Stable population structure in Europe since the Iron Age, despite high mobility.

Ancient DNA research in the past decade has revealed that European population structure changed dramatically in the prehistoric period (14,000-3000 years before present, YBP), reflecting the widespread introduction of Neolithic farmer and Bronze Age Steppe ancestries. However, little is known about how population structure changed from the historical period onward (3000 YBP - present). To address this, we collected whole genomes from 204 individuals from Europe and the Mediterranean, many of which are the first historical period genomes from their region (e.g. Armenia and France). We found that most regions show remarkable inter-individual heterogeneity. At least 7% of historical individuals carry ancestry uncommon in the region where they were sampled, some indicating cross-Mediterranean contacts. Despite this high level of mobility, overall population structure across western Eurasia is relatively stable through the historical period up to the present, mirroring geography. We show that, under standard population genetics models with local panmixia, the observed level of dispersal would lead to a collapse of population structure. Persistent population structure thus suggests a lower effective migration rate than indicated by the observed dispersal. We hypothesize that this phenomenon can be explained by extensive transient dispersal arising from drastically improved transportation networks and the Roman Empire's mobilization of people for trade, labor, and military. This work highlights the utility of ancient DNA in elucidating finer scale human population dynamics in recent history.

Computer-aided craniofacial superimposition validation study: the identification of the leaders and participants of the Polish-Lithuanian January Uprising (1863-1864).

In 2017, a series of human remains corresponding to the executed leaders of the "January Uprising" of 1863-1864 were uncovered at the Upper Castle of Vilnius (Lithuania). During the archeological excavations, 14 inhumation pits with the human remains of 21 individuals were found at the site. The subsequent identification process was carried out, including the analysis and cross-comparison of post-mortem data obtained in situ and in the lab with ante-mortem data obtained from historical archives. In parallel, three anthropologists with diverse backgrounds in craniofacial identification and two students without previous experience attempted to identify 11 of these 21 individuals using the craniofacial superimposition technique. To do this, the five participants had access to 18 3D scanned skulls and 14 photographs of 11 different candidates. The participants faced a cross-comparison problem involving 252 skull-face overlay scenarios. The methodology follows the main agreements of the European project MEPROCS and uses the software Skeleton-ID™. Based on MEPROCS standard, a final decision was provided within a scale, assigning a value in terms of strong, moderate, or limited support to the claim that the skull and the facial image belonged (or not) to the same person for each case. The problem of binary classification, positive/negative, with an identification rate for each participant was revealed. The results obtained in this study make the authors think that both the quality of the materials used and the previous experience of the analyst play a fundamental role when reaching conclusions using the CFS technique.

On some paleopathological examples of amputation and the implications for healthcare in 13th-17th century Lithuania.

OBJECTIVE: To explore care that was likely provided to an adult male amputee from medieval Lithuania, positioning analysis within what is known of contemporary amputation practices. MATERIALS: Three sets of skeletal remains with evidence for amputation, dating to between the 13th-17th centuries AD and recovered during different archaeological excavations in Vilnius, Lithuania. METHODS: Macroscopic inspection of lesions, with additional X-ray analysis of the main subject. The Index of Care was used to investigate possible caregiving. RESULTS: Two individuals experienced amputation of a single element, and the third experienced bilateral hand amputation. Only one individual displayed healing. Historic sources suggest use of amputation for punitive purposes during this period, and judicial punishment is proposed as the most likely reason for amputation in at least two cases. CONCLUSIONS: Evidence of long-term healing in one individual suggests receipt of care. This individual likely relied on family and/or community members for survival immediately following amputation, and subsequently for support in managing disability. SIGNIFICANCE: Successfully combining osteology with history in a framework for analyzing care provision in past Eastern European society, this study underlines the critical importance of context in undertaking bioarchaeology of care analyses. It also adds two examples of perimortem abscissions in this region to the paleopathological record. LIMITATIONS: Our approach relied on skeletal interpretation. Soft tissue was lost to decomposition and no relevant archaeological evidence was found in association with the remains. SUGGESTIONS FOR FURTHER RESEARCH: A review of skeletal collections may allow identification of overlooked cases of amputation (and care).

An integrative skeletal and paleogenomic analysis of stature variation suggests relatively reduced health for early European farmers.

Human culture, biology, and health were shaped dramatically by the onset of agriculture ∼12,000 y B.P. This shift is hypothesized to have resulted in increased individual fitness and population growth as evidenced by archaeological and population genomic data alongside a decline in physiological health as inferred from skeletal remains. Here, we consider osteological and ancient DNA data from the same prehistoric individuals to study human stature variation as a proxy for health across a transition to agriculture. Specifically, we compared "predicted" genetic contributions to height from paleogenomic data and "achieved" adult osteological height estimated from long bone measurements for 167 individuals across Europe spanning the Upper Paleolithic to Iron Age (∼38,000 to 2,400 B.P.). We found that individuals from the Neolithic were shorter than expected (given their individual polygenic height scores) by an average of −3.82 cm relative to individuals from the Upper Paleolithic and Mesolithic (P = 0.040) and −2.21 cm shorter relative to post-Neolithic individuals (P = 0.068), with osteological vs. expected stature steadily increasing across the Copper (+1.95 cm relative to the Neolithic), Bronze (+2.70 cm), and Iron (+3.27 cm) Ages. These results were attenuated when we additionally accounted for genome-wide genetic ancestry variation: for example, with Neolithic individuals −2.82 cm shorter than expected on average relative to pre-Neolithic individuals (P = 0.120). We also incorporated observations of paleopathological indicators of nonspecific stress that can persist from childhood to adulthood in skeletal remains into our model. Overall, our work highlights the potential of integrating disparate datasets to explore proxies of health in prehistory.

A possible case of paralysis in early modern Vilnius and the implications for social care.

OBJECTIVE: To present a case of possible paralysis from early modern Vilnius and to discuss the potential level of care that was provided in the society of that time. MATERIALS: A partially disturbed skeleton of a young female from a 16th-17th century Orthodox Christian cemetery. METHODS: Macroscopic, osteometric and X-ray examinations coupled with a literature review aimed at providing a differential diagnosis. RESULTS: The skeletal remains showed signs of disuse atrophy most probably due to a neurological disorder acquired in the woman's late teens. Differentials suggest that the observed limb atrophy was most likely a consequence of poliomyelitis. CONCLUSIONS: The case of a young female with paralysis presented in this paper could serve as an example of care provided by her household. SIGNIFICANCE: This study substantially contributes to further understanding of the nature and quality of care provided to disabled individuals in their households even in the absence of written sources. LIMITATIONS: There is a degree of diagnostic ambiguity due to the application of routine clinical criteria to paleopathological cases. SUGGESTIONS FOR FURTHER RESEARCH: The article makes several recommendations for future research, e.g., systematic investigation of possible cases of bone atrophy in a broader sociocultural context, as well as searching for evidence of gastrointestinal infections, especially poliomyelitis, supplemented by the application of biomolecular technologies.

Insights into biogenic and diagenetic lead exposure in experimentally altered modern and archaeological bone: Synchrotron radiation X-ray fluorescence imaging.

Bones represent a valuable biological archive of environmental lead (Pb) exposure for modern and archaeological populations. Synchrotron radiation X-ray fluorescence imaging (SR-XFI) generates maps of Pb in bone on a microstructural scale, potentially providing insights into an individual's history of Pb exposure and, in the context of archaeological bone, the biogenic or diagenetic nature of its uptake. The aims of this study were to (1) examine biogenic spatial patterns for Pb from bone samples of modern cadavers compared with patterns observed archaeologically, and (2) test the hypothesis that there are spatial differences in the distribution of Pb for diagenetic and biogenic modes of uptake in bone. To address these aims, this study used inductively coupled plasma-mass spectrometry (ICP-MS) and SR-XFI on unaltered and experimentally altered cadaveric bone samples (University of Saskatchewan, Saskatoon, SK) and archaeological bone samples from 18th to 19th century archaeological sites from Antigua and Lithuania. Bone concentrations of modern individuals are relatively low compared to those of archaeological individuals. SR-XFI results provide insights into modern Saskatchewan Pb exposure with some samples demonstrating a pattern of relatively low Pb exposure with higher levels of Pb exposure occurring in bone structures of a relatively older age that formed earlier in life, likely during the era of leaded gasoline (pre-1980s), and other samples demonstrating a pattern of fairly consistent, low-level exposure. Results support hypotheses for the spatial distribution of Pb corresponding to biogenic vs. diagenetic uptake. Diagenetic Pb is mainly confined to the periosteal surface of each sample with some enrichment of cracks and sub-periosteal canals. This may be useful in the future for differentiating diagenetic from biogenic Pb accumulation, analyzing environmental contamination, and informing sampling strategies in archaeological or fossil bone.

An integrative approach to studying plasticity in growth disruption and outcomes: A bioarchaeological case study of Napoleonic soldiers.

OBJECTIVES: The aim of this study was to investigate how much variation in adult stature and body mass can be explained by growth disruption among soldiers who served in Napoleon's Grand Army during the Russian Campaign of 1812. METHODS: Linear enamel hypoplasia (LEH) were recorded as representations of early life growth disruption, while the impact on future growth was assessed using maximum femur length (n = 73) as a proxy for stature and maximum femoral head diameter (n = 25) as a proxy for body mass. LEH frequency, severity, age at first formation, and age at last formation served as explanatory variables in a multiple regression analysis to test the effect of these variables on maximum femur length and maximum femoral head diameter. RESULTS: The multiple regression model produced statistically significant results for maximum femur length (F-statistic = 3.05, df = 5 and 67, P = .02), with some variation in stature (adjusted r2 = 0.13) attributable to variation in growth disruption. The multiple regression model for maximum femoral head diameter was not statistically significant (F-statistic = 1.87, df = 5 and 19, P = .15). CONCLUSIONS: We hypothesized stress events during early life growth and development would have significant, negative, and cumulative effects on growth outcomes in adulthood. The results did not support our hypothesis. Instead, some variables and interactions had negative effects on stature, whereas others had positive effects. This is likely due to catch-up growth, the relationship between acute and chronic stress and growth, resilience, and plasticity in human growth over the life course.

Diet patterns in medieval to early modern (14th-early 20th c.) coastal communities in Lithuania.

Coastal residents are quite often expected to consume a significant amount of aquatic resources, though historical evidence often reveals a rather complex diet. To better understand the actual consumption and the distribution of various foods, stable isotope (δ13C and δ15N) analyses were employed to skeletal remains from three coastal communities, Palanga, Kretinga and Smelte, ranging in date from the medieval period to the early modern ages (14th-early 20th c.) near the Curonian Lagoon and the Baltic Sea in Lithuania. Animal bones from the region, covering the same time periods, were also analysed. Stable isotope results were compared with historical records. According to historical sources different types of diet were prevalent during that period of time: Medieval Prussian-Lithuanian peasant, Lithuanian fisherman, German urban, and religious-based. Elevated δ15N values for Smelte samples suggest a diet consisting of considerable amounts of freshwater fish protein, which is in contrast to historical sources. There were no significant differences in stable isotope values between males and females, while subadult δ15N values were significantly higher than adult ones, indicating that those children were breastfed for an extended period. Meanwhile, Palanga and Kretinga samples had isotope values suggesting a high reliance on terrestrial resources and a peasant type of diet.

A treponemal genome from an historic plague victim supports a recent emergence of yaws and its presence in 15th century Europe.

Developments in techniques for identification of pathogen DNA in archaeological samples can expand our resolution of disease detection. Our application of a non-targeted molecular screening tool for the parallel detection of pathogens in historical plague victims from post-medieval Lithuania revealed the presence of more than one active disease in one individual. In addition to Yersinia pestis, we detected and genomically characterized a septic infection of Treponema pallidum pertenue, a subtype of the treponemal disease family recognised as the cause of the tropical disease yaws. Our finding in northern Europe of a disease that is currently restricted to equatorial regions is interpreted within an historical framework of intercontinental trade and potential disease movements. Through this we offer an alternative hypothesis for the history and evolution of the treponemal diseases, and posit that yaws be considered an important contributor to the sudden epidemic of late 15th century Europe that is widely ascribed to syphilis.

Stable carbon and nitrogen isotope inter- and intra-individual dietary reconstruction from the late 14th to early 18th century site of Alytus, Lithuania.

OBJECTIVES: The Baltic region, particularly Lithuania, was a politically vibrant area of Eastern Europe during the medieval and early modern period. To better understand the diet of Lithuanians during the late 14th to early 18th century, we examine stable carbon and nitrogen isotope ratios from bone and dentin samples from the site of Alytus. We investigate possible dietary differences based on sex, age, and religious practice, as well as dietary changes throughout an individual's lifetime, within the broader European milieu. MATERIALS AND METHODS: Stable carbon and nitrogen isotope analysis was conducted on bone (n = 35) and dentin (n = 38) collagen samples from a total of 39 individuals buried in the cemetery at Alytus (late 14th to early 18th centuries). RESULTS: Results indicate individuals at Alytus consumed a C3 terrestrial based diet. The δ13 C and δ15 N values are not significantly different between bone and dentin, and did not vary by sex. DISCUSSION: The diet at Alytus was homogeneous between males and females and between tissue types. The lack of evidence indicating substantial consumption of fish is unexpected given widespread Catholic fasting practices and marine resource trade throughout Europe. Comparisons with other populations indicate that individuals from Alytus differ in diet from contemporaneous Polish-Lithuanian Commonwealth elites. Comparison of the diets of non-elite individuals in the Eastern Baltic region also reveals dietary variability.

A rare case of child abuse from early modern Lithuania.

The aim of this article was to describe the outcome of a probable case of physical abuse in the remains of a child dating from the 16th-18th centuries CE. The skeleton of a subadult was recovered during archaeological excavations carried out in the village of Uzubaliai, located in Alytus in southern Lithuania, and subsequently curated in the Faculty of Medicine at Vilnius University. The bones of this child were observed macroscopically and then submitted for radiological investigation. In order to speculate on the presence of abuse, features such as the presence, quantity, and type of injuries were considered. Stages of healing and the occurrence of additional nonspecific stress markers, such as linear enamel hypoplasia or Harris lines, were also recorded. The remains revealed the presence of lesions showing three stages of healing, including antemortem and perimortem fractures. Periosteal reactions were also observed on many of the bones. Traumas with high specificity for abuse, such as rib and scapular fractures, were assessed. Finally, endocranial new bone formation was also noted as a possible sign of neglect. The pattern of observed injuries showed evidence that was compatible with a case of physical abuse.

Four cases of beheading from 14th-17th century Lithuania.

ABSTRACT: Skeletal evidence of beheading in early modern Lithuania has been scarce, despite historical documentation indicating it as a popular practice. This study presents the first bioarchaeological cases of decapitation in early modern Lithuania, with four adult male individuals from the 14th-17th centuries A.D. that reveal evidence of perimortem lesions in the cervical and thoracic vertebrae. The osteological and radiological study of the affected bones suggests that the individuals might have been victims of episodes of interpersonal violence, rather than subjected to capital punishment.

Author Correction: The genetic prehistory of the Baltic Sea region.

The original version of this Article omitted references to previous work, which are detailed in the associated Author Correction. These omissions have been corrected in both the PDF and HTML versions of the Article.

Eastern Baltic region vs. Western Europe: modelling age related changes in the pubic symphysis and the auricular surface.

ABSTRACT: Objectives. The present study addresses the following two main questions: a) Is the pattern of skeletal ageing observed in well-known western European reference collections applicable to modern eastern Baltic populations, or are population-specific standards needed? b) What are the consequences for estimating the age-at-death distribution in the target population when differences in the estimates from reference data are not taken into account? Materials and methods. The dataset consists of a modern Lithuanian osteological reference collection, which is the only collection of this type in the eastern Baltic countries (n = 381); and two major western European reference collections, Coimbra (n = 264) and Spitalfields (n = 239). The age-related changes were evaluated using the scoring systems of Suchey-Brooks (Brooks & Suchey 1990) and Lovejoy et al. (1985), and were modelled via regression models for multinomial responses. A controlled experiment based on simulations and the Rostock Manifesto estimation protocol (Wood et al. 2002) was then carried out to assess the effect of using estimates from different reference samples and different regression models on estimates of the age-at-death distribution in the hypothetical target population. Results. The following key results were obtained in this study. a) The morphological alterations in the pubic symphysis were much faster among women than among men at comparable ages in all three reference samples. In contrast, we found no strong evidence in any of the reference samples that sex is an important factor to explain rate of changes in the auricular surface. b) The rate of ageing in the pubic symphysis seems to be similar across the three reference samples, but there is little evidence of a similar pattern in the auricular surface. That is, the estimated rate of age-related changes in the auricular surface was much faster in the LORC and the Coimbra samples than in the Spitalfields sample. c) The results of simulations showed that the differences in the estimates from the reference data result in noticeably different age-at-death distributions in the target population. Thus, a degree bias may be expected if estimates from the western European reference data are used to collect information on ages at death in the eastern Baltic region based on the changes in the auricular surface. d) Moreover, the bias is expected to be more pronounced if the fitted regression model improperly describes the reference data. Conclusions. Differences in the timing of age-related changes in skeletal traits are to be expected among European reference samples, and cannot be ignored when seeking to reliably estimate an age-at-death distribution in the target population. This form of bias should be taken into consideration in further studies of skeletal samples from the eastern Baltic region.

The genetic prehistory of the Baltic Sea region.

While the series of events that shaped the transition between foraging societies and food producers are well described for Central and Southern Europe, genetic evidence from Northern Europe surrounding the Baltic Sea is still sparse. Here, we report genome-wide DNA data from 38 ancient North Europeans ranging from ~9500 to 2200 years before present. Our analysis provides genetic evidence that hunter-gatherers settled Scandinavia via two routes. We reveal that the first Scandinavian farmers derive their ancestry from Anatolia 1000 years earlier than previously demonstrated. The range of Mesolithic Western hunter-gatherers extended to the east of the Baltic Sea, where these populations persisted without gene-flow from Central European farmers during the Early and Middle Neolithic. The arrival of steppe pastoralists in the Late Neolithic introduced a major shift in economy and mediated the spread of a new ancestry associated with the Corded Ware Complex in Northern Europe.

The Stone Age Plague and Its Persistence in Eurasia.

Yersinia pestis, the etiologic agent of plague, is a bacterium associated with wild rodents and their fleas. Historically it was responsible for three pandemics: the Plague of Justinian in the 6th century AD, which persisted until the 8th century [1]; the renowned Black Death of the 14th century [2, 3], with recurrent outbreaks until the 18th century [4]; and the most recent 19th century pandemic, in which Y. pestis spread worldwide [5] and became endemic in several regions [6]. The discovery of molecular signatures of Y. pestis in prehistoric Eurasian individuals and two genomes from Southern Siberia suggest that Y. pestis caused some form of disease in humans prior to the first historically documented pandemic [7]. Here, we present six new European Y. pestis genomes spanning the Late Neolithic to the Bronze Age (LNBA; 4,800 to 3,700 calibrated years before present). This time period is characterized by major transformative cultural and social changes that led to cross-European networks of contact and exchange [8, 9]. We show that all known LNBA strains form a single putatively extinct clade in the Y. pestis phylogeny. Interpreting our data within the context of recent ancient human genomic evidence that suggests an increase in human mobility during the LNBA, we propose a possible scenario for the early spread of Y. pestis: the pathogen may have entered Europe from Central Eurasia following an expansion of people from the steppe, persisted within Europe until the mid-Bronze Age, and moved back toward Central Eurasia in parallel with human populations.

Checklist and Scoring System for the Assessment of Soft Tissue Preservation in CT Examinations of Human Mummies: Application to the Tyrolean Iceman.

Purpose Soft tissues make a skeleton into a mummy and they allow for a diagnosis beyond osteology. Following the approach of structured reporting in clinical radiology, a recently developed checklist was used to evaluate the soft tissue preservation status of the Tyrolean Iceman using computed tomography (CT). The purpose of this study was to apply the "Checklist and Scoring System for the Assessment of Soft Tissue Preservation in CT Examinations of Human Mummies" to the Tyrolean Iceman, and to compare the Iceman's soft tissue preservation score to the scores calculated for other mummies. Materials and Methods A whole-body (CT) (SOMATOM Definition Flash, Siemens, Forchheim, Germany) consisting of five scans, performed in January 2013 in the Department of Radiodiagnostics, Central Hospital, Bolzano, was used (slice thickness 0.6 mm; kilovolt ranging from 80 to 140). For standardized evaluation the "CT Checklist and Scoring System for the Assessment of Soft Tissue Preservation in Human Mummies" was used. Results All checkpoints under category "A. Soft Tissues of Head and Musculoskeletal System" and more than half in category "B. Organs and Organ Systems" were observed. The scoring system accounted for a total score of 153 (out of 200). The comparison of the scores between the Iceman and three mummy collections from Vilnius, Lithuania, and Palermo, Sicily, as well as one Egyptian mummy resulted in overall higher soft tissue preservation scores for the Iceman. Conclusion Application of the checklist allowed for standardized assessment and documentation of the Iceman's soft tissue preservation status. The scoring system allowed for a quantitative comparison between the Iceman and other mummies. The Iceman showed remarkable soft tissue preservation. Key Points · The approach of structured reporting can be transferred to paleoradiology.. · The checklist allowed for standardized soft tissue assessment and documentation.. · The scoring system facilitated a quantitative comparison among mummies.. · Based on CT, the Tyrolean Iceman demonstrated remarkable soft tissue preservation.. Citation Format · Panzer S, Pernter P, Piombino-Mascali D et al. Checklist and Scoring System for the Assessment of Soft Tissue Preservation in CT Examinations of Human Mummies: Application to the Tyrolean Iceman. Fortschr Röntgenstr 2017; 189: 1152 - 1160.

Reconstructing diet in Napoleon's Grand Army using stable carbon and nitrogen isotope analysis.

OBJECTIVES: Historical evidence has provided information regarding disease and mortality in Napoleon Bonaparte's Grand Army, but dietary information beyond individual soldier accounts remains scarce. The purpose of this research is to reconstruct the diets of Napoleon's multiethnic army who were associated with the Russian Campaign of 1812. MATERIALS AND METHODS: We conducted stable carbon and nitrogen isotope ratio analysis on femoral bone collagen of 78 individuals recovered from a salvage excavation at the mass gravesite of Siaures miestelis in Vilnius, Lithuania. These individuals were later discovered to be Napoleonic soldiers and camp followers who participated in the 1812 Russian Campaign. RESULTS: Stable carbon isotope ratios range from -19.2‰ to -11.8‰, with a mean of -17.8‰ ± 1.5‰ (1 σ). Stable nitrogen isotope ratios range from 7.1‰ to 13.6‰, with a mean of 10.5‰ ± 1.4‰ (1 σ). Both δ13 C and δ15 N values show a wide range of variation. DISCUSSION: Stable isotope data indicate considerable dietary variation in this population associated with a multiethnic and socially stratified military population. Diets ranged from predominantly C3 -based to predominantly C4 -based, with varying inputs of terrestrial, freshwater, and marine animal protein. Comparison with other European populations further denotes the exceptional range of dietary variation of soldiers and camp followers in Napoleon's army.

Paleoimaging of a modern mummy from Lithuania (circa 19th-20th century).

An anthropogenic human mummy curated in the Museum of the History of Medicine, Vilnius University, was recently examined by means of computed tomography. Although the mummy lacked data regarding its specific context and historical information on its identity and chronology, the investigation focused on the embalming method adopted to preserve it. Some pathological alterations were also recorded. This research appears to suggest that this body was prepared for educational and/or scientific purposes rather than funerary purposes. Hence, the case could be categorized as a "medical mummy" prepared between the mid-19th and the mid-20th centuries.

17th Century Variola Virus Reveals the Recent History of Smallpox.

Smallpox holds a unique position in the history of medicine. It was the first disease for which a vaccine was developed and remains the only human disease eradicated by vaccination. Although there have been claims of smallpox in Egypt, India, and China dating back millennia [1-4], the timescale of emergence of the causative agent, variola virus (VARV), and how it evolved in the context of increasingly widespread immunization, have proven controversial [4-9]. In particular, some molecular-clock-based studies have suggested that key events in VARV evolution only occurred during the last two centuries [4-6] and hence in apparent conflict with anecdotal historical reports, although it is difficult to distinguish smallpox from other pustular rashes by description alone. To address these issues, we captured, sequenced, and reconstructed a draft genome of an ancient strain of VARV, sampled from a Lithuanian child mummy dating between 1643 and 1665 and close to the time of several documented European epidemics [1, 2, 10]. When compared to vaccinia virus, this archival strain contained the same pattern of gene degradation as 20th century VARVs, indicating that such loss of gene function had occurred before ca. 1650. Strikingly, the mummy sequence fell basal to all currently sequenced strains of VARV on phylogenetic trees. Molecular-clock analyses revealed a strong clock-like structure and that the timescale of smallpox evolution is more recent than often supposed, with the diversification of major viral lineages only occurring within the 18th and 19th centuries, concomitant with the development of modern vaccination.