brief communications
A lost Neanderthal neonate found
A remarkable discovery in a French museum answers some long-standing questions.
F
ossil remains of adult Neanderthals are
well documented, but juvenile speci-
mens are rare and information about
them is scant. Here we identify a beautifully
preserved skeleton that has been lost to sci-
ence for almost 90 years as the Neanderthal
neonate known as ‘Le Moustier 2’, which was
originally found at Le Moustier in the Dor-
dogne, southwest France. This find will be a
P. JUGIE/MUSÉE NATIONAL DE PRÉHISTOIRE
Figure 1 The neonatal skeleton Le Moustier 2, one of the most
complete Neanderthal individuals ever discovered. The parietals
and the posterior part of the hemifrontals are broken into small
fragments. The left temporal, both scapulae and the pubis are
missing. Most of the cranial base and face, deciduous tooth
germs, cervical vertebrae and long bones are preserved. The right
humerus and femur, formerly misidentified as La Ferrassie 4, are
not shown. Many Neanderthal derived traits can be seen in the
lateral and basal portions of the occipital bones and the petrous
portion of the temporal bone. Derived traits are also evident in the
postcranial skeleton in the medio–lateral curvature of the radius,
the relative proportion of the thumb phalanxes, the orientation of
the ulna head and the curvature of the ribs. The thickness and
mass of the bones are noteworthy.
NATURE | VOL 419 | 5 SEPTEMBER 2002 | www.nature.com/nature
rich source of data for studying the evolution
of human ontogeny1 as well as the phylo-
genetic relationship between these extinct
hominids and anatomically modern humans.
The burial site of Le Moustier 2 was
discovered at the lower rock shelter in Le
Moustier by Peyrony in 1914 (ref. 2). The
burial pit originated in archaeological level
J (ref. 3), a Mousterian cultural layer that
has been dated to 40,30052,600 years BP by
thermoluminescence4. Although at the time
the bones were thought to belong to a
neonate2,3, the specimen was not investigated
further and the remains were thought to
have been lost in Paris5.
During a survey in 1996 of collections in
the Musée National de Préhistoire in Les
Eyzies-de-Tayac-Sireuil, France, the remains
of a neonatal skeleton were found among the
lithic sample from Le Moustier. Some bones
were isolated, but others were still embedded
in blocks of sediment. The possibility that
these might be the bones of Le Moustier 2
prompted a re-examination of Peyrony’s
notes, which made no mention of the speci-
men ever having been sent to Paris.
Several crucial details emerged during
the cleaning and restoration of the speci-
men (Fig. 1) that confirm that this skeleton
is indeed the missing Le Moustier 2. The
fossil was embedded in a sediment of sand
containing green hornblendes, garnets and
mica particles of muscovite type, a compo-
sition that has always been associated with
the deposition of the Vézère river, which
flows past the Le Moustier cliff. Preliminary
sedimentological analysis revealed many
similarities with layers I and J of Le Mous-
tier. Furthermore, flint flakes and faunal
remains (Rangifer tarandus, Cervus sp.,
Capra hircus ibex and a large Bovinae) in
the sediment around the skeleton are iden-
tical to those found in the Mousterian levels
at the Le Moustier inferior rock shelter.
Estimations of stature based on the length
of the long bones6 confirmed that the age at
death was no more than four months.
Although Le Moustier 2 is one of the
most complete Neanderthal individuals to
have been discovered, both scapulae and the
pubis are missing. However, the right femur
and right humerus, which were previously
also missing, have now been found, having
been wrongly attributed to the Neanderthal
La Ferrassie 4 (LF4).
It had been assumed that the right
femur and right humerus came from the
burial pit of the La Ferrassie 4bis Neander-
thal (LF4bis), even though their colour
and fossilization differ markedly from those
of LF4bis (ref. 7). Moreover, the sediment
© 2002 Nature Publishing Group
adhering to the LF4 bones contains mus-
covite, a mineral that is not present at La
Ferrassie but which is prevalent at Le
Moustier. Besides the matching fossilization
and colour, the bones’ morphology, vari-
ability, muscular insertions, discrete traits
and measurements are identical to those of
the left humerus and femur of the specimen
from the Musée National de Préhistoire.
It is likely that the two bones attributed
to LF4 are in fact samples from Le Moustier
2 that were sent by Peyrony to Boule for
an age diagnosis. The attribution of the
LF4 limb bones to Le Moustier 2 eliminates
from the archaeological record the only
double burial associated with Neanderthals.
The morphology of Le Moustier 2 dif-
fers greatly from that of living human
neonates, but shows many similarities with
the features of juvenile and adult Neander-
thals. For example, the left stapes has
asymmetrical limbs7; there is no infra-
orbital depression on the maxilla8; and the
premaxillary suture is fully open on both
the palatal surface (accompanied by two
interincisive sinuses) and the nasal floor9.
Also, the zygomatic bone is short in
comparison with the length of the frontal
process, as in immature Neanderthals10; the
sagittal profile of the nasal bones is the
same as that in adult Neanderthals8,11; and
in the dentition, the crowns of the central
and lateral upper deciduous incisors are
shovel-shaped and biconvex, features that
are often evident in Neanderthals12.
This taxonomic diagnosis awaits final
confirmation through atomic mass spectro-
scopic carbon-dating of the skeleton.
Nevertheless, the differences between the
modern human skeleton and that of Le
Moustier 2 bear out the proposed extensive
genetic variation between Neanderthals
and extant humans13.
Bruno Maureille
UMR 5809 CNRS, Laboratoire d’Anthropologie des
Populations du Passé, Université Bordeaux,
1 avenue des Facultés, 33405 Talence, France
e-mail: b.maureille@anthropologie.u-bordeaux.fr
1. Ponce de Léon, M. S. & Zollikofer, C. P. E. Nature 412,
534–538 (2001).
2. Peyrony, D. Rev. d’Anthropol. 40, 48–76, 155–176 (1930).
3. Peyrony, D. Les Moustériens Inhumaient: Ils Leurs Morts?
(Ribes, Périgueux, 1921).
4. Valladas, H. et al. Nature 322, 452–454 (1986).
5. Heim, J.-L. Les Hommes Fossiles de La Ferrassie Vol. 1 (Masson,
Paris, 1976).
6. Sellier, P., Tillier, A.-M. & Bruzek, J. Am. J. Phys. Anthropol. 22
(suppl.), 208 (1997).
7. Heim, J.-L. Les Enfants Néandertaliens de La Ferrassie (Masson,
Paris, 1982).
8. Maureille, B. La face chez Homo erectus et Homo sapiens:
Recherche sur la variabilité morphologique et métrique. Thesis,
Univ. Bordeaux 1, France (1994).
9. Maureille, B. & Bar, D. J. Hum. Evol. 37, 137–152 (1999).
33
brief communications

RSPCA PHOTO LIBRARY

10. Dodo, Y., Kondo, O., Muhesen, S. & Akasawa, T. in
Neanderthals and Modern Humans in Western Asia
(eds Akasawa, T., Aoki, K. & Bar-Yosef, O.) 323–338 (Plenum,
New York, 1998).
11. Trinkaus, E. The Shanidar Neanderthals (Academic,
New York, 1983).
12. Patte E. La Dentition des Néandertaliens (Masson,
Paris, 1962).
13. Krings, M. et al. Nature Genet. 26, 144–146 (2000).
Competing financial interests: declared none.

Ecology combination with other culling practices, so

a ban would lead to a significant population
Effect of British hunting increase with consequential economic dam-
age and loss of biodiversity4.
ban on fox numbers The main fox-hunting season in Britain

P
ressure to ban the hunting of foxes with
hounds in Britain has fuelled debate
about its contribution to the control of
fox populations. We took advantage of a
nationwide one-year ban on fox-hunting
during the outbreak of foot-and-mouth
disease (FMD) in 2001 to examine this issue
and found that the ban had no measurable
impact on fox numbers in randomly selected
areas. Our results argue against suggestions
that fox populations would increase markedly
in the event of a permanent ban on hunting.
In the 2000–2001 season, foxes were hunt-
ed (Fig. 1) by 196 registered foxhound packs
and 7 harrier packs1, and by an unknown
number of unregistered packs, the latter
mainly in southwest England and Wales2.
However, their contribution to controlling
fox numbers is unclear. The arguments are
that the number of foxes killed by hunting is
relatively small, and therefore that hunting
has little influence in regulating population
size3, and conversely that hunting is impor-
tant for limiting fox numbers, particularly in
a 0.3
0.2
Absolute change
in faecal density
runs from November to March/April; cub-
hunting lasts from August to October, and
‘problem’ foxes are targeted after the main
season. From 23 February to 17 December
2001, the Foot-and-Mouth Disease Declara-
tory (Controlled Area) Order 2001 banned
hunting. From 17 December, some hunts
were licensed to hunt in FMD-free areas, and
from 11 February 2002, any hunt could be
licensed if certain restrictions were observed.
Hunting was thus banned for ten months and
severely curtailed for two further months.
With 240,000 adult foxes, and 425,000
cubs born in Britain each year5, 64% mor-
tality per annum is required to prevent
growth of the fox population6. A one-year
ban should therefore lead to a detectable
population increase if hunting has a signifi-
cant impact on fox numbers.
To quantify the changes in fox numbers
during the hunting ban, we surveyed 160
one-kilometre squares between 1 February
and 17 March in 1999 and 2000 (pre-FMD)
and 2002 (post-FMD) — that is, towards
the end of the main culling period and just
before the cubs are born. We estimated
relative fox density by using faecal-accumu-
Figure 1 Hunting with hounds. This centuries-old British tradition
may not be particularly effective in controlling fox populations.
(Kolmogorov–Smirnov goodness-of-fit test,
Z40.71, P40.70), with homogenous vari-
ances between regions (Levene statistic,
F8,15141.84, P40.07). There was a signifi-
cant difference between nine regions in the
relative magnitude of the change in faecal
density (ANOVA, F8,14542.69, P40.01).
Tukey’s HSD post hoc comparisons showed
a significant increase in eastern England
(one sample t-test, t2242.52, P40.02) and
a significant decrease in southern England
(t37412.65, P40.01). No other region devi-
ated significantly from zero in test values.
We evaluated the relative change in
faecal density against a regional index of
hunting pressure (number of days hunted
by registered packs per week per km2 in the
2000–2001 season1). There was no associa-
tion between the reduction in hunting pres-
sure in each region and the relative change
in faecal density (linear regression with
replication, r 240.007, F1,15841.18, P40.28;
Fig. 2b). We conclude that there was no
significant change in fox numbers during

0.1 lation rates7 along transects (mean length, the one-year hunting ban, and that in most
0 6.9 km) that followed linear features; we regions the average faecal density had
–0.1 surveyed each transect twice in each period declined. Our results therefore support the
–0.2
(pre- and post-FMD). We removed all fox view taken by the Committee of Inquiry
faeces on the first visit, and counted fresh into Hunting with Dogs2 that a permanent
–0.3
EE CE NS M NE SE SWE W SS faeces 2–6 weeks later. We calculated faecal ban on hunting is unlikely to result in a
Region density for each transect in each period (F) dramatic increase in fox numbers.
b EE as F4S/KD, where S is the number of faeces Philip J. Baker, Stephen Harris,
in faecal density (R')

0.8 NE NE
on the second visit, K is the transect length Charlotte C. Webbon
Relative change

CE WM SWE
0.4 SS SE in kilometres, and D is the number of days School of Biological Sciences, University of Bristol,
0 between visits. Absolute changes in faecal Woodland Road, Bristol BS8 1UG, UK
–0.4
density were calculated as Fpost-FMD1Fpre-FMD, e-mail: s.harris@bristol.ac.uk
and relative changes (R8) as log(R&1), 1. Anon. Baily’s Hunting Directory 2000–2001 (Pearson,
–0.8
where R4(Fpost-FMD1Fpre-FMD)/(Fpre-FMD&1). Cambridge, 2000).
Overall, faecal density declined by 4.7% 2. Burns, L., Edwards, V., Marsh, J., Soulsby, L. & Winter, M.
0. 0
25
0. 0

0. 5
20
10

45
0. 0
35
05

3
0

4
00
00
00

00

00
00

00
00
00
00

Report of the Committee of Inquiry into Hunting with Dogs in

0.

(Fig. 2a), with no significant difference in

0.
0.

0.
0.

0.

England and Wales (Stationery Office, London, 2000).

Regional hunting pressure
in 2000–2001 season
Figure 2 Changes in faecal density before and after the ban on
fox-hunting during the 2001 outbreak of foot-and-mouth disease
(FMD) in Britain. a, Mean (5s.d.) change in absolute faecal
density by region (post-FMD minus pre-FMD). b, Relationship
between regional reduction in hunting pressure and relative
change in faecal density. CE, central England, n416; EE, eastern
England, n423; M, Midlands, n421; NE, northern England,
n415; NS, northern Scotland, n48; SE, southern England,
n438; SS, southern Scotland, n49; SWE, southwest England,
n425; W, Wales, n45. The results of this survey are posted by
The Mammal Society at www.mammal.org.uk.
the number of squares where faecal density
increased (n483) compared with those
that decreased or remained unchanged
(n477) in ‘hunted’ (n4118) as oppposed
to ‘not-hunted’ (n442) squares (x 2140.19,
P40.66). Moreover, neither the absolute
(t15840.60, P40.55) nor the relative
(t15840.51, P40.61) change in faecal density
differed between ‘hunted’ and ‘not-hunted’
squares. ‘Hunted’ squares lay within the
area covered by a hunt, as designated on a
map issued in 1996 (ref. 8).
R8 values were normally distributed
3. Macdonald, D. W. & Johnson, P. J. in The Exploitation of
Mammal Populations (eds Taylor, V. J. & Dunstone, N.)
160–207 (Chapman & Hall, London, 1996).
4. Tapper, S. A Question of Balance: Game Animals and their Role
in the British Countryside (Game Conservancy Trust,
Fordingbridge, Hampshire, 1999).
5. Harris, S., Morris, P., Wray, S. & Yalden, D. A Review of British
Mammals: Population Estimates and Conservation Status of
British Mammals other than Cetaceans (Joint Nat. Conserv.
Commit., Peterborough, 1995).
6. Harris, S. & Saunders, G. Symp. Zool. Soc. Lond. 65,
441–464 (1993).
7. Putman, R. J. Mammal Rev. 14, 79–97 (1984).
8. Alexander, K. Baily’s Hunting Directory 1996–1997 (Pearson,
Cambridge, 1996).
Competing interests: financial, declared none; P.J.B. and C.C.W.
were funded by the International Fund for Animal Welfare.

34 © 2002 Nature Publishing Group NATURE | VOL 419 | 5 SEPTEMBER 2002 | www.nature.com/nature