The Iberomaurusian enigma: north African progenitor or dead end?

Data obtained during an ongoing dental investigation of African populations address two long-standing, hotly debated questions. First, was there genetic continuity between Late Pleistocene Iberomaurusians and later northwest Africans (e.g., Capsians, Berbers, Guanche)? Second, were skeletally-robust Iberomaurusians and northeast African Nubians variants of the same population? Iberomaurusians from Taforalt in Morocco and Afalou-Bou-Rhummel in Algeria, Nubians from Jebel Sahaba in Sudan, post-Pleistocene Capsians from Algeria and Tunisia, and a series of other samples were statistically compared using 29 discrete dental traits to help estimate diachronic local and regional affinities. Results revealed: (1) a relationship between the Iberomaurusians, particularly those from Taforalt, and later Maghreb and other North African samples, and (2) a divergence among contemporaneous Iberomaurusians and Nubian samples. Thus, some measure of long-term population continuity in the Maghreb and surrounding region is supported, whereas greater North African population heterogenity during the Late Pleistocene is implied.     

Canary islands-north African population affinities: measures of divergence based on dental morphology

This investigation addresses two related questions about the origins and biological affinities of the Canary Islands' aboriginal inhabitants. First: With which North African populations do the pre-conquest inhabitants of the Canary Islands have their greatest affinities? Second: Does inter-island biological variability among the Canary Islanders, as has been suggested by other researchers (Hooton 1925, Schwidetzky 1963), imply that potentially different founding populations remained distinct during the pre-conquest period? This study employs dental morphology data derived from pre-conquest skeletons to answer these questions. Non-metric dental traits appear to be controlled by polygenic systems with a low to moderate environmental contribution to the resulting phenotype (Berry 1978, Harris & Bailit 1980, Nichol 1990) and can thus be assumed to reflect genetic relationships. The dental morphology of a sample of Canary Islanders (n = 397) is compared to that of Northwest African samples of Algerian Shawia Berbers (n = 26), Kabyle Berbers (n = 32), Bedouin Arabs (n = 49) and Punic Carthaginians (n = 28) as well as to six samples from Northeast Africa (n = 307) included for the purpose of understanding Canary Islanders' affinities within a wider context. The analysis employs 28 dental traits, quantifying differences in their expression among the various samples through a summary statistic, CAB Smith's Mean Measure of Divergence (MMD). The MMD analysis indicates that the Canary Island sample is most similar to the four samples from Northwest Africa: the Shawia Berbers, Kabyle Berbers, Bedouin Arabs and Carthaginians, less similar to the three Egyptian samples and least like the three Nubian samples. An intra-island comparison among samples from La Gomera, Gran Canaria and Tenerife reveals low, insignificant MMD values in all cases, implying that inter-island dental morphology differences are not so great as to require hypotheses of separate founding populations.     

Post-glacial recolonization of the Great Lakes region by the common gartersnake (Thamnophis sirtalis) inferred from mtDNA sequences

Pleistocene events played an important role in the differentiation of North American vertebrate populations. Michigan, in particular, and the Great Lakes region, in general, were greatly influenced by the last glaciation. While several hypotheses regarding the recolonization of this region have been advanced, none have been strongly supported. We generated 148 complete ND2 mitochondrial DNA (mtDNA) sequences from common gartersnake (Thamnophis sirtalis) populations throughout the Great Lakes region to evaluate phylogeographic patterns and population structure and to determine whether the distribution of haplotypic variants is related to the post-Pleistocene retreat of the Wisconsinan glacier. The common gartersnake was utilized, as it is believed to have been one of the primary vertebrate invaders of the Great Lakes region following the most recent period of glacial retreat and because it has been a model species for a variety of evolutionary, ecological, behavioral, and physiological studies. Several genetically distinct evolutionary lineages were supported by both genealogical and molecular population genetic analyses, although to different degrees. The geographic distribution of the majority of these lineages is interpreted as reflecting post-glacial recolonization dynamics during the late Pleistocene. These findings generally support previous hypotheses of range expansion in this region.     

Questions of khoesan continuity: Dental affinities among the indigenous holocene peoples of South Africa

The present report follows up on the findings of previous research, including recent bioarchaeological study of well‐dated Khoesan skeletal remains, that posits long term biological continuity among the indigenous peoples of South Africa after the Pleistocene. The Arizona State University Dental Anthropology System was used to record key crown, root, and intraoral osseous nonmetric traits in six early‐through‐late Holocene samples from the Cape coasts. Based on these data, phenetic affinities and an identification of traits most important in driving intersample variation were determined using principal components analysis and the mean measure of divergence distance statistic. To expand biological affinity comparisons into more recent times, and thus preliminarily assess the dental impact of disproportionate non‐Khoesan gene flow into local peoples, dental data from historic Khoekhoe and San were also included. Results from the prehistoric comparisons are supportive of population continuity, though a sample from Matjes River Rockshelter exhibits slight phenetic distance from other early samples. This and some insignificant regional divergence among these coastal samples may be related to environmental and cultural factors that drove low‐level reproductive isolation. Finally, a close affinity of historic San to all samples, and a significant difference of Khoekhoe from most early samples is reflective of documented population history following immigration of Bantu‐speakers and, later, Europeans into South Africa. Am J Phys Anthropol 155:33–44, 2014. © 2014 Wiley Periodicals, Inc.     

The accretion model of Neandertal evolution

The Accretion model of Neandertal evolution specifies that this group of Late Pleistocene hominids evolved in partial or complete genetic isolation from the rest of humanity through the gradual accumulation of distinctive morphological traits in European populations. As they became more common, these traits also became less variable, according to those workers who developed the model. Its supporters propose that genetic drift caused this evolution, resulting from an initial small European population size and either complete isolation or drastic reduction in gene flow between this deme and contemporary human populations elsewhere. Here, we test an evolutionary model of gene flow between regions against fossil data from the European population of the Middle and Late Pleistocene. The results of the analysis clearly show that the European population was not significantly divergent from its contemporaries, even in a subset of traits chosen to show the maximum differences between Europeans and other populations. The pattern of changes, over time within Europe of the traits in this subset, does not support the Accretion model, either because the characters did not change in the manner specified by the model or because the characters did not change at all. From these data, we can conclude that special phenomena such as near-complete isolation of the European population during the Pleistocene are not required to explain the pattern of evolution in this region.     

Who were the ancient Egyptians? Dental affinities among Neolithic through postdynastic peoples

Qualitative and quantitative methods are employed to describe and compare up to 36 dental morphological variants in 15 Neolithic through Roman-period Egyptian samples. Trait frequencies are determined, and phenetic affinities are calculated using the mean measure of divergence and Mahalanobis D2 statistics for discrete traits; the most important traits in generating this intersample variation are identified with correspondence analysis. Assuming that the samples are representative of the populations from which they derive, and that phenetic similarity provides an estimate of genetic relatedness, these affinities are suggestive of overall population continuity. That is, other than a few outliers exhibiting extreme frequencies of nine influential traits, the dental samples appear to be largely homogenous and can be characterized as having morphologically simple, mass-reduced teeth. These findings are contrasted with those resulting from previous skeletal and other studies, and are used to appraise the viability of five Egyptian peopling scenarios. Specifically, affinities among the 15 time-successive samples suggest that: 1) there may be a connection between Neolithic and subsequent predynastic Egyptians, 2) predynastic Badarian and Naqada peoples may be closely related, 3) the dynastic period is likely an indigenous continuation of the Naqada culture, 4) there is support for overall biological uniformity through the dynastic period, and 5) this uniformity may continue into postdynastic times.     

A Hominin Femur with Archaic Affinities from the Late Pleistocene of Southwest China

The number of Late Pleistocene hominin species and the timing of their extinction are issues receiving renewed attention following genomic evidence for interbreeding between the ancestors of some living humans and archaic taxa. Yet, major gaps in the fossil record and uncertainties surrounding the age of key fossils have meant that these questions remain poorly understood. Here we describe and compare a highly unusual femur from Late Pleistocene sediments at Maludong (Yunnan), Southwest China, recovered along with cranial remains that exhibit a mixture of anatomically modern human and archaic traits. Our studies show that the Maludong femur has affinities to archaic hominins, especially Lower Pleistocene femora. However, the scarcity of later Middle and Late Pleistocene archaic remains in East Asia makes an assessment of systematically relevant character states difficult, warranting caution in assigning the specimen to a species at this time. The Maludong fossil probably samples an archaic population that survived until around 14,000 years ago in the biogeographically complex region of Southwest China.     

Dental affinities of the C-group inhabitants of Hierakonpolis, Egypt: Nubian, Egyptian, or both?

By c. 2050 BC a small community of C-Group Nubians was present deep within Egyptian territory at the city of Hierakonpolis. Their descendants stayed for the next 400 years. Today, the site of Hierakonpolis, 113 km north of Aswan, is known for its Egyptian deposits; however, it also contains a C-Group cemetery, which documents the northernmost occurrence of this culture. Sixty skeletons were excavated. Tombs feature Nubian architecture and goods, including leather garments, although the use of Egyptian mortuary practices and artifacts increased through time. Dates range from the early 11th Dynasty into the Second Intermediate period. During this time the Egyptian empire occupied Lower Nubia, and their state ideology vilified Nubians. Yet, at least in death, the C-Group inhabitants of Hierakonpolis proudly displayed their cultural heritage.Beyond discerning the reason(s) for their presence at the site (e.g., mercenaries, leather-workers, entertainers?), the focus of this report is to estimate their biological affinity. Were they akin to other Nubians, Egyptians, or both? And, was increasing ‘Egyptianization’ evident in the mortuary ritual accompanied by concomitant genetic influence? To address these queries, up to 36 dental morphological traits in the recovered individuals were compared to those in 26 regional comparative samples. The most influential traits were identified and phenetic affinities were calculated using the mean measure of divergence and other multivariate analyses. Assuming phenetic similarity provides an estimate of genetic relatedness, these affinities suggest the individuals comprising the C-Group sample were, and remained Nubian during their tenure at Hierakonpolis.     

Microsatellite analysis of the phylogeography, Pleistocene history and secondary contact hypotheses for the killifish, Fundulus heteroclitus

The mummichog, Fundulus heteroclitus, exhibits extensive latitudinal clinal variation in a number of physiological and biochemical traits, coupled with phylogeographical patterns at mitochondrial and nuclear DNA loci that suggest a complicated history of spatially variable selection and secondary intergradation. This species continues to serve as a model for understanding local and regional adaptation to variable environments. Resolving the influences of historical processes on the distribution of genetic variation within and among extant populations of F. heteroclitus is crucial to a better understanding of how populations evolve in the context of contemporary environments. In this study, we analysed geographical patterns of genetic variation at eight microsatellite loci among 15 populations of F. heteroclitus distributed throughout the North American range of the species from Nova Scotia to Georgia. Genetic variation in Northern populations was lower than in Southern populations and was strongly correlated with latitude throughout the species range. The most common Northern alleles at all eight loci exhibited concordant latitudinal clinal patterns, and the existence of an abrupt transition zone in allele frequencies between Northern and Southern populations was similar to that observed for mitochondrial DNA and allozyme loci. A significant pattern of isolation by distance was observed both within and between northern and southern regions. This pattern was unexpected, particularly for northern populations, given the recent colonization history of post-Pleistocene habitats, and was inconsistent with either a recent northward population expansion or a geographically restricted northern Pleistocene refugium. The data provided no evidence for recent population bottlenecks, and estimates of historical effective population sizes suggest that post-Pleistocene populations have been large throughout the species distribution. These results suggest that F. heteroclitus was broadly distributed throughout most of its current range during the last glacial event and that the abrupt transition in allele frequencies that separate Northern and Southern populations may reflect regional disequilibrium conditions associated with the post-Pleistocene colonization history of habitats in that region.     

Post-Pleistocene changes in the human dentition

Published evidence indicates sharp reductions in the hominid dentition following the end of the Pleistocene. These reductions, both in size and in morphological complexity, have proceeded farthest in those areas where culture change has occurred most rapidly. The model proposed here suggests that post-Pleistocene dental reduction may be the result of the change in selective forces consequent from the invention and use of pottery and the changes in food-preparation techniques after the end of the Pleistocene. Models for testing this hypothesis are discussed.     

The Xujiayao 14 Mandibular Ramus and Pleistocene Homo Mandibular Variation

The earlier Late Pleistocene mandibular ramus from Xujiayao (northern China) preserves traits that vary distributionally among western Old World Pleistocene Homo samples and between Early/Middle Pleistocene archaic humans and Late Pleistocene modern humans in eastern Eurasia. Xujiayao 14 presents a lateral mandibular notch crest, an open mandibular foramen, a wide ramus, an asymmetrical mandibular notch, an enlarged superior medial pterygoid tubercle, (probably) a retromolar space, and gonial eversion, as well as an unusual depression in the planum triangulare. The first two traits appear ancestral for Later Pleistocene and recent Homo and are dominant among modern humans. The second two traits largely separate Xujiayao 14 and archaic Homo from modern humans. The next two traits are found in the highest frequency among the Neandertals, although gonial eversion contrasts with Late Pleistocene Neandertals. Xujiayao 14, in the context of Pleistocene and recent Homo samples and the other Xujiayao human remains, therefore provides a morphological mosaic, highlighting regional variation through the Pleistocene.     

Phylogeography of the tree lizard, Urosaurus ornatus: responses of populations to past climate change

Isolation due to both geological barriers and range contractions during the Pleistocene glacial maxima has been an important cause of diversification of arid-adapted species in the North American deserts. Tree lizards, Urosaurus ornatus, are distributed across much of the southwestern arid regions and can tolerate a wide range of environments. Thus, they may have avoided large-scale shifts in distribution caused by Pleistocene climate change and any subsequent evolutionary impacts. Cytochrome b sequences were sampled from U. ornatus across the northern part of their range to test if current structure of these populations resulted from post-Pleistocene range expansion and habitat fragmentation, or prior geological isolation. Phylogenetic analyses found geographical structuring of populations consistent with a model of long-term geographical isolation corresponding to each of the desert regions. The two post-Pleistocene hypotheses were not well supported as estimated times of divergence predated the retreat of the last continental ice sheet. Populations in different regions were impacted by different processes. Southern populations of U. ornatus appear to have remained largely independent of more derived northern and eastern populations during Pleistocene climate change, while populations in regions containing more derived populations showed evidence of more recent range expansion (Colorado Plateau). As populations of U. ornatus attest to, the complex and dynamic history of the southwestern USA has left a deep-rooted and multifaceted imprint on genetic and phylogeographical structure of the species living there.     

Phylogeographic and demographic effects of Pleistocene climatic fluctuations in a montane salamander, Plethodon fourchensis

Climatic changes associated with Pleistocene glacial cycles profoundly affected species distributions, patterns of interpopulation gene flow, and demography. In species restricted to montane habitats, ranges may expand and contract along an elevational gradients in response to environmental fluctuations and create high levels of genetic variation among populations on different mountains. The salamander Plethodon fourchensis is restricted to high-elevation, mesic forest on five montane isolates in the Ouachita Mountains. We used DNA sequence data along with ecological niche modelling and coalescent simulations to test several hypotheses related to the effects of Pleistocene climatic fluctuations on species in montane habitats. Our results revealed that P. fourchensis is composed of four well-supported, geographically structured lineages. Geographic breaks between lineages occurred in the vicinity of major valleys and a narrow high-elevation pass. Ecological niche modelling predicted that environmental conditions in valleys separating most mountains are suitable; however, interglacial periods like the present are predicted to be times of range expansion in P. fourchensis . Divergence dating and coalescent simulations indicated that lineage diversification occurred during the Middle Pleistocene via the fragmentation of a wide-ranging ancestor. Bayesian skyline plots showed gradual decreases in population size in three of four lineages over the most recent glacial period and a slight to moderate amount of population growth during the Holocene. Our results not only demonstrate that climatic changes during the Pleistocene had profound effects on species restricted to montane habitats, but comparison of our results for P. fourchensis with its parapatric, sister taxon, P. ouachitae , also emphasizes how responses can vary substantially even among closely related, similarly distributed taxa.     

Mitochondrial diversity of Musca domestica housefly populations in the Asian and western Pacific biogeographical regions

Houseflies (Musca domestica L., Diptera: Muscidae) are cosmopolitan, colonizing, and eusynanthropic. Their distribution in the Malaysian archipelago provides an opportunity to study successive waves of colonization and extinction during the Pleistocene and Recent epochs. We scored single-strand conformation polymorphisms (SSCPs) at 16S2 and COII mitochondrial loci in 47 housefly samples from the Australian, Austro-Malayan, Indo-Malayan, Manchurian and Indo-Chinese subregions of Wallace's zoogeographical classification. We discuss the results in light of the Pleistocene vs. post-Pleistocene dispersal and faunal exchange in the Asia-Pacific area. Fourteen haplotypes were detected, of which 10 were confined to a single subregion. No haplotype was ubiquitous and only one was found in four subregions. Population diversity, HS, was greatest in the Indo-Malayan (0.36) and heterogeneous among subregions. The mean subregional diversity was 0.21 +/- 0.03, representing the probability that two randomly chosen flies, from any subregion, had different haplotypes. The hierarchical partition of diversity indicated restricted maternal gene flow among subregions (GRT = 0.60, Nm approximately 0.32). These results suggest long-standing genetic isolation of houseflies in the Malaysian archipelago and support the hypothesis that they dispersed widely during the Pleistocene. Haplotypes common among mainland populations but shared with island groups in low frequencies (<1%) indicate surprisingly little recent gene flow.     

Testing nested phylogenetic and phylogeographic hypotheses in the Plethodon vandykei species group

Mesic forests in the North American Pacific Northwest occur in two disjunct areas: along the coastal and Cascade ranges of Oregon, Washington, and British Columbia as well as the Northern Rocky Mountains of Idaho, Montana, and British Columbia. Over 150 species or species complexes have disjunct populations in each area, and a priori hypotheses based on phytogeography and geology potentially explain the disjunction via either dispersal or vicariance. Here, we test these hypotheses in the disjunct salamander complex Plethodon vandykei and P. idahoensisby collecting genetic data (669 bp of Cyt b) from 262 individuals. Maximum likelihood analysis indicated reciprocal monophyly of these species, supporting the ancient vicariance hypothesis, whereas parametric bootstrap and Bayesian hypothesis testing allow rejection of the dispersal hypothesis. The coalescent estimate of the time since population divergence (estimated using MDIV) is 3.75 x 106 years, and the 95%credibility interval of this value overlaps with the geological estimate of vicariance, but not the hypothesized dispersal. These results are congruent with the pattern seen in other mesic forest amphibian lineages and suggest disjunction in amphibians may be a concerted response to a geological/climatological event. WithinP. idahoensis, we tested the corollary hypothesis of an inland Pleistocene refugium in the Clearwater drainage with nested clade analysis and coalescent estimates of population growth rate (g). Both analyses support post-Pleistocene expansion from the Clearwater refugium. We corroborated this result by calculating Tajima's Dand mismatch distribution within each drainage, showing strong evidence for recent population expansion within most drainages. This work demonstrates the utility of statistical phylogeography and contributes two novel analytical tools: tests of stationarity with respect to topology in the Bayesian estimation, and the use of coalescent simulations to test the significance of the population growth-rate parameter.     

Phylogeographic and population genetic analyses reveal Pleistocene isolation followed by high gene flow in a wide ranging,but endangered,freshwater mussel

Freshwater organisms of North America have had their contemporary genetic structure shaped by vicariant events, especially Pleistocene glaciations. Life history traits promoting dispersal and gene flow continue to shape population genetic structure. Cumberlandia monodonta, a widespread but imperiled (IUCN listed as endangered) freshwater mussel, was examined to determine genetic diversity and population genetic structure throughout its range. Mitochondrial DNA sequences and microsatellite loci were used to measure genetic diversity and simulate demographic events during the Pleistocene using approximate Bayesian computation (ABC) to test explicit hypotheses explaining the evolutionary history of current populations. A phylogeny and molecular clock suggested past isolation created two mtDNA lineages during the Pleistocene that are now widespread. Two distinct groups were also detected with microsatellites. ABC simulations indicated the presence of two glacial refugia and post-glacial admixture of them followed by simultaneous dispersal throughout the current range of the species. The Ouachita population is distinct from others and has the lowest genetic diversity, indicating that this is a peripheral population of the species. Gene flow within this species has maintained high levels of genetic diversity in most populations; however, all populations have experienced fragmentation. Extirpation from the center of its range likely has isolated remaining populations due to the geographic distances among them.     

MITOCHONDRIAL DNA VARIATION WITHIN AND BETWEEN SPECIES OF THE PAPILIO MACHAON GROUP OF SWALLOWTAIL BUTTERFLIES

Species limits and phylogenetic relationships in the Papilio machaon species group are potentially confounded by a complex pattern of Pleistocene range fragmentation, hybridization, and ecological race formation. Mitochondrial DNA (mtDNA) restriction-site analysis has been used to define genetic affinities and genetic population structure within this species group. The distribution of mtDNA haplotypes generally confirms prior phylogenetic hypotheses and species delineations, but there is poor correspondence between ecological races and mtDNA haplotypes. The amount and distribution of mtDNA sequence variation within species vary among species, reflecting differences in current patterns of gene flow and/or historical population structure. In spite of wing pattern characters that ally them with P. polyxenes, both P. joanae and P. brevicauda have mtDNA that is closely related to that of P. machaon. We suggest that P. joanae and P. brevicauda are of hybrid origin.     

Molecular evidence for Pleistocene glacial cycles driving diversification of a North American desert spider, Agelenopsis aperta

The influence of historical climatic vs. geological changes on species diversification patterns was investigated in a widely distributed North American desert spider, Agelenopsis aperta (Araneae: Agelenidae), with particular reference to Pleistocene glacial cycles and earlier patterns of mountain building. Levels of sequence divergence obtained from the mitochondrial gene, cytochrome oxidase I, dated to the Pleistocene, eliminating Rocky Mountain orogeny as a cause of diversification, as orogeny ended 4 million years ago. The results of phylogenetic and network analyses showed the presence of three geographically defined clades, which were consistent with the presence of at least three glacial refugia: (i) east of the Rocky Mountains; (ii) between the Rocky Mountains and Sierra Nevadas; and (iii) west of the Sierra Nevadas. In addition, populations within the Rocky Mountains exhibited significantly lower genetic diversity than populations east of the Rocky Mountains and the haplotypes found within the Rockies were a subset of eastern haplotypes. These patterns suggest that a post-Pleistocene range expansion occurred out of an eastern glacial refugium into the Rocky Mountains. Examination of phylogeographical studies of other North American desert taxa indicated that mountain building explained diversification patterns more effectively for some taxa but Pleistocene climate change was more important for others, including A. aperta.