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Supplementary Information For Supplementary Information for An Abundance of Developmental Anomalies and Abnormalities in Pleistocene People Erik Trinkaus Department of Anthropology, Washington University, Saint Louis MO 63130 Corresponding author: Erik Trinkaus Email: [email protected] This PDF file includes: Supplementary text Figures S1 to S57 Table S1 References 1 to 421 for SI reference citations Introduction Although they have been considered to be an inconvenience for the morphological analysis of human paleontological remains, it has become appreciated that various pathological lesions and other abnormalities or rare variants in human fossil remains might provide insights into Pleistocene human biology and behavior (following similar trends in Holocene bioarcheology). In this context, even though there were earlier paleopathological assessments in monographic treatments of human remains (e.g., 1-3), it has become common to provide details on abnormalities in primary descriptions of human fossils (e.g., 4-12), as well as assessments of specific lesions on known and novel remains [see references in Wu et al. (13, 14) and below]. These works have been joined by doctoral dissertation assessments of patterns of Pleistocene human lesions (e.g., 15-18). The paleopathological attention has been primarily on the documentation and differential diagnosis of the abnormalities of individual fossil remains, leading to the growing paleopathological literature on Pleistocene specimens and their lesions. There have been some considerations of the overall patterns of the lesions, but those assessments have been concerned primarily with non-specific stress indicators and traumatic lesions (e.g., 13, 15, 19-21), with variable considerations of issues of survival 1 w ww.pnas.org/cgi/doi/10.1073/pnas.1814989115 and especially the inferred social support of the afflicted (e.g., 22-27). At the same time, it has been noted that there appears to be an elevated level of abnormalities among Pleistocene humans, both ones that are clearly pathological and others that are either rare variants or of unknown etiology (14, 28). The apparently high frequency of these abnormalities has been linked to differential mortuary behavior in the Upper Paleolithic (8, 29) and to possibly high levels of consanguinity among Pleistocene humans (14, 30- 32). These assessments of overall patterns have tended to combine abnormalities that are developmental versus degenerative, of clear etiology versus unknown cause, of functional significance versus probably unknown to the individual involved, and/or from associated skeletons versus isolated remains. In this context, and especially with the steadily increasing paleopathological record for Pleistocene humans, it is appropriate to reconsider one general class of abnormalities among these past foraging populations – those that derived from irregularities of developmental processes (other than non- specific stress indicators, which are primarily reflected in the ubiquitous dental enamel hypoplasias). In general, these abnormalities can be divided into those that were systemic versus anatomically localized, to the extent that the distinction can be assessed on incomplete paleontological remains. They can be separated into those for which at least a proximate etiology is evident and those currently of unknown cause, the former being separable into ones whose ultimate etiology is evident versus those which may have multiple causes. The ones that are evidently pathological can be separated from those that appear as unusual and rare variants on “normal” developmental processes, the latter limited to ones that are infrequently found in large recent human samples (and hence would be unexpected in the small available fossils sample sizes for the given element, especially if those samples are limited in time and space) or exceptional in terms of size and/or proportions for the relevant paleontological sample. Given these considerations, following is an annotated review of the currently documented developmental abnormalities among Pleistocene humans, excluding isolated non-specific stress indicators. It is likely that it represents a minimum inventory, given the recent (primarily since the 1990s) documentation and/or reassessment of a substantial number of abnormalities, a number of them noticed only recently on long known specimens. They are grouped in regional anatomical terms, from systemic ones through cranial and mandibular, dental, vertebral, shoulder and arm, pelvis and leg, and hand and foot ones. Within each anatomical group, they are arranged largely alphabetically by sites, independent of the time period/morphological group of the specimen(s) involved, with a few grouped by the form of the anomaly. Materials and Methods Almost all of the abnormalities presented here have been described in varying levels of detail and many subjected to differential diagnosis. The lesions or variants are therefore summarized, with appropriate references, although the interpretations here do not universally follow those of the original assessments. The descriptions and interpretations are based on the published details, supplemented when possible by personal assessments. Images of the primary abnormalities are provided, as available through personal research and the generosity of colleagues; published figures are referenced for specimens for which images are not available. As appropriate for issues of body, cranial or dental size and proportions, measurements and indices of the remains are compared to relevant samples of Pleistocene human remains. All of the measurements follow the Martin system (33) or Howells (34). Percent asymmetry is calculated as [(maximum – minimum)/minimum] (35). Within the context of anatomical regions, the abnormalities are primarily presented by individual. Eight specimens (Arene Candide 2, Dolní Věstonice 15 and 16, Kebara 2, Shanidar 3 and 4, Sunghir 2, and Villabruna 1) appear more than once, given more than one apparently independent anomalies. Two additional specimens (Lazaret 18/19 and Pech-de-l’Azé 1) have two abnormalities each, presented together. 2 The specimens are limited to Pleistocene members of the genus Homo, not including initial Pleistocene remains variably included in Homo versus Australopithecus (see 36). In addition, the Middle and Late Pleistocene sample from Liang Bua and Mata Menge, Flores, Indonesia (37-39) and the Middle Pleistocene one from Dinaledi and Lesedi, South Africa (40, 41) are not included. The status of each sample relative to other Pleistocene Homo fossils is unresolved, especially since each sample exhibits morphological features that cannot be easily explained phylogenetically in a broader hominid context (42, 43). Many of the abnormalities can be referred to clinically known conditions, even if the precise natures of the defects or dysplasias are not known. Several of the variants are unique or at least not documented in the clinical or paleopathological literature, and care has been taken to assure that they are abnormalities and not merely morphological variants characteristic of Pleistocene populations. When the anomalies are clinically known and data are available on their incidence in sufficiently large recent human samples, summaries of those data are included. However, it remains unclear whether the incidences in recent human samples are directly applicable to Pleistocene specimens or samples. Many of the variants with known incidences in recent human samples have unknown ultimate etiologies or can result through diverse etiologies; the relevant clinical literature is replete with vague etiologies that invoke genetic causes (without specific variants), inherited predispositions, and/or forms of stress during development. Moreover, the susceptibilities among Pleistocene people may have been different, given contrasting levels of systemic stress, skeletal hypertrophy and/or craniofacial or body proportions. The recent human data are often the only reference available and are therefore employed as available. As a result, the abnormalities for which there are available data, whether they are systemic or individual, are ascribed general levels of probability (<5.0%, <1.0%, <0.1% and <0.01%) based on the incidence data from recent human samples (Table 1). In the few cases in which the unusual aspect is related to size or proportions, the associated probabilities are based on relevant human paleontological comparative distributions, determined from the general paleontological group from which the specimen derives. Ideally, these probabilities would be adjusted for the number of paleontological specimens available and preserving the relevant portions of the skeleton or dentition. However, it is unclear how the reference samples for such adjustments should be delimited (by site, by time period, by geography, by taxonomic group, etc.?). In most cases, the adjustment would not change the assignment to one of the levels of probability, given the modest sample sizes available for most relevant Pleistocene groups. Systemic Conditions Arene Candide 3 The final Late Upper Paleolithic (Epigravettian, MIS 2) site of Arene Candide in northwestern Italy has yielded ≈20 individuals, two-thirds of which derive from primary burials (32, 44, 45). The Arene Candide 3 partial skeleton retains the cranium and mandible, the L5, the complete pelvis, the right and left humeri, ulnae and fibulae, and one each of the scapulae, radii, femora and tibiae; additional costal, manual and pedal remains are mixed with those of
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