Evidence for interpersonal violence in the
St. Césaire Neanderthal
Christoph P. E. Zollikofer*†, Marcia S. Ponce de León*, Bernard Vandermeersch‡, and François Lévêque§
*Anthropological Institute and MultiMedia Laboratory兾Department of Computer Science, University of Zürich, 8057 Zürich, Switzerland; ‡Laboratoire
d’Anthropologie, Université de Bordeaux I, 33405 Talence, France; and §Résidence Charles Perrault, 3, Rue de Provence, 86000 Poitiers, France
Communicated by Erik Trinkaus, Washington University, St. Louis, MO, February 26, 2002 (received for review December 1, 2001)
The St. Césaire 1 Neanderthal skeleton of a young adult individual
is unique in its association with Châtelperronian artifacts from a
level dated to ca. 36,000 years ago. Computer-tomographic imag-
ing and computer-assisted reconstruction of the skull revealed a
healed fracture in the cranial vault. When paleopathological and
forensic diagnostic standards are applied, the bony scar bears
direct evidence for the impact of a sharp implement, which was
presumably directed toward the individual during an act of inter-
personal violence. These findings add to the evidence that Nean-
derthals used implements not only for hunting and food process-
ing, but also in other behavioral contexts. It is hypothesized that
the high intra-group damage potential inherent to weapons might
have represented a major factor during the evolution of hominid
social behavior.
Châtelperronian 兩 paleopathology 兩 tool use 兩 trauma 兩 computer
tomography
T he St. Césaire 1 Neanderthal partial skeleton was discovered
in 1979 at the site of La Roche à Pierrot (near the village of
St. Césaire, Charente Maritime, France), a collapsed rock shelter
comprising a sequence of Mousterian, Châtelperronian, and
Aurignacian deposits (1). The skeleton was recovered from level
EJ0P (1), which contains a Châtelperronian assemblage ther-
moluminescence-dated to ⬇36,000 years ago (2, 3). This ensem-
ble represented the first direct evidence for the association of
Neanderthals with Châtelperronian implements. Together with
a similar association from the site of Arcy-sur-Cure (4), these
finds spurred an intense and ongoing debate over the evolution-
ary, paleodemographic, and cultural relationships between local
Neanderthal populations and the early modern human (EMH)
newcomers during the early Upper Paleolithic in Europe.
The St. Césaire 1 skeleton is fragmented and partially eroded,
but the reconstructed craniomandibular and long bone diaphy-
seal morphology permitted significant inferences regarding the
phyletic status and behavioral specializations of this individual.
The craniomandibular morphology of the specimen largely
corresponds to the ‘‘classical’’ Neanderthal type (5). Tooth
microwear analysis suggests a meat-rich diet comparable to that
of earlier Neanderthals and modern hunting societies (6). The
morphology of the well-preserved right femoral shaft indicates
Neanderthal-type hyperarctic body proportions, but cross-
sectional biomechanical analysis suggests locomotor patterns
closer to those of EMH than of classic Neanderthals (7).
Here we report on recently discovered paleopathological
aspects of the morphology of the fossil. During computer-
assisted reconstruction of the skull, we detected a healed frac-
ture in the cranial vault. When paleopathological diagnostic
standards (8, 9) are applied, this bony scar bears direct evidence
for the impact of a sharp implement, which may have been
directed toward the individual during an act of interpersonal
violence. We discuss the possible behavioral context of this
evidence and its implications for hominid behavior during the
Middle-to-Upper Paleolithic transition in Europe.
Materials and Methods
The St. Césaire skeletal remains belong to a young adult, possibly
male, individual. All preserved cranial and postcranial elements
6444 – 6448 兩 PNAS 兩 April 30, 2002 兩 vol. 99 兩 no. 9
derive from a spatially confined area with a diameter of no more
than 70 cm (1). The excavation yielded no signs of a burial pit;
yet the local inhomogeneity in the distribution of rocks and
implements at the spot where the skeleton was found, as well as
its association with Dentalium shells (10), may be indicative of an
intentional burial (11). The postcranium is represented by
fragments of the axial skeleton and the limb bones, some of
which were found in anatomical connection. The skull was lying
on its right side, with the upper and lower jaws in anatomical
association. Most of the preserved cranial structures come from
the right side and comprise the mandible and maxillae (up to the
left lateral incisors), the face, and the right anterolateral region
of the braincase. The internal lamina of the cranial vault bones
is partially eroded, and with the exception of several isolated
teeth, the left cranial half is missing (Fig. 1). Deterioration and
loss of these elements is probably because of temporary exposure
and weathering of the upper layers of the sediment in which the
fossil was embedded (1).
The physical reconstruction of the fragmented right hemi-
mandible and skull was performed by one of us (B.V.) and
revealed taphonomic deformation of the face relative to the
braincase, resulting in an everted position of the cranial vault
relative to the midplane of the skull. As an effect of the general
flattening of the cranial morphology, the anatomical connec-
tions between larger reconstructed pieces remained inconclu-
sive. Among these was an apical vault fragment representing
substantial portions of the right frontal and parietal bones joined
along the coronal suture. This fragment is delimited by post-
mortem fractures on its anterior, posterior, and lateral sides (Fig.
1). The medial margin, however, exhibits a smooth border, which
extends in an anteroposterior direction across the coronal suture
and was initially interpreted as representing the partially fused
sagittal (interparietal) and metopic (interfrontal) sutures.
To correct the taphonomic deformation of the skull and
re-assess the anatomical position of this piece, we performed a
computerized reconstruction of the skull. Virtual reconstruction
was indispensable because the brittleness of the original fossil
material prevented physical disassembly and manipulation of the
specimen. Following three-dimensional data acquisition with
computer tomography (CT), all fragments were isolated elec-
tronically from filling material and then recomposed on the
computer screen, according to procedures and criteria described
in ref. 12 (a detailed account of the new reconstruction will be
given elsewhere). One result was that the medial border of the
isolated cranial vault piece clearly does not correspond to the
midsagittal plane of the skull (Fig. 1), nor does it represent
normal cranial anatomy at its true position lateral of the
midsagittal plane.
We established a differential diagnostic scheme, according to
which this unusual morphology, its underlying causes, and
Abbreviations: EMH, early modern human(s); CT, computer tomography.
†To whom reprint requests should be addressed. E-mail: zolli@ifi.unizh.ch.
The publication costs of this article were defrayed in part by page charge payment. This
article must therefore be hereby marked “advertisement” in accordance with 18 U.S.C.
§1734 solely to indicate this fact.
www.pnas.org兾cgi兾doi兾10.1073兾pnas.082111899

Fig. 1. Computerized reconstruction of the St. Césaire 1 Neanderthal skull
(mirror-imaged completed parts are transparent) showing the bony scar in the
right apical cranial vault. The skull is seen from the direction in which the
hypothetical blow was exerted. Arrows indicate the anteroposterior (a and p)
extent of the preserved lateral border of the injury (c, coronal suture; b,
bregma; *, location of the coronal cross section through the injury (see Fig. 2b);
ⴛ, location of the parasagittal cross section through the coronal suture (see
Fig. 3c); the dotted line indicates the reconstructed position of the midsagittal
plane of the cranial vault). Note oblique, off-midline position of the scar.
(Scale bar ⫽ 5 cm.)
behavioral context could be studied at increasing levels of detail
(Table 1). For the comparative analysis of the external and
internal structure of the bone, we used a sample of adult
specimens comprising (i) an archeological specimen exhibiting a
healed slash resulting from sharp trauma to the cranial vault
(13), (ii) three specimens with healed trepanations [one neolithic
(14) and two modern skulls], and (iii) four specimens exhibiting
metopic sutures and兾or persisting fontanelles.
Paleopathology
The medial border of the cranial vault fragment of the St. Césaire
Neanderthal deviates in its external and internal structure from
both postmortem fractures and interosseous sutures. The exter-
nal lamina of the bone is rounded off toward the medial margin,
and the diploic region is covered with cortical bone (Fig. 2). This
morphology is characteristic of in vivo apposition of bone matrix
and excludes postmortem abrasion and erosion as potential
mechanisms. Bone apposition at structural boundaries may
occur under two different circumstances: normal periosteal
growth in sutural tissue and bone regeneration following an
injury. The off-axis position and oblique orientation of the
Zollikofer et al.
margin (Fig. 1) exclude that it represents parts of the interpa-
rietal and a supposed metopic suture. On the other hand, a
presumed suture in parasagittal position would imply the exis-
tence of a large persisting fontanelle or a bregmatic ossicle (15).
However, the fairly straight margin does not correspond to a
border around a fontanelle or a suture around a bregmatic
ossicle. Furthermore, the cross-sectional structure of the bone
clearly differs from that of an interosseous suture, which exhibits
a corrugated border with less dense cortical bone than the border
of a healed wound (Fig. 3). The most probable cause of the
observed morphology is therefore bone regeneration following
a lesion, a scenario that is corroborated by the close match of the
cross-sectional morphology of the St. Césaire fragment (Fig. 3a)
with that of comparative specimens exhibiting healed trepana-
tions and scars (Fig. 3 b, e, and f ).
In a next step, we analyzed the morphology of the injury with
the aim to infer the proximate mechanical causes of the lesion
and to reconstruct its posttraumatic history. As evidenced by the
nearly straight border of the scar, the individual most probably
suffered a lesion from a blade-shaped object. The resulting slash
in the cranial vault is preserved on its right (lateral) side over a
length of 68 mm; it was probably slightly longer in vivo. The left
(medial) side of the groove was lost during fossilization. Judging
from the degree of bone remodeling along the margin, the injury
ANTHROPOLOGY
reached its greatest depth near the coronal suture, correspond-
ing to the presumed primary site of impact. Toward the anterior
and posterior ends, the groove tapers off. This morphology most
closely matches the pattern of direct, sharp trauma. Under these
biomechanical conditions, the high but localized stresses caused
by the impacting object lead to puncturing兾cutting of the scalp
and the external lamina of the bone, comminution of the diploë,
and separation兾displacement of bone fragments from the in-
ternal lamina (9). The St. Césaire specimen in fact bears
evidence that at the primary site of the impact the internal
lamina was fractured and parts of it were dislocated whereas,
toward the periphery of the slash, it was only partially severed
(Figs. 1 and 2). The interpretation of the slash as a linear
fracture resulting from blunt trauma is a less likely scenario
because the compressive forces induced by blunt objects lead
to nonlocal deformation of the cranial vault, typically resulting
in multiple radial and tangential fractures around the center of
impact (9, 16).
Comparison with an archeological specimen exhibiting a
healed slash on the frontal bone permits further inferences
regarding the severity of the injury and the subsequent healing
process (Fig. 2). Given the moderate depth of the St. Césaire
scar, it appears that the lesion was relatively mild. Cross-sectional
CT images (Figs. 2 and 3) show that the injured region was
extensively remodeled during lifetime; the severed external
lamina was smoothed out by bone resorption and deposition, and
the dislocated parts of the internal lamina were fixated, probably
through formation of connective tissue in the diploic region. The
external and cross-sectional morphology of the injured region
does not show any signs of post-traumatic infection (e.g., peri-
ostitis or osteomyelitis), and the degree of healing is relatively
advanced. Considering that bone healing is visible only 2–3
weeks after a traumatic event (9), it can be concluded that the
individual survived the injury for at least some months, such that
a direct causal connection between the trauma and the individ-
ual’s death is unlikely.
Behavioral Context
By using forensic criteria originally developed for trauma anal-
ysis in extant and archeological populations (9), it is possible to
conceive of various scenarios under which the St. Césaire injury
could have occurred, assess their relative likelihood, and discuss
their behavioral and motivational implications. The anteropos-
terior orientation of the slash as well as its apical position are
PNAS 兩 April 30, 2002 兩 vol. 99 兩 no. 9 兩 6445
Table 1. Differential diagnostic scheme for trauma analysis in fossil hominids
Criterion* Range of possible alternatives

Paleopathology, proximate causes

Incident In vivo • Postmortem
Etiology Trauma • Pathology
Epigenetic variant
Type of trauma Direct • Indirect
Type of lesion Penetrating, sharp • Blunt
Severity of trauma Mild • Fatal
Post-traumatic Complete healing • Chronic impairment
Object causing lesion Implement • Natural object
Implement type Stone ? Wood
Forensics, ultimate causes
Behavioral context Intentional • Accidental
Motivation Aggression • Ritual
Medical
Action Casual (short-term) • Premeditated (long-term)
Actor Opponent • Self-infliction
Social context Intragroup • Intergroup (intra-兾interspecific)
Recovery Autonomous • Supported (nursing)

The position of the • represents the inferred status of St. Césaire between alternatives.
*Compiled after refs. 8, 9, 16, 18, and 19.

indicative of a blow or a thrust exerted against the individual
from the front or from behind, assuming that the individual was
in an upright position. These spatial relationships indicate an
intentional action, effected with an implement rather than a
natural object. Accidental injury, such as falling onto a sharp
edge, a rockfall, or an unintentional blow, such as resulting from
a hunting incident, are less likely explanations; comparative
forensic evidence suggests that accidental trauma typically af-
fects the sides of the cranial vault, as opposed to the apical
location of intentional injuries (16).
Inferences regarding the nature of the implement, the agents
causing the lesion, and the behavioral context must remain
tentative. From a mechanical point of view, the severity of sharp
trauma depends on the mass, impact velocity, and incisiveness of
the weapon. The wide range of possible combinations of these

Fig. 2. Anatomy of a blow. (a) Mediolateral view of the right border of the
St. Césaire 1 cranial vault injury (same symbols as in Fig. 1: a and p, anterior and
posterior delimitations of the scar, c, position of the coronal suture, and *,
location of the cross section in b. The drawing indicates areas of bone
remodeling (gray) and areas affected by postmortem fracturing (hatched). (b
and c) Comparative morphology of the injured vault bones in St. Césaire 1 (b)
and a specimen from a medieval graveyard (13) exhibiting a healed scar
resulting from sharp trauma on the left frontal bone (c) (both specimens in
anteroposterior view, coronal cross sections); the CT sections show bone
remodeling at the margin of the injured external lamina (*) and a space (‚)
between the dislocated internal lamina and the diploë, probably filled by
connective tissue. (Scale bar ⫽ 5 cm.)
Fig. 3. Comparative cross-sectional morphology of bone injuries and su-
tures. (a and b) Healed scars of St. Césaire 1 and a medieval specimen (same
coronal sections as in Fig. 2 b and c). (c) St. Césaire 1 coronal suture (parasag-
ittal section through ⴛ in Fig. 1). (d) Metopic suture in a recent specimen
(coronal section). (e) Large occipital trepanation during early phases of heal-
ing [Ensisheim skull (14), coronal section]. ( f ) Healed occipital trepanation in
a recent specimen (coronal section). Note apposition of dense cortical bone at
the injured borders, resulting in a smooth contour, as opposed to the less
dense and corrugated aspect of the sutural borders. (Scale bar ⫽ 5 cm.)

6446 兩 www.pnas.org兾cgi兾doi兾10.1073兾pnas.082111899 Zollikofer et al.

parameters encompasses a correspondingly wide spectrum of
implements that might have been used to inflict the wound. For
example, the Châtelperronian stone tools recovered from the
site of St. Césaire are relatively small and exhibit retouched,
blunt edges. To attain the kinetic energy necessary to penetrate
bone, considerable acceleration, probably through hafting (17),
would have been essential. Implications regarding the nature of
the weaponry are complicated by the fact that the function of
Châtelperronian implements is poorly understood and the pre-
served stone industry probably represents a small fraction of the
actual in vivo spectrum of available tools.
Comparative evidence from historical populations suggests
that interpersonal conflict behavior resulting in cranial vault
fractures is relatively frequent (18, 19). Given the characteristic
apical location of the injury in St. Césaire, direct interaction with
another individual is the most parsimonious scenario. Theoret-
ically, the injury may have resulted from intragroup, intergroup,
or even interspecific conflict behavior. The first scenario is the
most likely one because in socially organized species the vast
majority of interpersonal interactions occur at the within-group
level (20). Population densities were low during the Late Pleis-
tocene, such that mutual avoidance during the rare encounters
between different groups (both intra- and interspecific) might
have represented the optimum strategy for the resolution of
potential conflict (21). Nevertheless, it must be considered that
patchy resource distribution might have induced temporary
between-group competition.
Another aspect that requires consideration is the motivational
background of the conflict in which the St. Césaire Neanderthal
was involved. Motivations may range from a premeditated
assault to a brief argument emerging from a temporary conflict
between individuals, such as over social status, access to poten-
tial mates, or intragroup resources. Overall, a likely scenario for
the interpersonal violence in St. Césaire is intragroup tempers
with available ‘‘weaponry.’’ The immediate effects of the trauma
were probably serious, implying heavy bleeding, cerebral com-
motion, and temporary impairment. Although it is possible that
the individual sustained these adverse effects autonomously, it
can be assumed that it had benefited at least to some extent from
initial intragroup assistance.
Discussion
The St. Césaire cranial injury adds to the extremely small sample
of specimens bearing direct evidence that Neanderthals used
implements during acts of interpersonal violence. The only other
clearly documented example, probably older than 50,000 years,
comes from the Shanidar 3 Neanderthal. This specimen exhibits
a slash in the superior margin of the ninth left rib, resulting from
a penetrating implement, which remained stuck between neigh-
boring ribs until the individual’s death, but which was lost
postmortem (22, 23). These rare cases of tool-mediated wounds
must be interpreted in an evolutionary, behavioral, and cultural
context, using comparative data on trauma in Neanderthals and
fossil EMH, as well as modern evidence on the relationship
among trauma, interpersonal violence, and tool use in human
and nonhuman primates.
Neanderthals were shown to differ from fossil and extant
EMH populations in both the frequency and pattern of skeletal
injury: The overall incidence of trauma was comparatively high
and concentrated to the head and neck (21). Differences in
pattern of trauma between Neanderthals and the EMH from the
Upper Paleolithic have not yet been investigated systematically,
but it appears that the overall frequency of trauma was lower in
the EMH populations, whereas the proportion of head and neck
injuries was probably as elevated as in the Neanderthals (E.
Trinkaus, personal communication). Taking into account that
injuries tend to accumulate over an individual’s lifetime, such
that older individuals typically exhibit a greater number of
Zollikofer et al.
lesions (21, 24), it is worth noting that immature specimens of
Neanderthals [Devil’s Tower and Le Moustier (25, 26) and EMH
(Qafzeh 11)] also bear evidence for craniofacial trauma. The
elevated frequency of trauma might therefore reflect not only a
high risk of injury during close-quarter hunting of medium to
large-sized game (21), but a physically demanding and stressful
life from early ontogenetic stages (27, 28).
Against this comparative background, the dearth of direct
evidence for tool involvement in fossil hominid trauma may be
interpreted in two different ways. On one hand, it may reflect a
low in vivo frequency of intentional or accidental tool-inflicted
injuries relative to other types of trauma. It also may reflect the
limits of paleopathological diagnosis; in a fossil, a lesion induced
by an implement can be recognized only if evidence of healing
is still present postmortem and the morphology of the scar
permits inferences regarding the responsible weapon. The prob-
ability of detecting such injuries is further reduced through the
effects of diagenesis, which tends to degrade the paleopatho-
logical evidence (9). Following this line of argument, a relevant
proportion of the nondiagnostic injuries may represent tool-
induced trauma and兾or may be seen in the context of interper-
sonal violence rather than accident.
Data from nonhuman primates indicate that interpersonal
violence within social networks represents a major cause of
ANTHROPOLOGY
trauma. For example, in the Gombe chimpanzees more than
one-half of the observed skeletal lesions are fractures and bite
wounds resulting from intragroup violence (24), and several
cases of fatal outcome of intragroup clashes have been reported
in both wild and captive chimpanzee groups (29). A question
arises whether there is evidence of tool use in interpersonal
conflict situations in nonhuman primates. The cultural variabil-
ity and contextual diversity of tool production and use have been
studied extensively in various primate communities (30–32),
revealing a general behavioral basis to aim objects, but not tools,
toward conspecifics in the context of conflict behavior. However,
although object-throwing to intimidate group members is fairly
common (30, 33), the purposeful and directed use of tools
designed for a distinct function during acts of interpersonal
violence has not been reported. On the basis of the currently
available evidence, it appears therefore that hominids differ
from nonhuman primates in their ability to produce and use tools
in an expanded, multifunctional context, including conflict be-
havior. Accordingly, the St. Césaire 1 and Shanidar 3 wounds
indicate that Neanderthals transformed a ‘‘tool’’ into a ‘‘weap-
on’’; in other words, they used an implement in a functional
context which differed from that for which it was originally
designed.
The cognitive ability to use tools multifunctionally was most
probably acquired earlier during hominid evolution. Direct
evidence for this hypothesis is scant, but it appears that the
cognitive capacities and technical skills of early hominids are
typically underestimated because relatively little is known about
the in vivo level of complexity of tool production and utilization.
Nevertheless, evidence for hafting of Mousterian stone imple-
ments with heat-processed bitumen (17, 34) and for the use of
aerodynamically designed wooden hunting spears (35) suggest
that the variety, effectiveness, and material sophistication of
Middle and Late Pleistocene implements, and therefore of their
utilization, was considerable. The intentional use of implements
in the context of intragroup conflict must have had a major
impact during hominid evolution because the availability of
highly effective hunting and兾or food-processing tools in inter-
personal conflict created a new and considerable potential for
intragroup damage, a potential that required specific behavioral
adjustments with which to cope. Intragroup aggression in pri-
mate societies must be understood as one specific behavioral
option in a complex network of social interactions, which is
typically balanced by active reconciliatory behavior (20) and兾or
PNAS 兩 April 30, 2002 兩 vol. 99 兩 no. 9 兩 6447
the minimization of social interactions under crowding
conditions (36).
If we adhere to the hypothesis that the St. Césaire individual
was injured in an act of intragroup violence and was later assisted
to some extent during healing, this fossil lesion sheds light on
both the disruptive兾deleterious and integrative兾supportive as-
pects of Neanderthal behavior. This fits well into the picture that
Neanderthals were capable of sustaining severely impaired
individuals over extended periods of time (21), and that this
behavioral competence was already present in the Middle Pleis-
tocene (37). We must therefore conceive that Neanderthals,
depending on the context, inflicted wounds to conspecifics and
nursed the injured, using aggressive and integrative behavioral
elements as tools in a network of social interactions. Within this
basic hominid pattern of behavior, implements probably played
a crucial role because of their high effectiveness in interpersonal
violence and because they represented an additional level of
complexity of social interactions.
The difficulty to infer an indisputable behavioral context from
the observed skeletal traumatic alterations is therefore not only
because of the limitations of paleodiagnosis but also may be
indicative of the behavioral polyvalence of tool use in Neander-
thals. Accordingly, the links among form, purpose, and effective
function of a tool might have been relatively loose. The most
prominent behavioral context of Neanderthal tool use directed
toward conspecifics is cannibalism, for which evidence has been
advanced since the early days of Neanderthal research (38).
However, even the clearest evidence for butchering of conspe-
cifics from the site of Moula-Guercy (39, 40) ultimately remains
unresolved in terms of its behavioral and motivational context.
Implications that Neanderthal tool use was far more sophisti-
cated than generally assumed also comes from other sources,
1. Lévêque, F. & Vandermeersch, B. (1980) C. R. Acad. Sci. 291, 187–189.
2. Valladas, H., Geneste, J.-M., Joron, J.-L. & Chadelle, J.-P. (1986) Nature
(London) 322, 335–344.
3. Mercier, N., Valladas, H., Joron, J. L., Reyss, J. L., Lévêque, F. & Vander-
meersch, B. (1991) Nature (London) 351, 737–739.
4. Hublin, J. J., Spoor, F., Braun, M., Zonneveld, F. & Condemi, S. (1996) Nature
(London) 381, 224–226.
5. Vandermeersch, B. (1984) Bull. Mém. Soc. Anthropol. Paris 1, 191–196.
6. Lalueza, C., Pérez-Pérez, A. & Turbón, D. (1996) Am. J. Phys. Anthropol. 100,
367–387.
7. Trinkaus, E., Ruff, C. B., Churchill, S. E. & Vandermeersch, B. (1998) Proc.
Natl. Acad. Sci. USA 95, 5836–5840.
8. Courville, C. B. (1967) in Diseases in Antiquity, eds. Brothwell, D., Sandison,
A. T. & Dawson, W. R. (Thomas, Springfield, IL), pp. 606–622.
9. Lovell, N. C. (1997) Yearbook Phys. Anthropol. 40, 139–170.
10. Lévêque, F., Backer, A. M. & Guilbaud, M., eds. (1993) Context of a Late
Neandertal (Prehistory Press, Madison, WI).
11. Vandermeersch, B. (1993) in Context of a Late Neandertal, eds. Lévêque, F.,
Backer, A. M. & Guilbaud, M. (Prehistory Press, Madison, WI).
12. Zollikofer, C. P. E., Ponce de León, M. S. & Martin, R. D. (1998) Evol.
Anthropol. 6, 41–54.
13. Lanz, C. (1998) Schweiz. Ärzteztg. 79, 935–938.
14. Alt, K. W., Jeunesse, C., Buitrago-Tellez, C. H., Wächter, R., Böes, E. &
Pichler, S. (1997) Nature (London) 387, 360 (lett.).
15. Hauser, G. & De Stefano, G. F. (1989) Epigenetic Variants of the Human Skull
(E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart).
16. Fischer, H. (1972) in Handbuch der Speziellen Pathologischen Anatomie und
Histologie, ed. Uehlinger, E. (Springer, Berlin), Vol. 9, pp. 353–424.
17. Boëda, E., Connan, J., Dessort, D., Muhesen, S., Mercier, N., Valladas, H. &
Tisnérat, N. (1996) Nature (London) 380, 336–338.
18. Walker, P. L. (1989) Am. J. Phys. Anthropol. 80, 313–324.
19. Weber, J. & Czarnetzki, A. (2001) Am. J. Phys. Anthropol. 114, 352–356.
20. de Waal, F. B. (2000) Science 289, 586–590.
21. Berger, T. D. & Trinkaus, E. (1995) J. Archaeol. Sci. 22, 841–852.
22. Trinkaus, E. & Zimmerman, M. R. (1982) Am. J. Phys. Anthropol. 57, 61–67.
23. Trinkaus, E. (1983) The Shanidar Neanderthals (Academic, New York).
24. Jurmain, R. (1989) Am. J. Phys. Anthropol. 80, 229–237.
6448 兩 www.pnas.org兾cgi兾doi兾10.1073兾pnas.082111899
notably the evidence of burial and a hypothetical case of cranial
vault deformation (41).
Conclusions
To reconstruct the causes and consequences of the traumatic
event that affected the St. Césaire individual, we followed a
maximum-likelihood approach based on comparative fossil and
actualistic data. We come to the conclusion that the cranial
injury was inflicted with a tool during an act of intragroup
interpersonal violence. In a wider context, we suggest that the
basic behavioral and cognitive abilities to use implements during
interpersonal conflict were probably already present early during
hominid evolution and may represent a significant aspect of the
evolution of social tool use. Accordingly, it can be assumed that
in this specific respect, no major ‘‘transition’’ from Neanderthal-
specific to EMH-specific behavioral patterns during the early
Upper Paleolithic took place. This process most likely went
across species and was eminently patterned, both spatially and
temporally. Although genetic, developmental, and morpholog-
ical data suggest that Neanderthals and EMH are separated at
the species level (42–44), their ways to balance between aggres-
sive and cooperative tool-mediated behavioral patterns were
largely similar.
This study has greatly benefited from discussions with E. Trinkaus, T.
Böhni, R. Majcen, C. van Schaik, J. Zilhão, H. Thieme, and M. Schultz,
as well as from the comments and suggestions of three anonymous
reviewers. The constant support of P. Stucki is gratefully acknowledged.
We thank K. Alt, U. Bochsler, and Ch. Lanz for kindly providing the
archeological specimens. We also thank Gea Bijl for technical assistance
during CT scanning of the comparative sample. This work was supported
by the Swiss National Science Foundation.
25. Zollikofer, C. P. E., Ponce de León, M. S., Martin, R. D. & Stucki, P. (1995)
Nature (London) 375, 283–285.
26. Ponce de León, M. S. & Zollikofer, C. P. E. (1999) Anat. Rec. 254, 474–489.
27. Ogilvie, M. D., Curran, B. K. & Trinkaus, E. (1989) Am. J. Phys. Anthropol. 79,
25–41.
28. Trinkaus, E. (1995) J. Archaeol. Sci. 22, 121–142.
29. Fawcett, K. & Muhumuza, G. (2000) Am. J. Primatol. 51, 243–247.
30. McGrew, W. C. (1992) Chimpanzee Material Culture: Implications for Human
Evolution (Cambridge Univ. Press, Cambridge, U.K.).
31. Whiten, A., Goodall, J., McGrew, W. C., Nishida, T., Reynolds, V., Sugiyama,
Y., Tutin, C. E., Wrangham, R. W. & Boesch, C. (1999) Nature (London) 399,
682–685.
32. van Schaik, C. P. & Knott, C. D. (2001) Am. J. Phys. Anthropol. 114, 331–342.
33. van Schaik, C. P., Deaner, R. O. & Merrill, M. Y. (1999) J. Hum. Evol. 36,
719–741.
34. Mania, D. & Toepfer, V. (1973) Königsaue: Gliederung, Oekologie und Mittel-
paläolithische Funde der letzten Eiszeit (Deutscher Verlag der Wissenschaften
VEB, Berlin).
35. Thieme, H. (1997) Nature (London) 385, 807–810.
36. Aureli, F. & de Waal, F. B. (1997) Am. J. Primatol. 41, 213–228.
37. Lebel, S., Trinkaus, E., Faure, M., Fernandez, P., Guerin, C., Richter, D.,
Mercier, N., Valladas, H. & Wagner, G. A. (2001) Proc. Natl. Acad. Sci. USA
98, 11097–11102.
38. Gorjanovič-Kramberger, D. (1908) Korr. Deutsch. Ges. Anthropol. Ethnol.
Urgesch. 39, 108–112.
39. Defleur, A., Dutour, O., Valladas, H. & Vandermeersch, B. (1993) Nature
(London) 362, 214 (lett.).
40. Defleur, A., White, T., Valensi, P., Slimak, L. & Crégut-Bonnoure, E. (1999)
Science 286, 128–131.
41. Pap, I., Tillier, A.-M., Arensburg, B. & Chech, M. (1996) Ann. Hist.-Nat. Hung.
Nat. Mus. 88, 233–270.
42. Krings, M., Capelli, C., Tschentscher, F., Geisert, H., Meyer, S., von Haeseler,
A., Grossschmidt, K., Possnert, G., Paunovič, M. & Pääbo, S. (2000) Nat. Genet.
26, 144–146.
43. Ponce de León, M. S. & Zollikofer, C. P. E. (2001) Nature (London) 412,
534–538.
44. Stringer, C. B. & Gamble, C. (1993) In Search of the Neanderthals: Solving the
Puzzle of Human Origins (Thames and Hudson, London).
Zollikofer et al.