Teeth, prenatal growth rates, and the evolution of human-like pregnancy in later Homo.

Evidence of how gestational parameters evolved is essential to understanding this fundamental stage of human life. Until now, these data seemed elusive given the skeletal bias of the fossil record. We demonstrate that dentition provides a window into the life of neonates. Teeth begin to form in utero and are intimately associated with gestational development. We measured the molar dentition for 608 catarrhine primates and collected data on prenatal growth rate (PGR) and endocranial volume (ECV) for 19 primate genera from the literature. We found that PGR and ECV are highly correlated (R2 = 0.93, P < 0.001). Additionally, we demonstrated that molar proportions are significantly correlated with PGR (P = 0.004) and log-transformed ECV (P = 0.001). From these correlations, we developed two methods for reconstructing PGR in the fossil record, one using ECV and one using molar proportions. Dental proportions reconstruct hominid ECV (R2 = 0.81, P < 0.001), a result that can be extrapolated to PGR. As teeth dominate fossil assemblages, our findings greatly expand our ability to investigate life history in the fossil record. Fossil ECVs and dental measurements from 13 hominid species both support significantly increasing PGR throughout the terminal Miocene and Plio-Pleistocene, reflecting known evolutionary changes. Together with pelvic and endocranial morphology, reconstructed PGRs indicate the need for increasing maternal energetics during pregnancy over the last 6 million years, reaching a human-like PGR (i.e., more similar to humans than to other extant apes) and ECV in later Homo less than 1 million years ago.

Uncovering the mosaic evolution of the carnivoran skeletal system.

The diversity of vertebrate skeletons is often attributed to adaptations to distinct ecological factors such as diet, locomotion, and sensory environment. Although the adaptive evolution of skull, appendicular skeleton, and vertebral column is well studied in vertebrates, comprehensive investigations of all skeletal components simultaneously are rarely performed. Consequently, we know little of how modes of evolution differ among skeletal components. Here, we tested if ecological and phylogenetic effects led to distinct modes of evolution among the cranial, appendicular and vertebral regions in extant carnivoran skeletons. Using multivariate evolutionary models, we found mosaic evolution in which only the mandible, hindlimb and posterior (i.e. last thoracic and lumbar) vertebrae showed evidence of adaptation towards ecological regimes whereas the remaining skeletal components reflect clade-specific evolutionary shifts. We hypothesize that the decoupled evolution of individual skeletal components may have led to the origination of distinct adaptive zones and morphologies among extant carnivoran families that reflect phylogenetic hierarchies. Overall, our work highlights the importance of examining multiple skeletal components simultaneously in ecomorphological analyses. Ongoing work integrating the fossil and palaeoenvironmental record will further clarify deep-time drivers that govern the carnivoran diversity we see today and reveal the complexity of evolutionary processes in multicomponent systems.

Beringia and the peopling of the Western Hemisphere.

Did Beringian environments represent an ecological barrier to humans until less than 15 000 years ago or was access to the Americas controlled by the spatial-temporal distribution of North American ice sheets? Beringian environments varied with respect to climate and biota, especially in the two major areas of exposed continental shelf. The East Siberian Arctic Shelf ('Great Arctic Plain' (GAP)) supported a dry steppe-tundra biome inhabited by a diverse large-mammal community, while the southern Bering-Chukchi Platform ('Bering Land Bridge' (BLB)) supported mesic tundra and probably a lower large-mammal biomass. A human population with west Eurasian roots occupied the GAP before the Last Glacial Maximum (LGM) and may have accessed mid-latitude North America via an interior ice-free corridor. Re-opening of the corridor less than 14 000 years ago indicates that the primary ancestors of living First Peoples, who already had spread widely in the Americas at this time, probably dispersed from the NW Pacific coast. A genetic 'arctic signal' in non-arctic First Peoples suggests that their parent population inhabited the GAP during the LGM, before their split from the former. We infer a shift from GAP terrestrial to a subarctic maritime economy on the southern BLB coast before dispersal in the Americas from the NW Pacific coast.

Evolutionary adaptation highlights the interconnection of fatty acids, sunlight, inflammation and epithelial adhesion.

Gene variants that influence human biology today reflect thousands of years of evolution. Genetic effects on infant health are a major point of selective pressure, given that childhood survival is essential to evolutionary success. Knowledge of this evolutionary history can have implications for paediatric research. CONCLUSION: An episode of human adaptation to the extremely low ultraviolet radiation environment of the Arctic 20,000 years ago implicates the Ectodysplasin A Receptor (EDAR) and the Fatty Acid Desaturases (FADS) in human lactation and epithelial inflammation.

Effects of hybridization on pelvic morphology: A macaque model.

Ancient DNA analyses have shown that interbreeding between hominin taxa occurred multiple times. Although admixture is often reflected in skeletal phenotype, the relationship between the two remains poorly understood, hampering interpretation of the hominin fossil record. Direct study of this relationship is often impossible due to the paucity of hominin fossils and difficulties retrieving ancient genetic material. Here, we use a sample of known ancestry hybrids between two closely related nonhuman primate taxa (Indian and Chinese Macaca mulatta) to investigate the effect of admixture on skeletal morphology. We focus on pelvic shape, which has potential fitness implications in hybrids, as mismatches between maternal pelvic and fetal cranial morphology are often fatal to mother and offspring. As the pelvis is also one of the skeletal regions that differs most between Homo sapiens and Neanderthals, investigating the pelvic consequences of interbreeding could be informative regarding the viability of their hybrids. We find that the effect of admixture in M. mulatta is small and proportional to the relatively small morphological difference between the parent taxa. Sexual dimorphism appears to be the main determinant of pelvic shape in M. mulatta. The lack of difference in pelvic shape between Chinese and Indian M. mulatta is in contrast to that between Neanderthals and H. sapiens, despite a similar split time (in generations) between the hybridizing pairs. Greater phenotypic divergence between hominins may relate to adaptations to disparate environments but may also highlight how the unique degree of cultural buffering in hominins allowed for greater neutral divergence. In contrast to some previous work identifying extreme morphologies in first- and second-generation hybrids, here the relationship between pelvic shape and admixture is linear. This linearity may be because most sampled animals have a multigenerational admixture history or because of relatively high constraints on the pelvis compared with other skeletal regions.

Environmental selection during the last ice age on the mother-to-infant transmission of vitamin D and fatty acids through breast milk.

Because of the ubiquitous adaptability of our material culture, some human populations have occupied extreme environments that intensified selection on existing genomic variation. By 32,000 years ago, people were living in Arctic Beringia, and during the Last Glacial Maximum (LGM; 28,000-18,000 y ago), they likely persisted in the Beringian refugium. Such high latitudes provide only very low levels of UV radiation, and can thereby lead to dangerously low levels of biosynthesized vitamin D. The physiological effects of vitamin D deficiency range from reduced dietary absorption of calcium to a compromised immune system and modified adipose tissue function. The ectodysplasin A receptor (EDAR) gene has a range of pleiotropic effects, including sweat gland density, incisor shoveling, and mammary gland ductal branching. The frequency of the human-specific EDAR V370A allele appears to be uniquely elevated in North and East Asian and New World populations due to a bout of positive selection likely to have occurred circa 20,000 y ago. The dental pleiotropic effects of this allele suggest an even higher occurrence among indigenous people in the Western Hemisphere before European colonization. We hypothesize that selection on EDAR V370A occurred in the Beringian refugium because it increases mammary ductal branching, and thereby may amplify the transfer of critical nutrients in vitamin D-deficient conditions to infants via mothers' milk. This hypothesized selective context for EDAR V370A was likely intertwined with selection on the fatty acid desaturase (FADS) gene cluster because it is known to modulate lipid profiles transmitted to milk from a vitamin D-rich diet high in omega-3 fatty acids.

A genotype:phenotype approach to testing taxonomic hypotheses in hominids.

Paleontology has long relied on assumptions about the genetic and developmental influences on skeletal variation. The last few decades of developmental genetics have elucidated the genetic pathways involved in making teeth and patterning the dentition. Quantitative genetic analyses have refined this genotype:phenotype map even more, especially for primates. We now have the ability to define dental traits with a fair degree of fidelity to the underlying genetic architecture; for example, the molar module component (MMC) and the premolar-molar module (PMM) that have been defined through quantitative genetic analyses. We leverage an extensive dataset of extant and extinct hominoid dental variation to explore how these two genetically patterned phenotypes have evolved through time. We assess MMC and PMM to test the hypothesis that these two traits reveal a more biologically informed taxonomy at the genus and species levels than do more traditional measurements. Our results indicate that MMC values for hominids fall into two categories and that Homo is derived compared with earlier taxa. We find a more variable, species-level pattern for PMM. These results, in combination with previous research, demonstrate that MMC reflects the phenotypic output of a more evolutionarily stable, or phylogenetically congruent, genetic mechanism, and PMM is a reflection of a more evolutionarily labile mechanism. These results suggest that the human lineage since the split with chimpanzees may not represent as much genus-level variation as has been inferred from traits whose etiologies are not understood.

The integration of quantitative genetics, paleontology, and neontology reveals genetic underpinnings of primate dental evolution.

Developmental genetics research on mice provides a relatively sound understanding of the genes necessary and sufficient to make mammalian teeth. However, mouse dentitions are highly derived compared with human dentitions, complicating the application of these insights to human biology. We used quantitative genetic analyses of data from living nonhuman primates and extensive osteological and paleontological collections to refine our assessment of dental phenotypes so that they better represent how the underlying genetic mechanisms actually influence anatomical variation. We identify ratios that better characterize the output of two dental genetic patterning mechanisms for primate dentitions. These two newly defined phenotypes are heritable with no measurable pleiotropic effects. When we consider how these two phenotypes vary across neontological and paleontological datasets, we find that the major Middle Miocene taxonomic shift in primate diversity is characterized by a shift in these two genetic outputs. Our results build on the mouse model by combining quantitative genetics and paleontology, and thereby elucidate how genetic mechanisms likely underlie major events in primate evolution.

Breaking the rules: Phylogeny, not life history, explains dental eruption sequence in primates.

OBJECTIVES: Although a great deal is known about the biology of tooth development and eruption, there remains disagreement about the factors driving the evolution of dental eruption sequence. We assessed postcanine eruption sequence across a large sample of primates to test two hypotheses: (1) Dental eruption sequence is significantly correlated with life history and body size variables that capture postnatal growth and longevity (Schultz's Rule), and (2) Dental eruption sequence is conserved phylogenetically. MATERIALS AND METHODS: We assessed postcanine dental eruption sequence for 194 individuals representing 21 primate genera spanning eight families. With the inclusion of an additional 29 primate genera from the literature, this is the most comprehensive report on dental eruption sequence in primates to date. We used a series of phylogenetic analyses to statistically compare dental eruption sequence to life history and body size and test for phylogenetic signal in these traits. RESULTS: Dental eruption sequence is conserved phylogenetically in primates, and body and brain size are both significantly associated with dental eruption sequence. Ancestral state reconstruction supports the hypothesis that the third molar erupted before one or more of the premolars in the ancestor of primates and derived clades within primates evolved an eruption sequence in which the third molar erupts after the premolars. DISCUSSION: Schultz's Rule, as it is currently written and applied, is not supported by this extended data set. Our results demonstrate that dental eruption sequence is a far better predictor of phylogeny and will likely prove useful in phylogenetic hypotheses about relationships between extinct and extant mammalian taxa. The evolution of dental eruption sequence is likely driven by factors that significantly influence body size and mandibular symphyseal fusion.

OH 83: A new early modern human fossil cranium from the Ndutu beds of Olduvai Gorge, Tanzania.

OBJECTIVE: Herein we introduce a newly recovered partial calvaria, OH 83, from the upper Ndutu Beds of Olduvai Gorge, Tanzania. We present the geological context of its discovery and a comparative analysis of its morphology, placing OH 83 within the context of our current understanding of the origins and evolution of Homo sapiens. MATERIALS AND METHODS: We comparatively assessed the morphology of OH 83 using quantitative and qualitative data from penecontemporaneous fossils and the W.W. Howells modern human craniometric dataset. RESULTS: OH 83 is geologically dated to ca. 60-32 ka. Its morphology is indicative of an early modern human, falling at the low end of the range of variation for post-orbital cranial breadth, the high end of the range for bifrontal breadth, and near average in frontal length. DISCUSSION: There have been numerous attempts to use cranial anatomy to define the species Homo sapiens and identify it in the fossil record. These efforts have not met wide agreement by the scientific community due, in part, to the mosaic patterns of cranial variation represented by the fossils. The variable, mosaic pattern of trait expression in the crania of Middle and Late Pleistocene fossils implies that morphological modernity did not occur at once. However, OH 83 demonstrates that by ca. 60-32 ka modern humans in Africa included individuals that are at the fairly small and gracile range of modern human cranial variation.

A comparative examination of odontogenic gene expression in both toothed and toothless amniotes.

A well-known tenet of murine tooth development is that BMP4 and FGF8 antagonistically initiate odontogenesis, but whether this tenet is conserved across amniotes is largely unexplored. Moreover, changes in BMP4-signaling have previously been implicated in evolutionary tooth loss in Aves. Here we demonstrate that Bmp4, Msx1, and Msx2 expression is limited proximally in the red-eared slider turtle (Trachemys scripta) mandible at stages equivalent to those at which odontogenesis is initiated in mice, a similar finding to previously reported results in chicks. To address whether the limited domains in the turtle and the chicken indicate an evolutionary molecular parallelism, or whether the domains simply constitute an ancestral phenotype, we assessed gene expression in a toothed reptile (the American alligator, Alligator mississippiensis) and a toothed non-placental mammal (the gray short-tailed opossum, Monodelphis domestica). We demonstrate that the Bmp4 domain is limited proximally in M. domestica and that the Fgf8 domain is limited distally in A. mississippiensis just preceding odontogenesis. Additionally, we show that Msx1 and Msx2 expression patterns in these species differ from those found in mice. Our data suggest that a limited Bmp4 domain does not necessarily correlate with edentulism, and reveal that the initiation of odontogenesis in non-murine amniotes is more complex than previously imagined. Our data also suggest a partially conserved odontogenic program in T. scripta, as indicated by conserved Pitx2, Pax9, and Barx1 expression patterns and by the presence of a Shh-expressing palatal epithelium, which we hypothesize may represent potential dental rudiments based on the Testudinata fossil record.

A one-million-year-old hominid distal ulna from Olduvai Gorge, Tanzania.

OBJECTIVE: Our aim was to recover new evidence of the evolution of the hominid lineage. METHODS: We undertook paleontological fieldwork at Olduvai Gorge, Tanzania, in one of the richest paleoanthropological sites in the world, documenting the evolution of our lineage and its environmental contexts over the last 2 million years. RESULTS: During field work in 2012, the Olduvai Vertebrate Paleontology Project discovered the distal end of a hominid ulna (OH 82) on the north side of Olduvai Gorge a few meters west of the Third Fault, eroding from Bed III sediments that are ∼1 million years in age. DISCUSSION: The size and morphology of this distal ulna falls within the normal range of variation seen in humans, although at the larger end of the distribution.

Identification of a derived dental trait in the papionini relative to other old world monkeys.

Variation in the shape of teeth provides an immense amount of information about the evolutionary history and adaptive strategy of a mammalian lineage. Here, we explore variation in the expression of a purported molar lingual remnant (the interconulus) across the Old World Monkeys (Primates: Cercopithecidae) with the aim of elucidating a component of the adaptive radiation of this family. This radiation is characterized by a wide geographic range (Asia and Africa) as well as diverse dietary niches. While all of the cercopithecids are distinguished by their derived bilophodont molars, the colobines have evolved taller and more pointed cusps compared with the cercopithecines. We investigate whether the interconulus also correlates with phylogenetic affinity and/or dietary adaptation. We assess the frequency and range of interconulus expression in 522 specimens representing seven species of Old World Monkeys (Cercopithecus mitis, n = 78; Macaca fascicularis, n = 85; Macaca mulatta, n = 70; Papio hamadryas, n = 55; Colobus guereza, n = 76; Presbytis melalophos, n = 82; Presbytis rubicunda, n = 76). Results show that the interconulus has a significantly higher frequency and degree of expression in Tribe Papionini and exhibits ordered metameric variation with greatest expression in the third molars. Given the rarity of the interconulus in other closely related taxa, and its morphological distinction from the purportedly homologous features in other primates, we interpret the high degree of expression of the interconulus to be a trait derived in papionins that originated in the Miocene baboon/macaque ancestor.

Assessing tooth wear progression in non-human primates: a longitudinal study using intraoral scanning technology.

Intraoral scanners are widely used in a clinical setting for orthodontic treatments and tooth restorations, and are also useful for assessing dental wear and pathology progression. In this study, we assess the utility of using an intraoral scanner and associated software for quantifying dental tissue loss in non-human primates. An upper and lower second molar for 31 captive hamadryas baboons (Papio hamadryas) were assessed for dental tissue loss progression, giving a total sample of 62 teeth. The animals are part of the Southwest National Primate Research Center and were all fed the same monkey-chow diet over their lifetimes. Two molds of each dentition were taken at either two- or three-year intervals, and the associated casts scanned using an intraoral scanner (Medit i700). Tissue loss was calculated in WearCompare by superimposition of the two scans followed by subtraction analysis. Four individuals had dental caries, and were assessed separately. The results demonstrate the reliability of these techniques in capturing tissue loss data, evidenced by the alignment consistency between scans, lack of erroneous tissue gain between scans, and uniformity of tissue loss patterns among individuals (e.g., functional cusps showing the highest degree of wear). The average loss per mm2 per year for all samples combined was 0.05 mm3 (0.04 mm3 for females and 0.08 mm3 for males). There was no significant difference in wear progression between upper and lower molars. Substantial variation in the amount of tissue loss among individuals was found, despite their uniform diet. These findings foster multiple avenues for future research, including the exploration of wear progression across dental crowns and arcades, correlation between different types of tissue loss (e.g., attrition, erosion, fractures, caries), interplay between tissue loss and microwear/topographic analysis, and the genetic underpinnings of tissue loss variation.

Technical note: Micro-computed tomography calibration using dental tissue for bone mineral research.

Computed tomography (CT) and microcomputed tomography (µCT) require calibration against density phantoms scanned with specimens or during routine internal calibration for assessment of mineral concentration (MC) and density. In clinical studies involving bone, alternative calibration methods using bodily tissues and fluids ("phantomless" calibration) have been suggested. However, such tissues are seldom available in archeological and osteological research. This study investigates the potential of dental tissue as internal reference for calibration of µCT scans, facilitating the analysis of bone MC. We analyzed 70 molars from 24 extant primate species, including eight human teeth, each scanned with density phantoms for calibration. Our findings indicate that sampling specific regions of molars (lateral aspects of the mesial cusps) yields low variation in enamel and dentine MC values, averaging 1.27 g/cm3 (±0.03) for dentine and 2.25 g/cm3 (±0.03) for enamel. No significant differences were observed across molar types or among scanning procedures, including scanner model, resolution, and filters. An ad hoc test on 12 mandibles revealed low variance in MC between the conventional phantom and dental tissue calibration methods; all 36 measurements (low, medium, and high MC for each mandible) were within 0.05 g/cm3 of each other -81% were within 0.03 g/cm3 and 94% within 0.04 g/cm3. Based on these results, we propose a new "phantomless" calibration technique using these mean enamel and dentine MC values. The presented phantomless calibration method could aid in the assessment of bone pathology and enhance the scope of studies investigating bone structure and physical property variations in archeological, osteological, and laboratory-based research.

Artificial neural networks reconstruct missing perikymata in worn teeth.

Dental evolutionary studies in hominins are key to understanding how our ancestors and close fossil relatives grew from the early stages of embryogenesis into adults. In a sense, teeth are like an airplane's 'black box' as they record important variables for assessing developmental timing, enabling comparisons within and between populations, species, and genera. The ability to discern this type of nuanced information is embedded in the nature of how tooth enamel and dentin form: incrementally and over years. This incremental growth leaves chronological indicators in the histological structure of enamel, visible on the crown surface as perikymata. These structures are used in the process of reconstructing the rate and timing of tooth formation. Unfortunately, the developmentally earliest growth lines in lateral enamel are quickly lost to wear once the tooth crown erupts. We developed a method to reconstruct these earliest, missing perilymata from worn teeth through knowledge of the later-developed, visible perikymata for all tooth types (incisors, canines, premolars, and molars) using a modern human dataset. Building on our previous research using polynomial regressions, here we describe an artificial neural networks (ANN) method. This new ANN method mostly predicts within 2 counts the number of perikymata present in each of the first three deciles of the crown height for all tooth types. Our ANN method for estimating perikymata lost through wear has two immediate benefits: more accurate values can be produced and worn teeth can be included in dental research. This tool is available on the open-source platform R within the package teethR released under GPL v3.0 license, enabling other researchers the opportunity to expand their datasets for studies of periodicity in histological growth, dental development, and evolution.

Neanderthal teeth from Moula-Guercy, Ardèche, France.

Here we describe dental remains from a Neanderthal fossil assemblage from Moula-Guercy, France. Our report demonstrates that the Moula-Guercy hominid remains contribute important morphological, developmental, and behavioral data to understanding Neanderthal evolutionary history. We include gross comparative morphological descriptions and enamel surface microstructure and microwear data. These teeth reveal numerous characteristics that are diagnostic of Neanderthals and provide no evidence for the presence of any other hominid taxa. Enamel growth increment data from the Moula-Guercy specimens yield evidence of a Neanderthal pattern of development, although at the lower end of the range of variation. The presence of a significant number of linear enamel hypoplasias indicates that these individuals were stressed during childhood. Molar microwear data suggest that these Neanderthals did not differ significantly from modern humans in terms of the fracture properties of the food they were consuming. The incisor microwear and macro striations provide evidence that these individuals may have been using their anterior teeth as tools, similar to the practices of several modern human populations such as the Inuit, Ipiutak, and Australian Aboriginals, and reminiscent of evidence from other Neanderthals from Krapina, Croatia, as well as the 600,000 year old hominids from Sima de los Huesos, Spain. Am J Phys Anthropol 151:477-491, 2013.© 2013 Wiley Periodicals, Inc.

A Pleistocene assemblage of near-modern Papio hamadryas from the Middle Awash study area, Afar Rift, Ethiopia.

OBJECTIVES: The aim of this study is to assess a new assemblage of papionin fossils (n = 143) recovered from later Pleistocene sediments in the Middle Awash study area in the Afar Rift of Ethiopia. MATERIALS AND METHODS: We collected metric and qualitative data to compare the craniodental and postcranial anatomy of the papionin fossils with subspecies of modern Papio hamadryas and with Plio-Pleistocene African papionins. We also estimated sex and ontogenetic age. RESULTS: The new fossils fit well within the range of morphological variation observed for extant P. hamadryas, overlapping most closely in dental size and proportions with the P. h. cynocephalus individuals in our extant samples, and well within the ranges of P. h. anubis and P. h. hamadryas. The considerable overlap in craniodental anatomy with multiple subspecies precludes subspecific diagnosis. We therefore referred 143 individuals to P. hamadryas ssp. The majority of the individuals assessed for ontogenetic age fell into middle- and old-adult age categories based on the degree of dental wear. Males (26%) were better represented than females (12%) among individuals preserving the canine-premolar honing complex. DISCUSSION: These new near-modern P. hamadryas fossils provide a window into population-level variation in the later Pleistocene. Our findings echo previous suggestions from genomic studies that the papionin family tree may have included a ghost population and provide a basis for future testing of hypotheses regarding hybridization in the recent evolutionary history of this taxon.

Peopling of the Americas: A new approach to assessing dental morphological variation in Asian and Native American populations.

OBJECTIVES: Through biodistance analyses, anthropologists have used dental morphology to elucidate how people moved into and throughout the Americas. Here, we apply a method that focuses on individuals rather than sample frequencies through the application rASUDAS2, based on a naïve Bayes' algorithm. MATERIALS AND METHODS: Using the database of C.G. Turner II, we calculated the probability that an individual could be assigned to one of seven biogeographic groups (American Arctic, North & South America, East Asia, Southeast Asia & Polynesia, Australo-Melanesia, Western Eurasia, & Sub-Saharan Africa) through rASUDAS2. The frequency of classifications for each biogeographic group was determined for 1418 individuals from six regions across Asia and the Americas. RESULTS: Southeast Asians show mixed assignments but rarely to American Arctic or "American Indian." East Asians are assigned to East Asia half the time while 30% are assigned as Native American. People from the American Arctic and North & South America are assigned to Arctic America or non-Arctic America 75%-80% of the time, with 10%-15% classified as East Asian. DISCUSSION: All Native American groups have a similar degree of morphological affinity to East Asia, as 10%-15% are classified as East Asian. East Asians are classified as Native American in 30% of cases. Individuals in the Western Hemisphere are decreasingly classified as Arctic the farther south they are located. Equivalent levels of classification as East Asian across all Native American groups suggests one divergence between East Asians and the population ancestral to all Native Americans. Non-arctic Native American groups are derived from the Arctic population, which represents the Native American founder group.