A previous study of human craniometrics suggested that approximately 50 individuals was necessary to ensure the stability of the P matrix estimations [84] using a similar number of variables—14—as this study (16–17 PCs for the A–C test). Summary. Humans’ most recent ancestor, the species that predated our kind, remains shrouded in mystery. Additional hominin fossils from the crucial time period of 4-3 million years ago must be discovered to conclusively determine the place of platyops in our evolution. Indeed, tests of neutral evolution revealed a role for natural selection in frontal bone diversification among H. erectus, but not H. sapiens populations. The hypotheses were assessed separately for the frontal and occipital bones both to maximal fossil sample sizes and to assess mosaic cranial evolution. This was assessed by calculating the angle as the arccosine of the dot product of the normalized vectors for pmax and PC 1 of a between-group PCA of the H. erectus palaeodeme means or H. sapiens population means. information for hominin species accumulates we are discovering that they can also have distinctive life histories that do not conform to any living model. The evolution of human intelligence is closely tied to the evolution of the human brain and to the origin of language.The timeline of human evolution spans approximately 9 million years, from the separation of the genus Pan until the emergence of behavioral modernity by 50,000 years ago. The H. erectus sample consisted of n = 23 for the occipital analysis and n = 22 for the frontal analysis (n = 27 total) subdivided into six spatially and temporally circumscribed palaeodemes (sensu [67]) of 2–6 individuals each (table 1 and figure 1). Conventional wisdom holds that H. erectus originated around 1.9 Ma in East Africa [16], where there are some of the oldest H. erectus sites and abundant evidence of more ancient Homo species (figure 1). I thank Monica Castro for help with data processing. Low genetic variation in recent non-African H. sapiens is attributed to a major population bottleneck during the recent human diaspora out of Africa [4,11], so higher genetic variation in H. erectus implies that this species did not experience the same drastic population bottleneck during its migration. Abbreviations in legend are from table 1. The extent to which recent H. sapiens and H. erectus converged in their population history has not been explored previously. Homo erectus shares with H. sapiens some key features of evolutionary history, including migration out of Africa to occupy a range of more seasonal and temperate habitats across Eurasia and Asia. This prediction is consistent with long-standing ideas about geographical and genetic isolation of regional H. erectus populations [32] and empirical evidence for neutral cranial diversification in other Homo species, including H. sapiens [56]. Figure 2. The timeline of human evolution outlines the major events in the evolutionary lineage of the modern human species, Homo sapiens, throughout the history of life, beginning some 4.2 billion years ago down to recent evolution within H. sapiens during and since the Last Glacial Period.. (Online version in colour. They existed for about 3.5 – 2.45… The Paleolithic preceded the Middle Stone Age, or Mesolithic Period; this nomenclature sometimes causes … Under a pure drift model, the regression coefficient (β) is expected to be 1, and deviations from this expectation can indicate non-random processes. At least one extinct hominin subclade, Paranthropus, has a pattern of ... package, or did the components evolve independently and incrementally? This angle measures divergence between the primary axes of within- and between-group variation. Inset: landmarks and semilandmarks from the frontal bone (yellow) and occipital bone (green) superimposed on the Sambungmacan 3 crania. If the latter two groups split prior to 500–700 ka [1–3], then this postdates the vast majority of fossils assigned to the other candidate for this position, Homo heidelbergensis s.l. Prediction 2: H. erectus populations are more divergent than recent H. sapiens populations. Ordination of the first two principal components of within-population variation in (a) occipital and (b) frontal bone shape. However, analysis of endocranial dimensions across African, Chinese and Indonesian H. erectus was unable to identify regional differences in brain shape [44], making this an unlikely explanation. The Hominidae (/ h ɒ ˈ m ɪ n ɪ d iː /), whose members are known as great apes or hominids (/ ˈ h ɒ m ɪ n ɪ d z /), are a taxonomic family of primates that includes eight extant species in four genera: Pongo, the Bornean, Sumatran and Tapanuli orangutan; Gorilla, the eastern and western gorilla; Pan, the common chimpanzee and the bonobo; and Homo, of which only modern humans remain.. Several revisions in … Shape changes from the negative to positive ends of PC 1 and PC 2 are illustrated for (c,d) occipital and (e,f) frontal bones. The question mark indicates some uncertainty regarding the attribution of Daka (BOU-VP-2/66) to H. erectus. Therefore, the recent H. sapiens pooled within-population P matrices for the occipital and frontal bones were used as proxies for H. erectus. Semilandmarks were slid to maximize geometric homology by minimizing the bending energy matrix [61] and all landmarks/semilandmarks were superimposed by generalized Procrustes analysis separately for each bone [62]. Human evolution - Human evolution - Increasing brain size: Because more complete fossil heads than hands are available, it is easier to model increased brain size in parallel with the rich record of artifacts from the Paleolithic Period (c. 3.3 million to 10,000 years ago), popularly known as the Old Stone Age. Most workers, however, view H. erectus as either a broadly distributed and polytypic species [13,14,23,44] or a specialized Asian lineage, in which case the earlier African and possibly Georgian fossils are assigned to a separate species, H. ergaster [16,45,46]. The results highlight distinct evolutionary histories for the frontal and occipital bones in H. erectus. How did hominids change as they evolved? Many scenarios imply periods of regional isolation that facilitated in situ phyletic transformation through genetic drift and/or environmental adaptation but with sufficient gene flow to maintain species cohesion [15,23,25–34]. The first PC (horizontal axis) corresponds to the direction of greatest within-population variation (or the line of least evolutionary resistance) (pmax). The evidence for selection on the frontal bone is contingent on the underlying taxonomy and applies only to the broader definition of H. erectus which includes Asian, African and Eurasian groups, but not to a more restrictive, Asian-only definition of H. erectus. Colours for H. erectus are as in figure 2; the grey represents the three human populations. )Download figureOpen in new tabDownload powerPoint, Figure 3. Landmarks are indicated by larger spheres than semilandmarks. Key H. erectus cranial fossils are labelled, with bold signifying those included in the current study. The magnitude of cranial variation among populations is higher in H. erectus than recent H. sapiens occupying a similar geographical range. The timeline reflects the … Black rectangles indicate time ranges of samples included in each palaeodeme, while thinner grey rectangles indicate the full time range of fossils assigned to H. erectus in those regions. Ordination of the first two principal components of individual variation in (a) occipital and (b) frontal bone shape for geographically comparable samples of Homo erectus and Homo sapiens. Table 1. aBold indicates that a fossil was included in the occipital and frontal bone analyses. (but not when H. erectus is restricted to only the Asian lineages). Enter your email address below and we will send you your username, If the address matches an existing account you will receive an email with instructions to retrieve your username, Department of Anatomy, College of Graduate Studies, Midwestern University, Glendale, AZ 85308, USA. This evolutionary analysis is designed to provide a basic account of the evolution of language in our species. Human evolution took place as new genetic variations in early ancestor populations favored new abilities to adapt to environmental change and so altered the human way of life. Homo erectus is the first hominin species with a truly cosmopolitan distribution and resembles recent humans in its broad spatial distribution. Figure 3. First, the centroids of all six H. erectus palaeodemes (and all six H. sapiens populations) were projected onto those PCs calculated from the pooled within-population covariance matrix of recent H. sapiens accounting for greater than or equal to 1% of the variation. H. antecessor, which lived from just over a million to around 800,000 years ago in Africa and Europe, was actually a sister lineage to our own, closely related but not our ancestor. However, evolutionary change can follow an axis of high intrapopulation variability that is not perfectly aligned with pmax if the variance is spread across several dimensions [88]. Results indicate that H. erectus had higher individual and group variation than Homo sapiens, probably reflecting different levels of genetic diversity and population history in these spatially disperse species. If you are a Zinio, Nook, Kindle, Apple, or Google Play subscriber, you can enter your website access code to gain subscriber access. The first hominin species, a line that eventually leads to humans, may have emerged in Europe 7.2 million years ago and not Africa—the most widely accepted starting point for our ancestors. Untold Homo species contributed to the eventual emergence of both Neanderthals and Homo sapiens. genetic drift) given the greater time since divergence among H. erectus populations [55]. Both pmax and the A–C test assume that the P matrices of H. erectus and H. sapiens are proportional. Evolutionary morphological analyses adapted from quantitative genetics are already yielding valuable insights into human evolution, including recognition of a greater role for neutral processes (e.g. Hominins are primate species that are more closely related to humans than chimps so they are classified on the human branch, they did not evolve in a straight line to homo sapiens, several hominin species co existed at certain points, many lineages died out. This story is part of an ongoing series exploring questions about human origins. This measure scales linearly with neutral genetic variation across hominoids [54]. genetic drift) in human evolution than previously thought [49–51], but also support for selection in the evolution of the hominin face and postcranial skeleton [52,53]. Published by the Royal Society. The origin of the genus Homo in Africa signals the beginning of the shift from increasingly bipedal apes to primitive, large-brained, stone tool-making, meat-eaters that traveled far and wide. )Download figureOpen in new tabDownload powerPoint, Figure 2. Shape differences reflect well-documented distinctions between the species: H. erectus has a flatter frontal squama with a taller supraorbital torus and greater constriction posterior to the supraorbital torus, while the occipital bone is relatively wider but more tightly angled in its midline profile (figure 2c,e). Colours for H. erectus are as in figure 2; the grey represents the three human populations. Convex hulls encompass individual variation for H. erectus palaeodemes and H. sapiens populations and outlined circles are the centroids of each group. Here’s the Top 10 Science Stories You Missed This Year (While You Were Distracted by COVID-19), Science Board Game Reviews: Wingspan, Terraforming Mars, Endangered, and Neanderthal, America's Oldest City Is Not Where You'd Expect. Your website access code is located in the upper right corner of the Table of Contents page of your digital edition. Small fossil samples introduce uncertainty into estimates of palaeodeme averages, but relatively strong morphological homogeneity within palaeodemes [15] suggests that we are sampling near the group centroids in most cases. While placed in the Homo genus, they have not yet been given a species classification as no physical description exists. The A–C test was applied to 17 (occipital) or 16 (frontal) PC axes, accounting for greater than 90% of total shape variation in each bone. Homo sapiens therefore serves as a useful model for thinking about H. erectus population history. The quantitative genetics tests for Prediction 3 require an estimate of the G (or P) matrix, which contains the within-population additive genetic (or phenotypic) variances and covariances of traits and is essential for formulating predictions regarding the magnitude and direction of between-population variation under certain microevolutionary conditions [79]. All Homo erectus and the three recent human populations used to calculate the pooled within-population covariance matrix were projected on to these axes. Read our privacy policy. Next, we’ll summarize the current state of the evidence as it pertains to hominin evolution, and place the origins of our own species in that context. PNAS. This conjecture finds some support in the generally conserved nature of the G matrix within Homo [89,90], although some minor differences in integration between archaic and modern Homo have been documented [89,91]. (Online version in colour.). For the current study, cranial fossils were sorted into six major palaeodemes from sites across Africa and Asia spanning 1.8–0.1 Ma. This may imply a more prominent role of natural selection in H. erectus. Stratified resampling without replacement was applied to the geographically matched H. sapiens sample to produce 1000 samples of the same size (n = 23 for occipital bone and n = 27 for frontal bone) and population composition as the H. erectus sample (electronic supplementary material, Supplemental Methods). Abbreviations in legend are from table 1. Abbreviations for labelled fossils are from table 1. A slope less than 1.0 implies stabilizing selection in directions of high within-population variance and/or directional selection in directions of limited within-population variance. The evolution of Homo sapiens, its ancestors, and closely related species from the last common ancestor with chimpanzees onward (~7 million years ago to present). This H. erectus value was compared to the empirical distribution of 1000 estimates of between-population variation for humans to determine the probability that the H. erectus value exceeded that of H. sapiens. Less than a quarter of the total variation in occipital and frontal shape is concentrated onto their respective pmax vectors. Some scientists subscribe to the theory of species mate recognition, in which members of the same species “recognize” one another as mates through courtship rituals, breeding seasons, or protein compatibility. Hominids are believed to be an evolution of catarrhines (a primate parvord with the nose down), and is made up of four genera and seven species. Chronology of Homo erectus from west to east: East Africa, West Asia, Southeast and East Asia. This is supported by the stronger signal of natural selection in frontal bone diversification among populations compared to the occipital bone, but also the discordant patterns of shape differentiation among demes for these two bones (electronic supplementary material, figure S1). This hominin sympatry (multiple species in the same geographic space and time) eventually ended ... FOSSIL HOMININS | Genus Homo – Pleistocene Hominin Evolution Lab 209 Cranial Characteristics of Homo erectus 1. Human pelvis and long bones reveal differential preservation of ancient population history and migration out of Africa, Evolutionary processes shaping diversity across the Homo lineage, Divergent patterns of integration and reduced constraint in the human hip and the origins of bipedalism, Hominoid intraspecific cranial variation mirrors neutral genetic diversity, Unconstrained cranial evolution in Neandertals and modern humans compared to common chimpanzees, Grist for Riedl's mill: A network model perspective on the integration and modularity of the human skull, The evolutionary role of modularity and integration in the hominoid cranium, The evolution of modularity in the mammalian skull I: morphological integration patterns and magnitudes, Principles for the virtual reconstruction of hominin crania, Extensions of the Procrustes method for the optimal superimposition of landmarks, geomorph: an r package for the collection and analysis of geometric morphometric shape data, The R package sampling, a software tool for training in official statistics and survey sampling 2006, Paleo-demes, species clades, and extinctions in the Pleistocene hominin record, Chronostratigraphy of KNM-ER 3733 and other Area 104 hominins from Koobi Fora, New single crystal 40Ar/39Ar ages improve time scale for deposition of the Omo Group, Omo–Turkana Basin, East Africa, Stratigraphic context of fossil hominids from the Omo group deposits, northern Turkana Basin, Kenya and Ethiopia, High-resolution record of the Matuyama–Brunhes transition constrains the age of Javanese, Age control of the first appearance datum for Javanese, Age of the 20 Meter Solo River Terrace, Java, Indonesia and the survival of, 40Ar/39Ar dating and phytolith analysis of the Early Pleistocene sequence of Kvemo-Orozmani (Republic of Georgia): chronological and palaeoecological implications for the hominin site of Dmanisi, Global and local perspectives on cranial shape variation in Indonesian, Understanding the evolution and stability of the G-matrix, Evolvability and craniofacial diversification in genus, Comparison of genotypic and phenotypic correlations: Cheverud's conjecture in humans, Pervasive genetic integration directs the evolution of human skull shape, A comparison of genetic and phenotypic correlations, Testing the equivalence of modern human cranial covariance structure: implications for bioarchaeological applications, Adaptive radiation along genetic lines of least resistance, Size as a line of least evolutionary resistance: diet and adaptive morphological radiation in New World monkeys, Conserved phenotypic variation patterns, evolution along lines of least resistance, and departure due to selection in fossil rodents, Measuring and comparing evolvability and constraint in multivariate characters. The question mark indicates some uncertainty regarding the attribution of Daka (BOU-VP-2/66) to H. erectus. The H. erectus value is compared to the empirical distribution of modern H. sapiens values to determine the probability that H. erectus exhibits similar intraspecific variation as H. sapiens. Three-dimensional shape data from the occipital and frontal bones were used to compare intraspecific variation and test evolutionary hypotheses. Article CAS … Three-dimensional landmarks and semilandmarks acquired from the occipital and frontal bones (see figure 1 and electronic supplementary material, Supplemental Methods) in H. erectus and recent H. sapiens are the raw data used in subsequent analyses. References. Quantitative genetics models provide a means of interrogating aspects of long-standing H. erectus population history narratives. Therefore, the Ackerman–Cheverud (A–C) test was used to evaluate whether divergence among populations (B) is proportional to within-population variation (P) across multiple dimensions [49]. Two tests were used to evaluate Prediction 3. Intraspecific variation in H. erectus (red dashed line) compared to resampled values for recent H. sapiens using a histogram with a probability density curve for (c) occipital and (d) frontal bone shape. Recent H. sapiens, in contrast, exhibited a consistently neutral pattern of between-population divergence in the shape of both bones in agreement with previous studies arguing that the human cranium preserves a strong population history signal [95,96]. One scenario consistent with these findings is climate-driven facial adaptation in H. erectus, which is reflected in the frontal bone through integration with the orbits. But what bridged H. erectus and our own species is unclear. Between-population variation in H. erectus (red dashed line) compared to resampled values for recent H. sapiens using a histogram with a probability density curve for (g) occipital and (h) frontal bone shape. The H. erectus occipital SEV value was 27% larger than the average H. sapiens SEV and was higher than 98% of those values. Thus, local selection in H. erectus, perhaps related to climatic adaptation in the face, could have produced both the overall higher variation, greater population differentiation and the signal of selection in the analyses presented here. O and F indicate inclusion only in the occipital bone or frontal bone analysis, respectively. Origin of hominids . The H. erectus frontal bone SEV was 67% higher than the average SEV for recent H. sapiens and exceeded 100% of the resampled H. sapiens values (figure 3c,d). The four Asian palaeodemes (EAS, ESA, LSA-N, LSA-S) showed greater affinity in their frontal bone shape and there was greater variation between the oldest palaeodemes (WAS and EAF) (electronic supplementary material, figure S1b). Discover more. The sum of eigenvalues (SEV) was calculated from both the original H. erectus sample and the 1000 H. sapiens samples to test Prediction 1. Between-population variation in H. erectus (red dashed line) compared to resampled values for recent H. sapiens using a histogram with a probability density curve for (g) occipital and (h) frontal bone shape. Ordination of the first two principal components of individual variation in (a) occipital and (b) frontal bone shape for geographically comparable samples of Homo erectus and Homo sapiens. The Neanderthal ‘chignon’: variation, integration, and homology, Geometric variation of the frontal squama in the genus, Phenotypic variance inflation in fossil samples: an empirical assessment, Congruence of individual cranial bone morphology and neutral molecular affinity patterns in modern humans, The relative role of drift and selection in shaping the human skull, Population-specific deviations of global human craniometric variation from a neutral model, Random genetic drift, natural selection, and noise in human cranial evolution, Measuring the effects of farming on human skull morphology, Changes in human skull morphology across the agricultural transition are consistent with softer diets in preindustrial farming groups, Human cranial anatomy and the differential preservation of population history and climate signatures, Interspecific perspective on mechanical and nonmechanical models of primate circumorbital morphology, Shape analysis of spatial relationships between orbito-ocular and endocranial structures in modern humans and fossil hominids, https://dx.doi.org/10.5061/dryad.5qfttdz32, https://doi.org/10.6084/m9.figshare.c.5252529, doi:10.1666/0094-8373(2004)030<0487:PVIIFS>2.0.CO;2. Neutral evolution cannot be rejected for the occipital bone, but selection is implicated in the evolution of frontal bone shape for H. erectus s.l. These same population history events left an imprint on the species' cranial phenotype [5,10–12]. “Independent evolution of knuckle-walking in African apes shows that humans did not evolve from a knuckle-walking ancestor ... Leakey (2001) proposes that the fossil represents an entirely new hominin species and genus, while others classify it as a separate species of Australopithecus, Australopithecus platyops, and yet others interpret it as an individual of Australopithecus afarensis. These species are unique among hominins in experiencing a major migration out of Africa to occupy more diverse habitats across Eurasia and East and Southeast Asia. It has not been uncommon, for example, to identify a taxon of living mammals which face challenges similar to those faced We challenge the view that our species, Homo sapiens, evolved within a single population and/or region of Africa. Hence Homo erectus appears at a time in which multiple hominin species existed in Africa. … This early part of the human genus is represented by three species: H. habilis, H. rudolfensis, and H. erectus. Want it all? The SEV is equivalent to the sum of all trait variances and is used here to measure intraspecific shape variation. Ch 10 Early Hominin Origins and Evolution questionSahelanthropus tchadensis answerThe earliest pre-australopithecine species found in central Africa with possible evidence of bipedalism. (Online version in colour. (Online version in colour.). Your email address is used to log in and will not be shared or sold. Modularity between the cranial vault and the face may have facilitated these different evolutionary paths [92]. 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