Journal of Human Evolution xxx (2013) 1e8

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Journal of Human Evolution

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Extreme mobility in the Late Pleistocene? Comparing limb biomechanics among

fossil Homo, varsity athletes and Holocene foragers
Colin N. Shaw a, b, *, Jay T. Stock a
a
PAVE Research Group, Department of Archaeology and Anthropology, University of Cambridge, Downing Street, Cambridge, Cambridgeshire CB2 3ER, UK
b
McDonald Institute for Archaeological Research, Department of Archaeology and Anthropology, University of Cambridge, Cambridge, UK

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

Article history: Descriptions of Pleistocene activity patterns often derive from comparisons of long bone diaphyseal
Received 31 July 2012 robusticity across contemporaneous fossilized hominins. The purpose of this study is to augment existing
Accepted 9 January 2013 understanding of Pleistocene hominin mobility patterns by interpreting fossil variation through com-
Available online xxx
parisons with a) living human athletes with known activity patterns, and b) Holocene foragers where
descriptions of group-level activity patterns are available. Relative tibial rigidity (midshaft tibial rigidity
Keywords:
(J)/midshaft humeral rigidity (J)) was compared amongst Levantine and European Neandertals, Levantine
Peripheral Quantitative Computed
and Upper Palaeolithic Homo sapiens, Holocene foragers and living human athletes and controls. Cross-
Tomography (pQCT)
Fossil hominin
country runners exhibit significantly (p < 0.05) greater relative tibial rigidity compared with swimmers,
Diaphyseal cross-section and higher values compared with controls. In contrast, swimmers displayed significantly (p < 0.05) lower
Humerus relative tibial rigidity than both runners and controls. While variation exists among all Holocene
Tibia H. sapiens, highly terrestrially mobile Later Stone Age (LSA) southern Africans and cross-country runners
display the highest relative tibial rigidity, while maritime Andaman Islanders and swimmers display the
lowest, with controls falling between. All fossil hominins displayed relative tibial rigidity that exceeded,
or was similar to, the highly terrestrially mobile Later Stone Age southern Africans and modern human
cross-country runners. The more extreme skeletal structure of most Neandertals and Levantine
H. sapiens, as well as the odd Upper Palaeolithic individual, appears to reflect adaptation to intense and/
or highly repetitive lower limb (relative to upper limb) loading. This loading may have been associated
with bipedal travel, and appears to have been more strenuous than that encountered by even university
varsity runners, and Holocene foragers with hunting grounds 2000e3000 square miles in size. Skeletal
variation among the athletes and foraging groups is consistent with known or inferred activity profiles,
which support the position that the Pleistocene remains reflect adaptation to extremely active and
mobile lives.
Ó 2013 Elsevier Ltd. All rights reserved.

Introduction
The relationship between biomechanically imposed loads and
skeletal adaptation is quite complex (Pearson and Lieberman,
2004). Nevertheless, experimental research supports the general
concept of bone functional adaptation, and the use of diaphyseal
cross-sectional properties to explore patterns of habitual behavior
among groups (Currey, 1984; Lanyon and Rubin, 1984; Martin and
Burr, 1989; Rubin et al., 1990; Lanyon, 1992; Ruff et al., 2006). An-
alyses of the femoral and tibial diaphyseal structure of Pleistocene
Homo have concluded that while morphological variation is appa-
rent, there has been little shift in locomotor ranging levels amongst
* Corresponding author.
E-mail addresses: cns30@cam.ac.uk, cshaw111@mac.com (C.N. Shaw).
the foraging populations from this period (c.f. Trinkaus and Ruff,
2012). Interpretations of Pleistocene mobility patterns, however,
might be augmented through direct comparisons with more recent
foragers and living humans where activity patterns have been
better documented.
Early analyses concluded that the Neandertal lower limb was
better adapted to extended bouts of heavy physical activity com-
pared with those of early modern humans (Trinkaus, 1989). This
finding led to the suggestion that Neandertals spent comparatively
more time moving continuously and/or vigorously across the
landscape. This interpretation was later reconsidered (c.f. Trinkaus,
1997) following the implementation of methodology that better
controlled for the influence of body size on diaphyseal geometric
properties (Ruff, 2000a). Later analyses of Neandertals and Levan-
tine anatomically modern Homo sapiens lower limb diaphyseal
strength concluded that, once properly standardised, differences

0047-2484/$ e see front matter Ó 2013 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.jhevol.2013.01.004

Please cite this article in press as: Shaw, C.N., Stock, J.T., Extreme mobility in the Late Pleistocene? Comparing limb biomechanics among fossil
Homo, varsity athletes and Holocene foragers, Journal of Human Evolution (2013), http://dx.doi.org/10.1016/j.jhevol.2013.01.004
2 C.N. Shaw, J.T. Stock / Journal of Human Evolution xxx (2013) 1e8
between these groups were negligible (Trinkaus and Ruff, 1999a,b).
More recently, Trinkaus and Ruff’s (2012) detailed assessment of all
known Pleistocene hominins concluded that despite a significant
change in lower limb diaphyseal morphology throughout the
Pleistocene, there appears to have been little change in the overall
robustness of the femur or tibia within Homo. When scaled
appropriately, femoral midshaft rigidity did not show a significant
trend from the Early to the Late Pleistocene, although there was
a suggestion of a slight reduction with the earlier Upper Palae-
olithic. In the tibial diaphysis, there were decreases in relative
cortical area, especially between the Early Pleistocene and later
human groups. Increased tibial diaphyseal circularity (antero-pos-
terior versus medio-lateral) was found between the Early Pleisto-
cene and later samples, and then again with the earlier Upper
Palaeolithic. The authors suggest that these results reflect the
importance of mobility among all Pleistocene foragers, and that
variation in femoral diaphysis shape may reflect differential rang-
ing patterns among later Upper Palaeolithic and Holocene groups
(e.g., Holt, 2003; Holt and Formicola, 2008), but not across Pleis-
tocene Homo.
Interpretations of prehistoric hominin activity patterns often
depend primarily upon comparisons of fossilized skeletal structure
(Trinkaus et al., 1994; Churchill et al., 1996; Holt, 2003; Holt and
Formicola, 2008; Marchi, 2008; Ruff, 2008a, 2009). In partial con-
trast, studies of pre- and protohistoric Holocene skeletal pop-
ulations often benefit from a greater breadth of associated
archaeological and ethnographic evidence for behavior, and larger
sample sizes with which to interpret within-population morpho-
logical variation. This type of contextual information allows for
more detailed descriptions of general group-level activity patterns
(Larsen et al., 1995; Stock and Pfeiffer, 2001, 2004; Auerbach and
Ruff, 2006; Stock, 2006; Wanner et al., 2007). A third, and rela-
tively unexplored, approach (see Trinkaus et al., 1994; Ruff et al.,
1998; Ruff, 2000b) compares fossilized hominins against groups
of living humans with known activity patterns and well docu-
mented skeletal morphology (i.e., Shaw and Stock, 2009a,b; Shaw,
2010, 2011). To inform previous interpretations and provide a fur-
ther test of Pleistocene mobility patterns, this study compares the
robusticity and ‘pattern’ of robusticity (tibial J/humeral J) in Pleis-
tocene hominins, highly terrestrially mobile and marine-adapted
Holocene foragers, and living human cross-country runners and
swimmers.
Two main hypotheses are tested: (1) populations with known
and inferred high terrestrial mobility (runners and Later Stone Age
(LSA) southern Africans, respectively) will share greater robusticity
of lower limb elements relative to upper; and (2) comparisons of
relative tibial rigidity will reveal that Pleistocene fossil hominins
were adapted to levels of terrestrial mobility that are comparable
with, or even exceed, that of highly trained living cross-country
runners and highly terrestrially mobile Holocene foragers.
Materials and methods
Modern human athletes and controls
A total of 50 modern human males between the ages of 19 and
30 participated in this study. Subgroups included varsity-level
distance runners (15) and swimmers (15), and non-athletic con-
trols (20). All were recruited from the University of Cambridge, UK.
The mean age of all subjects (runners (23.2
3.2), swimmers
(21.9
2.5), controls (21.6
2.5)) was 22.2 (
2.8) years. All athletes
began their competitive playing and training career during late
childhood/early adolescence (runners; 13.7 years of age, swim-
mers; 10.5 years of age), and had been competing in their respec-
tive sport for an average of 10 years (runners; 9.7 years, swimmers;
Please cite this article in press as: Shaw, C.N., Stock, J.T., Extreme mobility in the Late Pleistocene? Comparing limb biomechanics among fossil
Homo, varsity athletes and Holocene foragers, Journal of Human Evolution (2013), http://dx.doi.org/10.1016/j.jhevol.2013.01.004
10.3 years). Athletic, lifestyle, and medical history was obtained
through a detailed questionnaire (Shaw and Stock, 2009b). A single
scan from each of the arms and shanks of each participant resulted
in a total of 200 cross-sectional pQCT (peripheral Quantitative
Computed Tomography) images captured at 50% of limb segment
length using an XCT-2000 (Stratec Medizintechnik GmbH, Pforz-
heim, Germany). Raw cross-sectional pQCT images were imported
into Image J (http://rsb.info.nih.gov/ij/) and analyzed using
Moment Macro (http://www.hopkinsmedicine.org/FAE/mmacro.
htm).
Holocene skeletal material
The skeletal remains of two Holocene forager populations were
included in this study: 16 LSA southern African males and 15 pro-
tohistoric Andaman Islander (AI) males. The methods used to cal-
culate cross-sectional diaphyseal properties for these populations
have been described previously (Stock and Pfeiffer, 2001). Briefly,
cross-sectional dimensions were calculated using two separate
methods, the first involving CT images taken perpendicular to the
diaphysis at the section location, the second, a procedure that
combined information from casts of the periosteal contour and
biplanar radiographs (Trinkaus and Ruff, 1989).
Detailed descriptions of both populations are available in Stock
and Pfeiffer (2001). In summary, the LSA southern Africans date
from between 11,000 and 2000 BP (Before Present), and are
described as highly terrestrially mobile. Subsistence was charac-
terized by the hunting of small game, terrestrial foraging, and the
intensive exploitation of coastal marine resources (Deacon, 1993).
An early model predicted that the area over which a band would
hunt may have been as large as 2000e3000 square miles (Clarke,
1959). Resources were broadly distributed, and exploitation,
whether terrestrial or marine, was land-based. The terrain of the
southern African region is generally rocky, with considerable ver-
tical relief. Therefore, the exploitation of diverse terrestrial and
marine resources throughout this landscape would have required
the negotiation of rugged terrain.
The Andaman Islander foragers date to the late-1800s and had
mobility patterns that were constrained terrestrially and were
generally marine based. Dietary staples included various fruits and
yams in addition to wild pigs and honey, while marine hunting
provided fish, dugong and sea turtles (Myka, 1993). The Andama-
nese frequently used the canoe for transportation and food pro-
curement. Swimming was a common activity in both sexes, with
children learning to swim almost as soon as they could walk, and all
Andamanese, regardless of age and sex, spending up to several
hours in the water at a time (Man, 1883).
Pleistocene fossil hominin material
Table 1 lists the hominin fossil material included in this study, as
well as relevant provenance information. Analyses were performed
on data from three male and one female anatomically modern
Levantine H. sapiens (AMHS; Qafzeh 8, 9, Skhul IV and V ), three
male and two female Homo neanderthalensis (Spy 2, La Chapelle 1,
La Ferrassie 1, 2, Tabun C1), five male European Early Upper
Palaeolithic (EUP) H. sapiens (Paviland 1, Grotte des Enfants 4, Dolni
Vestonici 13, 14, 16), six male European (Gough’s Cave 1, Neussing 1,
Obercassel 1, Romanelli 1, Romito 3, Veyrier 1) and 20 male North
African (Wadi Halfa 2, 3, 24, 26, 31, 37; Jebel Sahaba 21, 29, 38, 39,
40, 42; Afalou 2, 13, 25, 27, 28; El Wad 10259, 10260, 10263) Late
Upper Palaeolithic (LUP) H. sapiens. Cross-sectional geometric
measurements for these fossil hominins, taken at the midshaft of
the tibia and humerus, were obtained from the relevant literature
(see Table 2 for citations).
 C.N. Shaw, J.T. Stock / Journal of Human Evolution xxx (2013) 1e8 3

Comparisons Results

Cross-sectional geometric properties used in this investigation Modern human athletes, controls, and Holocene foragers
are limited to polar second moments of area (J). J is an estimate of the
bones ability to resist diaphyseal torsion and (twice) average bending Table 2 displays raw tibial J and humeral J values, and relative
forces (Ruff, 2008b) that account for a high proportion of midshaft tibial rigidity (tibial J/humeral J) calculations, for each group. The
strains (Bertram and Biewener,1988). As such, J is a relevant indicator LSA southern Africans and runners display the highest relative
of a bone’s overall mechanical performance (Ruff et al., 1993). Raw tibial rigidity (4.56 and 2.76, respectively), while Andaman Is-
measures of J taken at the tibial and humeral midshaft were used to landers and swimmers display the lowest (2.36 and 1.84, respec-
calculate relative tibial rigidity (tibial J/humeral J). When right and tively). The relative tibial rigidity displayed by controls (2.63) falls
left elements were available, relative tibial J was calculated using the in the middle of all other groups. Differences in relative tibial ri-
equation ((L þ R tibia)/2)/((L þ R humerus)/2), if either element was gidity between these groups are presented in Table 3, and also Fig. 1.
unavailable, the value for the available side alone was used. Using The results indicate a separation of groups at the extremes; the
estimates of relative tibial rigidity, one can assess differences in the groups with the highest (LSA southern Africans) and lowest
distribution of robusticity in the upper and lower limbs that may not (swimmers) relative tibial rigidity are both significantly different
be apparent when comparing upper or lower limbs in isolation. High from all other groups (in all cases p < 0.05). No significant differ-
ratio values can reflect either high relative tibial robusticity or low ences were found among the groups (runners, controls and Anda-
humeral robusticity. Assessment of this data through bivariate plots man Islanders) that fall between these extremes. Table 4 provides
allows one to determine which factor is driving the ratio for different slope characteristics for the relationship between raw tibial J and
individuals and populations. Additionally, ratios avoid difficulties humeral J for all groups. The slope of the line that describes the LSA
associated with standardising for differences in body size across southern Africans is significantly steeper (falling outside of the 95%
species where mass and stature can be quite large. confidence interval, therefore p < 0.05), compared with all other
The first analysis utilized univariate analysis of variance (ANOVA)
to compare relative tibial rigidity among modern human athletes,
controls, and Holocene foragers. Differences in relative tibial rigidity Table 2
were contrasted using either Hochberg’s GT2 or Games Howell post- Relative tibial rigidity (tibial J:humeral J), and raw tibial and humeral J.
hoc tests, based on the results of Levene’s Test for equality of var-
n Mass Tibial J: Tibial J Humeral J
iance. Significance was recognized at p  0.05. Slope values, calcu- (kg) Humeral J (mm4) (mm4)
lated using Least Squares Linear Regression, were also compared Holocene
among these populations. The second set of analyses included living LSA SA 19 52.35 4.56 30,685 (6997) 6736 (2405)
human athletes and controls, Holocene foragers and fossil hominins. (4.94)
Restrictions imposed by the small number of individuals that make Runner 15 68.21 2.76 51,427 (10,824) 18,642 (3892)
(5.94)
up most Pleistocene groups necessitated that non-statistical com-
Control 20 69.68 2.63 39,091 (7893) 14,849 (3478)
parisons were performed. Ratios were ordered and plotted, and (11.67)
from this descriptive comparisons were produced. Andaman Is. 15 48.52 2.36 17,326 (2396) 7342 (2077)
(4.81)
Table 1 Swimmer 15 74.27 1.84 41,771 (7474) 22,658 (5779)
Fossil hominin provenance. (7.72)
Neandertals
Sex Date Provenance La Ferrassie 2 1 66.4a 5.03 30,631a 6084b,R
(ca. years BP) La Ferrassie 1 1 84.9a 4.63 62,285a 13,447b, R&L
Neandertal Tabun C1 1 63.3a 3.88 27,216a 7015c, R&L
Spy 2 M 36,000a Spy, Belgium Spy 2 1 85.5a 3.81 52,606a 13,795b, R&L
La Chapelle 1 M 47e56,000e Corrèze, France La Chapelle 1 1 81.1a 3.33 58,502a 17,578b, R&L
La Ferrassie 1 M 72,000d Dordogne, France AMHS
La Ferrassie 2 F 72,000d Dordogne, France Skhul V 1 68.8a 6.55 86,569a 13,215c, R&L

Tabun C1 F 122,000b Western Israel Skhul IV 1 69.0a 5.07 63,838a 12,590c, L

AMHS Qafzeh 9 1 63.3a 4.25 69,506a 16,346c, R

Qafzeh 8 M 100,000c Western Israel Qafzeh 8 1 e 2.81 65,702a 23,400c, R

Qafzeh 9 F 100,000c Western Israel Upper Palaeolithic

Skhul IV M 100,000c Western Israel EUP-Europe 5
75.76 3.28 47,807 (13,692)a,e 15,312 (4341)d,e,f,g
Skhul V M 100,000c Western Israel (5.02)a
Upper Palaeolithic LUP-Europe 6 68.18 2.91 48,279 (13,876)e 16,853 (5747)e,d
EUP-Europe M 24e29,000f Wales, Italy, Czech (8.07)e
Republic LUP-N. Africa 20 66.30 3.03 51,956 (11,483)e,h 18,152 (4709)e,h
LUP-Europe M 9100e18,000g France, Italy, (5.68)e,i
Germany
Data is presented as: mean (standard deviation).
LUP-N. Africa M EpiPalaeolithic e Natufianh Sudan, Israel,
J: polar second moment of area, Andaman Is.: Andaman Islander, LSA SA: Later Stone
Algeria
Age southern African, AMHS; anatomically modern Homo sapiens.
a
Semal et al. (2009). Fossil specimen humeral J calculated from skeletal elements from R (right-side), L
b
Grün and Stringer (2000). (left-side), R&L (average of right & left).
c
McDermott et al. (1993). Cross-sectional geometric property data for fossil hominins obtained from:
d a
Delporte (1984). Trinkaus and Ruff (2012).
e b
Verneau (1906). Trinkaus et al. (1994).
f c
Raynal (1990), Svoboda (1995), Trinkaus and Svoboda (2006), Jacobi and Trinkaus and Churchill (1999).
d
Higham (2008). Churchill (1994).
g e
Davies (1904), Bonnet (1913e14), Seligman and Parsons (1914), Verworn et al. Laura Shackelford (Personal communication).
f
(1919), Pittard and Sauter (1945), Narr (1977), Mallegni and Fabbri (1995), Alciati Trinkaus (2000).
g
et al. (2005). Sladek et al. (2000).
h h
Boule et al. (1934), McCown (1939), Anderson (1968), Greene and Armelagos Shackelford (2007).
i
(1972), Chamla (1978). Mass estimates for Jebel Sahaba 21 and 42 were not available for inclusion.

Please cite this article in press as: Shaw, C.N., Stock, J.T., Extreme mobility in the Late Pleistocene? Comparing limb biomechanics among fossil
Homo, varsity athletes and Holocene foragers, Journal of Human Evolution (2013), http://dx.doi.org/10.1016/j.jhevol.2013.01.004
4 C.N. Shaw, J.T. Stock / Journal of Human Evolution xxx (2013) 1e8

Table 3 Table 4
P-values for ANOVA comparisons of relative tibia torsional rigidity (tibial J:humeral Slope characteristics for tibial J versus humeral J among athletes, controls and Ho-
J) among modern human athletes and controls, and Holocene foragers. locene foragers.

Groups Runners Swimmers Controls LSA SA Andaman Is. Slope Std. error 95% confidence interval
Runners *.024 1.000 *<.001 1.000 Lower limit Upper limit
Swimmers *.031 *.001 *.007
Runners 1.515a 0.647 0.221 2.809
Controls *.001 1.000
Swimmers 1.110 0.184 0.742 1.478
LSA SA *.001
Controls 1.178 0.457 0.264 2.092
Andaman Is.
Andaman Is. 1.313 0.192 0.929 1.697
*Significant difference based upon ANOVA with Hochberg’s GT2 or Games Howell LSA SA 2.973b 0.353 2.267 3.679
post-hoc test, p  0.05. a
Slope for runners is significantly steeper compared against swimmers only.
LSA SA: Later Stone Age southern Africans, Andaman Is.: Andaman Islanders. b
Slope for LSA SA is significantly steeper compared with all other groups.

groups. Additionally, the slope of the line that describes runners is

significantly steeper than that which describes swimmers.
Pleistocene fossil hominins
Compared with all other groups, the relative tibial rigidity of the
Neandertals and Anatomically Modern H. sapiens (AMHS) are most
similar to the LSA southern Africans, while in general the Upper
Palaeolithic H. sapiens are more similar to the runners (Table 2). All
Neandertals and AMHS display higher relative tibial rigidity com-
pared with runners, while four of 11 of these individuals (Skhul IV
and V and La Ferrassie 1 and 2) display relative tibial rigidity (4.63e
6.55) that is higher than the LSA southern Africans (4.56). Addi-
tionally, unlike all other Upper Palaeolithic individuals, Paviland 1,
a European-EUP H. sapiens, displays relative tibial rigidity that is
also exceptionally high (6.10). When Paviland 1 is removed from
the calculation of relative tibial rigidity for the EUP-Europe
H. sapiens the average decreases from 3.28 to 2.73. Similarly,
three of the 20 LUP-North African specimens also display levels of
relative tibial rigidity that exceed or are comparable with the LSA
southern African sample: El Wad 10260 (5.54), El Wad 10252 (4.40)
and Jebel Sahaba 21 (4.90). Fig. 2, a bivariate plot of raw tibial ri-
gidity against raw humeral rigidity, provides a visualization of all
Holocene H. sapiens, Neandertals, AMHS, and the four highly
asymmetric Upper Palaeolithic specimens noted above. This plot
allows one to evaluate whether tibial or humeral properties are
driving relative tibial rigidity. The higher slope values (Table 4) that
describe the LSA southern African sample reflect a greater increase
in tibial rigidity relative to increases in humeral rigidity. The higher
relative tibial rigidity of the eight Pleistocene fossil hominins
identified above indicate that these individuals have even more
hypertrophied lower limbs, relative to their upper limbs, compared
with all other groups, including the LSA southern Africans. The
humeral rigidity of these Pleistocene hominins is not out of place
amongst modern humans, falling between the small-bodied Ho-
locene foragers and the athletes. Their tibial morphology, in con-
trast, is at the high end, indicating that the relative tibial rigidity in
the fossil specimens is unique, driven by high tibial rigidity rather
than unusually weak humeri (however, see the end of the
Discussion section for more on the potential problems associated
with interpreting raw J values).
Table 5 provides predictions of tibial J for each of the Pleistocene
fossil hominins using equations derived from the relationship of
raw tibial J and raw humeral J constructed using the swimmer,
runner, LSA southern African, and Andaman Islander samples. For
virtually all Neandertals and AMHS, tibial rigidity was most accu-
rately predicted using the equation calculated from the LSA
southern African sample. The exception to this trend is that the
most accurate predictions were made for La Ferrassie 2, Qafzeh 8
and the Upper Palaeolithic H. sapiens using the equation derived
from the runner sample. The most accurate predictions of tibial
rigidity for Neandertals and Upper Palaeolithic individuals are
relatively close to the actual measured values. By contrast, pre-
dictions for the AMHS appear well below the actual measured
value.

Figure 1. Relative tibial strength (tibial J/humeral J) for modern human athletes and
controls, and Holocene foragers. Boxes represent the 25the75th percentile range, with
whiskers extending to the maximum and minimum values within 1.5 box lengths. Figure 2. Bivariate plot of tibial rigidity (J) versus humeral rigidity (J) for modern
Outliers are indicated with an o. human athletes, Holocene foragers, and Pleistocene hominins.

Please cite this article in press as: Shaw, C.N., Stock, J.T., Extreme mobility in the Late Pleistocene? Comparing limb biomechanics among fossil
Homo, varsity athletes and Holocene foragers, Journal of Human Evolution (2013), http://dx.doi.org/10.1016/j.jhevol.2013.01.004
 C.N. Shaw, J.T. Stock / Journal of Human Evolution xxx (2013) 1e8 5

Table 5
Predicted tibial torsional rigidity (J) from humeral torsional rigidity (J) for fossil hominins.

Fossil Tibial J Tibial J (predicted)

hominins (measured) Runner Swimmer Andaman is. LSA SA
equationa equationb equationc equationd
Neandertals
Spy 2 52,606 44,076 31,933 25,415 48,328
La Chapelle 1 58,502 49,807 36,132 30,382 59,574
La Ferassie 1 62,285 43,548 31,546 24,958 47,292
La Ferassie 2 30,631 32,393 23,373 15,290 25,401
Tabun C1 27,216 33,804 24,407 16,513 28,171
AMHS
Qafzeh 9 69,506 47,940 34,764 28,764 55,911
Skhul IV 63,838 42,251 30,596 23,833 44,746
Skhul V 86,569 43,197 31,289 24,653 46,602
Qafzeh 8 65,702 58,628 42,595 38,027 76,884
Upper Palaeolithic
EUP-Europe 47,807 46,374 33,617 27,407 52,837
LUP-Europe 48,279 48,708 35,327 29,430 57,418
LUP-N. Africa 51,956 50,676 36,769 31,136 61,280

Bold: most accurate prediction of tibia torsional rigidity (J). Prediction equations derived using Least Squares Linear Regression.
a
1.515 * Humeral J þ 23176.071.
b
1.110 * Humeral J þ 16620.256.
c
1.313 * Humeral J þ 7301.957.
d
2.973 * Humeral J þ 7314.087.

Discussion same intensity or repetitiveness) while locomoting bipedally.

Athletes and controls
Comparisons of relative tibial rigidity (tibial rigidity (J)/humeral
rigidity (J)) among modern human athletes and controls reveal that
despite similarities in body size and age, significant differences
exist. As expected, the distribution of skeletal rigidity between the
upper (humeral) and lower (tibial) limbs among athletes and
controls is consistent with their different athletic loading histories.
The main factors separating the two athlete groups are the bio-
mechanical patterns involved in cross-country running and
swimming. Regardless of the swimming stroke chosen, in order to
propel oneself near the top of the water, adequate force must be
generated for forward movement to be achieved. Although pro-
pulsion is profoundly aided through the action of kicking legs,
a large proportion of the work, and therefore a large proportion of
the biomechanical loading, is performed by the upper limbs. The
upper limb biomechanics of swimming contrast with those of
running where the arms are essentially ‘unloaded’, moving in op-
position to the legs to aid balance in the upper body and head. In
contrast, the striding gait of human bipedal runners requires the
lower limbs to generate sufficient force to counteract the force of
gravity on body weight and propel the individual forward at
a chosen speed. It is likely that the biomechanics of swimming
require a greater degree of upper limb loading compared with that
imposed on the lower limbs. Conversely, bipedal running loads the
lower limbs to a greater degree than it does the upper limbs.
Therefore, the opposing lower to upper limb bone strength patterns
of swimmers and cross-country runners are in keeping with the
expected influences of the habitual activity patterns performed
throughout adolescence.
Additionally, swimmers displayed significantly lower relative
tibial rigidity compared with controls, while no significant differ-
ences were found between runners and controls for the same
measurement. Given the variety of behavioral differences between
both runners and swimmers compared with controls, this latter
result is initially surprising. While controls did not engage in
competitive cross-country running they would have loaded their
lower limbs in a manner comparable to runners (yet not with the
Controls, however, did not perform upper limb loading activities
comparable with habitual competitive swimming. Given the rela-
tive lower limb loading similarities between controls and runners,
and the lack of upper limb loading similarities between controls
and swimmers, the differences described here are in keeping with
what should be expected.
Athletes, controls, and Holocene foragers
Ethnographic descriptions of the behavioral patterns of the
Holocene forager groups show general similarities with the sport-
ing histories of the modern human runners and swimmers. Spe-
cifically, the activity patterns of the swimming and watercraft using
Andaman Islanders are, in a very general sense, comparable with
those of the modern human swimmers. In contrast, the activity
patterns of the highly terrestrially mobile LSA southern Africans are
assumed to be roughly comparable with the runners. If skeletal
adaptations to general distance running and swimming activities
are somewhat consistent among groups, one might expect that the
distribution of diaphyseal rigidity between the lower and upper
limbs of foragers to be generally comparable to the athletes.
The finding of significantly greater relative tibial rigidity in the
LSA southern Africans compared to runners does not support the
presumption of similar morphological patterning in two ‘terres-
trially mobile’ groups, and may in fact reflect divergent mobility
patterns. Nevertheless, because the LSA southern Africans and
runners display the two highest relative tibial rigidity values the
differences between these groups likely reflect variation in degree
and not necessarily kind. Throughout adolescence, the training of
these athletes often involved running six or more days per week
(5.3 total hours and up to 80e100 total miles; Shaw and Stock,
2009b, unpublished data). The skeletal remains of the LSA south-
ern Africans may reflect a level of terrestrial mobility that was
comparatively more intense and/or more frequently performed.
Additionally, differences in terrain relief might have resulted in
greater lower limb loading in LSA southern Africa compared with
modern Britain. Marchi (2008) has suggested that the high tibial
and femoral robusticity found in Neolithic Ligurian populations was
a consequence of terrestrial locomotion over uneven terrain. It has

Please cite this article in press as: Shaw, C.N., Stock, J.T., Extreme mobility in the Late Pleistocene? Comparing limb biomechanics among fossil
Homo, varsity athletes and Holocene foragers, Journal of Human Evolution (2013), http://dx.doi.org/10.1016/j.jhevol.2013.01.004
6 C.N. Shaw, J.T. Stock / Journal of Human Evolution xxx (2013) 1e8
been shown that tibial strain in humans is significantly higher
during running performed on uneven ground compared with
treadmill running (Milgrom et al., 2003), and also during uphill and
downhill walking and running compared with travel over even
surfaces (Burr et al., 1996). If the age at which runners began seri-
ously training (w13 years of age, Shaw and Stock, 2009b) was later
than when LSA children began undertaking ‘long distance’ bipedal
travel, this could also go some way to explaining differences in
relative tibial rigidity between these groups.
Results comparing swimmers and Andaman Islanders are
similarly complicated. While the Andamanese have been descri-
bed as habitual paddlers and swimmers ethnographic accounts
suggest that these people also foraged terrestrially when the op-
portunity arose. By contrast, swimmers trained for an average of
9.3 hours per week throughout adolescence (Shaw and Stock,
2009a). It is therefore not overly surprising that significant dif-
ferences were found between these two groups regardless of the
fact that they displayed the two lowest relative tibial rigidity
values.
Athletes, controls, Holocene foragers, and Pleistocene fossil hominins
The relative tibial strength of the Pleistocene fossil hominins is
quite high, and in a number of cases far exceeds the upper to lower
limb asymmetry displayed by all Holocene foragers and athletes
included in this study. Variation among the fossil hominins and all
other groups is evident in Fig. 2, where all Pleistocene individuals
fall near or above the line of best fit for the LSA southern Africans,
the most asymmetric Holocene population included in this study.
La Ferrassie 1 and 2, Skhul IV and V exemplify the exceptional
relative tibial strength of the Neandertals and Levantine anatomi-
cally modern H. sapiens. This patterning was also displayed in
a single, atypical, European Early Upper Palaeolithic individual
(Paviland 1), and three North African Late Upper Palaeolithic in-
dividuals (El Wad 10260 and 10252, and Jebel Sahaba 21). Taken
together, these findings indicate that these individuals are adapted
to quite high levels of lower limb loading (relative to upper limb
loading). The hypertrophy of the lower limb may reflect adaptation
to very high levels of terrestrial mobility, perhaps considerably
greater than the level of habitual walking or running among highly
trained modern human cross-country runners and highly terres-
trially mobile early Holocene foragers. An alternative interpretation
is that the Pleistocene hominin skeleton reflects adaptation to
travel over highly undulating terrain (Marchi, 2008). However, to
support this alternative interpretation one would have to argue
that terrain was comparable among the different regions that these
fossil hominins inhabited.
An assessment of whether the more pronounced relative tibial
rigidity of the fossil hominins is sex-specific is possible by focusing
on the three females in the sample. The relative tibial rigidity of
Tabun C1, a female Neandertal, is comparable with that of other
male and female Pleistocene hominins included in the study.
However, raw tibial rigidity and raw humeral rigidity for Tabun C1
are considerably lower than most other fossil hominins, likely
reflective of a comparatively small body size. The second female
Neandertal, La Ferrassie 2, displays similarly low raw tibial rigidity,
yet particularly low humeral rigidity, similar to the mean for the
small-bodied LSA foragers. As a result, La Ferrassie 2 displays one of
the highest relative tibial rigidity scores of all individuals included
in this study. Compared with Tabun C1, the higher relative tibial
strength of La Ferrassie 2 likely indicates variation in the level of
relative lower limb loading between these two Neandertals, despite
displaying comparable estimated body mass (Table 2). Interest-
ingly, Qafzeh 9, a female anatomically modern H. sapiens, displays
a level of relative tibial rigidity similar to most other fossil
Please cite this article in press as: Shaw, C.N., Stock, J.T., Extreme mobility in the Late Pleistocene? Comparing limb biomechanics among fossil
Homo, varsity athletes and Holocene foragers, Journal of Human Evolution (2013), http://dx.doi.org/10.1016/j.jhevol.2013.01.004
hominins, yet levels of raw tibial and humeral rigidity that exceeds
many of the Pleistocene males.
Lieberman and Shea (1994) proposed mobility pattern variation
as one of the behavioral differences between Levantine Neandertals
and early H. sapiens. Using cementum annulation of gazelle teeth,
Lieberman (1993, 1998) concluded that early modern humans
occupied sites (Qafzeh Units XVIIeXXIV, Tabun C) on a seasonal
basis while Neandertals occupied sites (Amud B, Kebara IXeXII,
Tabun B) on a more continuous basis. Lieberman and Shea (1994;
Shea, 1998) concluded that early H. sapiens practiced a circulating
mobility strategy and emphasized intercept hunting while Nean-
dertals practiced a radiating strategy and focused on ambush
hunting. Other investigators have argued against some of these
assertions (Hovers, 1997; Stutz, 2002). More recent analyses of
Middle Palaeolithic lithic artifacts have hinted at differences in
residential mobility between these two Middle Palaeolithic pop-
ulations (Wallace and Shea, 2006), while zooarchaeological ana-
lyses have provided additional evidence of large-game hunting in
Neandertals (Niven et al., 2012). Regardless of potential differences,
the archaeological and zooarchaeological analyses discussed above
all concluded that both Levantine Neandertals and AMHS engaged
in high levels of terrestrial mobility.
A limitation to the present study is the use of raw cross-sectional
diaphyseal data. Merely constructing ratios of lower to upper limb
properties, as was done here, may not adequately control for the
influence of body mass on diaphyseal structure. Variation in rela-
tive tibial rigidity appears to be driven primarily by differences in
raw tibial rigidity and less so by difference in raw humeral rigidity
(Table 2). One might then argue that, rather than reflecting adap-
tation to differing mobility patterns, variation in relative tibial ri-
gidity primarily reflects the influence of body mass on tibial
diaphysis structure. If correct, this assumes that heavier individuals
should display more rigid tibiae, and a tendency towards higher
relative tibial rigidity. However, using data standardised by body
mass and bone length, as might be prescribed, to calculate relative
tibial rigidity is problematic. By standardising both the humerus
and the tibia by body mass the effect of standardisation is effec-
tively nullified following the subsequent calculation of the relative
tibial rigidity ratio. Additionally, not standardising tibial and hu-
meral rigidity by biomechanical lengths (squared) (Ruff, 2008b)
avoids artificially reducing relative tibial rigidity ratios, due to dif-
ferences in limb segment lengths between tibiae and humeri,
providing a closer match with locomotor expectations (see Ruff,
2000a, 2002; Shaw and Ryan, 2012). In support of the approach
implemented in this study, two observations are relevant. First,
because humans are bipedal, it is reasonable to expect that the
effect of body mass on the lower limbs relative to the upper limbs
will be at least comparable for all individuals. Second, the fossil
hominins that display the highest relative tibial rigidity (La Fer-
rassie 2 (66.4 kg), Skhul 4 (69.0 kg), Skhul 5 (68.8 kg) Paviland 1
(72.4 kg)) have estimated body masses that are similar to, or below,
the body mass average for the living human samples (68.2e
74.3 kg). By contrast, two of the three Neandertals and AMHS
with the highest estimated body mass (Spy 2 (85.5 kg), La Chapelle
1 (81.1 kg)) display relative tibial rigidities far below the average for
the LSA southern Africans, whose estimated body mass is up to
one-third less than those particular fossil hominin individuals.
These findings do not discount the potential influence that body
mass may be having on relative tibial rigidity, as calculated here.
However, the point is made that despite the influence of body mass
on tibial diaphysis structure, the distribution of diaphyseal rigidity
between the upper and lower limbs appears to correspond with
stated mobility patterns. Additionally, variation between the LSA
southern Africans and the Pleistocene sample may be related to
difference in body size. If the LSA sample was scaled-up to
 C.N. Shaw, J.T. Stock / Journal of Human Evolution xxx (2013) 1e8
a comparable body mass a positive allometric effect may occur
creating a relative tibial rigidity that is more similar to the fossil
hominins. Finally, variation described in this study may also have
been influenced by inter-specific differences in bone sensitivity to
biomechanical loading (Wallace et al., 2012).
Conclusion
Comparisons of relative tibial rigidity among modern human
athletes (cross-country runners, swimmers) and controls, Holocene
foragers (LSA southern Africans, Andaman Islanders) and Pleisto-
cene fossil hominins (Neandertals, anatomically modern and Upper
Palaeolithic H. sapiens) allow for three general conclusions:
1. As would be expected from their known activity patterns,
runners display quite high levels of tibial rigidity relative to
humeral rigidity. When compared with swimmers and con-
trols, this appears to be reflective of skeletal adaptation to
habitual training for competitive cross-country running.
2. Marine-adapted Andaman Islanders and highly terrestrially
mobile LSA southern Africans display patterns of relative tibial
rigidity closest to the athlete groups with more comparable
habitual activity patterns (swimmers and runners, respec-
tively). Nevertheless, significant differences between the LSA
southern Africans and runners, and swimmers and the Anda-
man Islanders, limit how accurately one can infer the duration
and intensity of specific activity patterns from skeletal remains
alone.
3. Modern human cross-country athletes reported running up to
80e100 miles per week, and the total hunting area of the LSA
southern Africans may have been as large as 2000e3000
square miles. The relative tibial strength of the Neandertals
and Anatomically Modern H. sapiens included in this study are
most similar to the LSA southern Africans, while almost half of
these individuals display even more pronounced relative tibial
rigidity. If relative tibial rigidity reflects adaptation to terrestrial
mobility, the results presented here suggest that these Nean-
dertals, the Skhul and Qafzeh H. sapiens and a few rare Upper
Palaeolithic H. sapiens were walking or running to a far greater
degree than even highly trained modern athletes and highly
terrestrially mobile early Holocene foragers.
Acknowledgements
Grant sponsorship: Royal Anthropological Institute, UK & Cush-
ing Anthropological Research Fund (CNS), The Leverhulme Trust
and NERC, UK (JTS). The authors wish to thank Eric Trinkaus and
Laura Shackelford for very generously providing much of the fossil
hominin data analysed in this manuscript, and members of the
PAVE Research Group (University of Cambridge) who provided
valuable intellectual input. Finally, this manuscript was greatly
improved by the efforts of Editor in Chief David Begun and two
anonymous reviewers.
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