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Trauma in the Krapina Neandertals

Chapter · January 2014

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David W Frayer Virginia Estabrook University of Kansas Armstrong State University

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1 2 3 4 THE ROUTLEDGE 5 6 7 HANDBOOK OF THE 8 9 BIOARCHAEOLOGY OF 10 11 HUMAN CONFLICT 12 13 14 15 16 17 18 19 20 21 Edited by 22 23 Christopher Knüsel and Martin J. Smith 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 Template: Royal A, Font: , Date: 26/07/2013; 3B2 version: 9.1.460/W Unicode (Apr 1 2008) (APS_OT) Dir: E:/TAFUK/3B2/KNUSEL-SMITH -131047/130001/APPFile/BK-TAF-KNUSEL-SMITH-131047-130001.3d

1 2 3 4 4 5 6 7 Trauma in the Krapina Neandertals 8 9 Violence in the Middle Palaeolithic? 10 11 12 V. Hutton Estabrook and David W. Frayer 13 14 15 16 17 18 Since their discovery, the Krapina Neandertal bones have served as models for Neandertal 19 morphology and behaviour (Radovcˇic´ 1988). The Krapina rock shelter, dated ~130 kyrs ago 20 (Rink et al. 1995), was excavated under the direction of Gorjanovic´-Kramberger from 1899 to 21 1905, who published extensively on the discoveries at the site (Gorjanovic´-Kramberger 1906; 22 Frayer 2007). In addition to extensive discussion of the anatomy of the remains, he described 23 evidence for trauma and speculated on its causes (Gorjanovic´-Kramberger 1908, 1913; Radovcˇic´ 24 1988). At the site, five cranial fragments preserve blunt force trauma lesions with two showing 25 extensive healing. In addition to the cranial trauma, one clavicle has a long-healed fracture, an 26 ulna shaft has a large fracture callus, and one ulna terminates in what appears to be an amputation 27 stump. Most modern interpretations have focused on occupational hazards, lifestyle and/or 28 accidents as likely explanations for many cases of Neandertal trauma (Berger and Trinkaus 1995; 29 Underdown 2006; Mann and Monge 2006). Older arguments (Roper 1969; Keith 1928; Dart 30 1953) claimed head wounds and other bone traumatic injuries were signatures of homicides, but 31 all are extensively healed and the injuries represent traumatic episodes that occurred long before 32 death. 33 One way to address the presence and possible causes of trauma is through the direct obser- 34 vation of the lesions, and here we include detailed descriptions and images of the trauma 35 observed at Krapina. Another approach is to compare the patterns of skeletal trauma to other 36 more recent populations in which interpersonal violence is recognized as part of a hazardous 37 way of life. If interpersonal violence was not a source of trauma in the Krapina Neandertals, 38 we should expect to see statistically different frequencies of incidents of trauma distributed 39 throughout the cranial and post-cranial bones compared to more recent skeletal samples. We 40 use this approach to compare the Krapina skeletal remains to those of other more recent 41 groups, mostly hunter-gatherers and foragers, to test the claim that trauma observed in 42 Neandertal skeletal remains is less often of violent origin than it is in other hunter-gatherer 43 groups. 44 45 The Krapina Neandertal sample 46 47 The Neandertal remains from Krapina represent individuals from one geographical location and 48 a time span of no more than 20 kyrs (Rink et al. 1995; Wolpoff 1989). They occupied more

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1 uniform, generally warmer environmental conditions in OIS 5e than a more general sample of 2 Neandertals from many sites spread over Europe who often survived harsher, glacial times. The 3 sample from Krapina contains more than 800 crania and post-crania (Radovcˇic´ 1988) repre- 4 senting the largest Neandertal sample in Europe. Its minimum number of individuals (MNI) 5 estimates range from 14 (Malez 1971) to 75–82 (Wolpoff 1979). The sample includes multiple 6 elements from almost every bone in the body. Most importantly, it is excellently curated and 7 catalogued (Radovcˇic´ et al. 1988; Radovcˇic´ and Wolpoff forthcoming; Kricun et al. 1999) so 8 that the total number of skeletal elements represented has been published. Recent work has 9 demonstrated that the demographic sample found at Krapina is consistent with other those of 10 other Palaeolithic populations (Wolpoff and Caspari 2006), and its patterns of trauma are 11 consistent with those observed in Neandertals in general (Estabrook 2009). 12 Over the past 100-plus years, several hypotheses have been offered to explain the distribu- 13 tion of human remains at Krapina. These explanations include: Krapina as regularly occupied 14 living site (Gorjanovic´-Kramberger 1899, 1906, 1913; Malez 1971, 1978); Krapina as the site 15 of cannibalism and/or massacre (Gorjanovic´-Kramberger 1901, 1906, 1909; Škerlj 1939, 1958; 16 Tomic´-Karovic´ 1970); Krapina as a death trap for a group of runaway juveniles (Bocquet- 17 Appell and Arsuaga 1999); Krapina as a site of purposeful burials (Trinkaus 1985); and Krapina 18 as a secondary burial site following mortuary processing (Russell 1987a, 1987b; Ullrich 2006). 19 Studies of the anatomical distributions of the fragments at Krapina, such as those of Trinkaus 20 (1985) and Van Arsdale (2007), generate radically different conclusions about the random 21 nature of the preservation of various elements at Krapina. The original site no longer exists, 22 field notes are limited and cave bear denning likely mixed some of the bones, so it is extremely 23 difficult to explain the accumulation of bones at the site. However, Ullrich (2006: 506) sum- 24 marized Gorjanovic´-Kramberger’s initial interpretation: “He also pointed out that skull frag- 25 ments have never been discovered in connection with postcranial remains of the same 26 individual and postcranial remains were never found in anatomical connections or anatomical 27 positions.” This is in contrast to the faunal remains at the site, which preserve occasional cases 28 of articulated limbs, carpals, tarsals and vertebrae (Miracle 2007). The high level of human 29 bone disarticulation and fragmentation seems to refute the argument that the human bones 30 originated as purposeful, primary burials. Moreover, not all the bones show evidence of can- 31 nibalism or defleshing, so the taphonomic conditions that led to the deposition of the human 32 bones remain unexplained. 33 The fragmentary preservation of the skeletal remains at Krapina precludes a good under- 34 standing of the demographic profile of the site. What is clear about the Krapina sample is that 35 there are many elements missing per individual. This is in some part due the public use of the 36 site as a sand-pit before the fossils were identified (Radovcˇic´ 1988), so some missing elements 37 may have been carried away long after the site was abandoned. But many bones show ancient 38 fragmentation and cutmarks, so some disarticulation and fragmentation occurred before the 39 bones were put into the ground (Russell 1987a, 1987b; Ullrich 2006). The combination of 40 these two factors probably means the current sample of remains from Krapina represents a few 41 pieces from many individuals rather than many pieces from a few individuals as is common in 42 most well-known Neandertals sites. If so, formal estimations of MNI would likely be low 43 because they do not come remotely close to actual number of individuals represented at 44 the site. 45 Becauseitisdifficult to get a sense of how many individuals are preserved at Krapina and 46 consequently what the age-at-death and sex profiles look like for the site, it is impossible to 47 directly assess whether or not Krapina is a representative sample of the population of 48 Neandertals living at this site. Wolpoff and Caspari (2006) use Libben (Ohio, USA)

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1 demography to estimate a 69% survivorship for the “missing” (i.e. not well represented in the 2 Krapina skeletal sample) infants and children under the age of five years under the assump- 3 tion that both populations were stationary with a stable age distribution. They then added 4 these 37 estimated juveniles to the survivorship distributions to make it comparable to the 5 samples from Libben, Atapuerca (Spain), Australopithecus specimens and the Aché (Paraguay). 6 Wolpoff and Caspari found that the sample from Krapina, Atapuerca and the aus- 7 tralopithecines had very similar survivorship distributions, which differed from Libben and 8 the Aché distributions. They concluded that the age-at-death distributions at Krapina and 9 Atapuerca were consistent with the findings of Caspari and Lee (2004, 2006) in that the 10 increased number of old individuals in the population comes later in time among Homo 11 sapiens sapiens. They also argued that the Krapina skeletal element frequencies do not reflect 12 catastrophic profiles. We know that some of the individuals in the sample are related, based 13 on shared morphological features such as the deviated internasal suture (Smith 1976), so 14 some specimens must have lived close in time. Here, we treat the Krapina cranial and post- 15 cranial sample as a population, but are aware that the bones are sampled over several thousands 16 of years. 17 18 More recent comparative samples 19 20 Comparative samples come from prehistoric North America and Italy, and from protohistoric 21 graves in Australia. These were chosen because they represent non-urban, hunter-gatherer, 22 forager or nomadic populations. They likely resemble (but do not duplicate) lifestyle stresses of 23 the Krapina Neandertals. None are drawn from samples that likely lived or were buried in caves. 24 For the cranial analysis, we added a Portuguese medieval sample from Aljubarrota. 25 The Libben site represents a late Woodland (AD 500–1000) Native American sedentary, 26 foraging population from northern Ohio from which only complete long bones were tabu- 27 lated (Lovejoy and Heiple 1981). SC1–038 is drawn from a hunter-gatherer population from 28 the Santa Clara Valley in central California, dating from between 240 BC to AD 1770, from 29 which only complete (defined in this case as at least two-thirds complete with “all major 30 articular areas preserved”) long bones were considered (Jurmain 2001). The sample from Ala- 31 329 is from the southeastern shore of the San Francisco Bay (California) dating from between 32 500 BC through the 1700s (Jurmain 1991; Jurmain and Bellifemine 1997). Pozzilli, from the 33 mountainous zone of south-central Italy, dates from 600–300 BC and is associated with the 34 “shepherd-warriors” of the Samnite culture (Brasili et al. 2004) 35 The five Australian samples derive from Webb (1989, 1995). All skeletons have been repa- 36 triated and were composed primarily of recent material. Two samples are from southeast 37 Australia and the Central Murray graves come from scattered cemeteries along the banks and 38 floodplains of the Murray River (Webb 1995: 13–14). The Rufus River collection samples 39 skeletal material from the area around Lake Victoria, Lindsay Creek, Chowilla and the Rufus 40 River (Webb 1995: 17). Other skeletal samples were based on ecological zones (Webb 1995), 41 and divided into Desert (arid central and western Australia), South Coast (coastal Victoria) and 42 East Coast (coastal New South Wales and Queensland). 43 We also used a recent European collection, which included fragmentary remains from a 44 common grave. This ossuary was composed of Portuguese and Castilian soldiers lost in the 45 massacre of the Aljubarrota battle in 1385 (Cunha et al. 2001; Estabrook 2009). According to 46 legend, the remains were left on the battlefield for seven years, before they were collected and 47 deposited in the common grave (Cunha et al. 2001; pers. comm.). Consequently, the bones 48 are fragmentary and mostly disarticulated, like the Krapina sample. Traumatic lesions recorded

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1 are only those received prior to death and represent injuries received during the course of 2 their lives. 3 4 Trauma at Krapina 5 6 There are 68 major cranial pieces and vault fragments from the frontal and parietal known from 7 the site (Radovcˇic´ and Wolpoff forthcoming). Of these, four show evidence of ante-mortem 8 blunt force trauma with depressed fractures (Gardner and Smith 2006) and another has partially 9 healed massive head trauma (Mann and Monge 2006). A total of 279 upper limb and 177 lower 10 limb fragments occur at the site and three from the upper limb and torso show trauma related 11 pathology. No ribs, vertebrae or lower limb bones show evidence of trauma, other than signs of 12 degenerative joint disease on a few specimens (Gardner and Smith 2006). We cannot conclude 13 these are trauma-related. Thus, evidence for trauma at Krapina is found only in bones above the 14 waist. 15 16 Healed cranial wounds 17 18 Krapina 4 (Figure 4.1) 19 Krapina 4 received a blunt force injury to the left frontal, just medial of the temporal line and 20 approximately 36mm distant from the mid-sagittal plane. On the ectocranial surface the injured 21 site is preserved as an oval depression (15mm æ 9mm) about ~2.5mm deep at the most medial 22 margin. Irregular, pitted and remodelled granular bone partly fills the oval depression, typical of 23 healing of the initial trauma (Courville 1962). Erosions and irregularities, particularly evident 24 laterally and inferiorly, are indicated by a depressed region measuring 11mm in the paracoronal 25 direction and 16mm transversely. This region has smoothed, sclerotic diplöe and its location 26 adjacent to the initial trauma site suggests it is a fistulous track from a purulent infection 27 (Figure 4.1). The lesion ends just superior to the temporal line where it reaches the original 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 Figure 4.1 (a) External surface of Krapina 4 showing head wound with healed reactive track inferior to it. 48 (b) Internal view of Krapina 4, showing stellate fracture typical of a heavy blunt force trauma.

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1 ectocranial surface. The head trauma was infected and a diﬀuse periostitis extends onto the 2 parietal and across the midline to the right frontal. Endocranially, the surface shows a stellate 3 fracture of the internal table with four thin fracture lines radiating from a single point. These 4 corresponding to the deepest part of the ectocranial wound are typical of massive blunt force 5 trauma (Courville 1962; Tung 2007). No endocranial bony reaction is present, so the heavy 6 blow never extended to the endosteal layer of the dura mater. The Krapina 4 lesion, particularly 7 the oval depression, closely resembles healed blunt force injuries found in other cranial fragments 8 from Krapina and in the prehistoric (e.g. Frayer et al. 2006; Pérez et al. 2004; Weidenreich 9 1943) and historic skeletal record (e.g. Judd 2004; Tung 2007; Walker 1989). Compared to the 10 other blunt force traumatic injuries at Krapina, this is the least healed and the only one associated 11 with what appears to be an infected head wound. 12 13 Krapina 5 (Figure 4.2) 14 15 Formerly known as the “D” skull, Krapina 5 preserves two areas of blunt force trauma. Above 16 the lambdoidal suture at the edge of the left parietal is a slightly recessed area that shows thinning 17 on its lateral, inferior-most margin. The depression is only a portion of the original injury 18 measuring ~16.5mm (anterior-posterior) and ~9.5mm (medio-laterally) and about 0.4mm deep. 19 Vault thickness at the deepest point on the fragment is 5mm compared to the parietal anterior to 20 the depression of about 8mm. All measurements are approximations because the depression is 21 clearly present, but its border is indistinct. Unlike the other depressions, there is no evidence of 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 Figure 4.2 Arrows indicate the old, healed blunt force traumatic injuries on Krapina 5.

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1 healing at the depth of the blunt force trauma, but the centre of the injury is likely more on the 2 occiput. This is not the most impressive of head wounds. In addition, there are two small 3 depressed areas on the right parietal, 35mm and 29mm posterior to the coronal suture. Both are 4 associated with some dark staining, but the specimen is heavily lacquered and, at present, the 5 cause is unknown. The depression, only slightly pressed into the ectocranial face, is oblong and 6 angled about 45 degrees from the sagittal suture with the long axis running antero-posteriorly. It 7 measures 7mm æ 5mm and is very shallow. The smaller, more medially placed depression is 8 round, ~2mm in diameter. We cannot be certain these depressions are healed injuries and have 9 not included them in the analysis. In fact, none of the depressions on Krapina 5 are deep, and 10 none show signs of infection and none are associated with any cutmarks, although the heavy 11 coat of lacquer may hide them. 12 There is a very marked periostitis on all the fragments making up Krapina 5, only absent on 13 the nuchal plane. For Krapina 5 the most distinctive feature is the diffuse periostitis over the 14 entire vault, probably associated with a low grade scalp inflammation, similar to infectious 15 reactions following hair plugs (Jones et al. 1980) or some cases of scalping survivals (Aufder- 16 heide and Rodríguez-Martin 1998; Ortner and Putschar, 1981). 17 18 Krapina 20 (Figure 4.3) 19 20 Krapina 20 is a left frontal and parietal fragment, preserving 59mm of a patent metopic suture 21 and approximately 79mm of the sagittal suture. There are two indications of blunt force trauma, 22 both appearing only on the frontal. One on the squama is ~25.5mm anterior to the coronal 23 suture and 16mm lateral to the metopic. It is oval and angled about 60 degrees to the para- 24 coronal plane. The maximum length, breadth and depth, respectively, are 10mm, 6.5mm and 25 1mm. The deepest parts of the fossa show some bone honeycombing and a ridge separates a 26 medial from a lateral compartment, giving the internal aspect a “double depressed” appearance. 27 Both are about the same depth, but the lateral is larger and more rounded and the medial more 28 triangular. The lateral chamber has more extensive pitting and bone remodelling, and there 29 appears to be a vesicular channel connecting the lesion’s base to the diploe. The compart- 30 mentalizing of the depression appears to be a function of the remodelling and not related to the 31 object responsible for the wound. The steep edges of the lesion and the rim are smooth and 32 polished with no signs of infection, so the lesion must have occurred well before the individual’s 33 death. There is a crack originating at deepest point of the lateral fossa, running for about 10mm 34 posteriorly and paralleling the sagittal suture. There are other ectocranial micro-cracks, but these 35 widen away from the blunt force trauma and are unrelated to it. Internally, there are no signs of 36 endocranial involvement or radiating fractures. 37 The second blunt force trauma is only partially preserved in that it continued across the 38 metopic suture on the right frontal. It is centred on the midline about 20.5mm anterior to the 39 intersection with the parietals. The ectocranial surface is crushed into the metopic suture so 40 that on either side of the suture the external table measures about 20mm and, at the depth of 41 the fossa, about 5mm. The blunt force trauma does not extend though the diploe. Compared 42 to the other blunt force trauma, it seems more rounded with the long axis running perpendi- 43 cular to the coronal direction. Its lateral length away from the metopic suture is ~7mm and its 44 breadth along the metopic suture is ~8.5mm. The deepest point preserved is on the suture and 45 measures ~0.8mm. There is some minor honeycombing of the floor of the lesion, especially in 46 the most anterior part. No ecto- or endocranial cracks are associated with the wound. 47 There is slight, diffuse periostitis on the frontal and less on the parietal, but the latter is 48 heavily lacquered. Thin cutmarks can be seen on the frontal, but all are post-mortem, and the

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 Figure 4.3 (a) The larger of the two blunt force traumatic injuries on the Krapina 20 left frontal. (b) A 42 rounded blunt force traumatic injury on the metopic suture. 43 44 45 ectocranial surface is not weathered or fragmented, as is the case for K4. The preserved diploë 46 is crushed and above it the ectocranial surface is completely healed, without involvement of 47 the inner table and any signs of infection. Like Krapina 5, there is a diﬀuse periostitis, pre- 48 sumably the result a subcutaneous infection.

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1 Both wounds have about the same degree of healing and apparently have the same shape, so 2 they may have occurred in the same incident. On the other hand, they are just as likely to 3 have been separate wounds. 4 5 6 Krapina 31 (Figure 4.4) 7 An adult frontal fragment shows a depression and small area of reactive bone on the left side, just 8 superior to the temporal line. The bone is slightly thinner than that of Krapina 5 and has a 9 pronounced, ridged temporal line. Just medial to the temporal line is an irregularly shaped 10 depression (it is neither oval or round) with a healed bone matrix overlying the diploe. The 11 well-healed injury shows no sign of infection and occurred long before death. Dimensions 12 are ~11mm from the temporal line and at least 12mm in the antero-posterior direction. The 13 exposed honeycomb-like bone is a triangular wedge, with the widest part (5mm) situated 14 anteriorly and is about 10mm long. Greatest depth of the lesion is about 0.8 mm. Along the 15 medial edge of the healed lesion is an interrupted groove, which resembles the frontal venous 16 markings. These seem not to be part of the lesion, but possibly are accentuated in their 17 expression by the injury. The pitting looks like completely healed bone with no further 18 involvement of the surrounding bone. Except for the honeycombed aspect of the lesion, there is 19 little resemblance to the Krapina 4 injury. There is some ectocranial pitting representing a mild 20 periostitis, mostly medial to the healed wound. There are also many ﬁne, thin scratches above 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 Figure 4.4 The Krapina 31 frontal squama comprising most of the bone left of the midline.

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1 the wound, overlapping each other and running lengthwise in a paracoronal direction. They are 2 all lateral to the midline, basically running from above the lesion to just lateral from the point 3 where the “10” is marked on the frontal. The lines are under the lacquer, presumably ancient 4 and not related to the wound. 5 6 7 Krapina 34.7 (Figure 4.5) 8 This right parietal fragment has been described in detail by Mann and Monge (2006) and consists 9 of a healed wound on the squama above the mastoid, near the lambdoidal suture. Mann and 10 Monge document a round, “quarter to half circle” blunt-force trauma about 50mm in length. 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 Figure 4.5 (a) Lateral view of the large healed injury on the right temporal of Krapina 34.7. (b) Superior 48 view of Krapina 34.7, showing extensive resorption of the superior border.

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1 As they observe, the wound must have extended onto the occipital and temporal and into the 2 dura mater, making this the largest, most extensive head trauma found in any Neandertal. The 3 fracture edge shows bone healing and area just inferior to it “is considerably thinner, with a 4 progressive thinning leading to the margin of the fragment” (2006: 263). There is minor pitting 5 of the external face, indicating some inflammation of the tissue surrounding the wound. 6 7 8 Post-cranial trauma at Krapina 9 Krapina 149 (Figure 4.6) 10 11 Of the 14 fragmentary clavicles at Krapina, this is the only one showing a traumatic injury. The 12 lateral angle of the acromial end, at the conoid tubercle, shows a high angle, suggested a well- ff 13 healed fracture a ecting the right shoulder in this individual. The lateral acromial facet is 14 missing, but the conoid tubercle is large and other muscle markings are patent. Internal bone 15 structure shows thick cortical walls, suggesting the injury occurred long before death and that ff 16 the individual su ered no long-term disability. 17 18 Krapina 180 (Figure 4.7) 19 20 Krapina 180 is a fragmentary right ulna shaft, preserving the distal aspect of semi-lunar notch and 21 the shaft below it for 133mm (Figure 4.7a). The lip of the coronoid process shows no sign of 22 trauma, but about midshaft the diaphysis ends in an uninfected, healed callus, representing either “ ” 23 an amputation stump or pseudoarthrosis. There are no signs of infection on the stump , which 24 has a remodelled surface with a smoothed appearance and no articular facets (Figure 4.7b). We 25 suspect this is an amputation, which occurred well before the death of the individual based on the complete healing of the distal end and lack of evidence for infection. In place of its typical 26 fi 27 angular shape, the ulnar surface is rounded lacking a well-de ned interosseus crest and muscle 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 Figure 4.6 (a) Krapina 149 radiograph. (b) Krapina 149 inferior view of the right clavicle showing the 48 distorted lateral angle.

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 Figure 4.7 (a) The proximal right ulna of Krapina 180, medial aspect. (b) Close-up of distal callus, which 47 we consider to be a likely terminus of the amputated ulna. (c) Superior image is CT scan 48 showing medullary cavity and thick cortical bone of the entire shaft.

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1 markings for brachialis, the flexors and extensors. Nevertheless, the cortical bone is thick 2 (Figure 4.7c) and shows no evidence of bone resorption or atrophy. It is clear that the individual 3 continued to use her/his forearm long after the severe trauma had occurred, maintaining the 4 integrity of the cortical bone structure. 5 6 7 Krapina 188.8 (Figure 4.8) 8 This specimen comes from an adult left proximal ulna, running from the distal coronoid process 9 to the approximate midshaft position. The trauma is in a similar position to that of Krapina 180, 10 although the bone is not completely severed and clearer definition of muscle markings is present. 11 There is no reason to suspect this is Krapina 180’s antimere. Externally, the bone shows a healing 12 fracture callus, with some remnants of bowing at the injury. The internal cortical bone is thick 13 above the fracture callus, and there are no signs of infection associated with the break. There is 14 no reason suspect the upper limb was dysfunctional following the injury, although two segments 15 were not properly aligned before they set. 16 17 Limitations of the Krapina sample 18 19 For trauma analysis, the Krapina sample has two major weaknesses. The first is the fragmentary 20 nature of the sample. For example, it is impossible to know the full extent of cranial trauma for 21 any neurocranium since no complete vaults exist. Facial bones are also rare, so the extent of face 22 wounds is indeterminable. The other drawback is the lack of association of any post-cranial 23 remains with any cranial fragments. There are no articulated limbs and no possibility to link most 24 bones with reliable estimates for sex or age-at-death. It is possible to determine sub-adult versus 25 adult in the sample, and we found no evidence for trauma in sub-adult remains, although there 26 is no way to determine if the trauma occurred in a sub-adult and the remnant of the injury 27 persisted into adulthood. The sub-adult and younger bones listed by Radovcˇic´ and Wolpoff 28 (forthcoming) show no signs of trauma. 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 Figure 4.8 Medial view of the left ulna (Krapina 188.8) showing fracture callus distal to the broken cor- 48 onoid process.

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1 Most palaeopathological standards for collecting information about injury in a sample 2 population require complete individual bones, and almost all studies of more recent human 3 collections are based on complete individual bones. The fragmentary nature of the Krapina 4 sample makes not only data collection under standard protocols (such as those of Lovejoy and 5 Heiple 1981) impossible, but it also makes comparisons of trauma from the Krapina sample 6 with trauma from other populations incomparable under standard protocols. 7 Because the Krapina remains do not represent discrete individuals, it is not possible to 8 address questions about the demographic patterning of trauma. Through seriation, estimates of 9 the sexual dimorphism of a group of isolated individual post-cranial elements may be made, 10 but in fragmentary remains this is difficult and fraught with potential inaccuracies. Although 11 age-at-death profiles have been examined for the Krapina collection as a whole, these profiles 12 come from dental remains that are not directly associated with any of the bones. The patterns 13 of trauma relating to the distribution by sex, age-at-death or by individual cannot be assessed 14 using the Krapina sample. However, it is possible to test hypotheses about the Krapina sample 15 through element-by-element comparative analysis. 16 17 Analysis of Contingency Tables Using Simulations (ACTUS) 18 19 The program ACTUS1 (Analysis of Contingency Tables Using Simulation; Estabrook and 20 Estabrook 1989) was designed to address the problem of statistical analyses of two-way tables 21 from small samples in determining whether to reject a null hypothesis of independence. Such 22 analyses are especially important in fields such as history and other social sciences (the context for 23 which this program was created) or in any instance where counts of differing character states are 24 being compared. It has been applied in ornithology (Marques 2003, 2004), entomology (Raguso 25 and Willis 2002) and ichthyology (Galhardo et al. 2008; Amorim et al. 2004; Almada et al. 1994; 26 Oliveira and Almada 1996). Because the approximation to classical statistical distributions (such 27 as chi-square) is poor when only a few cases are involved, the minimum number of cases 28 expected under the hypothesis of independence must exceed four for tabulated probabilities to 29 be accurate using classical statistical methods (Estabrook and Estabrook 1989: 5). With a small 30 sample, when the null hypothesis of independence is rejected for a contingency table using 31 classical statistics or other methods such as Fisher’s Exact test, it is difficult to tell which cells to 32 interpret as being the ones more and/or less frequent than predicted under the null hypothesis, 33 because the co-occurrences that merit a more substantive interpretation are not necessarily the 34 largest or smallest counts (Estabrook 2002: 23). 35 ACTUS calculates estimates of realized significance of each cell from small datasets. It does 36 this by comparing two classifications under the null hypothesis that they are independent. 37 Counts of both classifications are arrayed in a contingency table. The program then uses a 38 random number generator to simulate thousands of comparable datasets whose counts are 39 known to be samples of the null hypothesis, then it counts the number of simulated tables 40 with entries that are larger or smaller than the corresponding entries input by the user. The 41 results of an ACTUS analysis show not only whether the entire contingency table rejects the 42 null hypothesis, but also which cells are larger or smaller than predicted. ACTUS uses direct 43 comparison of the value of a statistic calculated from the observed table with a value calculated 44 in the same way from a simulated table to estimate its realized significance with the fraction of 45 simulated tables not less or not greater. This method works for any statistic, not just those that 46 approximate a known pre-calculated distribution. It provides an easy-to-understand, direct 47 measure of the extent to which an observed table differs from what might be predicted by a 48 hypothesis of independence.

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1 Results 2 3 Comparative analysis of trauma at Krapina with that of more recent groups 4 The amount of trauma observed in the Krapina sample was compared to the more recent 5 samples in two ways. The first way addresses the amount of trauma observed in each population 6 within the pooled context of the group of ten samples (Krapina and the nine comparative 7 samples examined for each element). In the second approach, for each element, pairwise 8 comparisons are made between the Krapina Neandertal sample and each comparative more 9 recent sample. The cranial elements tested were the frontals, parietals and occipitals; for the post- 10 cranial elements all major long bones. 11 For the pooled group of ten samples, the null hypothesis of independence was tested using 12 data from the Krapina sample and the other more recent samples. For each skeletal element, 13 incidence of trauma is hypothesized to be independent of the population observed. Each of 14 these columns represents a table used in the ACTUS simulation run. The number of times out 15 of the 10,000 simulated tables that the chi-squared values calculated from F-simulated tables 16 were equal to or exceeded the chi-squared value calculated from the observed table were 17 counted. These counts represent the significance (“p”-value) of the comparisons over the 18 group. For each element, the null hypothesis of independence of incidence of trauma and 19 population observed was rejected if p < 0.05. Any of the individual counts of incidences of 20 trauma in a population that are significantly higher or lower than predicted under the null 21 hypothesis (p < 0.05) are marked with “+” or “-”. This testing of independence of individual 22 cells, as well as the group as a whole, shows which cells represent statistically significant higher 23 or lower trauma counts than would be predicted within the context of the pooled group of 24 Krapina Neandertals and the comparative samples. In this way, we can observe whether the 25 Krapina Neandertals trauma counts are significantly smaller or larger than would be predicted 26 at random given the sample size compared to more recent comparative samples. 27 For the pairwise comparisons, the counts for each element in the Krapina Neandertal sample 28 were compared to each of the more recent samples using ACTUS as previously described. Out 29 of the 85 pairwise comparisons made, only a few rejected the null hypothesis of independence 30 between the amount of trauma observed and the collections sampled at p<0.05. 31 32 Results of comparisons of cranial elements 33 34 The results of the group comparisons are shown in Table 4.1. All cranial elements observed 35 (frontal, parietal and occipital) were inconsistent with the null hypothesis of independence of 36 incidence of trauma and population observed. This indicates a statistically significantly wide 37 range in the amount of cranial trauma observed in the groups, with the outliers indicated with 38 “+” or “-” signs. However, the Krapina Neandertal sample was never an outlier, and its trauma 39 counts did not represent significant deviations from what might be expected at random within 40 the context of the group as a whole. 41 Pairwise comparison results between the Krapina Neandertal sample and each of the more 42 recent samples are shown in Table 4.2. These results test if the trauma counts observed in the 43 Krapina Neandertal sample are significantly different from those observed in each more recent 44 comparative sample. For frontal trauma, the null hypothesis of independence was rejected in 45 five out of the nine comparisons with the amount of trauma at Krapina significantly higher 46 than the amount observed in two of the American samples, two of the Australian samples and 47 one of the European samples. For parietal trauma, the null hypothesis of independence was 48 rejected in three out of the nine comparisons with the amount of trauma at Krapina

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1 Table 4.1 Cranial trauma analysis by element for Krapina Neandertals and comparative populations (p 2 values listed for the entire group as a whole as well as for specific populations with significantly 3 smaller “-” or significantly larger “+” trauma counts than predicted under an hypothesis of 4 independence). 5 Frontal squama Parietal (posterior) Occipital squama 6 χ2: p=0.00for group χ2: p=0.00 for group χ2: p=0.00 for group 7 Skeletal population TOTAL # w/Trauma TOTAL # w/Trauma TOTAL # w/Trauma 8 Krapina Neandertals 20 3 29 2 14 0 9 Aljubarrota (Portugal) 93 7 91 4 24 4 10 SCl-038 (USA) 157 2 (p=0.03) - 159 4 (p=0.00) - 314 0 (p=0.02) - 11 Ala-329 (USA) 270 1 (p=0.00) - 268 7 (p=0.00) - 533 2 (p=0.03) - 12 Pozzilli (Italy) 50 1 53 3 (p=0.00) - 106 1 13 Central Murray (Australia) 384 14 390 62 734 8 14 Rufus River (Australia) 190 15 (p=0.04) + 204 53 (p=0.00) + 357 1 (p=0.03) - 15 South Coast (Australia) 246 15 253 56 466 8 16 Desert (Australia) 175 8 172 36 330 11 (p=0.01) + 17 East Coast (Australia) 202 17 (p=0.02) + 205 57 (p=0.00) 381 14 (p=0.00) + 18 19 20 Table 4.2 Results of pairwise comparisons of cranial elements for Krapina Neandertal sample versus each 21 comparative more recent population. 22 Element Null hypothesis rejected 23 24 Frontal Krapina v. SCl-038 (USA) [p =0.00] 25 Null hypothesis rejected for 5/9 comparisons Krapina v. Ala-329 (USA) [p =0.00] 26 Krapina v. Pozzilli (Italy) [p =0.04] 27 Krapina v. Central Murray (Aus.) [p =0.03] 28 Parietal Krapina v. SCl-038 (USA) [p =0.01] 29 Null hypothesis rejected for 3/9 comparisons Krapina v. Ala-329 (USA) [p =0.00] 30 Krapina v. Pozzilli (Italy) [p =0.01] Occipital 31 Null hypothesis not rejected for 9 comparisons 32 33 34 fi 35 signi cantly higher than the amount observed at two of the American samples and one of the 36 European samples. The null hypothesis of independence was not rejected for any of the pair- 37 wise comparisons of occipital trauma. 38 39 Results of comparisons of upper limb elements 40 41 The results of the group comparisons are shown in Table 4.3. The clavicle and the ulna were 42 inconsistent with the null hypothesis of independence of incidence of trauma and population 43 observed, however the null hypothesis was not rejected for the radius and humerus. This 44 indicates a statistically significantly wide range in the amount of ulnar and clavicular trauma 45 observed in the groups, with the outliers indicated with “+” or “-” signs. However, the 46 Krapina Neandertal sample was never an outlier, and its trauma counts did not represent 47 significant deviations from what might be expected at random within the context of the group 48 as a whole.

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1 2 0 3 6 2 1 4 13 =0.02) - =0.02) + 3 =0.05)- p p 4 p 5 6 0.02 for group

7 p= 62 12 48 : 180 1 ( 155 288 186 2 ( 163 2 Radius χ 8 TOTAL # w/Trauma 9 10 11 1 2 6 12 3 11 10 15 14 =0.02) - 349 20 ( =0.03) + 91 p 13 p 14 15 16 0.07 for group p= : values listed for the entire group as a whole as well as for

17 2 TOTAL # w/Trauma Ulna χ 18 p 19 20 02 61 170 31 134 275 3 3402 11 ( 128 0 16 12 79 47 10 ( 21 1 150 22

23 trauma counts than predicted under an hypothesis of independence). ”

24 “ + 0.40 for group 70 21 25 51 318 111 121 171 p=

26 : 2 Humerus χ 27 TOTAL # w/Trauma

28 cantly larger ﬁ 29 30 or signi 4 1 ” =0.01) -=0.01) - 139 299 31 =0.01) + 447 p p p 32 “ - 33

34 0.00 for group

35 p= 56 16 : 245 15 ( 159289 0 ( 2 ( 2 cantly smaller TOTAL # w/Trauma Clavicle χ No data No data No data No data 36 No data ﬁ 37 38 39 40 41 42

43 c populations with signi 44 ﬁ Upper limb trauma analysis by element for Krapina Neandertals and comparative populations ( 45 speci 46 47 Table 4.3 Skeletal population Pozzilli (Italy) Central Murray (Australia) Aljubarrota (Portugal) Rufus River (Australia) Krapina Neandertals SCl-038 (USA) Ala-329 (USA) South Coast (Australia) Desert (Australia) 48 East Coast (Australia) Template: Royal A, Font: , Date: 26/07/2013; 3B2 version: 9.1.460/W Unicode (Apr 1 2008) (APS_OT) Dir: E:/TAFUK/3B2/KNUSEL-SMITH -131047/130001/APPFile/BK-TAF-KNUSEL-SMITH-131047-130001.3d

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1 Pairwise comparison results between the Krapina Neandertal sample and each of the more 2 recent samples are shown in Table 4.4. These results show only one instance of a significantly 3 different amount of trauma in the Krapina Neandertal sample from that observed in any of the 4 comparative samples. In this case, the Krapina Neandertal sample was significantly higher in 5 one of the American clavicular samples. 6 7 8 Results of comparisons of lower limb elements 9 Although there was no lower limb trauma observed in the Krapina Neandertal sample, given the 10 size of the sample, the lack of observation of lower limb trauma is not significantly smaller than 11 expected. For all the groups together, the femur and the fibula were inconsistent with the null 12 hypothesis of independence of incidence of trauma and population observed, however the null 13 hypothesis was not rejected for the tibia (Table 4.5). This indicates a statistically significantly, 14 wide range in the amount of femoral and fibular trauma observed in more recent groups, with 15 the outliers indicated with “+” or “-” signs. However, despite the lack of lower limb trauma, 16 17 18 Table 4.4 Results of pairwise comparisons of upper limb elements for Krapina Neandertal sample versus 19 each comparative more recent population. 20 Element Null hypothesis rejected 21 22 Clavicle Krapina v. SCl-038 (USA) [p=0.02] 23 Null hypothesis rejected for 1/9 comparisons 24 Humerus Null hypothesis not rejected for 9 comparisons 25 Radius 26 Null hypothesis not rejected for 9 comparisons 27 Ulna 28 Null hypothesis not rejected for 9 comparisons 29 30 31 32 Table 4.5 Lower limb trauma analysis by element for Krapina Neandertals and comparative populations (p values listed for the entire group as a whole as well as for specific populations with significantly 33 smaller “-” or significantly larger “+” trauma counts than predicted under an hypothesis of 34 independence). 35 36 Skeletal population Femur Tibia Fibula χ2 χ2 χ2 37 : p=0.00 for group : p=0.09 for group : p=0.01 for group 38 TOTAL # w/Trauma TOTAL # w/Trauma TOTAL # w/Trauma 39 Krapina 32 0 19 0 32 0 40 Libben (USA) 338 9 (p=0.00) + 344 5 348 9 41 SCl-038 (USA) 117 2 163 1 128 1 42 Ala-329 (USA) 313 0 310 5 237 0 (p=0.03) - 43 Pozzilli (Italy) 80 0 75 3 59 1 44 Central Murray (Australia) 336 0 254 0 177 0 45 Rufus River (Australia) 167 0 154 1 146 2 South Coast (Australia) 133 1 201 0 77 0 46 Desert (Australia) 45 1 62 1 38 3 (p=0.02) + 47 East Coast (Australia) 172 1 164 4 126 4 48

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1 Table 4.6 Results of pairwise comparisons of lower limb elements for Krapina 2 Neandertal sample versus each comparative more recent population. 3 Element Null hypothesis rejected 4 5 Femur 9 comparisons 6 Null hypothesis not rejected 7 Tibia 9 comparisons 8 Null hypothesis not rejected 9 Fibula 9 comparisons Null hypothesis not rejected 10 11 12 13 the Krapina Neandertal sample was never an outlier, and its counts did not represent significant 14 deviations from what might be expected at random within the context of the group as a whole. 15 Pairwise comparison results between the Krapina Neandertal sample and each of the more 16 recent samples are shown in Table 4.6. These results show no instance of a significantly dif- 17 ferent amount of trauma in the Krapina Neandertal sample from amounts observed in any of 18 the comparative samples. 19 20 Discussion and conclusions 21 22 Comparisons among all the samples of frontal, parietal and occipital injury reveal a wide range in 23 the amount of cranial trauma among the (mostly) hunter-gatherer groups. The Krapina sample 24 fits easily within these ranges. Thus, the amount cranial trauma in the Krapina Neandertals is not 25 statistically significantly different from the comparative more recent samples. Most of the 26 pairwise comparisons produced no significant differences for Krapina frontal, parietal and 27 occipital trauma. Frontal and parietal trauma counts are statistically significantly higher at Kra- 28 pina than for two of the American sites SCl-038 and Ala-329 and the Italian site Pozzilli. Walker 29 (2001: 588) noted that cranial trauma observed at SCl-038 and Ala-329 sites was low compared 30 to that of other prehistoric Californian groups, while Brasili et al. (2004: 189) argued the Pozzilli 31 lifestyle was “rather aggressive and violent” due to the presence of iron spears, knives and blades. 32 However, it is possible that the violence at Pozzilli did not manifest itself though cranial blunt 33 force trauma. Despite these differences, overall levels of Krapina cranial trauma are within the 34 range of the more recent comparative samples, indicating that the amount of frontal, parietal and 35 occipital trauma is typical for hunter-gatherer or nomadic groups. The Krapina Neandertals do 36 not stand out as especially violent, measured by blunt force traumatic injuries. 37 Determining the cause of these head injuries is complicated and full of assumptions about 38 the nature of the healed wound, the perpetrator or action involved in the injury. Clues to the 39 specific, prehistoric actions can be inferred from modern forensic analyses, but forensic scien- 40 tists are primarily concerned with fatal head wounds. Kanz and Grossschmidt (2006) studied 68 41 gladiators from a Turkish cemetery, and all 16 ante-mortem blunt force traumatic injuries were 42 positioned above the “hat-brim line”, roughly defined as the perimeter of the maximum 43 neurocranial circumference (Ehrlich and Maxeiner 2002). Healed injuries were only on the 44 frontal (69%) and parietal (31%) and more common on the right frontal than the left, but not 45 the parietal. In a German sample, Ehrlich and Maxeiner (2002: 773) reported “head injuries 46 from blows occur[ing] more often (55%) above the so-called hat-brim line … than injuries 47 from falls on flat surface”. In a recent sample from Montreal, Kremer et al. (2008) found 48 similar results with fatal wounds above the hat-brim line related to blunt force trauma (68%), as

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1 opposed to falls, either from standing or from falling down stairs. Both of the latter were more 2 commonly below the hat-brim line and typically on the right side (72%). 3 In modern skeletal populations cranial trauma is often regarded as the result of interpersonal 4 violence, especially healed trauma of the head and face (Smith 1976; Walker 1997; Alvrus 5 1999; Steadman 2008), and we suspect that human violence is the likely cause of healed blunt 6 force injuries at Krapina. Similar healed wounds occur in earlier (e.g. Caspari 1997; De 7 Lumley-Woodyear 1973; Weidenreich 1943, 1951) and later fossils (e.g. Frayer et al. 2006; 8 Roksandic et al. 2006 Franciscus and Vlcˇek 2006). Others have suggested that healed head 9 wounds could be the result of environmental hazards such as falling rocks as a consequence 10 living in caves (Mann and Monge 2006) or occupational hazards related to hunting large prey 11 (Berger and Trinkaus 1995; Pettitt 2000; Underdown 2006). Rarely, as in the case of Saint 12 Césaire I (Zollikofer et al. 2002), is interpersonal violence suggested as the source of the trau- 13 matic injury. Some discount the role of Neandertal interpersonal violence due to low popu- 14 lation density in the Middle Pleistocene. This view is articulated by Berger and Trinkaus 15 (1995: 849): “…their [Neandertal] undoubtedly low population densities probably made 16 dispersal a far more viable alternative to conflict resolution than the interpersonal violence 17 which so often characterizes conflict resolution among sedentary Recent human populations.” 18 Population density seems irrelevant since all it takes is two for interpersonal violence to take 19 place. Violence against women (Martin 1997) and cultural habits, like Yanomamö club fight- 20 ing, are also possible scenarios. These non-lethal fights produce many healed head wounds and 21 “enormous scars – [of] which they are very proud” (Chagnon 1968: 173). Webb (1995: 205) 22 speculates that some frontal depression fractures in native Australian women were self-inflicted 23 when they beat their foreheads with stones as part of a mourning ritual. Presuming most of 24 these women were right-handed, the head trauma should be primarily on the right side. Other 25 unpredictable, idiosyncratic cultural habits could account for the healed head wounds at Kra- 26 pina, which may or may not be associated with violence. Four of five are on the left side of 27 the neurocranium suggesting the possibility of a frontal-attack by a right-handed aggressor. We 28 favour this argument over accidental injuries, such as roof debris falling causing the injuries. 29 The odds of this would be low with the head making a poor “target” for a ceiling rock and 30 these injuries were unlikely to have happened during sleep, given the location of most near the 31 crown of the cranium. No similar wounds occur in cave bears (Miracle 2007; our unpublished 32 observations), which presumably hibernated at Krapina, although their crania are much thicker 33 than those of Neandertals. 34 Our results show that both the amount of trauma at Krapina and its distribution over the 35 frontals, parietals and occipitals fit within the range of modern hunter-gather populations 36 where interpersonal violence seems to be the main cause of cranial trauma. Because the 37 Neandertal skeletal remains at Krapina are fragmentary and disarticulated, it is not possible to 38 integrate information from other parts of the body in determining a cause for the cranial 39 trauma. However, we see no reason to assume that some of the cranial injuries, especially those 40 of Krapina 4, 5 and 20, are not the result of interpersonal violence, given the shape and loca- 41 tion of the lesions. Only Krapina 34.7 is below the “hat-brim line” and on the right side, 42 making it a better candidate for an accident. After 100,000-plus years, reconstruction of the 43 specific events causing these head wounds is problematic, beyond the typical guesswork of 44 palaeoanthropology and we offer them with caution. 45 Like the evidence from the cranium, the level of post-cranial trauma at Krapina is not 46 statistically significantly different within the context of the entire group of comparative more 47 recent samples. The comparisons among all the samples of claviculae, humeri, radii, ulnae, 48 femora, tibiae and fibulae reveal a wide range of post-cranial trauma among mostly hunter-

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1 gatherer groups, and Krapina fits easily within these ranges. Only in the comparison of clavicular 2 trauma between Krapina and SCl-038 was the null hypothesis rejected. Like the cranial trauma, 3 causes of the trauma are difficult to unravel. Both of the ulnar fractures at Krapina are located 4 near midshaft. Judd (2004: 47) notes that midshaft ulnar injuries are not commonly associated 5 with falls, as opposed to Colles’ fractures, suggesting that their etiology is likely aggression related. 6 But, we just do not have the ability to decipher the cause of these ulnar injuries, other than to 7 suggest they must be related to some traumatic event in the Krapina Neandertals’ past. 8 Previous studies of Neandertal trauma (Underdown 2006; Pettitt 2000; Trinkaus 1995) 9 emphasized the low levels of lower limb injuries and the special role locomotion may have 10 played in Neandertal social structure. However, given our results, the absence of lower limb 11 injuries at Krapina does not represent a significantly lower injury rate than that observed in any 12 of the more recent comparative samples. Although our results do not show unambiguous 13 evidence of trauma caused by interpersonal violence at Krapina, these results demonstrate that 14 the frequency and placement of traumatic lesions in this sample of Neandertals are completely 15 within the range of those observed in modern hunter-gather and nomadic groups where 16 interpersonal violence is a part of life. 17 18 Acknowledgements 19 20 We thank George F. Estabrook (University of Michigan, Ann Arbor), Eugenia Cunha (Uni- 21 versade de Coimbra, Coimbra), Jakov Radovcˇic´ (Croatian Natural History Museum, Zagreb), 22 Janet Monge (University of Pennsylvania, Philadelphia), Luka Mjeda (Croatian Photographic 23 Center, Zagreb), Ivana Fiore and Luca Bondioli (Museo Nazionale Preistorico Etnografico “L. 24 Pigorini”, Rome), Antonio Todero (University of Bologna) and the editors for assistance with 25 this chapter. 26 27 Note 28 29 1 ACTUS was written originally as a DOS program (Estabrook and Estabrook 1989); however, now 30 there is an updated ACTUS Excel, which runs as an Excel macro (included in Estabrook 2011) or can be obtained by contacting V. Hutton Estabrook ([email protected]). 31 32 33 References 34 Almada, V. C., E. J. Goncalves, A. J. Santos and C. Baptista. 1994. Breeding ecology and nest aggrega- 35 tions in a population of Salaria-Pavo (Pisces, Blenniidae) in an area where nest sites are very scarce. 36 Journal of Fish Biology 45: 819–30. 37 Alvrus, A. 1999. Fracture patterns among the Nubians of Semna South, Sudanese Nubia. International – 38 Journal of Osteoarchaeology 9: 417 29. Ambade, V. N. and H. V. Godbole. 2006. Comparison of wound patterns in homicide by sharp and 39 blunt force. Forensic Science International 156: 166–70. 40 Amorim, M. C. P., Y. Stratoudakis and A. D. Hawkins. 2004. Sound production during competitive 41 feeding in the grey gurnard. Journal of Fish Biology 55: 182–94. 42 Aufderheide, A. C. and C. Rodríguez-Martin. 1998. The Cambridge Encyclopedia of Human Paleopathology. 43 Cambridge: Cambridge University Press. Berger, T. D. E. and E. Trinkaus. 1995. Patterns of trauma among the Neandertals. Journal of Archaeological 44 Science 22: 841–52. 45 Bocquet-Appell, J.-P. and J. L. Arsuaga. 1999. Age distributions of hominid samples at Atapuerca (SH) 46 and Krapina indicate accumulation by catastrophe. Journal of Archaeological Sciences 26: 327–38. 47 Brasili, P., E. Bianchi and A. R. Ventrella. 2004. Traumatic events and life-style in ancient Italian – 48 populations. Collegium Anthropologicum 28: 179 91.

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