Holocene Khoesan health: a biocultural analysis of cranial pathology and trauma

Gibbon Victoria (Orcid ID: 0000-0001-7875-3297) Davies Bronte (Orcid ID: 0000-0003-4848-1192) Holocene Khoesan health: a biocultural analysis of cranial pathology and trauma Victoria E. Gibbon1, Bronte Davies1 1 Department of Human Biology, University of Cape Town, Observatory, 7935, South Africa. Corresponding Author: Victoria Elaine Gibbon Email: Victoria.Gibbon@uct.ac.za Tel: +27 21 650 4431 Fax: +27 21 448 7226 Department of Human Biology, Anatomy Building Level 5, Room 5.14 (recipient Dr. Victoria Gibbon) Faculty of Health Sciences, University of Cape Town Anzio Road, Observatory, 7925 Abstract A bioarchaeological assessment of human skeletons provides an objective opportunity to study and understand past peoples. Crania were analysed to evaluate health and well-being of a past Khoesan sample in southern Africa. These data were analysed temporally and spatially (coastal vs inland), and by sex and age. Cranial and dental pathology of 150 adult individuals from the Human Skeletal Collection at the University of Cape Town were assessed. Most individuals were of coastal origin, equally distributed between pre- and post-2000 BP. The sample was nearly equally distributed by sex and consisted primarily of middle-aged adults (35-50 years at time-of-death). Physiological stress was indicated by evidence of nutrient deficiencies and signs of metabolic stress, without significant differences by sex. Cribra orbitalia was more frequent post-2 000 BP, and linear enamel hypoposlasias were common in those who died at younger ages. Signs of infection were observed in dental tissues as carious lesions, abscess and antemortem tooth loss. Trauma was found in 52% of the sample, 19% was antemortem, with a significant increase post-2000 BP, 39% were caused perimortem and was significantly higher in women. Significant distributions of pathology indicate an increased stress load from 2000 BP onwards and it was higher among coastal individuals, indicating a critical period with an influx of people migrating into the region causing population displacement, forced internal migration and increased conflict at this time. Despite the seemingly high pathogen load, these individuals were resilient and living through health problems. Across this broad time-period, the southern African Khoesan lifestyle proves to be adequate to maintain good health and longevity in their social and environmental context. KEYWORDS: hunter-gatherer, bioarchaeology, trauma, nutritional stress, Later Stone Age This article has been accepted for publication and undergone full peer review but has not been through the copyediting, typesetting, pagination and proofreading process which may lead to differences between this version and the Version of Record. Please cite this article as doi: 10.1002/oa.2854 This article is protected by copyright. All rights reserved. INTRODUCTION Prior to written historical records, the evidence of how people lived has been elucidated through artefactual remnants. Biological information gained from human remains serve as an objective lens to understand the stressors and health of past people. The cohesive stress model describes a cyclical relationship between biophysical, socioeconomic and political conditions of daily life that alters biological homeostasis, inducing strain on individual and population levels (Buzon, 2006; Goodman et al.,1988). Stress in bioarchaeology is also defined as a response to unhealthy environmental conditions causing physical disruption, which may have deleterious consequences at both the individual and population level (Buzon, 2006). Using adult crania, we aim to look at health and wellness temporally and spatially among a sample of indigenous people from southern Africa. The Later Stone Age (LSA) began around 40000 years ago and was dominated by hunter-gatherers commonly known as the San. They originally occupied most of the region before the arrival of Bantu-speaking people in southern Africa (Soodyall & Jenkins, 1997; Morris, 2002) who colonised the region through a series of migrations around 2000 BP (Deacon, 1984; Sealy, 2010). The emergence of the Khoekhoe immediately preceded the arrival of the Bantu-speaking populations. Khoekhoe people were Khoe speaking hunter-herders or “those who kept cattle and sheep, [and] were seasonally transhumant” (Sadr 2008:179). Khoesan is a term used to group the culturally distinct (Smith, 1990) but biologically and morphologically homogeneous Khoekhoe and San groups from southern Africa (Soodyall & Jenkins, 1997; Stynder, 2009). The Khoesan are often grouped as a homogenous group of people and ethnographic records from the Kalahari dominate the literature. A broad assessment of health and wellness among Holocene foragers temporally and spatially will contribute to the existing body of knowledge and data obtained from previous archaeological, bioarchaeological and ethnohistorical analyses on these past peoples. Examining a larger sample of people across time and space provides the opportunity to examine the impact cultural and subsistence differences have on gender roles, cultural practices and health. The sociopolitical landscape in southern Africa changed dramatically 2000 BP and examining the skeletal remains before and after this time one can examine how these changes may have impacted biological health and stress for these people. This will be achieved by first defining the age and sex profile of the sample: are both sexes equally represented? What does the age-at-death distribution indicate about population health in the sample? Followed by an assessment of cranial pathology, trauma and variation in this sample temporally and spatially; are health, stress, and interpersonal violence more common amongst inland or coastal individuals? Were there socioeconomic implications for health post-2000 BP? Or more frequent by a specific sex and age? Combined, these data will inform a broad assessment of Holocene forager health. MATERIALS AND METHODS Adult crania of 150 Holocene Khoesan were examined from the UCT Human Skeletal Repository (Figure 1). Each were classified spatially by distance from the coast. Sites within the Fynbos biome between the Cape Fold Belt Mountains and the coast, and outside the Cape within 20 km of the coastline were classified as coastal. Outside these regions, individuals were classified as inland. Where radiocarbon dates were unavailable, individuals were temporally classified as pre- or post-2000 BP. All data were obtained and analysed by Davies and verified by Gibbon. This article is protected by copyright. All rights reserved. Sex and age estimation The methods for sex and age estimation followed protocols described by Buikstra and Ubelaker (1994). Os-coxae or cranial morphology were used for sex. Age-at-death was estimated by dental eruption, epiphyseal fusion, cranial suture closure, changes to the sternal rib ends and os-coxae. Age-at-death for leaner populations of shorter stature and mobile south African foragers is often underestimated (Merritt, 2015; Pfeiffer et al., 2019), therefore it is acknowledged that the age may be underestimated in this study. Individuals were classified broadly as young (20-35 years), middle-aged (35- 50 years), old (>50 years) adults, or as ‘unknown adult’. Health and stress Cranial bone loss and formation were recorded and classified according to physiological stress, disease or trauma (Figure 2). Cribra orbitalia (CO), porotic hyperstosis (PH) and linear enamel hypoplasia (LEH) were recorded. CO refers to pitting in the orbital roof with diploic thinning, and PH is characterised by diploic thickening and pitting on the cranial vault (Walker et al., 2009; Rivera & Lahr, 2017). Both conditions were classified by severity (Kent et al., 1994; Scott et al., 2018; Figure 2). Disruption of growth during development can be observed dentally by LEH. Infection can be observed by lytic pitting or proliferation, and lesions may be non-specific. Metabolic disruptions causing a lesion may increase individual susceptibility to secondary infections (Goodman et al., 1988). Dental analyses were included as dental disease has the potential to weaken an individual’s immunity to secondary infections, and can be caused by diet, trauma, cultural practices, microorganisms and metabolic stress (Hillson, 2005). Carbohydrate fermentation by oral bacteria and subsequent decrease in oral pH produce cavities through the progressive demineralization of teeth. There are several anatomical, genetic, dietary and degenerative reasons that increase and predispose a person or people to cavities. Chronic inflammatory responses also irritate soft tissues that lead to resorption of the alveolar walls (Larsen, 1991). Dental infection in the form of carious lesions that may lead to antemortem tooth loss (AMTL) (Gibbon & Grimoud, 2014). Where teeth were present, frequency of caries and AMTL were recorded by site for each permanent tooth. Analysis of trauma can be important for assessing misadventure of past people, indicating levels of accidental/intentional violence, ability to heal or fatal events (Lovell, 2008). Cranial deformation and fracturing as a result of trauma was recorded as occurring ante-, peri- or postmortem based on the visible remodeling of bone (see Figure 2). Data analyses Inter- and intra-observer tests agreed with acceptable values. Data were analysed using twosided Fisher’s Exact tests, chi-square tests of independence and Kruskal-Wallis tests. Statistical analyses were conducted in IBM SPSS software (v. 25) and significance was considered at p≤0.05. RESULTS Results are summarised in Table 1. The sample was 44% female (66/150) and 39% male (59/150), with 16% of unknown sex (25/150). Most (56%, 84/150) were middle-aged adults (35-50 years); with 20% being young adults (30/150) and 3% old adults (4/150). Adult individuals of indeterminate age were 21% of the sample (32/150). Temporally, 33% (49/150) of individuals were dated to pre-2000 BP and 29% (43/150) to post-2000 BP, with 39% undated (58/150). Spatial distribution of the sample was predominantly coastal (73%, 110/150) compared to 22% from inland sites (33/150) and 5% from unknown sites (7/150). Teeth were present in 68% of individuals (102/150), with a total of 2049 permanent teeth This article is protected by copyright. All rights reserved. assessed. At least one instance of pathology was observed in 95% of the individuals (143/150) and ranged between one and 11 per individual. Unless otherwise stated below no statistically significant differences were observed by tooth type, age, sex, temporally or spatially. Evidence of stress CO was present in 23% of individuals (35/150), all instances were healed at time-of-death, with an average severity score of 2 out of 4 indicating a moderate condition. Of these lesions, 74% were bilateral (26/35) and 26% unilateral (9/35). Individuals from coastal sites had significantly higher instances of CO compared to individuals from the interior (p= 0.003). Significant difference was also found for individuals post-2000 BP compared to fewer from pre-2000 BP (p= 0.048). PH was observed in 41% of the sample (62/150), with instances occurring predominantly at the anterior portion of the frontal bone and posterior parietal regions, with a mean severity score of 1 out of 4, indicating a mild condition. Instances of PH were nearly equally distributed temporally and spatially, not discriminating with regards to age or sex. There was dual presence of both PH and CO in 15% of the sample (23/150). Incidence of LEH (independent of frequency per tooth) was observed in 11% of the sample (16/102), most often on the canines, followed by incisors and premolars in mandibles and maxillae. Higher incidences of LEH were found on canines of individuals estimated to be young adults at time-of-death compared to middle-aged or older adults (p= 0.037). Dental pathology and temporomandibular joint degeneration Carious lesions were found in 52% of individuals (53/102); 168 lesions in total. Of these, 50% (51/168) were gross carious lesions, while the remaining caries (117/168) were predominantly observed on the tooth crown or at the cemento-enamel junction, at occlusal, mesial or distal surfaces in both the maxillae and mandibles. Infection was observed in alveolar bone as evidence of dental abscess or periapical infections in 30% of the sample (45/150). Infection was observed at the mandibular condyles evidenced by abscess in two coastal individuals: UCT 593 and UCT 596. At the infection sites, a range of one to three lesions were observed. AMTL with varying levels of resorption, was observed in 36% of individuals (52/150). Specifically, 9% of examined teeth were lost antemortem (185/2049). Most of these were molars (87/185), followed by incisors (47/185). Bi- and unilateral degeneration at the temporomandibular joint (TMJD) was observed in four individuals: three coastal and one inland. Interpersonal violence Dental trauma, including chipping and fracturing, were observed in 23% of the teeth examined (464/2049). Instances of ante- or peri- mortem trauma were observed in 52% of the sample (78/150). Antemortem trauma was observed in 19% of the sample (29/150), occurring most commonly on the parietal bones, followed by the frontal and occipital bones. The number of antemortem lesions in individuals ranged between one and three per individual. Significant differences in antemortem trauma frequencies were found temporally, being higher in individuals post-2000 BP (p= 0.008). Instances of perimortem trauma (39%, 58/150) were most commonly found in the lateral facial zygomatic regions and temporoparietal areas. Lesions indicative of perimortem trauma ranged between one to five per individual. Instances of perimortem trauma were significantly higher in females (p= 0.046). Individuals with both ante- and perimortem trauma were observed in 6% (9/150) of the sample, with 13% having only antemortem trauma (20/150) and 33% only perimortem trauma (49/150). This article is protected by copyright. All rights reserved. DISCUSSION This Khoesan sample is a broad sample through time and space, which allows for an assessment of the impact cultural, subsistence and the sociopolitical changes in southern Africa had on biological health and stress for these people. The results are interpreted acknowledging limitations to archaeological skeletal analyses (Wood et al., 1992; Larsen, 1995). Poorly preserved remains and fragmentation were barriers to sex and age estimations. Sex estimation proved challenging, as high activity of the Khoesan lifestyle (Churchill & Morris, 1998) may have diminished sexual dimorphism of the crania. Therefore, the pelvis was more reliable and when unavailable often resulted in indeterminant sex. Age-at-death for leaner populations of shorter stature and mobile south African foragers is often underestimated (Merritt, 2015; Pfeiffer et al., 2019). Therefore, the true age for these people is likely older, and the large proportion of middle-aged adults in the sample (56%) likely indicates the end-of-life period. This is supported by the small sample of older adults present, and most people being ‘middle-aged’ adults between 35-50 years at time-of-death. Research on living Khoesan groups showed an average lifespan to be 40 to 60 years-of-age (Truswell & Hansen, 1976), and living beyond 50 was relatively rare (Lee & DeVore, 1976). Other studies on archaeological assemblages of Khoesan have shown similar results (Botha & Steyn, 2016). These data show unsuitability of contemporary morphological sex and age methods, highlighting the need for developing population-specific sex and age estimation methods (Pfeiffer & Harrington, 2011; Kurki et al., 2012; Pfeiffer et al., 2019). Health and stress PH may indicate a vitamin deficiency both spatially and temporally. Low severity scores for both CO and PH, with all CO being healed, indicate that metabolic and physiological stress were prevalent. However, it also suggests individual resistance to stress instead of inherent weakness as individuals did not succumb to the conditions, which were unlikely severe enough to be a leading cause-of-death. Pfeiffer (2007) reported that infectious diseases were rare in mid-Holocene South African populations, similar results were found in this sample. While CO and PH may be caused by an underlying infectious disease, the non-specificity of these markers prevents direct diagnosis (Walker et al., 2009; Rivera & Lahr, 2017). It is also caused by imbalance of gut microbiota and/or pathogens, prolonged breastfeeding and infant weaning practices (Walker et al., 2009). Evidence of intestinal parasites and iron deficiency in past huntergatherers from a shell midden in Canada has been observed (Bathhurst, 2005). Parasitic infections are still common today in southern Africa (Samie et al., 2009) and thus the potential for parasites in the environment of past South Africans is high. Research of contemporary foragers shows enriched microbiomes were resistant to diarrheal infections (Schnorr et al., 2014). The LSA diet and lifestyle likely also produced an enriched microbiome, offering some protection against parasitic infections and diarrhea, which may account for the resistance to stress. LEH in the sample were mainly found on the canines, which form between six months and six years of age, indicating metabolic stress occurring during these ages (Goodman & Rose, 1990; Esan & Schepartz, 2018). This is consistent with the age of weaning that is known to occur among Khoesan between the ages of two and four (Konner & Worthman, 1980; Clayton et al., 2006); a period described as often traumatic, which can lead to caloric deficiency, impact development and increase susceptibility to infection or illness (Lee & DeVore, 1976; Truswell & Hansen, 1976; Stinson, 2002; Konner, 2017). Child and infant mortality among foragers were common (Stinson, 2002). This combined with the high This article is protected by copyright. All rights reserved. level of LEH, which are evidence of childhood stress (Goodman & Rose, 1990; Katzmarzyk & Leonard, 1998), suggest these people were biological stressed from a young age. As these were found in adults, it also indicates survival through stress-periods into adulthood. The presence of LEH in young adults was significant and may indicate individual frailty or that physiological stressed children became frail adults, increasing their susceptibility to infection or disease, resulting in premature death (Steckel, 2005; Buzon, 2006; Brown et al., 2009). The gestation and early child rearing years in a nomadic forager lifestyle are energy expensive on the mother. Prolonged breastfeeding is a known method of birth spacing used by the Khoesan and considered optimal for infant survival and group fitness (Jones, 1986; Clayton et al., 2006). It is suggested that lactating females require an extra 1000 calories a day (Kolata, 1974). Other research has shown increased mortality risk for Holocene forager women associated with childbirth (Pfeiffer et al., 2014). Therefore, in this study more evidence of stress among females was anticipated, but not found. Thus, the similarity in stress level among both sexes may indicate the ability for Khoesan women to recover from poor conditions, and the biological burden of reproduction and lactation. These results may be due to the well-documented biological stress buffer within females for reproduction to better resist biological stress and strain (Stinson, 2000; Wells, 2012). It has been shown that men are more adversely affected by stress and similarly have greater responses to improved conditions (Vercellotti et al., 2011). Ethnographic studies show food sharing and mutual aid were an integral component of Khoesan society and differences in marine versus terrestrial foods cannot be discerned between men and women (Sealy & Pfeiffer, 2000). Women may have used body fat stores during lactation periods (Kirchengast & Winkler, 1996), minimising reproductive nutritional strain. Despite the biological burdens of carrying children and lactation, women were no more biologically stressed than their male counterparts. The high prevalence of dental pathology found would have been a risk factor for general health and may have caused secondary infections. Such pathology was a leading cause of death in the past (Calcagno & Gibson, 1991). LSA people had heavy dental wear eventually leading to pulp exposure, which may lead to the formation of bacterial infection, septic lesions and allow infection to enter the blood stream. Flat molar wear as observed in this sample is a pattern functional in human history common until the advent of agriculture (Kaifu et al., 2003; Grimoud & Gibbon, 2017). Studies show that variation in cavity rate depends on region, with extremely low rates on the coast (van Reenen, 1966; Sealy et al., 1992; Botha & Steyn, 2015). Differences in cavity incidence between males and females have been shown (Sealy et al., 1992) and dental pathology and tooth loss are associated with increasing age (Larsen, 1995). Neither of these patterns were found in this study. Most dental abscesses and periapical lesions were found in posterior teeth: the first molar is commonly the most worn tooth, and horizontal wear is a caries risk as it reaches the pulp chamber. The high rates of infection and number of gross caries in this sample (50% of caries) indicate infection due to wear as the most probable cause of tooth loss. Interpersonal violence Case studies of trauma have generated varying hypotheses regarding violence and aggression among South African Khoesan. Some argue they were able to resolve disputes before resorting to violence (Draper, 1978; Thomas, 1989). This indicates that trauma would likely be the result of altercations with predators, or accidental injury infliction by tools (Morris et al., 1987; Morris, 2012; Pfeiffer, 2016). Alternatively, this ‘dispute resolution hypothesis’ is opposed through multiple instances of perimortem trauma and intentional wound infliction (Pfeiffer & van der Merwe, 2004; Morris, 2012; 2014). Violence is found in all modern human societies, it was an integral aspect used to contain and resolve social conflict and to This article is protected by copyright. All rights reserved. secure desired outcomes (McCall & Shields, 2008; Martin et al., 2010). The location of the ante- and perimortem trauma in this sample was predominantly on the cranial vault, indicating interpersonal violence, with some individuals showing up to five lesions. Trauma on the facial skeleton can be caused by either accidental or interpersonal violence (Lovell, 2008), however, relatively fewer instances of this were observed. Accidental trauma by tools or terrain cannot be ruled out, incidence and location of trauma in this sample arguably indicates that events of inter and intra-group conflict occurred. Lee (1979) describes violence in contemporary Khoesan as verbal disputes, fights with and without weapons and some as fatal. These descriptions are consistent with a knock on the head with a round or sharp object. This may be the result of inadequate conflict management strategies (Pfeiffer, 2009) or Vitamin B12 deficiency that has been associated with neurological and behavourial disorders (Walker et al., 2009). Antemortem trauma may indicate care for injured individuals and suggest interpersonal violence between bands rather than within bands (Pfeiffer, 2012; 2016). Perimortem trauma was significantly higher in females. These lesions are indicative of fatal events and their higher instances in females may suggest sources of conflict i.e. marriage, sex-based cultural practices, or daily foraging activities may make females more vulnerable to such trauma. Pfeiffer (2016) also noted that most victims of perimortem trauma from a sample pre-2000 BP were women and children. Lee (1979) and Christiansen and Winkler (1992) show violence was also perpetuated by female contemporary Khoesan, with emotional quarrels between males and females often starting suddenly and ending after a punch or sticks and stones thrown a short distance. Nomadic living !Kung women are shown to have had higher status and were directly involved in decision making (Thomas, 1989) and perhaps their social status and influence put them at risk. Although women are often described as victims of violence rather than perpetuators, research shows women use indirect violence such as verbal attacks, social manipulation and hitting, which have been linked to maintain social order (Martin et al., 2010). Dietary differences and cultural use of teeth as tools Dental trauma, including chipping and fracturing, were observed in 23% of the teeth examined (464/2049). In forager societies using teeth as a third hand was common, which causes heavier wear on anterior teeth and irregular chipping and fracturing patterns (Molnar, 1972; Gibbon and Grimoud, 2014). Van Reenen (1992) noted molar chipping found among South African foragers and their anterior teeth are shown to have high levels of wear (Van Reenen, 1966; 1992; Smith, 1984; Botha & Steyn, 2015). Therefore, these conditions likely suggest the use of teeth as tools. TMJD was observed in four individuals, wear was bilateral in three. Each had varying levels of AMTL, reflecting the observation that joint wear is associated with tooth loss and attrition (Hodges, 1991). Suby and Giberto (2018) suggest that dietary patterns may be an aetiological factor for TMJD. Complete wear of the condylar facet in the TMJ was observed in two coastal individuals from pre-2000 BP. Another coastal individual had left anterior unilateral arthritis. An inland female had bilateral posterior wear at the TMJ, which suggests hyperextension of the jaw, independent of AMTL and wear. Further exploration into its cause is required. TMJD may also be linked to the use of teeth as tools. Health at the coast The number of LSA human remains from the coastal region is rich (Deacon, 1984; Parkington, 1984; Sealy & van der Merwe, 1986), and knowledge about these people is biased towards coastal dwellers. This sample was primarily from the coast (73%). CO appeared to be predominant on the coast, most likely caused by hypocellular aplastic or protein deficiency anaemias, chronic, endocrine disorders and/or vitamin C deficiency This article is protected by copyright. All rights reserved. (Walker et al., 2009; Rivera & Lahr, 2017). This is consistent with the observation that marine-based diets are nutrient restrictive (Sealy & van der Merwe, 1986) although providing enough calories (De Vynck et al., 2016). However, inland foragers also experienced nutritional deficiencies (Truswell & Hansen, 1976). Studies of stable isotopes have shown diet variability with marine resource consumption (Sealy & Pfeiffer, 2000; Sealy, 2006). Therefore, the coastal lifestyle was not uniform across space or time. Perhaps the combination of a diet lacking nutritional variety and abrasive foodstuffs resulted in heavier dental wear exposing tooth pulp increasing the chances of infection (Sealy & van der Merwe, 1986; Botha & Steyn, 2015). Extreme forms of infection and TMJD were found in five different coastal individuals. Evidently, while the coast provided a steady source of food, it led to other risks. Socioeconomic implications for health post-2000 BP The sociopolitical and environmental landscape in southern Africa changed post-2000 BP when eastern African Bantu populations and later Europeans began to settle in the region, along with the advent of pastoralism among the Khoekhoen people (Deacon, 1984; Smith, 1986; Morris, 2002). Generally, these transitions cause population stress through resource competition, land occupation, subsistence transitions and colonial marginalisation (Smith, 1986: Larsen, 1995; Pfeiffer, 2007). Observed higher frequencies of CO and higher antemortem trauma post-2000 BP indicates levels of accidental or violent activity were higher, but also their ability to heal and live through such injuries and stress. Perimortem trauma was as frequent pre- and post-2000 BP, which provide information on fatal events. This violence is unlikely attributable to the major socioeconomic changes in the region, which is supported by Pfeiffer (2013; Pfeiffer et al., 1999). Interestingly, LSA people were biologically stressed prior to 2000 BP: previously, this has been shown through a decline in femoral length (Pfeiffer and Sealy, 2006) and reduction in body size (Pfeiffer & Sealy, 2006; Ginter, 2011). These conditions improve around 2000 BP, which cannot be attributed to milk and meat consumption offered by herding, but rather suggests these early stress periods as producing better and more flexible cultural and subsistence strategies (Sealy 2006; Ginter, 2011). Therefore, potential sources of conflict existed before 2000 BP as well as after (Morris et al., 1987; Smith, 1986; Pfeiffer et al., 1999) clearly suggesting increased interpersonal violence but less frequently fatal. CONCLUSION This investigation offers a broader perspective on Khoesan health status during the Holocene. For these people modern standards of sexing and ageing are inaccurate and require revision. The prevalence of stress markers shows their ability through immune strength or cultural adaptation to survive to adulthood. The ability to live through stress episodes, evidenced by LEH, CO, and PH especially during childhood, points to individual fitness and resistance. Evidence of trauma shows that interpersonal violence was prevalent, contrasting theories of a ‘harmless people’, and women were at higher risk for fatal injuries. The significant distribution of CO post-2000 BP and in coastal areas may suggest an intermediate level of compromised health during these periods of sociopolitical changes and nutritional strain. Further investigation is required into this. 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Temporal and spatial distribution of 150 Khoesan individuals from the University of Cape Town Human Skeletal collection. This article is protected by copyright. All rights reserved. Figure 2. Classification flow-chart of cranial and dental pathology, as described in materials and methods. This article is protected by copyright. All rights reserved. Figure 3. Cranial pathology observed, with an example of cribra orbitalia in the left orbital of UCT 88 (A); posterior degeneration on mandibular condyle of Tankwa skeleton (B); infection at mandibular condyle of UCT 596 (C); antemortem trauma on vault of UCT 372. 1 cm scale bar. This article is protected by copyright. All rights reserved. Table 1. Cranial and dental pathology of 150 Khoesan individuals with estimated sex, age, spatial and temporal distribution. Statistically significant differences determined by Fisher’s exact tests are marked by an asterisk (*) in bold (p ≤ 0.05). This article is protected by copyright. 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