AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 000:000–000 (2011)
Lumbar lordosis of Extinct Hominins
Ella Been,1* Asier Gómez-Olivencia,2,3 and Patricia A. Kramer4,5
1
Department of Anatomy and Anthropology, Sackler Faculty of Medicine, Tel Aviv University, Tel Aviv 69978, Israel
2
Division of Biological Anthropology, Department of Archaeology and Anthropology, University of Cambridge,
Fitzwilliam Street, Cambridge CB2 1QH, UK
3
Centro UCM-ISCIII de Investigación sobre Evolución y Comportamiento Humanos, Madrid, Avda. Monforte de
Lemos 5 (Pabellón 14), Madrid 28029, Spain
4
Department of Anthropology, University of Washington, Seattle, WA 98195-3100
5
Department of Orthopaedics and Sports Medicine, University of Washington, Seattle, WA 98195-3100
KEY WORDS posture; spine; Australopithecus; Neandertal; fossil H. sapiens
ABSTRACT The lordotic curvature of the lumbar spine
(lumbar lordosis) in humans is a critical component in the
ability to achieve upright posture and bipedal gait. Only
general estimates of the lordotic angle (LA) of extinct homi-
nins are currently available, most of which are based on
the wedging of the vertebral bodies. Recently, a new
method for calculating the LA in skeletal material has
become available. This method is based on the relationship
between the lordotic curvature and the orientation of the
inferior articular processes relative to vertebral bodies in
the lumbar spines of living primates. Using this relation-
ship, we developed new regression models in order to calcu-
late the LAs in hominins. The new models are based on pri-
mate group-means and were used to calculate the LAs in
The lordotic curvature of the lumbar spine (lumbar
lordosis) in humans is a critical component in the ability
to achieve upright posture. Along with pelvic tilt and the
angulation of the sacrum within the pelvis, lumbar lor-
dosis brings the upper body over the lower limbs and
allows the load from the upper body to be applied via
the vertebral column and sacrum to the pelvis and legs
(Gracovetsky and Iacono, 1987; Farfan, 1995; Whitcome
et al., 2007). These three characteristics act in tandem
to locate the line of gravity of the body close to the ace-
tabulum. Lumbar curvature influences, and is influenced
by, the morphology of the anterior pillar (formed by the
vertebral body and intervertebral disks) and posterior
pillar (formed by the articular processes and the lami-
nae) of the spine, the pelvis, and the thoracic cavity
(Sanders, 1998; Harrison et al., 2002; Lovejoy, 2005;
Jang et al., 2009; Been et al., 2010a).
When lordosis is out of sync with the other features,
allowing the line of gravity to deviate from its alignment
near the acetabulum, people can have substantial com-
plications, including functional issues such as a reduc-
tion in stride length and walking velocity and spinal
pathologies. For instance, people with LAs below the
normal range of 30–808 exhibit a short stride and slow
walking speed (Grasso et al., 2000; Sarwahi et al., 2002;
Hirose et al., 2004). Because of the flattening of their
normal lordosis, their line of gravity lies anterior to the
acetabulum, necessitating hip and knee flexion. On the
other side of the lordosis continuum, people with hyper-
lordosis (lordotic curvature greater than the normal
range) have a higher tendency to develop spondylolisthe-
sis (Antoniades et al., 2000), a potentially debilitating
C 2011
V WILEY-LISS, INC.
the spines of eight extinct hominins. The results were also
compared with the LAs of modern humans and modern
nonhuman apes. The lordotic angles of australopithecines
(418 6 4), H. erectus (458) and fossil H. sapiens (548 6 14)
are similar to those of modern humans (518 6 11). This
analysis confirms the assumption that human-like lordotic
curvature was a morphological change that took place dur-
ing the acquisition of erect posture and bipedalism as the
habitual form of locomotion. Neandertals have smaller lor-
dotic angles (LA 5 298 6 4) than modern humans, but
higher angles than nonhuman apes (228 6 3). This suggests
possible subtle differences in Neandertal posture and loco-
motion from that of modern humans. Am J Phys Anthropol
000:000–000, 2011. V 2011 Wiley-Liss, Inc.
C
condition where one vertebra slides forward relative to
adjacent vertebrae. This relative movement between ver-
tebrae can compromise spinal nerve function. Conse-
quently, it seems clear that an appropriate amount of
lumbar lordosis is important to normal function in
humans.
Lordotic curvature is defined herein as the angle
between the superior endplate of the first sacral vertebra
and the superior end plate of the fifth presacral verte-
bra. Only general estimates of the LA of extinct homi-
nins are available today, usually based on the wedging of
the vertebral bodies. These descriptions of the LA of the
fossils indicate where the specimen’s morphology
Grant sponsor: Ministerio of Educación (Programa Nacional de
Movilidad de Recursos Humanos del Plan Nacional de I1D1I 2008-
2011); Grant sponsor: Ministerio de Ciencia e Innovación (Proyecto);
Grant number: CGL2009-12703-C03-03; Grant sponsor: European
Community Research Infrastructure Action (SYNTHESYS Project;
http://www.synthesys.info/; FP6 ‘‘Structuring the European
Research Area’’ Programme).
*Correspondence to: Ella Been, Department of Anatomy and
Anthropology, Sackler Faculty of Medicine, Tel Aviv University, Tel
Aviv 69978, Israel. E-mail: beenella@post.tau.ac.il
Received 26 September 2010; accepted 15 September 2011
DOI 10.1002/ajpa.21633
Published online in Wiley Online Library
(wileyonlinelibrary.com).
2 E. BEEN ET AL.
resembled or differed from that of modern humans (Rob-
inson, 1972; Latimer and Ward, 1993; Sanders, 1998;
Ward, 2002; Whitcome et al., 2007; Weber and Pusch,
2008). For instance, the vertebral specimens of Australo-
pithecus africanus (STS 14) have been described as hav-
ing vertebral body wedging similar to that of humans
(Whitcome et al., 2007). On the basis of these estimates
from vertebral wedging, most researchers agree that
both australopithecines and H. erectus had lumbar LAs
within the normal range of modern humans (Robinson,
1972; Latimer and Ward, 1993; Sanders, 1998; Ward,
2002; Whitcome et al., 2007), presumably in response to
constraints similar to those apparent in humans.
However, since the discovery of the nearly complete
skeleton of La Chapelle-aux-Saints in 1908 (Bouyssonie
et al., 1908), an ongoing debate over the LA of Neander-
tals (H. neanderthalensis) has existed. On the one hand,
Trinkaus (1983), Arensburg (1991), and Cleuvenot (1999)
have argued that the LA of Neandertals is within the
normal range for modern humans. On the other hand,
other scholars have found that the LA of Neandertals is
smaller than that of modern humans (Boule, 1911–1913;
Weber and Pusch, 2008; Been et al., 2010a). Definitive
understanding of the degree of lumbar lordosis in Nean-
dertals requires more than simple description of verte-
bral wedging.
Recently, a new method for calculating the LA in skel-
etal material has become available (Been et al., 2010a).
This method is based on the relationship between the
lordotic curvature and the orientation of the inferior
articular processes in the lumbar spines of living human
and nonhuman primates. Because the method uses a
measurement taken within each vertebra, articulation of
the entire lumbar spine is not required. This work dem-
onstrated that 89% of the variation in the lordotic curva-
ture in living primates is related to the orientation of
the inferior articular process (Been et al., 2010a) and not
dependent on soft tissue morphology. Thus, this method
should be a reliable predictor of the lumbar LA in disar-
ticulated hominin spines, as long as extinct hominins
exhibit the same relationship as do living primates.
The aims of this study are, therefore, first, to calculate
a specific statistical model for calculating the lordotic
curvature for each extinct hominin specimen. This is
necessary because the fossil specimens vary in the num-
ber of preserved vertebrae. The second aim is to calcu-
late the lordotic curvature of the lumbar spines of
extinct hominins using these statistical models. Finally,
based on our results, we will examine the evolution of
lordotic curvature in hominins.
MATERIALS
Radiographic analysis
A total of 106 lateral radiographs of modern humans
(after Been et al., 2010a) and 76 lateral radiographs of
nonhuman primates were examined. All radiographs
met the following criteria: (1) from adult living subjects
with no detectable radiographic abnormalities (e.g.,
degeneration or reduced disk height) and (2) high resolu-
tion with a clearly visible lumbar spine. The modern
human sample included 106 adult humans (56 men and
50 women) from Israel. The radiographs were taken
when the subjects were standing with arms flexed across
their chests.
The nonhuman primates included the spinal (lumbar)
radiographs of Macaca fuscata and Macaca fasicularis
American Journal of Physical Anthropology
(52); nine apes including four chimpanzees (Pan troglo-
dytes), two gorillas (Gorilla gorilla), two orangutans
(Pongo pygmaeus), and one gibbon (Hylobates sp.);
Lemur (8); Saimiri (4); and Saguinus (3). For the nonhu-
man primates, standard lateral lumbar radiographs
were obtained with the anesthetized subjects lying on
their sides with flexed shoulders, hips, and knees in a
position as relaxed and as close to a natural posture as
possible. Preuschoft et al. (1988) demonstrated that
lateral spinal radiographs of anesthetized monkeys
(M. fuscata) exhibit similar spinal angulations to the lat-
eral radiographs of naturally standing (quadrupedal)
monkeys. Our naturally flexed position is similar to the
position described by Preuschoft et al. (1988).
Some researchers have claimed differences in the lum-
bar LA between men and women and that these were
also present in A. africanus (Whitcome et al., 2007). In
the orthopedic literature, no consensus exists regarding
potential differences in LAs between men and women.
Although some researchers found no differences in LAs
between the sexes (Lin et al., 1992; Takao et al., 2010),
others have found that females have slightly more lor-
dotic lumbar spines than do males, but the difference is
only 28–38 (Vialle et al., 2005). In our sample, the differ-
ence for the LA (LA) between the male and female
means is 18, a nonsignificant difference (Been et al.,
2007). Consequently, we have used a pooled sex sample.
Fossil samples
The lumbar vertebrae (PS1-PS5) of eight fossil homi-
nins were measured. These included two Pliocene
australopithecines (Sts-14 and Stw-431), one Lower Pleis-
tocene H. erectus (KNM–WT 15000), three Neandertals
from the Upper Pleistocene (Shanidar 3, La Chapelle-
aux-Saints, and Kebara 2), and two fossil H. sapiens from
the late Upper Pleistocene (Cro-Magnon 1 and 3). Details
regarding these fossils are included in Table 1. In two
cases, radiographs were not available to us (KNM-WT
15000 and Stw-431); so, we used lateral photographs
instead (See Table 2). These photographs clearly showed
all relevant anatomy. Isolated intact vertebra, such as
those from AL 288-1, Stw H8/41, and Régourdou 1, were
not included in this analysis because the standard error
of estimation of the regression equation exceeded 108
(See Methods section below). Other potentially interest-
ing specimens for our study like those from the early
Upper Pleistocene sites of Skhul and Qafzeh do not pre-
serve intact lumbar vertebrae, so we could not include
them in our analysis.
METHODS
Angular measurements
The modal number of lumbar vertebrae varies among
primate species. For example, the lumbar spine of Lemur
and Macaca has seven lumbar vertebrae, whereas the
lumbar spine of Pan has only four lumbar vertebrae
(Schultz, 1961). The difference in vertebral number is
not only restricted to the lumbar segment but also pres-
ent in the thoracic and sacral regions (Pilbeam, 2004).
Because LA is dependent on the number of vertebrae
included in the calculation (Harrison et al., 2001; Vialle
et al., 2005), comparison of results among genera with
different numbers of lumbar vertebra requires standardi-
zation. Thus, we have chosen a presacral segment of five
vertebrae which is the number of lumbar vertebrae in
 HOMININ LORDOSIS 3
TABLE 1. Hominin individuals studied in this research
Additional
Individual Species Vertebrae used Observations references
Stw 431 Australopithecus PS1-PS4 (n 5 4) Adult male. Fifth (PS1) and McHenry, 1992;
africanus fourth (PS2) lumbar McHenry and
vertebrae show pathological Berger, 1998a,b;
signs: spondylosis deformans Haeusler et al.,
following Staps (2002) also 2002
interpreted as brucellosis by
D’Anastasio et al. (2009).
Sts 14 Australopithecus PS2-PS4 (n 5 3) Young adult (Bonmatı́ et al., Haeusler, 2002
africanus 2008). Five (Haeusler et al.,
2002) or six lumbar
vertebrae (Robinson, 1972).
The inferior articular
processes in PS1 and PS5
are reconstructed.
KNM WT-15000 H. erectus/H. PS1-PS3 (n 5 3) Subadult male; five (Brown
ergaster et al., 1985, Haeusler et al.,
2002) or six (Walker and
Leakey, 1993; Latimer and
Ward 1993) lumbar vertebrae.
Possible spinal pathology,
related to infant malnutrition
(axial dysplasia); small
vertebral canal (Latimer and
Ohman, 2001; but see Schiess
et al., 2006).
Shanidar 3 H. neanderthalensis PS2, PS3, PS5 35–50–year-old male (Stewart,
(n 5 3) 1977; Trinkaus 1983). It
shows articular facets for
lumbar ribs attached to the
first lumbar vertebra,
Baastrup disease on the
spinous processes, and
marked degenerative
changes at the body of the
first and second lumbar
vertebra (PS4-PS5) (Ogilvie
et al., 1998).
Kebara 2 H. neanderthalensis PS1-PS5 (n 5 5) 20–30-year-old male; lumbar Arensburg, 1991
ribs attached to the first
lumbar vertebra and
nonossification
(pseudoarthrosis) of the
spinous processes of the last
four lumbar vertebrae
(PS1-PS4) (Duday and
Arensburg, 1991).Weber and
Pusch (2008) further report
mild degenerative changes
of the vertebral bodies
(PS3-PS2) and facet joint
(PS1).
La Chapelle- H. neanderthalensis PS1, PS2, PS5 [40 year old male; Baastrup Boule, 1911–1913;
aux-Saints (n 5 3) disease in the spinous Dawson and
processes (Ogilvie et al., Trinkaus, 1997.
1998).
Cro-Magnon 1 H. sapiens PS1, PS2, PS4, PS5 Middle aged male;
(n 5 4)a,b degenerative changes on PS5
and PS4.
Cro-Magnon 3 H. sapiens PS1-PS5 (n 5 5)b,c Adult male.
a
PS1(4289), PS2(4266), PS4(4286), PS5(4285). We attribute these four vertebrae to Cro-Magnon 1 based on the presence of degener-
ative pathology. The number between parentheses is the label of the individual vertebral specimens.
b
These associations of vertebrae belonging to the same individual differ from those published by Vallois and Billy (1965).
c
PS1(4281), PS2(4280), PS3(4287), PS4(4268), PS5(4282). We attribute these five vertebrae to a male, younger than Cro-Magnon 1,
and thus it could belong to Cro-Magnon 3 or 4. The number between parentheses is the label of the individual vertebral specimens.

modern and fossil H. sapiens, Neandertals, and H. hei- Gómez-Olivencia, 2009; Bonmatı́ et al., 2010). Australo-
delbergensis (McCown and Keith, 1939; Trinkaus, 1983; pithecus and H. erectus could arguably have six rather
Arensburg, 1991; Haeusler et al., 2002; Pilbeam 2004; than five lumbar vertebrae (Robinson, 1972; Walker and

American Journal of Physical Anthropology

American Journal of Physical Anthropology

4
TABLE 2. Inferior articular process angles (AP) and vertebral body wedge angles (B; in degrees) of modern humans, extinct hominins, and nonhuman primates
Articular process angle (AP) Vertebral body wedge angles (B)c
Species Individual Source PS5 PS4 PS3 PS2 PS1 SAP PS5 PS4 PS3 PS2 PS1 SB
H. sapiens Cro-Magnon 1 Radiograph 100 93 M 101 108 27 24 5
H. sapiens Cro-Magnon 3 Radiograph 99 108 107 98 126 538 22 0 4 5 11 18
H. sapiens average 99.5 100.5 107 99.5 117.0 24.5 22 4 5 8
H. neanderthalensis Kebara 2 Radiograph 88 87 89 98 103 465 27 27 27 22 9 215
H. neanderthalensis La Chapelle- Radiograph 87 M F 102 101 26 25 1 6
aux-Saints
H. neanderthalensis Shanidar 3 Radiograph 89 F 95 99 F 29 25 24 1 10 27

E. BEEN ET AL.
H. neanderthalensis 88.0 87 92.0 99.7 102 27 26 25.5 0 8 211
average
H. erectus KNM WT-15000 Photograph F F 109 101 100 25 5.5 11
Australopithecus STS - 14 Radiograph F 101 93 98 F 23 21 0 3 4 3
africanus
A. africanus Stw - 431 Photograph F 99 98 108 101 25 21 21 0 8 1
Australopithecus 100 96 103 101 24 21 20.5 1.5 6 2
average
Modern Homo x̄ 6 SD Radiograph 94 6 5 97 6 5 100 6 6 106 6 6 113 6 7 510 6 15 23 6 3 216 3 063 263 863 6 6 10
sapiensb (n 5 106)
Macacaa (n 5 52) x̄ 6 SD Radiograph 87 6 4 87 6 5 89 6 5 89 6 4 88 6 5 440 6 15 27 6 4 27 6 3 27 6 4 25 6 4 22 6 5 228 6 16
Apes (n 5 9)b x̄ 6 SD Radiograph 95 6 5 95 6 3 94 6 1 93 6 3 93 6 4 471 6 7 26 6 3 25 6 2 23 6 2 21 6 2 1.5 6 2 213 6 4
Lemur (n 5 8)a x̄ 6 SD Radiograph 83 6 3 85 6 3 86 6 3 87 6 2 88 6 2 430 6 10 24 6 2 23 6 2 23 6 2 23 6 2 22 6 2 216 6 5
Saimiri (n 5 4)a x̄ 6 SD Radiograph 91 6 3 90 6 4 89 6 6 93 6 3 94 6 10 456 6 23 23 6 1 23 6 2 22 6 2 21 6 2 21 6 1 29 6 8
Saguinus (n 5 3)a x̄ 6 SD Radiograph 83 6 6 88 6 7 85 6 4 89 6 2 95 6 11 440 6 28 24 6 1 26 6 0 25 6 2 23 6 2 23 6 4 221 6 8

M, vertebra missing; F, vertebra present but fragmentary.a After Been et al. (2010a).
b
The sample is composed of four chimpanzees (Pan troglodytes), two gorillas (Gorilla gorilla), two orangutans (Pongo pygmaeus), and one gibbon (Hylobates sp.).
c
Vertebral body wedge angles (B): Negative value indicates kyphotic wedging; Positive value indicates lordotic wedging.
 HOMININ LORDOSIS
Fig. 1. Lumbar lordotic angle (LA) shown on a spinal lateral
radiograph of modern human.
Leakey, 1993), although this vision is contested by other
scholars (Haeusler et al., 2002).
Nonetheless, in order to be able to compare LA among
species, we define LA as that of the last five presacral
vertebrae. This approach creates a lumbar region that is
intermediate in vertebral number between the longer
lumbar spines of monkeys and (potentially) early homi-
nins and the shorter lumbar spines of modern great
apes. This approach is justified, because no correlation
exists between the number of lumbar vertebrae and the
degree of lordosis in primate species. For example,
humans have five lumbar vertebrae with lordosis of
[508, apes have four lumbar vertebrae with 20–258 of
lordosis, and macaque monkeys have seven lumbar
vertebrae with lordosis of 148 (Been et al., 2010a).
To facilitate comparison among the different hominin
groups and between hominins and nonhuman primates,
we numbered the vertebrae in relation to the sacrum, as
have others that compared the lumbar vertebrae of non-
human primates, early hominins, and modern humans
(Latimer and Ward, 1993; Abitbol, 1995). The vertebra
just above the sacrum is denoted PS1 (first presacral) in
all of the spines; the vertebra above PS1 is denoted PS2,
and so forth.
As described previously (Been et al., 2010a), lateral ra-
diographs were used to measure LA of the five presacral
vertebrae, vertebral body wedging and the orientation of
the inferior articular processes relative to the vertebral
body. For each of the five vertebrae (PS1-PS5), three lines
were drawn (Figs. 1 and 2): along the cranial endplate of
the vertebral body (including the first sacral vertebra);
along the caudal endplate of the vertebral body; and
along the ventral border of the inferior articular process.
These lines were used to measure three angles:
1. The LA between the cranial endplate of PS5 and the
cranial endplate of S1.
2. The vertebral body wedge angle (B) between the
cranial and caudal end plates of the vertebral body.
5
Fig. 2. Measurement of the body wedge angle (B) and the
inferior articular process angle (AP) of the first lumbar (PS5) of
La Chapelle-aux-Saints.
3. The inferior articular process angle (AP) between the
cranial endplate and the ventral border of the inferior
articular process of the same vertebra, as seen in the
lateral radiograph in the facet joint space.
Measurements B and AP were made for each of the
five presacral segments. All measurements were taken
by the same investigator (EB) using a 25-cm Jamar goni-
ometer with a 3608 scale in 18 increments. Summations
of measurement B and AP were also calculated: SB
equals the sum of the body wedge angles of vertebrae
PS1-PS5; and SAP equals the sum of the inferior articu-
lar process angles of these vertebrae.
The body wedge angles were measured in order to
compare our results to previously published data,
because previous assessments of the LA in extinct homi-
nins usually refer to the wedging of the vertebral bodies.
The inferior articular process angles were measured to
develop new regression models for calculating the LA in
fossil hominins (Been et al., 2010a).
In order to determine if our measurements were repro-
ducible, five radiographs of modern humans (65 meas-
urements) were measured twice, 3 weeks apart, by one
of us (EB). The intraclass correlation coefficients were
[0.97 (P \ 0.001) for LA and for B and [0.88 (P \ 0.01)
for AP. The mean angular deviation between trials was
2.48 for LA, 1.98 for B, and 2.858 for AP.
Developing regression models
New regression formulae for calculating LA in fossil
hominins were developed based on primate group means
(Table 3 and Fig. 3), using bivariate fit models (fit Y by
X, JMP statistics software, SAS institute, Cary, NC). For
each primate group, the individual values for LA, B, and
AP were averaged to produce the mean for the group.
These models were specifically developed for each speci-
men based on the number of intact vertebrae present in
the specimen. Our method yields standard errors of esti-
mation (SEE) of greater than 108 when only one or two
vertebrae are present. Consequently, we did not calcu-
late the LAs for specimens with less than three intact
vertebrae, which limit the available fossil sample.
We grouped the primates loosely based on phylogenetic
relationship. For example, in the macaque group, we
American Journal of Physical Anthropology
6 E. BEEN ET AL.

combined the results for M. fasicularis and M. fuscata,

SAP 2 206.8.

in extant primates; Average sum of articular process angle (SAP) for different combinations of presacral vertebrae (to accommodate the intact vertebrae of the extinct hominins)
The individual species data for apes is provided for informational purposes only; it was not used in the regression analysis.In this table, we present the average lordotic angles
SAP8 (PS1-PS4)

LA 5 0.618 3
whereas in the ape group, we combined the results for

Stw - 431
P. troglodytes, G. gorilla, P. pygmaeus, and Hylobates sp.

0.95
416

353
376

347
365
357

3.9
(Table 3). We recognize that grouping nonhuman apes,
but separating Saimiri and Saguinus might at first
seem illogical, but we were constrained in our sampling
ability. All of the nonhuman primates were captive ani-
mals, and the radiographs were obtained during veteri-

SAP 2 231.7.
(PS2, PS3, PS4)

LA 5 0.923 3
nary exams, except for those from the macaques, which

STS - 14
were obtained for research purposes. Consequently, our
SAP8

0.92
303

265
283

259
271
261

4.7
sample is one of convenience and we could neither obtain
equal numbers of individuals of all species nor could we
obtain all species that we would have preferred to have.
Some genera (Gorilla, Pongo, and Hylobates) are repre-
sented by only one or two individuals, while other gen-
era have more. All of the nonhuman apes had similar
SAP 2 186.2.
(PS1, PS2, PS3)

LA 5 0.744 3

WT-15000
LAs (Table 3), but Saimiri and Saguinus are different.
KNM
We created the primate groups, then, based on phylog-
0.964
SAP8

3.36
319

266
280

261
276
269

eny, but influenced by the number of available individu-

als and the morphology of the lumbar region.
We evaluated the impact of the decision to use the pri-
mate group-mean method by eliminating the groupings
TABLE 3. Regression models for calculating lordosis in hominins

and using all the primate individual values to develop

SAP 2 226.7.

statistical models. This method has its own flaws; how-

LA 5 0.911 3
(PS2, 3, PS5)

Shanidar 3

ever, in that it over-represents individuals in Macaca

0.926
SAP8

and Homo relative to the nonhuman apes. The all pri-

300

265
282

256
273
257

4.8

mate, individual-based method produced the same

trends as the primate group-mean method (analysis not
shown), allowing us to conclude that our primate group-
mean models are robust.
Still, the possibility exists that SEEs and R2s that are
SAP 2 200.
LA 5 0.799 3

aux-Saints
La Chapelle-
(PS1, 2, PS5)

based on (6) group means could be misleading when pre-

dicting individual variation (Ruff, 2003). In order to test
SAP8

0.96
313

264
282

259
277
269

3.4

how well the group-mean models work to predict individ-

ual values, we calculated the LAs for all of our extant
individuals (human and nonhuman primates) and then
calculated the mean difference between the measured
and calculated LAs.
in extant primates; and the regression models developed based on these values.
SAP 2 215.6.

Cro-Magnon 1
LA 5 0.645 3

We performed individual and sample comparisons of

(PS1,2,PS4,5)

the calculated LAs between the fossil individuals and

SAP8

0.94
4.22
410

351
377

344
367
355

the modern human and ape sample using Z-scores and

Mann-Whitney’s U-test. Using the Bonferroni correction,
the threshold value would be 0.0125, which is similar to
that obtained by the Dunn-Šidák correction. The differ-
ence between these two correction methods is higher as
SAP 2 220.5.

the number of groups involved increases.

Cro-Magnon 3,
LA 5 0.528 3

Kebara2

Finally, because it is possible that LA is influenced by

(PS1-PS5)

0.948
SAP8

locomotor form, we calculated the LAs of the extinct

4.04
510

440
471
470
465
469
477
430
456
440

hominins using statistical models developed just from

the angles measured from the radiographs of modern
humans (See also Been et al., 2010a). As we had done
for the group-mean method, we calculated the mean dif-
ference between the measured and calculated LAs of the
Lordotic angle

humans in order to determine how well the statistical

14 6 10
51 6 11
x̄ 6 SD

22 6 3

23 6 5
9 6 17

model predicted individual values.

868
21
22
19
25

RESULTS
Regression models based on primate
group-mean method
Formula applied to:
estimation (SEE)
Modern H. sapiens

Hylobates (n 5 1)a

Regression models (that predict lordotic curvature)

Standard error of
Regression model
Saguinus (n 5 3)
Macaca (n 5 52)

Gorilla (n 5 2)a

Saimiri (n 5 4)

were developed from the sum of the inferior articular

Pongo (n 5 2)a

Lemur (n 5 8)
Average SAP8

Pan (n 5 4)a
Apes (n 5 9)

process angles (SAP) in primates, using the mean value

(n 5 106)

for each primate group. The regression models and the

data used to develop them are summarized in Table 3
and Figure 3. All of the group-mean models have coeffi-
R2

cients of determination R2 [ 0.9 between the sum of

a

American Journal of Physical Anthropology

 HOMININ LORDOSIS 7

Fig. 3. Correlation between lordotic angle (LA) and sum of inferior articular process (SAP), based on primate group means. Lin-
ear fit with 95% bivariate normal ellipse. These ellipses are both density contours and confidence curves. As confidence curves, they
show where a given percentage of the data is expected to lie. ~5 modern human; ! 5 Nonhuman apes; h 5 Macaca; l 5 Lemur;
* 5 Saimiri; * 5 Saguinus.

articular process wedging and the lordotic curvature and
low standard errors of estimation (SEE \ 5; Table 3 and
Fig. 3). The resultant lordotic curvatures of the fossil
hominins and the formulae used to calculate them are
summarized in Table 4.
Regression models based on modern
human-only method
Regression models (that predict the lordotic curvature)
were also developed from the sum of the inferior articular

American Journal of Physical Anthropology

American Journal of Physical Anthropology

8
TABLE 4. Calculated lumbar lordotic angles of extinct hominins
Regression model based on Calculated Regression model based on Calculated
Sample Individual Measured vertebrae SAP primate group-mean method lordosis modern human only method lordosis
Fossil H. sapiens Cro-Magnon 1 PS1, PS2, PS4, PS5 402 LA 5 0.645 3 402 2 215.6 44 LA 5 0.63 3 SAP 2 208 46
R2 5 0.94, SEE 5 4.22 R2 5 0.55, SEE 5 7.5

Fossil H. sapiens Cro-Magnon 3 PS1-PS5 538 LA 5 0.528 3 538 2 220.5 64 LA 5 0.565 3 SAP 2 237 67
R2 5 0.948, SEE 5 4.04 R2 5 0.62, SEE 5 6.8
Fossil H. sapiens x̄ 6 SD 54.0 6 14.1 56.5 6 15
average

E. BEEN ET AL.
H. neanderthalensis Kebara 2 PS1-PS5 465 LA 5 0.528 3 465 2 220.5 25 LA 5 0.565 3 SAP 2 237 26
R2 5 0.948, SEE 5 4.04 R2 5 0.62, SEE 5 6.8
H. neanderthalensis La Chapelle-aux-Saints PS1, PS2, PS5 290 LA 5 0.799 3 290 2 200 32 LA 5 0.693 3 SAP 2 167 34
R2 5 0.96, SEE 5 3.4 R2 5 0.40, SEE 5 8.6

H. neanderthalensis Shanidar 3 PS2, PS3, PS5 283 LA 5 0.911 3 283 2 226.7 31 LA 5 0.636 3 SAP 2 140 40
R2 5 0.926, SEE 5 4.8 R2 5 0.42, SEE 5 8.5
H. neanderthalensis x̄ 6 SD 29.3 6 3.8 33.3 6 7
average
H. erectus KNM-WT 15000 PS1, PS2, PS3 310 LA 5 0.744 3 310 2 186.2 45 LA 5 0.61 3 SAP 2 144 45
R2 5 0.964, SEE 5 3.36 R2 5 0.37, SEE 5 8.8
A. africanus STS - 14 PS2, PS3, PS4 292 LA 5 0.923 3 292 2 231.7 38 LA 5 0.62 3 SAP 2 136.4 44
R2 5 0.929, SEE 5 4.7 R2 5 0.45, SEE 5 8.2
A. africanus Stw - 431 PS1, PS2, PS3, PS4 406 LA 5 0.618 3 406 2 206.8 44 LA 5 0.61 3 SAP 2 203 45
R2 5 0.95, SEE 5 3.9 R2 5 0.54, SEE 5 7.5
Australopithecus x̄ 6 SD 41.0 6 4.2 44.5 6 1
average
 HOMININ LORDOSIS
Fig. 4. Lumbar lordotic angles of extinct hominin speci-
mens.
process angles (SAP) using only the modern human data
(Been et al., 2010a). These models have coefficients of
determination (R2) ranging from 0.37 to 0.62 between the
sum of articular process wedging and the lordotic curva-
ture and standard errors of estimation (SEE \ 9, Table 4).
The resultant lordotic curvatures of the fossil hominins
and the formulae used to calculate them are summarized
in Table 4.
Validity of statistical models for predicting
individual values
In order to determine the degree to which our statisti-
cal models were able to predict individual values, we
calculated the mean individual prediction errors. We cal-
culated the LA of all of our extant individuals (human
and nonhuman primates) using the primate group-mean
model and compared it with their measured LAs. The
mean difference between the calculated and measured
LA is 65.928. We repeated the process using the modern
human-only model. The mean difference between the cal-
culated and measured LAs in humans is 65.938.
Vertebral body wedging and articular
process angles
The body wedge angles (B; negative value indicates
kyphotic wedging; positive value indicates lordotic wedg-
ing) and the articular process angles (AP) for extant
primates and for extinct hominins are shown in Table 2.
Modern humans have a higher (more lordotic) sum of
vertebral body wedging (SB 5 6 6 10, x 6 SD) and a
higher sum of articular process wedging (SAP 5 510 6
15) compared with nonhuman apes (SB 5 213 6 4, and
SAP 5 471 6 7). Fossil H. sapiens show vertebral body
wedging (SB 5 18) and articular process wedging (SAP 5
538) similar to modern humans. H. neanderthalensis
shows vertebral body wedging (SB 5 211) and articular
process wedging (SAP 5 465) that are smaller than that
of modern humans and comparable with that of nonhu-
man apes. H. erectus shows vertebral body wedging and
articular process wedging that are comparable with that
of modern humans. Australopithecus shows vertebral
9
Fig. 5. Lumbar lordotic angles of extinct hominins, modern
humans and apes. Diamond 5 average; Bar 5 one standard
deviation.
body wedging (SB 5 2) and articular process wedging that
are comparable with modern humans, but below the aver-
age for modern humans throughout the spine (Table 2).
Although the overall articular process wedging of
Neandertals is low compared with modern humans and
similar to nonhuman apes, the Neandertal pattern of
distribution within the spine of articular process wedg-
ing is similar to modern humans. Taking the mean val-
ues, vertebra closer to the sacrum in modern humans
show an increase in articular process wedging, as do
those of Neandertals. On the contrary, nonhuman apes
show an approximately constant articular process wedg-
ing throughout the five presacral vertebrae. This pattern
is difficult to ascertain, however, from single hominin
individuals due to the fragmentary status of the record
and, potentially, large intrapopulational variation.
Lordotic angles
The calculated LAs of the individual specimens are
shown in Table 4 and Figure 4. The LA values for homi-
nin groups and nonhuman apes are illustrated in Figure
5. Based on the primate group-mean models, the largest
values of LA are in fossil H. sapiens (548 6 14; 95% CI
5 35–73). Australopithecus (418 6 5; 95% CI 5 35–47)
and H. erectus (458) show lower values. Neandertals
show LAs (298 6 5; 95% CI 5 24–34) that are lower
than all other hominins, but higher than modern nonhu-
man apes.
The modern human-only models show similar results:
the largest values of LA are in fossil H. sapiens (578 6
15). Australopithecus (44.58 6 1) and H. erectus (458)
show lower values. Neandertals exhibit LAs (338 6 7)
that are lower than the other hominins, but higher than
modern nonhuman apes. We should note that the mod-
ern human-only model calculates LAs that are on aver-
age 28 higher than the primate group-mean model.
Because of the similarity between the two methods,
we report below only those analyses that used the values
calculated from the primate group-mean method.
We performed individual and sample comparison of the
calculated LA between the fossil individuals and the
modern human and ape sample using Z-scores and
American Journal of Physical Anthropology
10 E. BEEN ET AL.
TABLE 5. Individual and sample comparison of the calculated lordotic angle between the fossil individuals and
the modern human and nonhuman ape sample using Z-scores and Mann-Whitney’s U-test
Z-score Mann–Whitney
Sample Individual Calculated lordosis Modern human Apes Modern human Apes
**
Fossil H. sapiens Cro-Magnon 1 44 20.65 6.44
Fossil H. sapiens Cro-Magnon 3 64 1.17 12.32**
Fossil H. sapiens Sample mean 6 SD 54.0 6 14.1 0.7844 0.01818*
H. neanderthalensis Kebara 2 25 22.37* 0.85
H. neanderthalensis La Chapelle-aux-Saints 32 21.74 2.91**
H. neanderthalensis Shanidar 3 31 21.83 2.62**
H. neanderthalensis Sample mean 6 SD 29.3 6 3.8 0.006626** 0.03182*
H. erectus KNM-WT 15000 45 20.55 6.74**
Australopithecus africanus STS - 14 38 21.19 4.68**
A. africanus Stw - 431 44 20.65 6.44**
A. africanus Sample mean 6 SD 41.0 6 4.2 0.1323 0.01818*
Modern human 51.1 6 11.0 (24–75)
mean 6 SD (range)
Nonhuman ape 22.1 6 3.4 (18–28)
mean 6 SD (range)

* 5 , P \ 0.05; **5 , P \ 0.01. We have underscored those LA values outside the range of the comparison sample.If we apply the
Dunn-Šidák correction for multiple comparisons [1 2 (1 2 a)1/n] with an a 5 0.05 and four groups considered (modern humans,
apes, Neandertals, and australopithecines) the threshold value obtained for significance 0.0127, and only Neandertals would signifi-
cantly depart from modern humans.

Mann-Whitney’s U-test (Table 5). The individual
specimens of fossil H. sapiens, H. erectus, and Australo-
pithecus have LAs that are within the normal range for
modern humans and above the normal range for extant
nonhuman apes. Two of the three Neandertal specimens
(La Chapelle-aux-Saints and Shanidar 3) have LAs that
are at the lower end of the normal range for modern
humans (Z-score between 1.5 and 1.96), but significantly
above modern nonhuman apes. One Neandertal speci-
men (Kebara 2) has a very small LA (258), which is
below the normal range for modern humans and within
the normal range for modern nonhuman apes.
Kruskal-Wallis tests between the LA values of modern
H. sapiens, modern nonhuman apes, H. neanderthalen-
sis, Australopithecus, and fossil H. sapiens indicate sig-
nificant differences among these groups. To localize these
significant differences, we performed Mann-Whitney
pairwise tests (See Table 5). Both fossil H. sapiens and
Australopithecus have LAs that are within the normal
range for modern humans and above the normal range
for modern nonhuman apes. It is interesting to note that
the LA values of Neandertals are significantly below the
values for modern Homo sapiens, but significantly above
nonhuman ape values (Table 5). If we apply the Dunn-
Šidák correction for multiple comparisons [1 2 (1 2 a)1/
n
] with an a 5 0.05 and four groups considered (modern
humans, nonhuman apes, Neandertals, and australopi-
thecines), the threshold value obtained for significance is
0.0127, and Neandertals are significantly different from
modern humans.
DISCUSSION
Regression models
The statistical models that were developed to calculate
lordosis in fossil hominins (based on the primate group-
mean method) have coefficients of determination
between inferior articular processes angles and LAs of
[0.9, as well as standard errors of estimation \78. The
LAs calculated with the primate group-mean method
were confirmed by calculating models considering all pri-
mates as individuals and are consistent with the LAs
calculated based on the modern human-only method.
Previous reports of the vertebral wedging, (Latimer
and Ward, 1993; Sanders, 1998; Ward, 2002; Whitcome
et al., 2007), are also consistent with the LAs found
using the primate group-means method. Nonhuman apes
have small LAs [228 6 2, (Abitbol, 1995); 228 6 3 current
study] and kyphotic lumbar vertebral bodies [SB 5 2198
(Latimer and Ward, 1993); SB 5 2138 6 4, current
study]. Neandertals also have small LAs (298 6 3) and
kyphotic lumbar vertebral bodies (SB 5 2118, current
study). Species with high LAs (LA [ 408) have lordotic
wedging of their vertebral bodies: modern humans [SB
5 68 (Vialle et al., 2005; SB 5 68 6 10 current study),
fossil H. sapiens (SB 5 188current study) and Australo-
pithecus (SB8 5 18 (Whitcome et al., 2007); SB 5 28, cur-
rent study].
All of these comparisons yield the same pattern among
the hominin species, providing us with confidence that
the pattern we have reported is robust. Although it is
possible that fossil hominins could exhibit a relationship
between articular process angle and LA that is different
from that found among all primates sampled or among
only modern humans, it seems unlikely.
Lordotic angle
This study is the first to publish angular values for
the LAs of extinct hominins. Australopithecines (LA 5
418 6 4, 95% CI 5 35–47), H. erectus (LA 5 458), and
fossil H. sapiens (LA 5 548 6 14, 95% CI 5 35–73) have
LAs that overlap the range of lordosis in our modern
human sample (LA 5 518 6 11, 95% CI 5 49–53). The
finding of human-like LAs in australopithecines and
H. erectus is in agreement with previously published
data (Robinson, 1972; Latimer and Ward, 1993; Sanders,
1998; Ward, 2002; Whitcome et al., 2007). Lordotic cur-
vature, in combination with pelvic tilt on the leg and
sacral tilt within the pelvis, allows the upper body to be
balanced above the acetabulum. It has also been sug-
gested, based on pelvic morphology, that the lordotic cur-
vature of Ardipithecus ramidus was already greater

American Journal of Physical Anthropology

 HOMININ LORDOSIS 11
than that of nonhuman apes (Lovejoy, 2009); however, no
lumbar vertebra of this species has been described. Our
analysis supports the assumption that human-like lor-
dotic curvature was a morphological change that took
place during the acquisition of erect posture and bipedal-
ism as the habitual form of locomotion.
Neandertals depart from this hominin trend, having
smaller LAs (LA 5 298 6 4, 95% CI 5 24–34) than do all
other hominins, but higher angles than do nonhuman
apes (LA 5 228 6 3, 95% CI 5 20–24). The relatively
small LAs of Neandertals are in agreement with Boule
(1911–1913), Weber and Pusch (2008), and Been et al.
(2010a), but contradict the results of Trinkaus (1983),
Arensburg (1991), and Cleuvenot (1999). The less lordoti-
cally curved lumbar column of Neandertals could be
inherited from their presumed middle Pleistocene ances-
tors—H. heidelbergensis—as is suggested by the
kyphotic lumbar vertebral body wedging and the hori-
zontal sacral endplate of the specimens from the Sima
de los Huesos site (Arsuaga et al., 1999; Gómez-Oliven-
cia, 2009; Bonmatı́ et al., 2010).

Limitations
Our study is limited by several issues. First, even
though we have used all the fossils available to us, the
fossil sample is small and that lessens the power of any
inferential statistical methods. Consequently, we have
had to rely mostly on descriptive analysis. New findings,
especially of lower Pleistocene H. erectus sensu lato,
should help us to better understand the evolution of
lumbar lordosis in hominins.
Second, some of the fossil specimen show marked
pathologies such as degenerative changes (Cro-Magnon 1,
Shanidar 3), spondylosis or brucellosis (Stw-431),
Baastrup disease (La Chapelle-aux-Saints and Shanidar 3),
and nonossification of the spinous processes (Kebara 2).
These pathologies are typically found in most modern
human populations, and there is no evidence to indicate
that these pathologies affect the degree of lordotic curva-
ture (Maes et al., 2008; Papadakis et al., 2010).
Third, one of our specimens (KNM-WT 15000) is a
juvenile with a suspected axial dysplasia (Latimer and
Ohman, 2001), and therefore, it might exhibit different
LAs than a fully adult specimen. We include it because the
fossil record of lumbar spinal elements is sparse for early
genus Homo but recognize that future work might render
any evidence from it moot. In any case, elimination of
it from our sample would not change our conclusions.
Finally, reconstructing LA without soft tissue requires
that we understand the relationship between the inter-
vertebral discs and lumbar lordosis. Fortunately, the
models we employ are based on radiographs of the lum-
bar spines of living individuals, that is, that possessed
both the vertebral bodies and the intervertebral discs
(Been et al., 2010a), and intervertebral discs do not con-
tribute to differences in overall lordosis as much as does
vertebral wedging (Been et al 2010b). This suggests that
the absence of intervertebral discs would not change the
conclusions presented herein.
Evolutionary Perspective on Lumbar Lordosis
Given that Neandertals exhibit less lumbar curvature
than other hominins, a critical question obtains: do
Neandertals retain the primitive configuration of small
LAs as seen in extant nonhuman apes? This proposition
Fig. 6. Schematic diagram of the evolution of the lumbar
curvature. Dramatic increase in lordotic angle from the ape
condition to australopithecines. Lordotic angle stays constant
for H. erectus and the common ancestor of modern humans and
Neandertals. Then two parallel changes occurred: the lordotic
curvature of H. sapiens slightly increased and the lordotic
curvature of Neandertals decreased.
seems unlikely as parsimonious considerations exclude
it. It is our contention that with the acquisition of biped-
alism lordosis increased—from a degree similar to that
of modern nonhuman apes (small lordosis) to the high
lordosis seen in australopithecines, H. erectus and
H. sapiens. The Neandertals, then, represent a reversal
in this morphological trend.
The LA shows an increase in lordosis from the primi-
tive condition, preserved in modern nonhuman apes
(208  258) to australopithecine and H. erectus (40  45)
and then to H. sapiens (50  55), while Neandertals
show a decreased lordosis (30) when compared with
other hominins (Fig. 6). The lack of fossil lumbar spines
from early genus Homo is frustrating, because their mor-
phology would provide important clues to understanding
the evolution of lumbar lordosis.
A normal lumbar lordosis (within the range of 308–808,
see Bernhardt and Bridwell, 1989; Jackson et al., 2000;
Kimura et al., 2001; Harrison et al. 2001) plays a major
role in bipedal walking in modern humans, as it serves as
a shock absorber, helps maintain minimal perturbations
of the head, and helps create the moment that moves the
pelvis and the legs during bipedal walking. Although
lumbar lordosis plays a role in the maintenance of an
efficient upright posture, it also adds a certain amount of
compliance for locomotion and protects the posterior

American Journal of Physical Anthropology

12 E. BEEN ET AL.
Fig. 7. Sacral and lumbar posture of modern humans (A) and Neandertals (B). Note the smaller lordotic angle and more hori-
zontal sacral endplate of the Neandertal spine.
spinal ligament system from excess strain (Gracovetsky
and Iacono, 1987; Bogduk, 1997; Vialle et al., 2005).
The events or selective pressures that generated
smaller lumbar lordosis in the Neandertal lineage com-
pared with other hominins are not immediately obvious.
We can hypothesize that the lumbar lordosis is related
to other distinctive Neandertal postcranial features like
the lower limb, pelvic and thoracic morphology (Rak,
1991; Franciscus and Churchill, 2002; Polk, 2004;
Gómez-Olivencia et al., 2009). These differences may
indicate postural and locomotive differences in the Nean-
dertal lineage compared with that of our species.
Lordotic curvature in humans appears to correlate
with pelvic and thoracic morphology. Small LAs usually
correlate with a vertical sacrum (horizontally oriented
sacral endplate) and small thoracic kyphosis (Fig. 7;
Harrison et al., 2002; Lovejoy, 2005; Boulay et al., 2006;
Jang et al., 2009), although there are also cases of small
LAs accompanied by normal or even hyperkyphotic
thorax (Harrison et al., 2002). Examining the wedging of
the Neandertal thoracic vertebrae may help to resolve
that discrepancy (Goh et al., 1999).
Neandertals are known to have a larger thorax than
modern humans. The shape of the thorax is also differ-
ent: the uppermost and lowermost ribs are similar to
those of modern humans but mid-thoracic ribs are signif-
icantly larger (Gómez-Olivencia et al., 2009). Some
authors propose that Neandertals show a larger antero-
posterior dimension compared with modern humans
based on the ribs of Shanidar 3 (Franciscus and Church-
ill, 2002) or based on the clavicular proportions of
several Neandertals. From the ribs of Tabun C1, Wein-
stein (2008) proposes that the thorax of this individual
was expanded mediolaterally. Because of taphonomical
distortion of ribs from Kebara 2 (and also Tabun C1),
Gómez-Olivencia et al. (2009) did not propose in which
American Journal of Physical Anthropology
direction (anteroposterior, mediolateral, or both) the
Neandertal thorax was larger. In any case, it is reasona-
ble to link the biomechanical differences that a larger
thorax would impose to the spinopelvic balance and the
presence of a less lordotic lumbar spine in Neandertals.
The Neandertal pelvis is wide (bilaterally) with a long
bi-iliac breadth, long and slender superior pubic rami,
and the acetabulum is posterior relative to the pelvic
inlet, compared with H. sapiens (Rak and Arensburg,
1987; Rak, 1990; 1993). Rak (1993) postulates that Nean-
dertal gait is less efficient than that of modern humans
and that the Neandertal pelvis lacks the shock-absorbing
mechanism with which H. sapiens is equipped. A posteri-
orly situated acetabulum requires smaller LAs in order
to adjust the weight of the upper body above the pelvis,
but small LAs are less efficient at shock absorption,
which is in accordance with Rak’s interpretation of the
Neandertal pelvis.
Small LAs in humans are correlated with short stride
length, slow walking velocity and an anteriorly flexed
trunk (Grasso et al., 2000; Sarwahi et al., 2002; Hirose
et al., 2004, Jang et al., 2009). Nonetheless, less-lordotic
spines are clearly more stable and less mobile than
more-lordotic lumbar spines (Scholten et al., 1988) and
can carry heavy loads without developing high shear
stress on the lumbar structures (Shirazi-Adl and Par-
nianpour, 1999; Meakin et al., 2008). If this holds true
for Neandertals, it might suggest, on the one hand, that
Neandertals were better adapted to carry heavy loads
and, potentially, to engage in generally more rigorous
upper body activities (Pearson, 2000; Weaver, 2009). On
the other hand, it might suggest that Neandertals had a
short stride length and slower walking velocity com-
pared with modern humans as has been suggested by
Polk (2004). Nonetheless, if the upper body is more dor-
sally oriented, then less lordosis would be needed to
 HOMININ LORDOSIS
keep the center of mass in the same relative position.
This would allow gait features such as knee flexion and
stride length to be unaffected. Future biomechanical
research should investigate the effect of these complex
relationships on Neandertal gait.
CONCLUSIONS
If nonhuman apes are representative of the lumbar
morphology of the last common ancestor between
humans and nonhuman apes, then lumbar LA increased
dramatically during hominin evolution from 208  258 as
seen in nonhuman apes to [508 in H. sapiens. This
change appeared early in hominin evolution, because the
lumbar spines of australopithecines and H. erectus show
human like LAs of 40  458. Neandertals are different
from all other hominins (with the possible exception of
the hominins from Sima de los Huesos, who are poten-
tially Neandertal ancestors), exhibiting relatively small
LAs (308). The reduced lordosis of Neandertals sug-
gests postural and locomotive differences from the other
hominins.
ACKNOWLEDGMENTS
The authors wish to thank Dr. Frances Thackeray (the
Transvaal Museum of National History, Pretoria, South
Africa), Prof. Jean-Jacques Hublin (Max Planck Institute
for Evolutionary Anthropology, Leipzig) and Dr. Philip
Mennecier (Musee de l’Homme Paris-MNHN) for ena-
bling us to study material in their care. Special thanks
to Prof. Yoel Rak for his useful comments, to Dr. Michal
Catz for her help, and to Ms. Anna Behar and Mr. Alon
Barash for the illustrations. Additional thanks to Ms.
Aurélie Fort and Prof. James C. Ohman for the help
with the radiographs, and to our colleagues from TAU,
LCHES, ISCIII and LEH. We would also like to thank
our reviewers for their instructive comments.
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