RESEARCH ARTICLE

Lethal Interpersonal Violence in the MiddlePleistoceneNohemi Sala1*, Juan Luis Arsuaga1,2, Ana Pantoja-Pérez1, Adrián Pablos3,1,Ignacio Martínez3,1, Rolf M. Quam1,4,5, Asier Gómez-Olivencia6,7,8,1, José María Bermúdezde Castro9, Eudald Carbonell10,11,12

1 Centro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos, Madrid, Spain, 2 Departamento dePaleontología, Facultad Ciencias Geológicas, Universidad Complutense de Madrid, Madrid, Spain, 3 Áreade Antropología Física, Departamento de Ciencias de la Vida, Universidad de Alcalá, Madrid, Spain,4 Department of Anthropology, Binghamton University (SUNY), Binghamton, New York, United States ofAmerica, 5 Division of Anthropology, American Museum of Natural History, New York, New York, UnitedStates of America, 6 Departamento Estratigrafía y Paleontología, Facultad de Ciencia y Tecnología, EuskalHerriko Unibertsitatea, Bilbao, Spain, 7 IKERBASQUE, Basque Foundation for Science, Bilbao, Spain,8 Équipe de Paléontologie Humaine, Département de Préhistoire, Muséum national d'Histoire naturelle,Musée de l’Homme, Paris, France, 9 Centro Nacional de Investigación sobre la Evolución Humana, Burgos,Spain, 10 Àrea de Prehistòria, Departamento d’Història i Història de l’Art, Universidad Rovira i Virgili,Tarragona, Spain, 11 Institut Català de Paleoecologia Humana i Evolució Social, Tarragona, Spain,12 Institute of Vertebrate Paleontology and Paleoanthropology of Beijing, Beijing, China

* nsala@isciii.es

AbstractEvidence of interpersonal violence has been documented previously in Pleistocene mem-

bers of the genus Homo, but only very rarely has this been posited as the possible manner

of death. Here we report the earliest evidence of lethal interpersonal violence in the hominin

fossil record. Cranium 17 recovered from the Sima de los Huesos Middle Pleistocene site

shows two clear perimortem depression fractures on the frontal bone, interpreted as being

produced by two episodes of localized blunt force trauma. The type of injuries, their location,

the strong similarity of the fractures in shape and size, and the different orientations and im-

plied trajectories of the two fractures suggest they were produced with the same object in

face-to-face interpersonal conflict. Given that either of the two traumatic events was likely

lethal, the presence of multiple blows implies an intention to kill. This finding shows that the

lethal interpersonal violence is an ancient human behavior and has important implications

for the accumulation of bodies at the site, supporting an anthropic origin.

IntroductionInterpersonal violence (lethal and nonlethal) in prehistory is of special interest since it providesa window into human social relations in the past and may be associated with subsistence con-texts such as competition for scarce resources, population density or territorial defense [1–2].Interpersonal violence can be manifested in different ways in the archaeological record, includ-ing trauma on hominin bones, which makes it susceptible to approach these questions in

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OPEN ACCESS

Citation: Sala N, Arsuaga JL, Pantoja-Pérez A,Pablos A, Martínez I, Quam RM, et al. (2015) LethalInterpersonal Violence in the Middle Pleistocene.PLoS ONE 10(5): e0126589. doi:10.1371/journal.pone.0126589

Academic Editor: Christopher Bae, University ofHawaii at Manoa, UNITED STATES

Received: March 9, 2015

Accepted: April 6, 2015

Published: May 27, 2015

Copyright: © 2015 Sala et al. This is an open accessarticle distributed under the terms of the CreativeCommons Attribution License, which permitsunrestricted use, distribution, and reproduction in anymedium, provided the original author and source arecredited.

Data Availability Statement: All relevant data arewithin the paper and its Supporting Information file.

Funding: This research was supported by theMinisterio de Economía y Competitividad of thegovernment of Spain (Project Nos. CGL2012-38434-C03-01, 02 & 03). CT scanning was carried out incollaboration with the Laboratorio de la EvoluciónHumana at the Universidad de Burgos (Spain) withfunding provided by the Junta de Castilla y LeónProject No. BU005A09. Fieldwork at the Atapuercasites was funded by the Junta de Castilla y León andthe Fundación Atapuerca. N.S and A.P.P. havereceived postdoctoral and predoctoral respectively

paleoanthropological contexts through the application of modern forensic methods of traumaanalysis. Interpersonal violence is well-documented since at least Neolithic times [3–5]. In re-cent prehistory, perimortem human manipulation in the form of cutmarks or bone breakagepatterns has often been interpreted as cannibalism [6–9] and could indicate violence betweenhuman social groups [10]. Evidence of cannibalism and defleshing is also present during thePaleolithic and has been documented in fossil hominins dating to at least the Early Pleistocene[11–14]. Nevertheless, there is no evidence of direct traumatic injury as a possible cause ofdeath in any of these Pleistocene cases.

Cranial and postcranial trauma are relatively common among Middle and Upper Pleisto-cene hominins and in most cases show signs of healing (S1 Table), indicating survival of the in-dividual [15–18]. Currently, there are only two examples in the fossil record that are tentativelyconsidered cases of lethal interpersonal violence. The Shanidar 3 Neandertal shows a penetrat-ing lesion to the left ninth rib consisting of a parallel-sided groove with exostoses along its mar-gins [19]. Nevertheless, some bone remodelling is apparent, suggesting this individual survivedfor several weeks after the lesion, and it is not clear that the final cause of death was related tothe rib injury. The Upper Paleolithic Homo sapiens individual Sunghir 1 shows a perimortemsharp trauma in the first thoracic vertebra that has been interpreted as the likely cause of death.While this would seem to represent a relatively clear case of lethal interpersonal violence, theauthors did not rule out the possibility of a hunting accident [20]. Here we report on the pres-ence of perimortem lethal cranial traumatic lesions in a Middle Pleistocene individual from theSima de los Huesos (SH) site, a singular case in the hominin fossil record.

The SH site has yielded an extraordinarily large sample of Middle Pleistocene (c. 430 kya)hominin fossils belonging to the Neandertal clade [21] and corresponding to a minimum of 28individuals [22]. During the time the hominin bones accumulated at the SH site, the only possi-ble access route to the site were through a deep (13 m) vertical chimney [23]. Given the skeletalpart representation in the collection [24], it is likely that entire bodies were deposited at thesite. There are no cutmarks on any of the 6700+ hominin bones recovered to date, and carni-vore manipulation (tooth marks) of the bones is rare [25]. The origin of the accumulation hasbeen highly debated, and four different hypotheses (carnivore activity, transport by geologicalagents, accidental falls, and intentional accumulation of bodies by hominins) have been pro-posed [24,26–29].

Recent taphonomic studies have ruled out carnivores and geological processes as accumula-tion agents [23,25,30]. The sediment of the hominin deposit (Lithostratigraphic Unit-6, LU-6)is indicative of a low-energy depositional environment (decantation) (Fig 1), and the small sizeof the SH site suggests only short-distance transport of the fossils within the site [21,23]. Fur-thermore, the red clay of LU-6 is devoid of extraclasts which provides strong evidence that thefossils were not subjected to long-distance transport, but likely accumulated in situ at the SHconduit [23]. The pattern of postcranial fractures in the assemblage indicates that the vast ma-jority occurred after burial, and were caused by the overlying sediment pressure [30]. Neverthe-less, a low proportion (around 4%) of postcranial perimortem fractures were found [30] andseveral cases of antemortem healed cranial injuries are also present in the SH sample [31] (S1Table).

Materials and MethodsCranium 17 (Cr-17) is a very complete specimen composed of 52 bone fragments preservingthe complete facial skeleton, including the partial upper dentition (the right C1 to M3 and theleft C1 to M2), the frontal bone, the sphenoid bone (missing only the body), the left parietalbone, the left temporal bone (lacking only the mastoid process) and most of the occipital bone.

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grants from the Fundación Atapuerca. R.M.Q. hasreceived financial support from BinghamtonUniversity (SUNY).

Competing Interests: The authors have declaredthat no competing interests exist.

The preserved right M3 is fully functional and shows only slight wear, indicating that Cr-17 be-longed to a young adult individual [21]. Junta de Castilla y León (Burgos, Spain) is the reposito-ry of the fossil (Cr-17). All necessary permits (excavation permit granted by the Junta deCastilla y León) were obtained for the described study, which complied with all relevantregulations.

Observations of the fractures were made with a Nikon SMZ800 Stereoscopic light micro-scope, as well as a DINO-LITE digital microscope. Detailed images of the fractures were madewith a Nikon Digital Sight DS-Fi1 camera.

Contour and trajectory analysis of the traumas3D imaging provides an opportunity to present critical aspects of cranial blunt force trauma[32], such as shape analysis [33], and to investigate plausible injury trajectories. Cr-17 was CTscanned in the coronal plane using an industrial XYLONMU 2000-CT scanner at the Univer-sidad de Burgos (Spain) with scanner energy of 180 kV and 4 mA. Slice thickness was collimat-ed to 0.5 mm, inter-slice spacing was 0.2 mm, and the approximate field of view was 225 mm.A total of 1108 slices was obtained as a 1024 x 1024 matrix of 32 bit Float format, with a finalpixel size of 0.219 x 0.219 mm. A virtual (3D CT) model of the cranium was generated fromthe resulting slices using the Mimics 16.0 (Materialise N.V.) software package.

Both, fracture angle and cortical delamination were measured on the virtual reconstructionsusing Mimics 16.0 software tools. Fracture angle is the angle formed by the fracture surface andthe bone cortical table, while cortical delamination or bevelling is the cleavage between the dip-loë and the inner/outer table.

Fig 1. Stratigraphy of the Sima de los Huesos site (modified from Arsuaga et al. [21]). The homininbones were recovered in Lithostratigraphic Unit 6 (LU-6) dated to c. 430ka [21]. This unit is composed of purered clays, filtering into the conduit system from overlying soils with little or no lateral transport, and very lowvelocity of sedimentation (decantation by dripping water) [23]. The figure also shows a detailed image of Cr-17 during its excavation at the site. Note the pure red clay that covers the cranial bones (partially cleaned insitu to enhance visualization) and the typical in situ postmortem (fossil diagenetic) fractures of the cranialvault. Photo credit: Javier Trueba (Madrid Scientific Films).

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The form of both of the fractures was analyzed in order to compare their contours. Relyingon the virtual reconstruction of the cranium, ten equidistant points were placed along the pre-served outlines of both traumatic fractures (T1 and T2). The first point was placed on thenotch in each fracture, and the 3-D coordinates of all the points were recorded. The coordi-nates were then transferred to the Morphologika 2.5 software program to perform the superpo-sition of the two outlines relying on the first landmark (the notch) as a reference point. Thetwo fractures outlines were rotated and superimposed but were not rescaled, to permit compar-ison of the true size of the lesions.

In forensic cases, the impact trajectories are estimated following the vectors of the entrance/exit wounds [34], especially for gunshot wounds. In the present case, the trajectory of the im-pact for each fracture was established by creating a normal vector to the plane of the fracturedefined by the points outlining its contour.

Results

SH Cranium-17 injuriesThe 52 fragments that comprise Cr-17 show clear postmortem fractures along their edges(Table 1, Figs 2 and 3) typical of a dry bone breakage pattern commonly found in fossils underdiagenetic conditions. No evidence of carnivore manipulation, such as tooth marks, is presenton this specimen [25]. On the left side of the frontal squama, this individual shows two sub-rectangular-shaped fractures, Trauma 1 (T1) and Trauma 2 (T2) (Fig 2).

The bone fragments that comprise the Cr-17 vault, as well as in the entire SH neurocranialsample, show a clear dominance of straight/curved fracture outlines, right angles between thecortical table and surface of the fracture, jagged surfaces and absence of cortical delaminationalong the fracture edges (Table 1). These are typical features of a dry bone breakage pattern,with fractures commonly considered as occurring postmortem in forensic cases (see Table 2)These criteria are applicable to the SH cranial collection, as well as the postcranial bones [30].The T1 and T2 fractures do not follow this dry bone breakage pattern.

Trauma 1 (T1) is located 16 mm lateral of bregma and 72 mm anterior to the coronal suture.Trauma 2 (T2) is located 30 mm lateral of bregma and 56 mm anterior to the coronal suture.Both T1 and T2 are connected and affect the inner and outer bone tables. On the ectocranialsurface (Fig 2B), the outline of T1 is sharp, with well-defined borders, and shows fracture linesradiating from the center of the trauma involving both the external and internal tables. Thefracture angle of T1 is oblique (32.5°-44.8°, Fig 4). A small, but distinct, notch is present alongthe superior border of the outline of T1 on the ectocranial surface (Fig 2). On the endocranialsurface there is a halo of cortical delamination (Fig 2) that varies from a minimum of about 4.9mm to a maximum of about 8.8 mm.

As in the case of T1, T2 exhibits an oblique fracture angle between the fracture edge and theexternal bone table (24°-49.2°, Fig 4), radiating fracture lines, a smooth fracture surface, corticaldelamination on the endocranial surface and a small, but distinct, notch along the superior bor-der of the fracture outline. Furthermore, just lateral to T2 on the ectocranial surface, there is alarge irregular cortical delamination (13.5 mm in maximum length) that exposes the diploë.The diploë is also exposed along the fracture edges in both T1 and T2. There are no signs ofbone remodeling (e.g. presence of woven bone) in any of these exposure areas (Figs 3 and 4).

Differential DiagnosisThe fracture pattern in both T1 and T2 is characterized by oblique angles, smooth surfaces,and the presence of cortical delamination (Table 1). This is the expected pattern for localizedperimortem blunt force trauma [35,39,44] and is consistent with penetration of the endocranial

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bone table. In contrast to postmortem fractures, perimortem fractures occur while the bone isstill fresh (i.e. surrounded by soft tissue and/or preserving the organic matrix), and the fractureproperties are well defined (Table 2). Furthermore, the morphology and characteristics of thetwo fractures in Cr-17 fit the criteria to be defined as depression fractures [1,45,46] that exhibitno evidence of healing (Fig 3). Depression fractures result from a concentration of energy suffi-cient to cause local failure to the bone and may be characterized as perforating and penetratingfractures with or without associated radial fractures [45,47].

The dimensions and contours of the two depression fractures were found to be almost indis-tinguishable (Fig 4) (including the presence of a similarly-placed notch in both fracture out-lines), strongly suggesting that both fractures were caused by the same object. Furthermore, thefractures show different orientations (T2 is rotated relative to T1, Figs 2 and 4) and differenttrajectories (Fig 4) implying that each fracture was caused by an independent impact.

Discussion and ConclusionsBecause soft tissue decomposition occurs sometime after the death of the individual, it is possi-ble the injuries in Cr-17 could have been produced either during the free-fall down the verticalshaft (the mode of entry of the hominin cadavers to the site) or inside the SH chamber after thebody arrived to the site. The few cases of perimortem fractures in the postcranial remainsmight be attributable to the corpse landing on a hard object (e.g. limestone block) at the bottomof the vertical shaft [30]. However, in the case of Cr-17, the same object likely produced thetwo fractures. Thus, any scenario related to the free-fall would require the highly improbableoccurrence of the same object striking the skull twice. The same criteria is valid to exclude lime-stone block-falls inside the SH chamber once the skull was deposited in the site.

Similarly, displacement of the skull over the sediments within the site is unlikely to accountfor the perimortem fractures in Cr-17. The sedimentological features of LU-6 (very low-energydepositional environment [21,23]), are incompatible with the high-energy processes necessaryto generate such modification in situ inside SH. Furthermore, it seems highly improbable thattaphonomic processes such as geological transport inside the SH chamber could have producedtwo episodes of identical blunt force trauma in the same individual, particularly given this sin-gular occurrence among the very large fossil sample recovered from the site.

If the taphonomic processes inside the site are discarded as the cause of the cranial trauma,other possible scenarios can be considered. The location and type of the injury are useful for

Table 1. Fracture properties of SH Cr-17.

Region Fracture outline Angle** Surface Corticaldelamination

N Straight Curved Depressed Mean SD Smooth Jagged Present Absent

Frontal* 6 5 1 0 95.6 7.6 0 6 0 6

Occipital 8 5 3 0 90.3 10.4 0 8 0 8

Left parietal 9 8 1 0 100.2 11.8 0 9 0 9

T1 1 0 0 1 38.7† - 1 0 1 0

T2 1 0 0 1 36.6† - 1 0 1 0

*T1 and T2 excluded.

**Angle between the cortical table and surface of the fracture (in degrees).† For T1 and T2, two angles were measured on opposite sides of each fracture (T1 = 32.52° and 44.81°; T2: 49.22° and 23.97°).

N = Number of fractures analyzed.

SD = Standard Deviation.

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distinguishing among the potential causes of cranial trauma (i.e. accidental vs violence-related)following forensic criteria [1]. Accidental or unintentional trauma typically affects the sides ofthe cranial vault, while intentional injuries are more commonly found in the facial region [48–50] (Table 3). Furthermore, falls are usually associated with generalized cranial trauma whichtends to produce large linear fractures, especially at the level of the “hat brim line” [44,48–50]

Fig 2. Cranium 17 bone traumatic fractures. (A) Frontal view of Cranium 17 showing the position of the traumatic events T1 (inferior) and T2 (superior); (B)Detailed ectocranial view of the traumatic fractures showing the two similar notches (black arrows) present along the superior border of the fracture outlines.Note that the orientation of the two traumatic events is different; (C) Detail of the notch in T1 under 2X magnification with a light microscope. (D) Endocranialview of T1 and T2 showing the large cortical delamination of the inner table (black arrows).

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(Table 3). Although cranial depression fractures can be a consequence of accidents, they aremore likely to be the result of interpersonal violence [2,48]. In the case of Cr-17 it is also possi-ble to rule out the injuries as either self-inflicted or resulting from an unintentional hunting ac-cident, mainly because the lesions involve multiple blows. Based on the absence of cut-marks,other potential post-mortem manipulations (e.g. cannibalism, ritual manipulations, etc.) seemeven less likely and more speculative.

Multiple cranial depression traumas in the frontal region above the hat brim line are com-patible with interpersonal violence injuries [1,2,48–50,53–55]. From their consistent size andshape, the Cr-17 blunt force traumas clearly are not unintentional, but, rather, they appear tohave been produced by the use of a tool of standardized size and shape. The location of the le-sions just to the left of the midline of the frontal squama in Cr-17 is also consistent with thegeneral pattern documented among recent humans, with most individuals showing lesions onthe left side of the skull reflecting the predominance of right-handedness during face-to-face

Fig 3. CT cross-sections comparing perimortem and postmortem fractures in the SH crania. (A) Perimortem fracture of the frontal bone in SH Cranium17; (B) Postmortem fracture in the occipital bone of SH Cranium 17 showing a right angle at the point of breakage. Scale bar = 2 cm.

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Table 2. Perimortem (fresh bone) vs postmortem (dry bone) fracture properties.

Feature Description Perimortem features Postmortemfeatures

Literature

Fracture texture Morphology of the broken bone surface Smooth* Jagged [32,35–40]

Fracture angle Angle between the cortical table and surface of thefracture

Oblique* (acute orobtuse)

Right [32,35–42]

Cortical delamination orbevelling

Cleavage between the diploë and the inner/outertable

Present * Absent [35,43]

Bone flakes Small bone fragments attached to the impact site Present* Absent [35,41]

* Present in Cr-17 traumatic fractures

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conflict [17,56]. Interestingly, the Sima de los Huesos population is considered mainly right-handed [57–59]. The severity of the injuries, with both blows to the head certainly involvingpenetration of the bone-brain barrier, and the absence of healing via bone remodeling (Fig 3)leads us to consider that this individual did not survive these cranial traumatic events. Indeed,either of the two traumatic events were likely mortal in and of themselves, and the presence ofrepeated blows might imply a clear intention to kill. Thus, the most plausible explanation forthe perimortem fractures on Cr-17 is as the result of intentional and repeated blows during alethal act of interpersonal violence. This represents the earliest clear case of deliberate, lethal

Fig 4. CT analysis of the Cranium 17 traumas. A) Sagittal cross-section of the T1 showing the acute fracture angles (32.5°-44.8°); B) Coronal cross-section of the T2 in detail showing the acute fracture angle (49.2°) and the large cortical delamination of the outer table (6.25mm); C) Virtual reconstruction ofthe SH Cranium 17 (left) showing the fractures on the left frontal squama and the landmarks used for the shape analysis (blue = T1 and red = T2). 3Dlandmarks of T1 and T2 in their original position (upper right), with arrows indicating the first landmark placed on the notch. Rotation and superposition of the3D landmarks of T1 and T2 using the first landmark of the two fractures as reference points (lower right). D) Reconstruction of the trajectory of the impactsrelying on the planar orientations of the external outlines of the fractures (blue = T1 and red = T2). Scale bars 2 cm.

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interpersonal aggression in the hominin fossil record, demonstrating that this is an ancienthuman behavior.

Finally, our results have important implications for the origin of the accumulation of homi-nin bodies at the SH site. As mentioned previously, geological events and carnivore activitywere discarded as the causal agents for the human fossil accumulation [23,25,30]. This leavesonly two possible explanations: i) accidental falls of 28 individuals through the vertical shaft, orii) intentional accumulation of bodies by other humans through the shaft [23].

The present study has established that the individual represented by Cr-17 was already deadbefore their arrival at the site, and it is possible to rule out an accidental fall as a possible expla-nation for the arrival of this individual to the SH chamber. The only possible manner by whicha deceased individual could have arrived at the SH site is if its cadaver were dropped down theshaft by other hominins. Thus, the interpretation of the SH site as a place where hominins de-posited deceased members of their social groups seems to be the most likely scenario to explainthe presence of human bodies at the site. This interpretation implies this was a social practiceamong this group of Middle Pleistocene hominins and may represent the earliest funerary be-havior in the human fossil record.

Supporting InformationS1 Table. Craniofacial and postcranial traumatic lesions in the PleistoceneHomo fossil re-cord(DOCX)

AcknowledgmentsThe authors wish to thank to the Atapuerca research and excavation team, especially those in-volved in the excavations at the Sima de los Huesos site. CT scanning was carried out in collab-oration with the Laboratorio de la Evolución Humana at the Universidad de Burgos (Spain).Thanks to Javier Trueba.

Table 3. Diagnosis of the trauma according to the mechanism of injury.

Factor Source

Severity of theforce

Less force Linear fractures [41,44,47]

More force Fracture lines radiating from the impact point* [41,44,47]

Objectcharacteristics

Blunt Radiating fractures are commonly produced with blunt force trauma*; Concentric fractures maypresent on internal bevel*; Multiple blunt trauma may be indicated by the presence of flakes ofbone spalled from fracture margins and the delamination of outer from inner tables*

[39,41,43,44]

Sharp Presence of incised wounds; Linear fractures often occur on the skull and result from out-bending of large thin portions of bone as the result of a direct blow of high velocity; Bonefracture lines leading to the sutures.

[35,51,52]

Cause of thefracture

Accidentalfalls

Linear and radial fractures; Lateral location; Fractures on Hat Brim Line†; Right side fractures [1,45,48–50,53,54]

Aggression Depressed fractures*; Frontal location*; Fractures above Hat Brim Line *†; Left side fractures*;Multiple blows*

[1,45,48–50,53,54]

*Present in Cr-17 traumatic fractures

†Hat brim line (HBL) corresponds to the area located between two lines parallel to a line inspired by the Frankfurt horizontal plane (horizontal plane

passing through right and left porion and the left orbitale), with the superior margin passing through the glabella (G line) and the inferior margin passing

through the center of the external auditory meatus (EAM line) [49].

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Author ContributionsConceived and designed the experiments: NS JLA APP AP IM. Performed the experiments: NSJLA APP AP IM. Analyzed the data: NS JLA APP AP IM RQ AGO. Contributed reagents/ma-terials/analysis tools: NS JLA APP AP IM RQ AGO. Wrote the paper: NS JLA APP AP IM RQAGO. Direction of the excavation and research project: JLA JMBC EC.

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1

Supporting Information

S1 Table. Craniofacial and postcranial traumatic lesions in the Pleistocene Homo fossil record (Updated from Wu et al. [1]).

Specimen Traumatic lesion Source

Early Pleistocene Gongwangling/ Lantian 1* Frontal healed lesion (Trauma or abscess/Postmortem erosion) [2,3] Sangiran 38 Two parietal depressions possibly post-traumatic [4] Middle Pleistocene Atapuerca-Trinchera Galería Parietal healed trauma [5,6]

Biache 1 Shallow exocranial depression [7]

Biache 2 Frontal minor lesion [7]

Broken Hill 1 Temporal trauma with associated infection [8] Casal de’ Pazzi 1 Depression on the parietal [9]

Ceprano 1 Frontal lesions [10] Ehringsdorf 2 Parietal lesion [11]

Florisbad 1

Multiple cranial vault and orbital lesions including infection post-trauma

[12]

Hulu/Nanjing 1 Healed ectocranial lesion (trauma or burning) [13] La Chaise BD-17 Parietal shallow depression [14]

Maba 1

Frontal healed trauma (probably interpersonal violence or accident)

[1]

Ngandong 7 Parietal depressions [15] Atapuerca-SH-Cranium 1 Occipital depression [16] Atapuerca-SH-Cranium 2 Parietal vault depression [16] Atapuerca-SH-Cranium 3 Parietal depression [16] Atapuerca-SH-Cranium 4 Frontal and occipital depressions. Parietal depressed groove [16]

Atapuerca-SH-Cranium 5

12 exocranial lesions, slight supratoral depression and postraumatic maxillary osteitis (traumatic origin)

[16,17]

Atapuerca-SH-Cranium 6 Slight exocranial depression [16] Atapuerca-SH-Cranium 7 Lambdoid suture healed external lesion [16] Atapuerca-SH-Cranium 8 Slight parietal exocranial depression [16] Atapuerca-SH-Cranium 11 Frontal healed depression [16,18] Swanscombe 1 Parietal depressions [19] Zhoukoudian skull X Frontal and parietal exocranial lesion [20] Zhoukoudian skull XII Parietal exocranial lesion [20] Zuttiyeh 1 Two depressed frontal lesions [21] Late Pleistocene Caviglione 1 Healed radial fracture [1] Cova Negra (Parietal I) Trauma with external remodeling [22,23] Dolní Věstonice 3 Healed traumatic impact on the face [24]

Dolní Věstonice 11/12

Healed depressed fracture of frontal (Possible interhuman violence)

[24]

Dolní Věstonice 13 Minor healed frontal trauma [24] Dolní Věstonice 15 Healed parietal lesion and ulnar fracture [24] Dolní Věstonice 16 Two external healed frontal lesions [24] Feldhofer 1 Healed supraorbital arch lesion. Fractured ulna. [25,26] Feldhofer 2 (NN59) Healed ulnar fracture [27] Kebara KMH2 Healed fractured thoracic vertebra and metacarpal [28] Kiik-Koba 1 Healed foot phalanx fracture [29] KNM-LH 1 Healed frontal trauma [30] Krapina 4 Healed scalp wound/depression fracture to frontal [31,32] Krapina 20 Healed lesion in the frontal [31,32] Krapina 31 Possible healed fracture near temporal line [31]

2

Krapina 34.7 Healed parietal injury [31,32] Krapina 149 Healed clavicular fracture [31,32] Krapina 180 Healed ulnar fracture and pseudoarthrosis [31,32] Krapina 188.8 Ulnar diaphyseal fracture with osseous callous [31,32]

La Chapelle-aux-Saints 1

Healed fractured rib; generalized osteoarthritis (post-traumatic event)

[33]

La Ferrassie 1 Healed displaced greater trochanter [33,34] La Ferrassie 2 Fibular fracture with infection [34] Le Moustier 1 Healed mandibular fracture [35] Mladeč 5 Frontal traumatic lesion [36] Pavlov 1 Minor healed exocranial alteration [24] Qafzeh 11 Healed depressed frontal fracture [37,38] Riparo Mezzena Mandible osteolytic lesion (infection by traumatic fracture) [39] Saint Césaire 1 Healed frontal trauma (interpersonal violence) [40] Šal’a 1 Healed frontal depression [41]

Shanidar 1

Facial fracture. Multiple arm fractures with possible amputation. Foot osteoarthritis and metatarsal fracture (accidental or crushing)

[42,43]

Shanidar 3

Rib penetrating wound (possible interpersonal violence). Post-traumatic talocrural osteoarthritis

[43,44]

Shanidar 4 Healed rib fracture [43] Shanidar 5 Healed frontal bony scar [43] Skhul 4 Healed metatarsal fracture [45] Subalyuk 1 Healed metatarsal fracture [46]

Sunghir 1

Perimortem incision to vertebra T1 (hunting accident or interpersonal violence)

[47,48]

Sunghir 2 Possible pelvis perimortem trauma [48] Sunghir 5 Healed frontal minor lesion [48] Tabun C1 Diaphyseal lesion on the fibula [42] Xujiayao 5a Healed parietal injury [49] Xujiayao 8 Healed parietal injury [49] Xujiayao 12 Healed occipital injury [49]

Five individuals of the Pleistocene hominin record (Maba 1, Dolní Věstonice 11/12, Saint Césaire 1, Shanidar 3 and Sunghir 1) have been interpreted as probable cases of interpersonal violence. The only traumas that have been interpreted as likely representing the cause of death of the individual are those of Sunghir 1 and Shanidar 3. Atapuerca SH-Cranium 5 suffered a chronic infection, probably as a result of an impact, that may have been the cause of death of this individual [17]. Nevertheless, an accidental cause cannot be discarded in any of these cases. *Caspari [2] identified a lesion in Gongwangling/Lantian 1 due to a trauma or abscess. However, Shang et al. [3] showed that the damage was due to postmortem alteration.

3

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