherwise be impossible to reach using conventional
archaeological methods.
Acknowledgments
This research would not have been possible without guidance
and support from the late Lewis Binford and Amber Johnson,
Truman State University. In addition to helping plan how to collect
and analyze clothing data, they provided financial support in the
summer following my undergraduate career. I would also like to
thank Daniel Adler, University of Connecticut, for encouraging me
to expand the scope of this study and for editing suggestions.
Special thanks to Randi Garcia for advice on statistics.
Appendix A to C. Supplementary data
Supplementary data related to this article can be found at http://
dx.doi.org/10.1016/j.jhevol.2012.08.006.
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