A Neanderthal partial femoral diaphysis from the “grotte de la Tour”, La Chaise-de-Vouthon (Charente, France): Outer morphology and endostructural organization

Abstract

We describe a human partial femoral shaft discovered during speleological exploration of the “grotte de la Tour”, near the prehistoric site of La Chaise-de-Vouthon (Charente, France). The context of discovery is compatible with a hyena den deposit; the associated mammal assemblage suggests a preliminary chronological attribution to MIS 3. Combined information from its outer morphology, cross-sectional geometric properties, and from the high-resolution 3D imaging and quantitative analysis of its inner structural organization shows that this specimen (CDV-Tour 1) is from an adult Neanderthal individual, more likely a male.RésuméNous décrivons une portion de diaphyse fémorale humaine découverte lors de l’exploration spéléologique de la « grotte de la Tour », près du site préhistorique de La Chaise-de-Vouthon (Charente, France). Le contexte de découverte est compatible avec un dépôt de tanière de hyène ; l’assemblage des restes mammifères associés suggère une attribution chronologique au MIS 3. La morphologie externe, les propriétés géométriques des sections et l’analyse quantitative tridimensionnelle à haute résolution de la structure interne montrent que ce spécimen (CDV-Tour 1) appartient à un individu néandertalien adulte, vraisemblablement de sexe masculin.

Full text

C.
R.
Palevol
11
(2012)
581–593
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Human
palaeontology
and
prehistory
(Palaeoanthropology)
A
Neanderthal
partial
femoral
diaphysis
from
the
“grotte
de
la
Tour”,
La
Chaise-de-Vouthon
(Charente,
France):
Outer
morphology
and
endostructural
organization
La
diaphyse
fémorale
néandertalienne
de
la
«
grotte
de
la
Tour
»,
La
Chaise-de-Vouthon
(Charente,
France)
:
morphologie
externe
et
organisation
endostructurale
Laurent
Puymeraila,∗,b,
Virginie
Volpatoc,
André
Debénathb,
Arnaud
Mazurierd,
Jean-Franc¸
ois
Tournepichee,f,
Roberto
Macchiarellib,g
aUnité
d’Anthropologie
bio-culturelle,
Droit,
Éthique
et
Santé
(ADÉS),
UMR
7268,
Université
d’Aix-Marseille-EFS-CNRS,
boulevard
Pierre-Dramard,
13344
Marseille,
France
bDépartement
de
Préhistoire,
Muséum
national
d’Histoire
naturelle,
UMR
7194,
75005
Paris,
France
cPaleoanthropology
and
Messel
Research,
Senckenberg
Research
Institute,
60325
Frankfurt
am
Main,
Germany
dSociété
Études
Recherches
Matériaux,
86000
Poitiers,
France
eMusée
d’Angoulême,
16000
Angoulême,
France
fIPGQ-UMR
5199,
université
Bordeaux-1,
33405
Talence,
France
gDépartement
Géosciences,
Université
de
Poitiers,
86022
Poitiers,
France
a
r
t
i
c
l
e
i
n
f
o
Article
history:
Received
23
May
2012
Accepted
after
revision
18
June
2012
Available
online
24
August
2012
Presented
by
Yves
Coppens
Keywords:
Grotte
de
la
Tour
La
Chaise-de-Vouthon
Femoral
shaft
Neanderthal
France
a
b
s
t
r
a
c
t
We
describe
a
human
partial
femoral
shaft
discovered
during
speleological
exploration
of
the
“grotte
de
la
Tour”,
near
the
prehistoric
site
of
La
Chaise-de-Vouthon
(Charente,
France).
The
context
of
discovery
is
compatible
with
a
hyena
den
deposit;
the
associated
mammal
assemblage
suggests
a
preliminary
chronological
attribution
to
MIS
3.
Combined
information
from
its
outer
morphology,
cross-sectional
geometric
properties,
and
from
the
high-resolution
3D
imaging
and
quantitative
analysis
of
its
inner
structural
organization
shows
that
this
specimen
(CDV-Tour
1)
is
from
an
adult
Neanderthal
individual,
more
likely
a
male.
©
2012
Académie
des
sciences.
Published
by
Elsevier
Masson
SAS.
All
rights
reserved.
Mots
clés
:
Grotte
de
la
Tour
La
Chaise-de-Vouthon
Diaphyse
fémorale
Néandertal
France
r
é
s
u
m
é
Nous
décrivons
une
portion
de
diaphyse
fémorale
humaine
découverte
lors
de
l’exploration
spéléologique
de
la
«
grotte
de
la
Tour
»,
près
du
site
préhistorique
de
La
Chaise-de-
Vouthon
(Charente,
France).
Le
contexte
de
découverte
est
compatible
avec
un
dépôt
de
tanière
de
hyène
;
l’assemblage
des
restes
mammifères
associés
suggère
une
attribu-
tion
chronologique
au
MIS
3.
La
morphologie
externe,
les
propriétés
géométriques
des
∗Corresponding
author.
E-mail
address:
puymerail@mnhn.fr
(L.
Puymerail).
1631-0683/$
–
see
front
matter
©
2012
Académie
des
sciences.
Published
by
Elsevier
Masson
SAS.
All
rights
reserved.
doi:10.1016/j.crpv.2012.07.001

582
L.
Puymerail
et
al.
/
C.
R.
Palevol
11
(2012)
581–593
sections
et
l’analyse
quantitative
tridimensionnelle
à
haute
résolution
de
la
structure
interne
montrent
que
ce
spécimen
(CDV-Tour
1)
appartient
à
un
individu
néandertalien
adulte,
vraisemblablement
de
sexe
masculin.
©
2012
Académie
des
sciences.
Publié
par
Elsevier
Masson
SAS.
Tous
droits
réservés.
1.
Introduction
The
“grotte
de
la
Tour”
is
a
new
cavity
of
paleoan-
thropological
interest
discovered
near
the
prehistoric
site
of
La
Chaise-de-Vouthon,
in
the
department
of
Charente
(Poitou-Charentes
region,
western
France).
This
famous
Middle
to
Late
Pleistocene
site
complex,
in
a
Middle
Juras-
sic
(Bajocian-Bathonian)
limestone
plateau,
consists
of
a
network
of
NW-oriented
karstic
caves
whose
opening
is
∼10
m
above
the
Tardoire
river
(Debénath,
2006).
Discovered
in
1850
and
extensively
excavated
between
1930
and
1963
(by
P.
David)
and
between
1967
and
1983
(by
A.D.),
La
Chaise-de-Vouthon
comprises
two
major
rock
shelters/caves,
both
having
provided
Neanderthal
remains:
the
Abri
Suard,
whose
deposits
were
originally
referred
to
a
period
ranging
from
Riss
II
to
Riss-Würm,
and
the
Abri
Bourgeois-Delaunay,
covering
a
period
from
Riss
III
to
Würm
III
(Blackwell
et
al.,
1983,
1992;
Debénath,
1974,
1977,
2006;
Schwarcz
and
Debénath,
1979).
According
to
a
new
set
of 230Th/234 U
speleothem
dates
(Couchoud,
2006;
Vieillevigne
et
al.,
2008),
the
human
remains
from
the
Abri
Suard,
all
from
MIS
6
and
associated
to
a
typically
cold
mammal
fauna
(Griggo,
1995),
are
younger
than
163
kyr,
while
the
age
of
those
from
Bourgeois-Delaunay
(MIS
5e,
Eemian),
associated
to
a
temperate
fauna
(Armand,
1998),
is
between
127
and
116
kyr
(Debénath,
2006).
To
date,
La
Chaise
has
provided
a
human
fossil
sample
consisting
of
75
subadult
and
adult
Neanderthal
remains
(52
from
Suard
and
23
from
Bourgeois-Delaunay),
includ-
ing
a
number
of
isolated
teeth
(Condemi,
2001;
Debénath,
2006;
Genet-Varcin,
1974,
1975a,
b,
1976;
Legoux,
1976;
Macchiarelli
et
al.,
2006;
Piveteau,
1953,
1955,
1970a,
b;
Piveteau
et
al.,
1982;
Teilhol,
2001,
2003;
Tillier
and
Genet-
Varcin,
1980).
Among
the
adult
postcranial
remains,
a
rather
slender
partial
right
femoral
shaft
of
239.5
mm,
BD
5,
was
recovered
at
the
Abri
Bourgeois-Delaunay
(Condemi,
2001:
pp.
137–152).
The
human
femoral
shaft
reported
in
this
study
was
collected
in
2006
by
D.
Augier
during
speleological
explo-
ration
of
a
cavity
named
“grotte
de
la
Tour”
(Tour
of
La
Chaise
cave),
whose
entrance
is
at
∼130
m
W
of
Abri
Suard
(0◦2648 E,
45◦4013 N)
(Fig.
1).
The
specimen,
CDV-Tour
1,
was
found
on
surface
associated
to
a
bovid
long
bone
fragment
among
the
soil
déblais,
likely
accumulated
by
a
badger
near
the
entrance
of
a
narrow
duct
leading
to
a
larger
inner
cavity.
About
10
m
from
this
spot,
a
concen-
tration
of
faunal
remains
showing
the
characteristics
of
a
hyena
den
deposit,
with
bones
broken
or
etched
from
digestion,
has
been
discovered
inside
the
duct.
A
prelimi-
nary
surface
survey
of
the
cavity
revealed
no
archaeological
evidence
of
human
activity,
which
supports
the
interpreta-
tion
of
a
carnivore
den,
similar
to
the
contexts
ascertained
for
the
cave
sites
of
Rochelot
(Tournepiche
et
al.,
1996)
and
Rochers-de-Villeneuve
(Beauval
et
al.,
2005),
from
the
same
region,
both
having
provided
Neanderthal
remains.
The
faunal
remains
from
Tour
of
La
Chaise
collected
so
far
represent
Equus
sp.
cf.
germanicus,
E.
hydruntinus,
Sus
scrofa,
Bovinae
(Bos
and
Bison),
Cervus
elaphus,
Capreolus
capreolus,
Megaloceros
sp.,
Crocuta
crocuta
spelaea,
Canis
lupus,
Panthera
spelaea,
P.
pardus,
Ursus
spelaeus,
Vulpes
vulpes.
As
a
whole,
this
Late
Pleistocene
mammal
assem-
blage,
suggesting
a
rather
temperate
climate,
overlaps
those
reported
for
the
nearby
site
of
La
Chauverie
(El
Albani
et
al.,
2011)
and
from
Rochers-de-Villeneuve,
in
the
neigh-
bouring
department
of
Vienne
(Beauval
et
al.,
2005),
both
dated
to
MIS
3.
Nonetheless,
this
preliminary
chronological
setting
deserves
confirmation
and,
based
on
the
available
information,
it
should
be
only
considered
as
the
most
likely.
CDV-Tour
1
(Fig.
2)
represents
a
shaft
portion
from
a
left
femur
whose
diaphyseal
dimensions
and
proportions,
subperiosteal
contour,
and
inner
structural
morphology
globally
differ
from
the
conditions
seen
in
fossil
and
extant
modern
humans
and
allow
its
attribution
to
an
adult
Nean-
derthal.
2.
Methods
of
analysis
CDV-Tour
1
has
been
imaged
in
2008
by
X-ray
microto-
mography
(␮CT)
at
the
Centre
de
Microtomographie
of
the
University
of
Poitiers.
Two
acquisitions
have
been
run
to
scan
the
entire
shaft
and
the
records
have
been
merged
after
volume
reconstruction.
Scans
were
realized
by
a
X8050-16
Viscom
AG
(image
intensifier
with
a
1004
×
1004
12
bit
camera
and
a
pixel
size
of
147
␮m)
according
to
the
following
parameters:
120
kV
voltage,
560
␮A
current,
32
integrations/projection,
and
1200
projections
(each
0.30◦).
The
final
volume
was
reconstructed
using
DigiCT
v.2.4.2
(Digisens)
in
8-bit
format,
with
an
isotropic
voxel
size
of
107.8
␮m.
Because
of
globally
good
outer
and
inner
bone
preser-
vation
quality
and
the
lack
of
any
sedimentary
infill
of
the
medullary
cavity,
a
semi-automatic
threshold-based
seg-
mentation
with
manual
corrections
has
been
successfully
carried
out
following
the
Half-Maximum
Height
method
(HMH;
Spoor
et
al.,
1993)
and
the
region
of
interest
thresh-
olding
protocol
(ROI-Tb;
Fajardo
et
al.,
2002)
and
by
taking
repeated
measurements
on
different
slices
of
the
virtual
stack
(Coleman
and
Colbert,
2007)
by
Avizo
v.6.2.
(Visual-
ization
Sciences
Group
Inc.)
and
ImageJ
v.1.46a
(Rasband,
2010).
To
orient
the
specimen
anatomically,
we
used
the
posi-
tion
of
the
linea
aspera.
The
approximate
position
of
the
midshaft
was
confidently
defined
on
comparative
ground
through
the
morphology
of
the
linea
aspera
and
the
shaft
outline
in
medial
and
lateral
views.
The
50%
cross-section
precise
location
is
usually
less
clearly
indicated
in
fossil
human
femora
displaying
subcircular
diaphyses
and
lack-
ing
a
true
pilaster,
such
as
CDV-Tour
1;
however,
this
region
shows
little
longitudinal
changes
in
morphology
along
several
centimetres,
making
modest
errors
in
the
location

L.
Puymerail
et
al.
/
C.
R.
Palevol
11
(2012)
581–593
583
Fig.
1.
Location
of
«grotte
de
la
Tour
»
(La
Chaise-de-Vouthon,
Charente,
France)
and
plan
of
the
site:
1:
place
of
discovery
of
the
human
femoral
shaft
near
the
entrance
of
the
duct;
2:
the
hyena
den
deposit.
Survey:
P.
Vauvilliers;
drawing:
J.-F.
Tournepiche.
Fig.
1.
Localisation
de
la
«
grotte
de
la
Tour
»
(La
Chaise-de-Vouthon,
Charente,
France)
et
plan
du
site
:
1
:
emplacement
de
la
découverte
de
la
diaphyse
fémorale
humaine
près
de
l’entrée
de
la
galerie
;
2
:
le
dépôt
de
la
tanière
d’hyène.
Relevé
:
P.
Vauvilliers
;
dessin
:
J.-F.
Tournepiche.
of
these
sections
not
critical
for
the
structural
characteri-
zation
of
the
specimen
(Sládek
et
al.,
2010;
Trinkaus
and
Ruff,
2012).
Cross-sectional
geometric
(CSG)
properties
of
CDV-Tour
1
were
measured
on
four
selected
virtual
sections
(A
to
D)
respectively
corresponding
to:
the
uppermost
(A,
62.4
mm
above
the
midshaft)
and
the
lowermost
(D,
33.4
mm
below
the
midshaft)
complete
sections
found
along
the
shaft
in
proximo-distal
direction;
the
midshaft
(C);
the
mid-
distance
between
the
uppermost
and
the
midshaft
sections
(B)
(Fig.
3).
As
CDV-Tour
1
misses
both
articular
ends,
we
could
not
directly
measure
its
biomechanical
length.
How-
ever,
if
the
average
value
of
∼407
mm
obtained
for
a
sample
of
eight
European
Neanderthals
(La
Chapelle-aux-Saints
1,
Feldhofer
1,
La
Ferrassie
1
and
2,
Fond-de-Forêt
1,
Palomas
92
and
96,
Spy
2;
in
Trinkaus
and
Ruff,
2012,
table
A1)
is
used
as
a
proxy
for
estimating
the
biomechanical
length
of
CDV-Tour
1,
the
segment
A-D
(95.8
mm)
should
approxi-
mately
correspond
to
the
shaft
portion
comprised
between
42%
(distal)
and
65%
(proximal)
of
its
biomechanical
length,
the
section
B
corresponding
to
∼57.5%.
In
addition
to
the
anteroposterior
(a-m)
and
mediolat-
eral
(m-l)
outer
diameters
(in
millimeter,
both
measured
directly
on
the
original
shaft
by
a
dial-equipped
vernier
caliper
to
the
nearest
0.1
mm,
and
also
on
the
virtually
reconstructed
specimen),
the
following
CSG
parameters
have
been
considered
by
using
a
custom
routine
developed
in
R
v.2.11.1
(R
Development
Core
Team,
2011):
total
area
(TA,
in
mm2);
cortical
area
(CA,
in
mm2);
percent
of
corti-
cal
area
(%CA);
second
moments
of
area
about
the
m-l
and
a-p
axes
(Ix,
Iy,
in
mm4);
polar
second
moment
of
area
(J,
in
mm4);
section
modulus
about
the
m-l
and
a-p
axes
(Zx,
Zy,
in
mm3);
polar
section
modulus
(Zp,
in
mm3);
maximal
and
minimal
second
moments
of
area
(Imax,
Imin,
in
mm4)
(Ruff,
2008).
In
order
to
place
CDV-Tour
1
within
the
pattern
of
variation
reported
for
the
European
Late
Pleistocene
fossil
record,
comparative
data
for
%CA
and
the
Ix/Iy
ratio
mea-
sured
at
50%
and
65%
cross-sectional
level
were
obtained
for
a
number
of
Neanderthal
(Neand.)
and
Upper
Pale-
olithic
(EUPH)
human
femora
from
the
database
recently
published
by
Trinkaus
and
Ruff
(2012),
integrated
by
fresh
evidence
on
a
Neanderthal
femoral
shaft
from
the
MIS
4
French
site
of
Les
Pradelles
(Mussini
et
al.,
2012)
(Table
1).
For
the
comparative
assessment
of
the
pilastric
index
(diameter
a-p/m-l
×
100),
we
used
the
same
samples
detailed
in
Table
1
integrated
by
data
from
the
follow-
ing
specimens:
Santa
Croce
1
and
Stadelhöhle
1,
among
the
Neanderthals,
and
Barma
Grande
6,
Caviglione
1,
and
Pˇ
redmostí
3,
4,
9,
10
and
14,
for
the
Upper
Paleolithic
sam-
ple
(Trinkaus
and
Ruff,
2012,
table
A13).
By
using
an
original
(␮)CT
record,
we
also
compara-
tively
detailed
cortical
bone
thickness
variation
at
eight
specific
sites
(anticlockwise:
posterior,
postero-lateral,
lateral,
antero-lateral,
anterior,
antero-medial,
medial,
postero-medial)
of
the
50%
and
65%
cross-sections
in

584
L.
Puymerail
et
al.
/
C.
R.
Palevol
11
(2012)
581–593
Fig.
2.
CDV-Tour
1
(left
femoral
shaft)
in
anterior
(a),
posterior
(b),
medial
(c),
and
lateral
(d)
views
(proximal
is
upper).
Scale
bar:
1.5
cm.
Fig.
2.
CDV-Tour
1
(diaphyse
fémorale
gauche)
en
vues
antérieure
(a),
postérieure
(b),
médiale
(c)
et
latérale
(d)
(proximale
vers
le
haut).
Échelle
:
1,5
cm.
CDV-Tour
1,
La
Ferrassie
1
(left),
Rochers-de-Villeneuve
1
(50%),
Spy
8
(Volpato
et
al.,
2012b),
Cro-Magnon
1
(left;
Puymerail,
2011;
Puymerail
and
Macchiarelli,
2009),
and
in
the
Magdalenian
skeleton
from
Chancelade
(right
femur;
Bondioli
et
al.,
2010;
Puymerail,
2011;
Puymerail
and
Macchiarelli,
2009),
as
well
as
in
an
extant
human
reference
sample.
In
our
study,
the
extant
human
condition
is
illustrated
by
20
femora
measured
from
medical
CT
records
per-
formed
at
spatial
resolutions
ranging
from
350
to
791
␮m
(Puymerail,
2011;
Puymerail
et
al.,
2012).
This
sample
(EH)
pools
adults
of
both
sexes
from
19th
century
France
and
the
Imperial
Roman
graveyard
of
Velia,
Italy,
stored
at
the
Musée
de
l’Homme
of
Paris,
the
University
of
Poitiers,
and
the
Museo
Nazionale
Preistorico
Etnografico
“L.
Pigorini”
of
Rome.
Three-dimensional
cortical
thickness
topographic
dis-
tribution
in
CDV-Tour
1
for
the
whole
portion
A-D
has
been
virtually
rendered
for
each
anatomical
projection
by
a
chromatic
scale
increasing
from
dark
blue
(thin)
to
red
(thick),
where
bone
thickness
is
defined
for
each
point
on
the
periosteal
surface
as
the
distance
to
the
closest
point
on
the
endosteal
surface
(Bayle
et
al.,
2011;
Bondioli
et
al.,
2010;
Mazurier
et
al.,
2010;
Puymerail
et
al.,
2012;
Volpato
et
al.,
2011,
2012a,
b).
Technical
advances
in
the
qualitative
and
quantitative
structural
characterization
through
functional
imaging
of
the
femoral
diaphyseal
organization
have
made
possible
the
realization
of
morphometric
maps
suitable
for
syn-
thetic
visual
appreciation
and
comparison
(Bondioli
et
al.,
2010;
Puymerail,
2011;
Puymerail
et
al.,
2012).
Accord-
ingly,
the
A-D
periosteal
surface
portion
of
CDV-Tour
1
has
been
mapped
onto
a
cylinder,
whose
diameter
corre-
sponds
to
the
maximum
width
of
the
original
surface
of
the
shaft,
virtually
unzipped
along
a
predefined
vertical
line
along
the
anterior
aspect,
and
then
unrolled
into
a
plane.
Thickness
values
have
been
standardized
between
0
and
1
(methodological
details
in
Bondioli
et
al.,
2010).
For
the
specific
purposes
of
the
present
study,
we
directly
com-
pared
the
structural
signature
characterizing
CDV-Tour
1
to
the
morphometric
maps
obtained
for
the
42–65%
shaft
portion
of
their
respective
biomechanical
lengths
for
the
Neanderthal
femora
La
Ferrassie
1
(left)
and
Spy
8
and
the
Gravettian
Cro-Magnon
1
(left).
With
this
respect,
we
also
provided
comments
about
the
␮CT-based
morphometric
map
already
generated
for
the
Chancelade
1
femur
(Bayle
et
al.,
2011,
fig.
6;
Bondioli
et
al.,
2010,
fig.
3;
Puymerail
and
Macchiarelli,
2009).
Finally,
since
it
is
possible
to
perform
generalized
additive
models
(GAM)
of
interpo-
lation
to
obtain
consensus
maps
by
merging
individual
morphometric
maps
into
a
single
dataset
(Wood,
2006),
Table
1
The
Late
Pleistocene
European
Neanderthal
(Neand.)
and
European
Upper
Paleolithic
(EUPH)
specimens
used
in
the
comparative
analysis
for
%CA
and
the
Ix/Iy
ratio
measured
at
50%
and
65%
cross-sectional
level
of
the
femoral
diaphysis.
Original
data
selected
from
Trinkaus
and
Ruff
(2012,
tables
A4
and
A5)
(see
also
Mussini
et
al.,
2012).
Tableau
1
Les
spécimens
néandertaliens
européens
du
Pléistocène
supérieur
(Neand.)
et
les
spécimens
du
Paléolithique
supérieur
(EUPH)
utilisés
dans
l’analyse
comparative
du
%CA
et
du
ratio
Ix/Iy
mesurés
à
50
%
et
à
65
%
de
la
diaphyse
fémorale.
Données
extraites
de
Trinkaus
et
Ruff
(2012,
tableaux
A4
et
A5)
(voir
aussi
Mussini
et
al.,
2012).
Sample
Cross-sectional
level
Specimens
Neand.
50%
and
65%
Feldhofer
1,
La
Ferrassie
1
and
2,
Fond-de-Forêt
1,
Spy
2
50%
only
La
Chapelle-aux-Saints
1,
Palomas
96,
La
Quina
5,
Rochers-de-Villeneuve
1,
Saint-Césaire
1,
Les
Pradelles
65%
only Krapina
257.32
and
257.33,
Palomas
52
and
92,
La
Quina
38
EUPH
50%
and
65%
Cro-Magnon
1,
4322
and
4324,
Dolní
V˘
estonice
3,
13,
14
and
16,
Paviland
1,
Pavlov
1,
Sunghir
1
and
4,
Willendorf
1
50%
only Arene
Candide
1,
Barma
Grande
2
(50%),
Dolní
V˘
estonice
35,
Grotte
des
Enfants
4,
Mladeˇ
c
27,
Paglicci
25,
Rochette
2,
Veneri
1
and
2,
65%
only
Dolní
V˘
estonice
41,
Mladeˇ
c
28,

L.
Puymerail
et
al.
/
C.
R.
Palevol
11
(2012)
581–593
585
Fig.
3.
CDV-Tour
1.
Microtomographic-based
reconstruction
of
the
femoral
shaft
in
anterior
(a),
posterior
(b),
medial
(c),
and
lateral
(d)
views
and
topo-
graphic
distribution
of
the
cortical
thickness
for
the
portion
approximately
comprised
between
42%
(distal,
level
D)
and
65%
(proximal,
level
A)
of
its
biomechanical
length
virtually
rendered
by
means
of
a
10-stepped
chromatic
scale
(dark
blue
is
thin,
red
is
thick).
The
cross-sectional
outlines
(posterior
is
below,
medial
is
to
the
right)
are
provided
at
four
levels
(A
to
D),
where
C
corresponds
to
the
midshaft
and
B
is
intermediate
between
A
and
C.
Scale
bar:
1.5
cm.
Fig.
3.
CDV-Tour
1.
Reconstruction
sur
base
microtomographique
de
la
diaphyse
fémorale
en
vues
antérieure
(a),
postérieure
(b),
médiale
(c)
et
latérale
(d)
et
variations
d’épaisseur
de
l’os
cortical
de
la
portion
comprise
entre
42
%
(distal,
niveau
D)
et
65
%
(proximal,
niveau
A)
de
sa
longueur
biomécanique
représentées
selon
une
échelle
chromatique
(en
bleu
les
valeurs
les
plus
fines,
en
rouge
les
plus
épaisses).
Les
sections
transversales
virtuelles
(côté
postérieur
vers
le
bas,
médial
à
gauche)
représentent
quatre
niveaux
(A
à
D),
où
C
correspond
à
la
mi-diaphyse
et
où
B
est
équidistant
de
A
et
C.
Échelle
:
1,5
cm.
we
also
included
in
the
comparative
analysis
the
con-
sensus
map
summarizing
the
whole
variation
expressed
by
our
recent
comparative
sample
(EH;
Puymerail,
2011;
Puymerail
et
al.,
2012).
To
generate
the
maps,
we
used
the
pattern
of
relative
thickness
repartition
(local
morphome-
tric
properties)
obtained
by
a
custom
routine
developed
in
Scilab
v.4.1.2
(The
Scilab
Consortium)
with
the
support
of
the
R
v.2.11.1
(R
Development
Core
Team,
2011)
packages
mcgv
(Wood,
2004,
2006,
2008),
spatstat
(Baddeley,
2008),
and
gstat
(Pebesma,
2004).
3.
The
CDV-Tour
1
human
femoral
shaft
3.1.
Outer
morphology
3.1.1.
Preservation
and
taphonomy
While
lacking
its
metaphyseal
ends,
the
femoral
diaph-
ysis
from
Tour
of
La
Chaise
is
from
a
fully
mature
individual
because
of
its
high
cross-sectional
diameter
values,
dense
and
smooth
subperiosteal
surface,
and
thick
cortical
bone
(Fig.
2).
Maximum
length
of
the
specimen
is
204.7
mm;
the
full
circumference
is
preserved
for
95.8
mm
(47%).
The
mid-
shaft
is
estimated
at
121
mm
from
the
proximal
fractured
end.
All
free
fractured
margins
of
CDV-Tour
1
are
not
fresh
and
show
the
same
patina
at
the
subperiosteal
surface.
The
slightly
spiral-shaped
breaks,
very
likely
originally
pro-
duced
on
the
fresh
bone,
show
relatively
clean
edges
at
the
distal
end
of
the
anterior
aspect
and,
to
a
minor
extent,
at
the
proximal
end
(postero-lateral
aspect).
The
oblique
break
opening
along
the
postero-lateral
aspect
of
the
upper
end
is
58.7
mm
long
and
27.3
mm
wide;
on
the
anterior
aspect,
maximum
length
and
breadth
of
the
distal
break
are
50.5
mm
and
33.31
mm,
respectively.
The
surface
texture,
lacking
distinct
indications
of
weathering
(Behrensmeyer
and
Hook,
1992),
does
not
sug-
gest
significant
displacement
of
the
specimen.
However,
some
multidirectional
cracks
resulting
from
compression
and
locally
affecting
the
entire
cortical
thickness
to
the
endosteal
surface
are
found
towards
the
proximal
end,
notably
on
its
postero-medial
aspect,
as
well
as
on
the
anterior
and
lateral
sides.
Some
superficial
and
faint
longi-
tudinal
striae,
discontinuously
extended
to
about
35
mm,
also
occur
along
the
distal
part
of
the
medial
aspect
and,
around
the
midshaft,
on
the
lateral
side.
No
evident
cortical
thickness
loss
is
found
along
the
shaft
or
at
the
upper
margins.
Only
towards
the
very
distal
edge
of
the
specimen,
the
bone
shows
moderate
thinning
posteriorly
in
association
to
the
presence
of
multiple
carni-
vore
tooth-marks.
Other
isolated
shallow
punctuations
are
also
present
towards
the
proximal
end
of
CDV-Tour
1,
near
the
linea
aspera.
However,
the
nature
of
these
latter
features

586
L.
Puymerail
et
al.
/
C.
R.
Palevol
11
(2012)
581–593
is
not
clear,
due
to
their
proximity
to
the
local
rough-
ness
of
the
linea
and,
mostly,
to
a
broad
and
porous
area
occurring
towards
the
proximal
half
of
the
medial/postero-
medial
aspect
of
the
specimen.
This
diffuse
porosity,
which
mimics
a
modest
periosteal
reaction,
more
likely
relates
to
chemical
dissolution,
a
phenomenon
also
observed
on
a
number
of
the
faunal
remains
from
the
same
den
cavity
and
common
in
similar
contexts
(Tournepiche
et
al.,
1996).
No
macroscopic
pathological
lesions/changes
or
anthropic
marks
are
present
on
this
partial
femoral
diaphysis.
3.1.2.
External
morphology
The
CDV-Tour
1
midshaft
region
shows
a
cross-sectional
morphology
from
slightly
transversally
expanded
to
sub-
circular.
The
antero-posterior
and
medio-lateral
diameters
at
midshaft
correspond
to
31.6
mm
and
33.7
mm,
respec-
tively
(Table
2),
with
a
pilastric
index
of
93.8.
Based
on
its
cross-sectional
values,
which
fall
near
the
upper
range
of
the
Neanderthal
variation
(Table
2),
we
estimate
that
this
globally
robust
shaft
is
more
likely
from
a
male
individual.
As
a
whole,
CDV-Tour
1
is
only
modestly
bowed
anteroposteriorly
(Fig.
2).
Compared
to
the
corresponding
portions
of
the
femora
from
the
Feldhofer
cave,
La
Fer-
rassie
1,
and
Spy
II,
it
is
also
poorly
bowed
latero-medially.
With
this
respect,
it
is
somewhat
intermediate
between
the
almost
straight
(poor
mesial
convexity)
partial
femur
BD
5
from
La
Chaise
Bourgeois-Delaunay
(Condemi,
2001)
and
the
right
femur
Spy
8,
from
Spy
II
(Volpato
et
al.,
2012b).
While
in
CDV-Tour
1
a
pilaster
is
missing,
the
linea
aspera
is
present,
even
if
modestly
salient
and
rather
smooth
along
its
preserved
portion,
with
no
adjacent
flat-
tening.
Proximally,
it
extends
to
the
base
of
the
gluteal
buttress.
Similarly
to
the
condition
described
for
the
RdV
1
Neanderthal
shaft
from
the
likely
penecontemporaneous
site
of
Rochers-de-Villeneuve
(Beauval
et
al.,
2005),
as
well
as
directly
observed
on
La
Ferrassie
1,
its
two
lips
coalesce
into
a
single
relief
locally
forming
a
smooth
crest.
The
max-
imum
breadth
of
this
relief,
encountered
at
medium-distal
level,
reaches
5.5
mm
(vs.
4.0
mm
measured
at
a
distance
of
about
25
mm
above
the
midshaft).
Distally,
the
linea
tends
to
disappear
without
really
splitting
into
a
medial
and
a
lateral
supracondylar
components,
both
present
but
very
poorly
expressed
in
CDV-Tour
1.
This
condition
slightly
differs
from
that
observed
on
La
Ferrassie
1
and
La
Chapelle-
aux-Saints
1,
where
the
lateral
supracondylar
line
is
more
salient
and
the
medial
one
hardly
discernible.
The
portion
above
the
midshaft,
where
the
linea
gently
turns
laterally,
shows
more
wrinkled
and
slightly
sharper
margins.
At
about
one
third
of
the
distance
between
the
midshaft
and
the
distal
end,
a
small
nutrient
foramen
is
present
within
a
shallow
and
narrow
gutter
leaning
against
the
medial
side
of
the
linea.
Just
above,
a
5–6
mm
wide
shal-
low
groove
corresponding
to
the
imprint
of
the
adductor
longus
slightly
obliquely
runs
for
25–30
mm.
Its
position
fits
that
of
similar
imprints
present
on
Feldhofer
1,
where
it
appears
more
vertically
oriented,
and
in
Spy
8,
similarly
ori-
ented
but
slightly
more
medially
displaced.
A
homologous
feature
is
also
present
on
La
Ferrassie
1
(right:
5
×
35
mm)
and
La
Chapelle-aux-Saints
1
(left:
3–4
×
45
mm).
Another
shallow
imprint,
likely
left
by
the
adductor
brevis,
is
found
on
CDV-Tour
1
immediately
medial
to
the
linea
aspera,
about
25–30
mm
above
the
midshaft.
This
feature
is
appar-
ent
on
both
femora
from
La
Chapelle-aux-Saints
1,
as
well
as
on
the
right
femur
of
La
Ferrassie
1.
Antero-medially,
the
corpus
shows
an
oval-shaped
area
of
faint
flattening
around
the
midshaft.
Similar
to
La
Ferrassie
1
and
La
Chapelle-aux-Saints
1,
a
moderately
developed
buttress,
running
slightly
obliquely
proximo-
distally,
is
also
present
medially.
3.2.
Cross-sectional
geometry
and
inner
structure
The
unstandardized
values
of
the
cross-sectional
geometric
properties
of
CDV-Tour
1
measured
(in
proximo-
distal
direction)
at
∼65%
(section
A),
∼57.5%
(B),
at
midshaft
(C)
and
at
∼42%
(D)
of
its
estimated
biomechanical
length
are
shown
in
Table
2.
The
cross-sectional
outlines
of
the
specimen
at
the
same
sites
are
shown
in
Fig.
3.
Percent
cortical
area
(%CA)
decreases
distally,
but
the
lower
value
measured
at
42%
could
reflect
the
mod-
erate
thinning
observed
on
the
posterior
outer
surface
in
association
with
some
carnivore
tooth-marks.
At
any
rate,
a
slight
periosteal
expansion
is
observed
along
the
∼31
mm
between
section
B
and
A.
As
revealed
by
the
second
moment
of
area
about
the
antero-posterior
axis
(Iy)
and
the
polar
section
modulus
(Zp),
a
similar
pattern
is
shown
in
medio-lateral
bending
rigidity
and
torsional
strength,
both
increasing
proximally
in
CDV-Tour
1
in
asso-
ciation
with
a
slight
but
constant
trend
towards
a
more
oval
shaft
(the
Imax/Imin
ratio
shifting
from
1.17,
at
D,
to
1.43,
at
A,
the
values
at
C
and
B
are
1.22
and
1.29,
respectively).
At
midshaft,
the
Zx/Zy
ratio,
which
reflects
antero-posterior
to
medio-lateral
bending
strength
of
the
diaphysis,
is
0.81.
CDV-Tour
1
shows
the
typical
Nean-
derthal
condition
of
medio-lateral
strengthening
(Trinkaus
and
Ruff,
2012;
Volpato
et
al.,
2012b).
At
midshaft,
site-specific
cortical
bone
thickness
follows
the
decreasing
pattern:
posterior
(9.3
mm)
≥
lateral
(9.1
mm)
>
postero-lateral
(8.1
mm)
>
medial
(7.7
mm)
≥
antero-medial
(7.4
mm)
>
anterior
(6.9
mm)
≥
postero-medial
(6.8
mm)
>
antero-lateral
(6.0
mm).
The
average
thickness
at
this
level
corresponding
to
7.7
mm.
At
∼65%
cross-sectional
level,
where
mean
cor-
tical
bone
is
thicker
(8.7
mm),
site-specific
distribution
shows
the
following
model:
lateral
(10.4
mm)
>
medial
(9.2
mm)
≥
posterior
(9.1
mm)
>
antero-lateral
(8.5
mm)
≥
antero-medial
(8.3
mm)
≥
postero-medial
(8.2
mm)
>
postero-lateral
(7.4
mm).
In
a
three-dimensional
perspective,
bone
topographic
variation
in
CDV-Tour
1
along
the
portion
A-D
is
rendered
in
pseudo-colours
in
Fig.
3,
distinctly
for
each
anatomical

L.
Puymerail
et
al.
/
C.
R.
Palevol
11
(2012)
581–593
587
Table
2
Comparative
values
of
the
outer
antero-posterior
(a-p)
and
medio-lateral
(m-l)
diameters
of
the
femoral
diaphysis
and
unstandardized
cross-sectional
geometric
properties
measured
at
four
sections
(A
to
D;
Fig.
3
for
localization)
in
CDV-Tour
1
and
(when
available)
in
three
samples
representing
Late
Pleistocene
European
Neanderthals
(Neand.;
n
=
17),
European
Upper
Paleolithic
(EUPH;
n
=
44)
and
extant
humans
(EH;
n
=
20).
In
italics,
the
standard
deviation.
Neand.
and
EUPH
data
from
Trinkaus
and
Ruff
(2012,
table
A13)
(see
also
Mussini
et
al.,
2012).
See
the
text
(Methods
of
analysis)
for
the
meaning
of
the
variables.
Tableau
2
Mesures
comparatives
des
diamètres
externes
antéropostérieur
(a-p)
et
médio-latéral
(m-l)
de
la
diaphyse
fémorale
et
valeurs
brutes
des
paramètres
de
section
mesurés
au
niveau
des
quatre
sections
(A
à
D
;
Fig.
3
pour
la
localisation)
pour
CDV-Tour
1
et
(si
disponible)
pour
trois
échantillons
de
Néandertaliens
européens
du
Pléistocène
supérieur
(Neand.,
n
=
17),
d’Européens
du
Paléolithique
supérieur
(EUPH,
n
=
44)
et
d’humains
modernes
(EH
;
n
=
20).
Les
valeurs
d’écart-type
sont
en
italique.
Les
valeurs
des
Neand.
et
EUPH
sont
extraites
de
Trinkaus
et
Ruff
(2012,
tableau
A13)
(voir
aussi
Mussini
et
al.,
2012).
Voir
le
texte
(Méthodes)
pour
la
signification
des
variables.
Section
Spec./sample
a-p
m-l
TA
CA
%CA
Ix
Iy
Ix/Iy
J
Zx
Zy
Zx/Zy
Zp
Imax
Imin
∼65%
(A)
CDV-Tour
1
31.6
33.7
745
600
80.5
39
294
48
004
0.81
87
298
2486
2846
0.87
5344
51
387
35
883
Neand.
603
108
504
84
84.0
6.4
26
193
8087
32
994
11
779
0.84
0.22
59
188
18
730
34
981
11
483
24
206
8037
EUPH
596
110
481
100
80.6
5.3
29
520
12
579
27
481
9781
1.01
0.23
57
002
21
331
32
590
12
477
24
411
9133
MH 28.4
2.3
28.2
2.6
594
99
461
77
77.9
7.5
27
798
8686
28
026
9187
1.01
0.14
55
824
17
327
1922
453
1945
476
0.99
0.08
3871
910
30
419
9192
25
405
8311
∼57.5%
(B) CDV-Tour
1
31.2
33.5
741
563
76.0
36
847
47
239
0.78
84
087
2364
2818
0.84
5199
47
354
36
706
MH
29.1
2.4
27.6
2.4
578
93
445
73
77.0
5.0
28
242
8925
25
569
8253
1.11
0.12
53
812
16
952
1907
453
1818
432
1.05
0.06
3730
879
30
061
8811
23
755
8279
50%
(C) CDV-Tour
1 31.2 33.3 744
566
76.2
37
844
46
344
0.81
84
188
2414
2799
0.86
5224
46
327
37
839
Neand.
29.2
2.2
29.4
1.9
656
85
525
87
79.6
5.3
32
755
8420
34
934
8406
0.94
0.14
67
613
15
917
2052
426
2192
449
0.94
0.13
3924
748
38
882
10
123
28
807
6540
EUPH
32.1
4.2
27.6
2.4
607
101
456
93
74.9
6.7
35
513
14
126
24
358
8171
1.45
0.23
59
872
21
748
1949
567
1659
397
1.17
0.14
3377
878
36
503
14
347
23
365
7672
MH
29.7
2.5
27.6
2.5
581
91
428
67
73.8
5.5
29
150
8896
24
713
7660
1.18
0.13
53
863
16
283
1928
442
1762
395
1.09
0.07
3699
829
30
428
8712
23
437
7727
∼42%
(D) CDV-Tour
1
31.5
32.6
760
510
67.2
39
011
43
618
0.89
82
630
2474
2672
0.92
5150
44
702
37