Pleistocene Hominins Research Articles

Early and Middle Pleistocene hominins from Atapuerca (Spain) show differences in dental developmental patterns

AbstractThe Bayesian statistical approach considers teeth as forming a developmental module, as opposed to a tooth‐by‐tooth analysis. This approach has been employed to analyze Upper Pleistocene hominins, including Neandertals and some anatomically modern humans, but never earlier populations. Here, we show its application on five hominins from the TD6.2 level of the Gran Dolina site (Homo antecessor, Early Pleistocene) and the Sima de los Huesos site (Middle Pleistocene) of the Sierra de Atapuerca (Burgos, northern Spain). Our results show an advanced development of the third molars in both populations with respect to modern Homo sapiens. In addition, the Sima de los Huesos hominins differ from H. sapiens and H. antecessor in the relatively advanced development of their second molar. The relative mineralization of I1/M1 in H. antecessor appears to be similar to that of modern humans, as opposed to that of Neandertals, which appear to be unique. These observations, combined with reduced enamel formation times and the advanced development of the third molars, appear to indicate a shorter ontogenetic period in the hominins from Gran Dolina and Sima de los Huesos in comparison to modern human average.

The earliest Pleistocene record of a large-bodied hominin from the Levant supports two out-of-Africa dispersal events

The paucity of early Pleistocene hominin fossils in Eurasia hinders an in-depth discussion on their paleobiology and paleoecology. Here we report on the earliest large-bodied hominin remains from the Levantine corridor: a juvenile vertebra (UB 10749) from the early Pleistocene site of ‘Ubeidiya, Israel, discovered during a reanalysis of the faunal remains. UB 10749 is a complete lower lumbar vertebral body, with morphological characteristics consistent with Homo sp. Our analysis indicates that UB-10749 was a 6- to 12-year-old child at death, displaying delayed ossification pattern compared with modern humans. Its predicted adult size is comparable to other early Pleistocene large-bodied hominins from Africa. Paleobiological differences between UB 10749 and other early Eurasian hominins supports at least two distinct out-of-Africa dispersal events. This observation corresponds with variants of lithic traditions (Oldowan; Acheulian) as well as various ecological niches across early Pleistocene sites in Eurasia.

Evidence of Middle Pleistocene hominin migration in the Qinling Mountains (central China) from the Miaokou Paleolithic site

Evidence of Middle Pleistocene hominin migration in the Qinling Mountains (central China) from the Miaokou Paleolithic site

Possible Pleistocene hominin tracks from South Africa’s west coast

Two probable tracks have been identified on the ceiling of a small overhang in the Pleistocene Langebaan Formation on South Africa’s west coast. They may have been made by a hominin trackmaker. They appear to have been registered at walking speed on a level, sandy dune substrate. Three tracks, attributed to Homo sapiens, were previously identified near Langebaan in 1995, and were popularly labelled ‘Eve’s Footprints’. The new identification of possible hominin tracks near Langebaan is the second from South Africa’s west coast. This discovery adds to the sparse but growing global record of possible hominin tracks preserved in aeolianites.Significance: Two probable fossil tracks have been identified on the ceiling of an overhang near Langebaan. The tracks may have been made by a human walking on a level dune surface during the Pleistocene. This discovery is the second of its kind on the west coast, and complements the 1995 identification near Langebaan of Pleistocene fossil tracks attributed to humans.

Exploring Late Pleistocene hominin dispersals, coexistence and extinction with agent-based multi-factor models

Toward the end of the Pleistocene, archaic humans in Eurasia such as the Neanderthals and Denisovans were completely replaced by anatomically modern humans dispersing from Africa. The causes underlying the replacement and extinction processes remain controversial, especially regarding the relative importance of random events, and anthropogenic and environmental factors. Here, we use the most comprehensive agent-based modeling framework to date for exploring Late Pleistocene human population dynamics under realistic time-evolving environmental conditions. Model simulations suggest multiple out-of-Africa dispersals. Most of these resulted in only partial replacement of Eurasians and long-term coexistence of spatially structured archaic and modern populations in Eurasia. Moreover, a comparison of empirical and model data suggests that the best-documented extinction process – that of the Neanderthals – did not have a single overarching cause, but spatially and temporally diverse causes and mechanisms, such as environmental fluctuations, and asymmetry in resource exploitation efficiency and reproductive rates. When viewed in isolation, various population properties have central importance for replacements, but their true importance can only be understood in comparison and with interactions with other properties.

Connections between the Levant and the Balkans in the late Middle Pleistocene: Archaeological findings from Velika and Mala Balanica Caves (Serbia)

Major changes in the technological, economic, and social behavior of Middle Pleistocene hominins occurred at the onset of the Middle Paleolithic, 400–200 ka. However, until recently it was not possible to establish when, where, and how certain forms of Middle Paleolithic behavior appeared and spread into Southeastern Europe, mainly owing to gaps in the Paleolithic record. Here we report new results of dating, material culture, and the archaeological context of finds from the Balanica Cave Complex in Sićevo (Serbia). Two methods—thermoluminescence and electron spin resonance—were used to date the sequence. The geoarchaeological context was examined through sedimentology, micromorphology, and spatial analysis. Microfaunal remains were used to constrain the dates within an ecological zone, whereas macrofauna was analyzed for taxonomy and taphonomy to examine the source of accumulation and hominin behavior. Technological and typological features of the lithic assemblage were used to characterize lithic production at the site. Materials recovered from Layer 3 in Velika Balanica and from Layer 2 in Mala Balanica, both dated to MIS 9–7, include a distinctive set of archaeological assemblages which resemble contemporaneous Yabrudian assemblages from the Levant in important ways, and which are unlike contemporary material from the surrounding regions. In Velika Balanica, the lithic assemblages are associated with a large fireplace containing evidence of human activities similar to those from Qesem Cave (Israel). Dental remains uncovered in the same layer are consistent with Neanderthals. These findings suggest that the end of the Middle Pleistocene (before 300–240 ka) saw population movement and/or cultural transmission between Southwest Asia and the Balkans, which led eventually to a transfer of technology between Middle Eastern and European hominin populations and contributed to the shaping of Neanderthal behaviors throughout the eastern and northern Mediterranean.

Late Acheulean Lithic Assemblages From Locality 010 at Gürgürbaba Hill (Eastern Anatolia)

The province of Van in north-eastern Turkey served as a land bridge between Africa and Eurasia during the Palaeolithic. The region is of particular relevance for understanding the movement of hominins between these continents. This study concerns the lithic remains from a locality at Gürgürbaba Hill, named Locality 010, north of the village of Ulupamir (Erciş district). Locality 010 was dated to 311±32 kya by terrestrial cosmogenic nuclides method, which coincides with Marine Isotope Stage 9 (MIS 9), a Middle Pleistocene interglacial period. The assemblage from this site is attributed to the Late Acheulean and resembles that of the southern Caucasus. This similarity indicates that the artefacts from Locality 010 were probably produced by late Lower Palaeolithic technology in a broad sense. These findings suggest local adaptations of late Middle Pleistocene hominins to high plateau environments.

Evolution of cranial capacity revisited: A view from the late Middle Pleistocene cranium from Xujiayao, China

The Late Middle Pleistocene hominin fossils from the Xujiayao site in northern China have been closely studied in light of their morphological variability. However, all previous studies have focused on separated cranial fragments. Here, we report the first reconstruction of a fairly complete posterior cranium, Xujiayao 6 (XJY 6), confidently dated to ∼200–160 ka, which facilitated an assessment of its overall cranial size. XJY 6 was reconstructed from three of the original fragments—the PA1486 (No.7/XJY 6a) occipital bone, PA1490 (No.10/XJY 6b) right parietal bone, and PA1498 (No.17/XJY 15) left temporal bone—which originated from the same young adult individual. The XJY 6 endocranial capacity, estimated by measuring endocranial volume, was estimated using multiple regression formulae derived from ectocranial and endocranial measurements on select samples of Pleistocene hominins and recent modern humans. The results indicate that the larger pooled sample of both Pleistocene and recent modern humans was more robust for the endocranial capacity estimate. Based on the pooled sample using the ectocranial and endocranial measurements, we conservatively estimate the XJY 6 endocranial volume to be ∼1700 cm3 with a 95% confidence interval of 1555–1781 cm3. This is close to Xuchang 1, which dates to 125–105 ka and whose endocranial volume is ∼1800 cm3. Thus, XJY 6 provides the earliest evidence of a brain size that falls in the upper range of Neanderthals and modern Homo sapiens. XJY 6, together with Xuchang 1, Homo floresiensis, Homo luzonensis, and Homo naledi, challenge the general pattern that brain size gradually increases over geological time. This study also finds that hominin brain size expansion occurred at different rates across time and space.

Early Pleistocene hominin subsistence behaviors in South Africa: Evidence from the hominin-bearing deposit of Cooper's D (Bloubank Valley, South Africa)

Evidence of the consumption of meat through hunting or scavenging by Early Pleistocene hominins is scarce, particularly in South Africa. Moreover, the interpretations of taphonomic evidence are subject to an important discussion commonly called the ‘hunting-vs-scavenging debate.’ Until today, only the Swartkrans Members 1–3 site has yielded a butchered bone assemblage large enough to permit reconstruction of carcass acquisition strategies by Early Pleistocene hominins in South Africa. This leaves an information gap between 1.4 and 1.0 Ma. Here, we provide the first evidence of meat consumption by hominins during this gap, based on the zooarchaeological study of the large mammal bone assemblage recovered from the Cooper's D site, South Africa. Based on skeletal part representation, our results show density-mediated attrition of bovid bones due to predepositional and postdepositional destruction. We argue that this attrition is the result of both abiotic (i.e., decalcification) and biotic (i.e., carnivore ravaging) processes. Bovid mortality profiles point out the involvement of ambush predators such as large felids. Bone surface modifications also indicate that the assemblage has been accumulated mostly by carnivores but with some hominin involvement as well. We observe all the stages of animal carcass processing (skinning, disarticulation, defleshing, marrow extraction) as well as the exploitation of a diversity of prey size classes at both Swartkrans Members 1–3 and Cooper's D. Thus, our study shows the importance of the Cooper's D bone assemblage for understanding Early Pleistocene hominin subsistence behaviors. Moreover, this article highlights the need for including long bone flake specimens in the analysis of large bone assemblages from South African caves to better understand the Early Pleistocene hominin bone damage record.

Mechanical loading of primate fingers on vertical rock surfaces

Mechanical loading of finger bones (phalanges) can induce angular curvature, which benefits arboreal primates by dissipating forces and economising the recruitment of muscles during climbing. The recent discovery of extremely curved phalanges in a hominin, Homo naledi, is puzzling, for it suggests life in an arboreal milieu, or, alternatively, habitual climbing on vertical rock surfaces. The importance of climbing rock walls is attested by several populations of baboons, one of which uses a 7-m vertical surface to enter and exit Dronkvlei Cave, De Hoop Nature Reserve, South Africa. This rock surface is an attractive model for estimating the probability of extreme mechanical loading on the phalanges of rock-climbing primates. Here we use three-dimensional photogrammetry to show that 82–91% of the climbable surface would generate high forces on the flexor tendon pulley system and severely load the phalanges of baboons and H. naledi. If such proportions are representative of vertical rock surfaces elsewhere, it may be sufficient to induce stress-mitigating curvature in the phalanges of primates.Significance: We present the first three-dimensional photogrammetric analysis of a vertical rock surface climbed by a non-human primate, the chacma baboon (Papio ursinus). Our results show that a large proportion of a vertical rock wall would compel crimp and slope hand positions during climbing – grips that could explain the extraordinary phalangeal curvature expressed by a Middle Pleistocene hominin, Homo naledi.

Geometric morphometric variability in the supraorbital and orbital region of Middle Pleistocene hominins: Implications for the taxonomy and evolution of later Homo

This study assessed variation in the supraorbital and orbital region of the Middle Pleistocene hominins (MPHs), sometimes called Homo heidelbergensis s.l., to test whether it matched the expectations of intraspecific variation. The morphological distinctiveness and relative variation of this region, which is relatively well represented in the hominin fossil record, was analyzed quantitatively in a comparative taxonomic framework. Coordinates of 230 3D landmarks (20) and sliding semilandmarks (210) were collected from 704 specimens from species of Homo, Australopithecus, Paranthropus, Gorilla, Pan, Papio, and Macaca. Results showed that the MPHs had expected levels of morphological distinctiveness and intragroup and intergroup variation in supraorbital and orbital morphology, relative to commonly recognized non-hominin catarrhine species. However, the Procrustes distances between this group and H. sapiens were significantly higher than expected for two closely related catarrhine species. Furthermore, this study showed that variation within the MPH could be similarly well contained within existing hypodigms of H. sapiens, H. neanderthalensis, and H. erectus s.l. Although quantitative assessment of supraorbital and orbital morphology did not allow differentiation between taxonomic hypotheses in later Homo, it could be used to test individual taxonomic affiliation and identify potentially anomalous individuals. This study confirmed a complicated pattern of supraorbital and orbital morphology in the MPH fossil record and raises further questions over our understanding of the speciation of H. sapiens and H. neanderthalensis and taxonomic diversity in later Homo.

Comparative dental study between Homo antecessor and Chinese Homo erectus: Nonmetric features and geometric morphometrics

The Chinese Middle Pleistocene fossils from Hexian, Xichuan, Yiyuan, and Zhoukoudian have been generally classified as Homo erectus s.s. These hominins share some primitive features with other Homo specimens, but they also display unique cranial and dental traits. Thus, the Chinese Middle Pleistocene hominins share with other European and Asian hominin populations the so-called ‘Eurasian dental pattern’. The late Early Pleistocene hominins from Gran Dolina-TD6.2 (Spain), representing the species Homo antecessor, also exhibit the Eurasian dental pattern, which may suggest common roots. To assess phylogenetic affinities of these two taxa, we evaluated and compared nonmetric and metric dental features and interpreted morphological differences within a comparative hominin framework. We determined that the robust roots of the molars, the shelf-like protostylid, the dendrite-like pattern of the enamel-dentine junction surface of the upper fourth premolars and molars, the strongly folded dentine of the labial surface of the upper incisors, and the rare occurrence of a mid-trigonid crest in the lower molars, are all characteristic of Chinese H. erectus. With regard to H. antecessor, we observed the consistent expression of a continuous mid-trigonid crest, the absence of a cingulum in the upper canines, a complex root pattern of the lower premolars, and a rhomboidal occlusal contour and occlusal polygon and protrusion in the external outline of a large a bulging hypocone in the first and second upper molars. Using two-dimensional geometric morphometrics, we further demonstrated that H. antecessor falls outside the range of variation of Chinese H. erectus for occlusal crown outline shape, the orientation of occlusal grooves, and relative locations of anterior and posterior foveae in the P4s, P3s, M1s, M2s, and M2s. Given their geographic and temporal separation, the differences between these two species suggest their divergence occurred at some point in the Early Pleistocene, and thereafter they followed different evolutionary paths.

Resolving the "muddle in the middle": The case for Homo bodoensis sp. nov.

Recent developments in the field of palaeoanthropology necessitate the suppression of two hominin taxa and the introduction of a new species of hominins to help resolve the current nebulous state of Middle Pleistocene (Chibanian) hominin taxonomy. In particular, the poorly defined and variably understood hominin taxa Homo heidelbergensis (both sensu stricto and sensu lato) and Homo rhodesiensis need to be abandoned as they fail to reflect the full range of hominin variability in the Middle Pleistocene. Instead, we propose: (1) introduction of a new taxon, Homo bodoensis sp. nov., as an early Middle Pleistocene ancestor of the Homo sapiens lineage, with a pan‐African distribution that extends into the eastern Mediterranean (Southeast Europe and the Levant); (2) that many of the fossils from Western Europe (e.g. Sima de los Huesos) currently assigned to H. heidelbergensis s.s. be reassigned to Homo neanderthalensis to reflect the early appearance of Neanderthal derived traits in the Middle Pleistocene in the region; and (3) that the Middle Pleistocene Asian fossils, particularly from China, likely represent a different lineage altogether.

First Evidence of Pleistocene Archaeology on the Neyshabur Plain and its Role in Reconstructing the Dispersal of Hominins on the Northeastern Iranian Plateau

The northeastern Iranian Plateau is considered a leading region in Paleolithic studies. The history of Paleolithic research in this region dates back to the mid-20th century. However, unlike the western and, to some extent, the central part of the Iranian Plateau, only a handful of sites have been identifi ed in the northeastern part. Field studies conducted on the Neyshabur plain have provided some of the only Paleolithic evidence at four locations in the foothills of the Binalud Mountains: Dar Behesht, Mushan Tappeh, Ali Abad, and Qezel Tappeh. Our research aims to assess this evidence, provide a revised typology of Pleistocene artifacts from the Neyshabur plain, and also study the role of these and other fi nds in the area and analyze their signifi cance in terms of the dispersal of Pleistocene hominin populations. We propose two main corridors on the northeastern Iranian Plateau assumed to have been infl uential in the dispersal of human ancestors.

A multi-proxy approach to exploring Homo sapiens\u2019 arrival, environments and adaptations in Southeast Asia

The capability of Pleistocene hominins to successfully adapt to different types of tropical forested environments has long been debated. In order to investigate environmental changes in Southeast Asia during a critical period for the turnover of hominin species, we analysed palaeoenvironmental proxies from five late Middle to Late Pleistocene faunas. Human teeth discoveries have been reported at Duoi U’Oi, Vietnam (70–60 ka) and Nam Lot, Laos (86–72 ka). However, the use of palaeoproteomics allowed us to discard the latter, and, to date, no human remains older than ~ 70 ka are documented in the area. Our findings indicate that tropical rainforests were highly sensitive to climatic changes over that period, with significant fluctuations of the canopy forests. Locally, large-bodied faunas were resilient to these fluctuations until the cooling period of the Marine Isotope Stage 4 (MIS 4; 74–59 ka) that transformed the overall biotope. Then, under strong selective pressures, populations with new phenotypic characteristics emerged while some other species disappeared. We argue that this climate-driven shift offered new foraging opportunities for hominins in a novel rainforest environment and was most likely a key factor in the settlement and dispersal of our species during MIS 4 in SE Asia.

Morphological description and evolutionary significance of 300 ka hominin facial bones from Hualongdong, China

Late Middle Pleistocene hominins in Africa displaying key modern morphologies by 315 ka are claimed as the earliest Homo sapiens. Evolutionary relationships among East Asian hominins appear complex due to a growing fossil record of late Middle Pleistocene hominins from the region, reflecting mosaic morphologies that contribute to a lack of consensus on when and how the transition to modern humans transpired. Newly discovered 300 ka hominin fossils from Hualongdong, China, provide further evidence to clarify these relationships in the region. In this study, facial morphology of the juvenile partial cranium (HLD 6) is described and qualitatively and quantitatively compared with that of other key Early, Middle, and Late Pleistocene hominins from East Asia, Africa, West Asia, and Europe and with a sample of modern humans. Qualitatively, facial morphology of HLD 6 resembles that of Early and Middle Pleistocene hominins from Zhoukoudian, Nanjing, Dali, and Jinniushan in China, as well as others from Java, Africa, and Europe in some of these features (e.g., supraorbital and malar regions), and Late Pleistocene hominins and modern humans from East Asia, Africa, and Europe in other features (e.g., weak prognathism, flat face and features in nasal and hard plate regions). Comparisons of HLD 6 measurements to group means and multivariate analyses support close affinities of HLD 6 to Late Pleistocene hominins and modern humans. Expression of a mosaic morphological pattern in the HLD 6 facial skeleton further complicates evolutionary interpretations of regional morphological diversity in East Asia. The prevalence of modern features in HLD 6 suggests that the hominin population to which HLD 6 belonged may represent the earliest pre-modern humans in East Asia. Thus, the transition from archaic to modern morphology in East Asian hominins may have occurred at least by 300 ka, which is 80,000 to 100,000 years earlier than previously recognized.

Middle Pleistocene environments, landscapes and tephrostratigraphy of the Armenian Highlands: evidence from Bird Farm 1, Hrazdan Valley

ABSTRACTThe significance of the southern Caucasus in understanding Pleistocene hominin expansions is well established. However, the palaeoenvironments in which Palaeolithic occupation of the region took place are presently poorly defined. The Hrazdan river valley, Armenian Highlands, contains a rich Palaeolithic record alongside Middle Pleistocene volcanic, fluvial and lacustrine strata, and thus offer exciting potential for palaeoenvironmental reconstruction. We present the first results of sedimentological, geochemical, tephrostratigraphical and biological (diatoms) study of the sequence of Bird Farm 1, located in the central part of the valley. These data show six phases of landscape development during the interval 440–200 ka. The sequence represents the first quantitative Pleistocene diatom record from the Armenian Highlands and the southern Caucasus, and indicates the persistence of a deep, stratified lacustrine system, with evidence for changing lake productivity that is tentatively linked to climate. Furthermore, major element chemical characterization of visible and crypto‐tephra horizons in the sequence enables the first stages of the development of a regional tephrostratigraphy. Together, the evidence from Bird Farm 1 demonstrates the importance of lacustrine archives in the region for palaeoenvironmental reconstruction and highlights the potential for linkages between archives on both a local and regional scale.

Multiple hominin dispersals into Southwest Asia over the past 400,000 years

Pleistocene hominin dispersals out of, and back into, Africa necessarily involved traversing the diverse and often challenging environments of Southwest Asia1–4. Archaeological and palaeontological records from the Levantine woodland zone document major biological and cultural shifts, such as alternating occupations by Homo sapiens and Neanderthals. However, Late Quaternary cultural, biological and environmental records from the vast arid zone that constitutes most of Southwest Asia remain scarce, limiting regional-scale insights into changes in hominin demography and behaviour1,2,5. Here we report a series of dated palaeolake sequences, associated with stone tool assemblages and vertebrate fossils, from the Khall Amayshan 4 and Jubbah basins in the Nefud Desert. These findings, including the oldest dated hominin occupations in Arabia, reveal at least five hominin expansions into the Arabian interior, coinciding with brief ‘green’ windows of reduced aridity approximately 400, 300, 200, 130–75 and 55 thousand years ago. Each occupation phase is characterized by a distinct form of material culture, indicating colonization by diverse hominin groups, and a lack of long-term Southwest Asian population continuity. Within a general pattern of African and Eurasian hominin groups being separated by Pleistocene Saharo-Arabian aridity, our findings reveal the tempo and character of climatically modulated windows for dispersal and admixture.

The bony labyrinth of Qafzeh 25 Homo sapiens from Israel

The bony labyrinth has received growing attention in the field of human evolution as it is a useful phylogenetic indicator in hominins and is particularly useful for distinguishing anatomically modern humans and Neanderthals. The partial adult skeleton of Qafzeh 25 dated to 92 ± 5 ka B.P. suffers from serious post-mortem taphonomic damage that has limited its anatomical description and metrical analysis. However, the two petrosal bones are preserved and the bony labyrinths are not affected by post-mortem deformations. In this study, the methodology developed by Spoor (1993) is used to analyze and compare the morphometric data from Qafzeh 25 semi-circular canals and cochlea to that of other fossils from the site and more generally to the published hominin sample (Middle and Late Pleistocene hominins). While this analysis reveals that the Qafzeh 25 bony labyrinth resembles that of other Qafzeh individuals, it extends the range of variation within the sample for some variables.

Early Pleistocene faunivorous hominins were not kleptoparasitic, and this impacted the evolution of human anatomy and socio-ecology

Humans are unique in their diet, physiology and socio-reproductive behavior compared to other primates. They are also unique in the ubiquitous adaptation to all biomes and habitats. From an evolutionary perspective, these trends seem to have started about two million years ago, coinciding with the emergence of encephalization, the reduction of the dental apparatus, the adoption of a fully terrestrial lifestyle, resulting in the emergence of the modern anatomical bauplan, the focalization of certain activities in the landscape, the use of stone tools, and the exit from Africa. It is in this period that clear taphonomic evidence of a switch in diet with respect to Pliocene hominins occurred, with the adoption of carnivory. Until now, the degree of carnivorism in early humans remained controversial. A persistent hypothesis is that hominins acquired meat irregularly (potentially as fallback food) and opportunistically through klepto-foraging. Here, we test this hypothesis and show, in contrast, that the butchery practices of early Pleistocene hominins (unveiled through systematic study of the patterning and intensity of cut marks on their prey) could not have resulted from having frequent secondary access to carcasses. We provide evidence of hominin primary access to animal resources and emphasize the role that meat played in their diets, their ecology and their anatomical evolution, ultimately resulting in the ecologically unrestricted terrestrial adaptation of our species. This has major implications to the evolution of human physiology and potentially for the evolution of the human brain.