A second possibility for Europe is that the fossils from the Sima de los Huesos date to only around 350 ka. Crania and jaws from Arago Cave and Petralona, and the spectacular assemblage from Sima de los Huesos, are particularly informative. There is less agreement about the deeper phylogeny of these lineages and related forms from the late Lower through early Middle Pleistocene, but that period predates the central focus of this paper. The extent and geographic patterning of morphological diversity among Late Middle Pleistocene (LMP) African hominins is largely unknown, thus precluding the definition of boundaries of variability in early H. sapiens and the interpretation of individual fossils. Describe the differences between middle Pleistocene hominins and Neandertals. These late, western Neanderthal mtDNA sequences have a coalescent age of 58 ka (the end of OIS 4) with a 95% CI 54-77 ka. A potential explanation for the apparent lack of a contraction in the effective population size of the ancestors of modern humans in Africa is that if there were in fact bottlenecks within a subdivided population in Africa, following the bottlenecks, members of dissimilar populations mixed extensively, restoring to the resulting population much of the genetic variation that existed before each bottleneck. Likewise, Rohling et al. Genetic drift provides an alternative explanation for morphological divergence (Howell). by indicating possible population expansions within Africa and possible expansions into Arabia during the windows of opportunity described by Rohling et al). A recent analysis of the ancient mtDNA of a Neanderthal from Valdegoba, Spain, dating to 48.5 ka, shows that all Neanderthal mtDNA sequences postdating this time formed a compact, monophyletic group within the known Neanderthal sequences (Dalén et al). synthesized paleoclimatic records for the whole of Africa using multiple proxies, including terrestrial, lacustrine, and oceanic data. von Koenigswald collected a number of Early-Middle Pleistocene hominid specimens in … Although this hypothesis has been marginalized historically, recent reviews have emphasized its potential importance (e.g., Hublin). The implications of these findings are that contrary to previous conclusions (Ruff), pelvic form appeared to have followed a pattern of largely neutral evolution like most human cranial dimensions (Betti et al; Roseman; Weaver, Roseman, and Stringer). Site ODP 659 also receives dust from Western Sahara and portions of the northern Sahara during these months. Likewise, by applying a population genetics model to expectations for (neutral) change in cranial dimensions, Weaver showed that crania that had dimensions that differed by one standard deviation from modern crania could be expected by around 165 ka, which corresponds reasonably well to when most researchers agree that modern (or nearly modern) humans appear in the East African fossil record. In Africa, especially in East Africa, biomass and human population size were much more dependent on the availability (and predictability) of precipitation, and the dust-flux data from deep-sea cores provide an accessible gauge of precipitation (deMenocal; Rohling et al.). Furthermore, all Neanderthal mtDNAs share a common ancestor approximately 100 ka and a common ancestor with modern humans ∼500 ka (Reich et al). A more slender physique typified Omo I from Africa (Pearson et al) and the early modern humans from Skhul and Qafzeh in Israel (Carretero et al; Ruff, Trinkaus, and Holliday). The Sima de los Huesos sample shows mosaics of Neanderthal and non-Neanderthal morphology in virtually all aspects of its morphology (Arsuaga et al). suggest that this population also had a strikingly low long-term effective population size of approximately individuals for the period between 400 and 100 ka (Li, Patterson, and Reich. In addition, some recent approaches to cultural innovations also emphasize the role of chance, especially if change is dependent on population size and density (e.g., Powell, Shennan, and Thomas; Shennan). In Middle Pleistocene China, the primary odic irruptions, followed by … The direct influence of climatic conditions on population sizes in Europe and Africa allows a series of predictions about the relative ability of selection and drift to produce changes in hominin populations. Ample precipitation over East Africa and the southern Sahara and Sahel promote the growth of vegetation, which decreases the amount of dust that winds scour off of the land. Major dry phases would have been guaranteed to produce greatly expanded Sahara and Kalahari deserts and unfavorable conditions for human habitation. Both African and European Middle Pleistocene hominins tended to be medium to tall in stature (Carretero et al) and very heavy for height relative to modern hunters and gatherers (Churchill et al; Kappelman). Rohling et al. Mellars and French refrained from proposing an estimate of the number of individuals this would have involved, but Bocquet-Appel et al.’s model for human population in Upper Paleolithic Europe produced an estimate of the population of Europe during the Aurignacian of 4,424 people (95% confidence interval [CI]: 1,738-28,359). In this paper, I argue that climate and population genetics are linked. These discoveries have decisively answered the question of whether interbreeding occurred between modern and archaic humans (it did in both cases) and opened new windows on which genes may have been involved in producing evolutionary novelties in both modern humans and Neanderthals. Copyright © 2013 Elsevier Ltd. All rights reserved. Hominins that differ from Homo erectus, the Neanderthals, and recent humans are known from Middle Pleistocene localities across the Old World. Periods of decreased precipitation diminish the amount of vegetation and dependent biomass (including humans) and produce more dust. If the Sima de los Huesos dates to 500-600 ka (Bischoff et al), then one can conclude the features characteristic of the later, “classic” Neanderthals dating to OIS 4-3 increased in frequency very slowly within the Neanderthal lineage. For example, in eastern Asia, these hominin fossils have been classified as archaic, early, or premodern H. sapiens. The taxonomic status of these populations has been clouded by controversy. As new morphologies may be effectively invisible in the fossil record during periods of contracted population size, they may, in fact, appear in the record only slightly later, once population sizes had rebounded. These features are similarly common in European fossils dating to 300-200 ka but much more rare (or absent) in earlier fossils from other sites in Europe. The two processes are not mutually exclusive, and both often act on a population at the same time. This could provide evidence that early modern humans had shifted to different niches than archaic humans and had experienced a substantial pulse of selection that tailored them for their new habits. 2012; Kappelman 1996). Christopher Bae. Many of the key events appear to date to periods in which population sizes were greatly reduced and genetic drift would have been rapid. (Multiple answers) H. heidlebergensis has a mix of H. erectus and more dervied features which included a larger brain case, but it retained thicker browridges. This period is associated with the first appearance of Homo heidelbergensis (or Homo rhodesiensis, if this name is to be preferred) in Africa (i.e., the Bodo cranium, dated to 600 ka; Clark et al; Rightmire) and, intriguingly, marked technological advances represented by precociously early blade production and core technology in the Kapthurin Formation at Lake Baringo (Johnson and McBrearty; Tryon and McBrearty). Periods of high dust flux correspond to less precipitation, less vegetation, fewer people, and thus rapid genetic drift. The problem may not be intractable, however, because during the Middle and Upper Pleistocene, recurrent 100,000-year-long glacial cycles drove climate change and almost certainly affected hominin populations. and divergence in cranial dimensions modeled as the result of neutral evolution (Weaver, Roseman, and Stringer). At the genetic level, two of the fundamental means by which evolution can occur are natural selection (referred to subsequently simply as “selection”) and genetic drift. Analysis of autosomal DNA indicates a divergence time between modern human and Neanderthal populations of 270-440 ka (Reich et al). Some of the reconstructed lake levels (e.g., for Lake Malawi) do not closely follow the dust curves, suggesting yet another layer of complexity in the climatic record. Wide hips may have also been inherited from Homo erectus (Simpson et al) rather than appearing as an evolutionary novelty in Middle Pleistocene hominins. In contrast, the mandibular dentition of Jebel Irhoud 3, a juvenile late archaic hominin from Morocco dating to 160 ka with affinities to modern humans (Hublin; Hublin and Tillier), preserves evidence of a slower, modern pace of dental development (Smith et al). The dust core also indicates marked dry periods in East Africa during OIS 8 (301-242 ka), OIS 6 (186-127 ka), and OIS 4-2 (71-12 ka), although Blome et al. During the same months the southerly Shamai winds scour dust off of the Arabian Peninsula and deposit it in the Arabian Sea. An interesting question is how these Middle Pleistocene hominins were related to those who lived in the Late Pleistocene epoch, in particular to Neanderthals in western Eurasia and to Denisovans, a sister group of Neanderthals so far known only from southern Siberia. 74 During the Middle Pleistocene both land and sea bridges were potential dispersal 75 routes for hominins (Derricourt, 2006). Climate affects ecological productivity and biomass, which in turn affects human population numbers. Patterns of covariation are compared in order to assess integration. 37 Full PDFs related to this paper. Fagundes et al. The record stems from long-standing patterns of atmospheric circulation. This suite of Neanderthal features had become common in European hominins by OIS 5, including the specimens from Krapina and Saccopatore, and they became even more frequent in OIS 4-3. present multiproxy data for the Mediterranean and Red Sea regions, two areas that were crucial for hominin dispersals from (and perhaps into) Africa during the last 500 kyr. Intriguingly, their data showed no evidence of a bottleneck between 200 and 100 ka. All extant mtDNA sequences coalesce to a common ancestor at 140-210 ka (Behar et al. ), although the pattern of wet and dry periods for Africa as a whole forms a complex mosaic that frequently departs from the pattern observed in Lakes Malawi and Tangyanika (Blome et al). The hominins from Bodo, Broken Hill, and Elandsfontein in Africa are quite similar to their Middle Pleistocene contemporaries in Europe. A further difficulty particular to Africa lies in the variability of dust-flux records: different patterns occur in different cores around Africa . In 1993 a human tibia was found at Boxgrove, from a sediment overlying freshwater deposits at Q1/B. In each case, populations can be inferred to have spread from regions with favorable climate and thus presumably comparatively high human population density into regions previously nearly devoid of people but with newly favorable climatic conditions. The last ENI, published in 2004, was 28. This paper. In addition, cores and seismology of several of the oldest East African great lakes, especially Lakes Malawi and Tanganyika, have shown that substantial portions of tropical Africa south of the equator experienced severe droughts over the last 200 kyr that would not have been inferred from the oxygen isotope curve (Burnett et al; Cohen et al; Scholz et al), although the effects of these droughts appear to have been mitigated or absent at the equator and cannot be generalized to the whole of Africa (Blome et al). The Middle Pleistocene is a crucial time period for studying human evolution in Europe, because it marks the appearance of both fossil hominins ancestral to the later Neandertals and the Acheulean technology. Comparison of the time lines of paleoclimate, the fossil record, and genetic divergences and bottlenecks provide a rough check of whether key events occur in periods favorable for large population numbers or in periods unfavorable for large populations. In fact, their results show growth in effective population size from ∼450 ka until 120-150 ka and very similar histories (and likely shared histories in an ancestral source population) of Yoruban, eastern Asian, and European population size before 60 ka. At the onset of the Quaternary, hominids identified as Homo erectus spread widely across the Old World. The stature of early modern humans from the Levalloiso-Mousterian of the Levant and the Gravettian of Europe is particularly striking relative to Neanderthals and almost all other samples from Europe before the twentieth century (Carretero et al). Toward the close of the Pleistocene, skulls appear increasingly similar to those of living humans. These results are exciting and motivate one to take a closer look at some of the recent genetic advances. If one accepts Bocquet-Appel et al.’s estimates for the Aurignacian and extends Mellars and French’s conclusions to the whole of Europe, it would imply that the Neanderthal population of Europe only totaled 492 individuals (95% CI: 193-3,151). Although Neandertals are the best-known fossil hominins, the tempo and evolutionary processes in their lineage are strongly debated. hominins show the following: (i) wide bodies, a plesiomorphic char-acter in the genus Homo inherited from their early hominin ancestors; (ii) statures that can be found in modern human middle-latitude pop- ulations that first appeared 1.6–1.5 Mya; and (iii) large femoral heads in some individuals, a trait that first appeared during the middle Pleistocene in Africa and Europe. proposed that this reduction in body mass may have been an evolutionary adaptation to a lifestyle that favored energy efficiency. This gradual increase in similarity to late Neanderthals has been dubbed the “accretion model” and may have unfolded as a single, long trend or perhaps in two pulses (Hublin). Indirect evidence from autosomal genes also supports the hypothesis that the African ancestors of modern humans experienced a major population bottleneck during this period. The completion of a draft of the Neanderthal nuclear genome (Green et al) and recovery and analysis of nuclear and mitochondrial DNA from “Denisovans,” a third lineage that separated from modern humans slightly before Neanderthals (Meyer et al; Reich et al), stand out as signal achievements. In sum, this evidence suggests a speciation event in which Discriminant functions facilitate the description of intergroup differences. Sexual dimorphism is an important component of the total variation seen in populations and plays a key role in taxonomic debates. Humans that are different from Homo erectus evolved first in Africa or western Eurasia. Many of the famous cultural advances associated with the Upper Paleolithic and Late Stone Age are also likely to have depended on population size and density (Powell, Shennan, and Thomas; Premo and Kuhn). The phylogenetic relationships between hominins of the Early Pleistocene epoch in Eurasia, such as Homo antecessor, and hominins that appear later in the fossil record during the Middle Pleistocene epoch, such as Homo sapiens, are highly debated 1-5 . Neanderthals may have only rarely experienced periods of population growth and range expansion. June 2011; Authors: D.R. In a review of 75 distinctive cranial, dental, and postcranial features of early modern humans and Neanderthals, Trinkaus) concluded that only one quarter were unique to Neanderthals while twice that many were unique to modern humans, a finding that means that Neanderthal morphology had remained fairly primitive while early moderns were much more derived. Copyright © 2021 Elsevier B.V. or its licensors or contributors. A similar approach is used with the Middle Pleistocene sample. Genetic drift would be expected to be the dominant factor during such a period, but it is worth reiterating that African climate was a complex and regionally variable mosaic (Blome et al). Views of the origin of modern humans and our divergence from Neanderthals have been profoundly and perhaps decisively influenced by genetic data from living humans as well as ancient DNA (aDNA) from Neanderthals. The results showed that the most probable scenario (isolation and drift in the western and eastern populations of Neanderthals at 48 ka) would have involved an effective population size (Ne) of western Neanderthals of only 300 females, a marked reduction from the estimate for the eastern subpopulation ( females). The effective population size for the autosomal genes in the entire population is expected to be four times that of mtDNA (i.e., and 8,000 in the western and eastern subpopulations, respectively). The ultimate source of new alleles is mutation, which occurs rarely. those of Middle Pleistocene hominins, and a substantially larger brain than those of most Middle Pleistocene hominins 1. The morphology of the Enamel-Dentine boundary is hidden beneath the enamel surface and so is not subjected to wear by the hominins continuous chewing throughout her or his lifetime. The picture that emerges is one of human population history that was highly (although almost certainly not exclusively) contingent on climatic changes. After that, all three populations experienced bottlenecks, although the one that affected the ancestors of the Yoruba appears to have been less severe and allowed an earlier recovery. During December, January, and February, the direction of air circulation reverses over East Africa, and the Trade Winds blow air onshore over East Africa and across the Sahel and much of the Sahara from a northeasterly counterclockwise direction. If these remarkably low population numbers are accurate, Neanderthals may never have been numerous enough to experience conditions in which there were enough individuals for favorable mutations to arise at a brisk pace. As a result of this comparison of records of paleoclimate, morphological change, and genetic change, it seems apparent that many of the observed changes leading to Neanderthals were more likely to have been the products of drift than selection, whereas both drift and selection may have been important in the emergence of modern humans. A severe population bottleneck (or selection, which may be less likely) could produce such a pattern. Both African and European Middle Pleistocene hominins tended to be medium to tall in stature (Carretero et al) and very heavy for height relative to modern hunters and gatherers (Churchill et al; Kappelman). In many cases, the relationships are highly suggestive, but problems remain. Recent alternative hypotheses that accept the greater antiquity for Sima de los Huesos have proposed the presence of two lineages in Europe until 300-400 ka (García and Arsuaga) or complicated scenarios of local extinction, recolonization, and admixture of two or more populations (Dennell, Martinón-Torres, and Bermúdez de Castro). Another population contraction in most of Africa in OIS 2 probably accounts for late (Holocene or terminal Pleistocene) appearance of crania that have cranial metrics that cannot be distinguished from one or more extant populations (De Villiers and Fatti; Habgood). Recent papers have produced a range of estimates for when the ancestors of Neanderthals and modern humans split, ranging from ∼835 ka for the average divergence for autosomal sequences (Green et al. Next, factor analysis is applied to the H. erectus specimens in an attempt to identify modules, tightly integrated traits that can evolve independently. This seems to have happened many times in the past, with conditions in OIS 2 serving as a case in point (Brooks and Robertshaw; Deacon and Lancaster). China is one example, on the other side of the of trees, which ‘‘on the Quaternary time-scale resemble the peri- Eurasian landmass. There is agreement that earlier Middle Pleistocene hominins share primitive traits with H. erectus. Recently Martinón-Torres and colleagues have shown that the sample has very Neanderthal-like teeth; some of the nonmetrical features are even more common in the Sima de los Huesos sample than in late, “classic” Neanderthals from OIS 4 to OIS 3, which casts doubt on simple models of a steady increase in Neanderthal features over time. After 34 years of research and findings in the Middle Pleistocene site of the Sima de los Huesos (SH) of the Sierra de Atapuerca (Burgos, Spain), we present an update of the estimation of the number of individuals (ENI) identified in the SH hominin assemblage. The time is ripe for a reconsideration of scenarios for adaptive change because of the accumulation of a critical mass of new evidence from paleoecology, genetics, anatomy, and chronology. Later populations display more derived features, and the skeletons from Qafzeh and Klasies River are near-modern in their morphology. Low lake levels in Lake Malawi and Lake Tanganyika and high levels of dust flux suggest generally unfavorable conditions for human population growth in tropical Africa during much of OIS 5 (Blome et al; Scholz et al). Using data from complete genomes of several modern men comprising two Yoruba, three Europeans, one Chinese, and one Korean, Li and Durbin applied population genetics models to infer changes in human effective population size over the last million years. Download Full PDF Package. If selection was the crucial factor driving change in the lineages of Neanderthals or modern humans, then major changes in anatomy in each lineage should emerge during periods that favor large population numbers. Wide hips and robust long bones were already present in the Sima de los Huesos sample (Arsuaga et al; Bonmatí et al) and may have been the primitive condition for Middle Pleistocene Homo (Arsuaga et al). The origin of modern humans dates to OIS 6. The complex mosaic of favorable climates over time in different parts of Africa reported by Blome et al. 2012) and very heavy for height relative to modern hunters and gatherers (Churchill et al. At the higher end of estimates, Sørensen proposed a population of less than 10,000 individuals during the Eemian interglacial (OIS 5e), when Neanderthal numbers and inhabited territory may have been at a peak. Periods of low dust flux indicate more precipitation, more vegetation, more animal biomass, and more people. The paleoclimate of Africa presents a more complicated picture but one that is ultimately related to orbital dynamics because of changes in air circulation and rainfall that arose as consequences of the amount of solar radiation (insolation) that reached the earth (Siddall et al; Trauth, Larrasoaña, and Mudelsee). Very wet and warm periods in Europe produced dense forests that may have also been unfavorable habitat (Roebroeks, Conard, and Van Kolfschoten), although interstadial periods seem to have been far more favorable for hominin populations than the coldest periods of glaciations. The scale of the effect of climatic changes on human populations is clearly apparent in the dramatic decrease in the number of sites in the Last Glacial Maximum in East Africa (Brooks and Robertshaw); difficulties for human populations likely continued even after that, including evidence of the desiccation of Lake Victoria until ca. To read the full-text of this research, you can request a copy directly from the author. This could have constrained human groups to migrate into such a propitious area. Reviewing previous estimates of Neanderthal population numbers, Dennell, Martinón-Torres, and Bermúdez de Castro proposed the Neanderthal population of Europe totaled 3,000-5,000 during interstadials and 1,500-2,500 during the depths of glacial advances, when Neanderthal populations were confined to refugia in Iberia, Italy, and the Balkans. Been clouded by controversy recent humans are known from Middle Pleistocene hominin fossils have classified... Help provide and enhance our service and tailor content and ads hominins in. 140-210 ka ( Behar et al. processes in their morphology, was 28 the... Ratified in January 2020 first fossils of recognizably modern form ( Bräuer ; Pearson ; )... By Rohling et al. long-standing patterns of atmospheric circulation look at some of the total variation seen populations... Dynamics, likely attributable to climatic cycles, affected archaic and modern populations in Africa or Western Eurasia Africa ca. 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Series of assumptions that are open to criticism Cave and Petralona, and more people in body mass have! Western Eurasia i.e., the tempo and evolutionary processes in their lineage are strongly debated a. Can override the signal of all but the strongest selective pressures lifeways during the Middle Pleistocene hominins! Odp 659 also receives dust from Western Europe indicates that hominins settled in different kinds of environments Serbia... Modeled as the result was likely relatively rapid genetic drift as a key role taxonomic. And other aspects of lifeways during the Middle Pleistocene localities across the Old World one more... S model, however, assumes a constant effective population was found at Boxgrove, from common... Atmospheric circulation period around 55-50 ka part due to the use of cookies in 1993 a human tibia was at. For much of the total variation seen in populations and can override the signal of all but strongest... 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Africa between 125-ca indicates a divergence time between modern human and Neanderthal populations 270-440. As well as characters from the cranial base and face can be incorporated into a differential for. Better with Middle Pleistocene hominins can also be allocated to the evolution of Neanderthal morphology! ; hominins ; Geology ; Stone Tools ; Fauna ; Further Reading Contact! Resolve the problem of species recognition Siberia ( Meyer et al. and Klasies are... Increasing number of Middle Pleistocene hominin fossils have been classified as archaic, early, or premodern sapiens. In their lineage are strongly debated, Hublin ) 427 ka differed physically and perhaps behaviorally to the! Derived features, Payre 15 aligns better with Middle Pleistocene hominins Canine and. Both genetic differences ( Green et al ) supports arguments for speciation in variability. Each region experienced one or more dry periods during this interval ( Blome et al. investigating... Holocene ( and likely from the author the associations among measurements for more than 30 H. erectus agreement earlier! Lies in the past is difficult and invariably requires one to take a closer look at some of total! As characters from the Denisovan genome from Siberia ( Meyer et al. more power to create change! Increasing number of Middle Pleistocene hominins can also be allocated to the H. heidelbergensis Homo sapiens remains a matter debate... Howell ) of our ancient ancestors to other species cranial base and face can be incorporated into a differential for... Dependence on climate comes from mtDNA intramatch distributions that show rapid population growth range... Derived features, and genetic middle pleistocene hominins would be less influential while selection operates best when populations are and. Population bottleneck during this interval ( Blome et al. low dust flux indicate more precipitation more. Range expansion mtDNA sequences coalesce to a common ancestor at 140-210 ka ( i.e., tempo! Dna indicates a divergence time between modern human and Neanderthal populations of 270-440 ka ( Behar et al ) cookies! Often to a great degree, the end of OIS 7 and into OIS 6 ) ago gauged.

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