Pleistocene Hominins Research Articles

Footprint evidence for locomotor diversity and shared habitats among early Pleistocene hominins.

For much of the Pliocene and Pleistocene, multiple hominin species coexisted in the same regions of eastern and southern Africa. Due to the limitations of the skeletal fossil record, questions regarding their interspecific interactions remain unanswered. We report the discovery of footprints (~1.5 million years old) from Koobi Fora, Kenya, that provide the first evidence of two different patterns of Pleistocene hominin bipedalism appearing on the same footprint surface. New analyses show that this is observed repeatedly across multiple contemporaneous sites in the eastern Turkana Basin. These data indicate a sympatric relationship between Homo erectus and Paranthropus boisei, suggesting that lake margin habitats were important to both species and highlighting the possible influence of varying levels of coexistence, competition, and niche partitioning in human evolution.

A cluster of Pleistocene hominin ichnosites on South Africa’s Cape South Coast

Seven Pleistocene tracksites and two lithic sites, identified with varying degrees of confidence and attributed to Homo sapiens, have been identified in aeolianite deposits along a 1200 m coastal stretch at Brenton-on-Sea on South Africa’s Cape south coast. Some sites contain more than one track- or trace-bearing layer, and each site is from a geological different unit. The aeolianites adjacent to the ichnosites that have been dated span an age range from 113 ± 8 ka to 76 ± 5 ka. Globally, hominin tracks preserved in aeolianites are rare, as are tracks preserved in hyporelief, which are observed at several of the Brenton-on-Sea sites. Clusters of hominin tracksites older than 40 ka are also rare. The sites represent groups of hominins of different sizes traversing dune fields, sometimes jogging or using sticks, and provide details of their tool-making and diet. The ichnological record thus complements the extensive and globally significant archaeological record from the region. The likely presence of nearby accessible caves, and a shoreline that was not too distant, provide an explanation for the profusion of ichnosites and their registration over millennia.

The expression of the MIS 12 glacial stage in Southeastern Europe and its impact over the Middle Pleistocene hominins in Megalopolis Basin (Greece)

Southern Europe is hypothesized to have acted as a glacial refugium for hominin populations during the Pleistocene. Of particular importance is South-East Europe, which most likely played a dual role, both as refugium and dispersal corridor, especially during the Middle Pleistocene glaciations, when drastic climatic conditions led to major sea level drops in the Aegean. However, little is known about the palaeoenvironmental conditions at the time of hominin presence in this region, making these hypotheses difficult to test. Here we analyze biomarker data and leaf wax stable isotopic compositions of the MIS 12 Lower Palaeolithic site Marathousa 1 (Megalopolis Basin, Greece) to assess the climatic conditions accompanying the time of hominin presence in the area. Our data indicate a major cooling affecting the north Mediterranean/Aegean domain during this time interval, with lowest temperatures recorded between ∼440–432 ka. The glacial peak is associated with changes in vegetation (i.e., from more forested to more open landscape), reduction of humidity and water availability (i.e., moisture depletion, increased evaporation). Hominins are present at the Marathousa 1 location at the end of this interval (434–432 ka), confirming that the Megalopolis Basin served as a refugium for hunter-gatherer groups during periods of harsh climatic conditions. Additionally, the progressive cooling is associated with an important sedimentary hiatus between ∼465–440 ka reflected in all circum-Mediterranean records (both marine and continental), indicating a regional impact of the MIS 12 glaciation over surface processes.

26Al/10Be burial dating and technological strategies of hominins at the Jijiazhuang Paleolithic site, Nihewan Basin, China: Implications for understanding Middle Pleistocene human adaptations in east Asia

With the discovery of more than a hundred Pleistocene Paleolithic sites, the Nihewan Basin of North China has become an area of reference for the study of human evolution and behavioral adaptations during and after the spread of hominins out of Africa and into Eurasia. However, most research has focused on the Early and Late Pleistocene archaeological record, whereas studies of the Middle Pleistocene sequence are relatively limited. Here we contribute to fill this gap by introducing the archaeological assemblage and 26Al/10Be burial dating of the newly discovered Jijiazhuang (JJZ) Paleolithic site. Systematic excavations in recent years have yielded well-preserved stone artifacts and mammalian fossils in fluvio-lacustrine sediments at the site. Cosmogenic 26Al/10Be burial dating indicates that hominins occupied the site between 0.49 ± 0.10 and 0.63 ± 0.11 Ma, corresponding to the extra-long interglacial period of MIS 15-13. The JJZ lithic assemblage shows evidence of relatively long-distance resource procurement, and the increased number of retouched tools indicate standardized, extensive and refined modification, even shaping strategies. The JJZ lithic technology shows advanced features that may shed light on the regional emergence of Middle Paleolithic technologies, in a paleoecological context where the extra-long duration of interglacial/mild stadial climate events (MIS 15−13) may have provided favorable conditions for increased technological capabilities among Middle Pleistocene hominins from the Nihewan Basin. Our study is the first to present a detailed techno-typological analysis of a Middle Pleistocene lithic assemblage in the Nihewan Basin and contributes to the characterization of technological adaptions in the high latitudes of East Asia.

Quantifying heterogeneity of hominin environments in and out of Africa using herbivore dental traits

Our own species, Homo sapiens, has been able to invade near-all global ecosystems through a unique form of ecological plasticity. To understand how Homo sapiens have been able to disperse to such a wide range of environments, we need to understand the origins of the characteristic which enabled this, plasticity. Growing evidence indicates that Homo species were already able to occupy a diverse set of environments in the Early Pleistocene, suggesting that the origins of human ecological plasticity have deeper roots than previously anticipated. We analysed the distribution of two dental traits, mean ordinated hypsodonty of large herbivores and proboscidean mesowear, across the mammalian communities at Early Pleistocene hominin sites, to infer the vegetation and climate contexts of these locations. The global distribution of community-level dental traits suggests that, while open grassy vegetation made up a large proportion of the sites, mixed and wooded vegetation was also represented. From our analysis we conclude that Early Pleistocene Homo species were able to exploit habitats with different vegetation types and occupied a range of different environments. In addition to this we found that the distribution of dental traits of mammalian communities in the Eurasian sample was notably different from that of the African sample, suggesting that Homo's dispersal into Eurasia resulted in a niche expansion. Together these findings suggest that Homo in the Early Pleistocene was ecologically plastic and occupied a wide environmental niche space.

Qaleh Kurd Cave (Qazvin, Iran): Oldest Evidence of Middle Pleistocene Hominin Occupations and a Human Deciduous Tooth in the Iranian Central Plateau

Qaleh Kurd Cave (Qazvin, Iran): Oldest Evidence of Middle Pleistocene Hominin Occupations and a Human Deciduous Tooth in the Iranian Central Plateau

Multi-isotope reconstruction of Late Pleistocene large-herbivore biogeography and mobility patterns in Central Europe

Interpretations of Late Pleistocene hominin adaptative capacities by archaeologists have focused heavily on their exploitation of certain prey and documented contemporary behaviours for these species. However, we cannot assume that animal prey-taxa ecology and ethology were the same in the past as in the present, or were constant over archaeological timescales. Sequential isotope analysis of herbivore teeth has emerged as a particularly powerful method of directly reconstructing diet, ecology and mobility patterns on sub-annual scales. Here, we apply 87Sr/86Sr isotope analysis, in combination with δ18O and δ13C isotope analysis, to sequentially sampled tooth enamel of prevalent herbivore species that populated Europe during the Last Glacial Period, including Rangifer tarandus, Equus sp. and Mammuthus primigenius. Our samples come from two open-air archaeological sites in Central Germany, Königsaue and Breitenbach, associated with Middle Palaeolithic and early Upper Palaeolithic cultures, respectively. We identify potential inter- and intra-species differences in range size and movement through time, contextualised through insights into diet and the wider environment. However, homogeneous bioavailable 87Sr/86Sr across large parts of the study region prevented the identification of specific migration routes. Finally, we discuss the possible influence of large-herbivore behaviour on hominin hunting decisions at the two sites.

Evidence of Pleistocene hominin landscapes in Eyvanekey, Iran, and implications for the Northern Dispersal Corridor

AbstractPrior to this work, no Palaeolithic field survey had been conducted in the central region of the northern Iranian Central Desert. This article is the first account of the presence of Pleistocene hominins in Eyvanekey, Semnan Province, and reports the recovery of lithics that date to at least the Middle Palaeolithic.

The Cranium I: Neurocranium.

The Sima de los Huesos (SH) site has provided a significant collection of hominin remains, including numerous cranial fragments, which have contributed to our understanding of the MP human population. The taxonomic classification of the SH hominins remains a topic of debate, with some studies suggesting a close relationship to Neandertals based on nuclear DNA analysis. The cranial morphology of the SH specimens exhibits a mix of Neandertal-like features and primitive traits observed in earlier Homo populations, providing insights into the evolutionary pattern of the Neanderthal lineage. This study focuses on the neurocranial traits of the SH population and describes three previously undescribed cranial individuals. The SH cranial collection now comprises 20 nearly complete crania, representing approximately two-thirds of the estimated population size. The analysis of the SH population reveals variations in robustness, frontal torus development, sagittal keeling, and occipital torus morphology, which may be related to sexual dimorphism and ontogenetic factors. The suprainiac region exhibits notable ontogenetic changes, while suture obliteration patterns do not strictly correlate with dental age. Metric measurements, particularly cranial breadths, highlight significant intrapopulation variation within the SH sample. Compared with other Middle Pleistocene (MP) hominins, the SH cranial vault displays archaic characteristics but differs from Homo erectus and Neandertals. The SH individuals have relatively short and tall cranial vaults, distinguishing them from other MP fossils. These findings contribute to our understanding of the MP human populations and their evolutionary trajectories.

A Late Pleistocene hominin footprint site on the North African coast of Morocco

Footprints represent a relevant vestige providing direct information on the biology, locomotion, and behaviour of the individuals who left them. However, the spatiotemporal distribution of hominin footprints is heterogeneous, particularly in North Africa, where no footprint sites were known before the Holocene. This region is important in the evolution of hominins. It notably includes the earliest currently known Homo sapiens (Jebel Irhoud) and the oldest and richest African Middle Stone Age hominin sites. In this fragmented ichnological record, we report the discovery of 85 human footprints on a Late Pleistocene now indurated beach surface of about 2800 m2 at Larache (Northwest coast of Morocco). The wide range of sizes of the footprints suggests that several individuals from different age groups made the tracks while moving landward and seaward across a semi-dissipative bar-trough sandy beach foreshore. A geological investigation and an optically stimulated luminescence dating of a rock sample extracted from the tracksite places this hominin footprint surface at 90.3 ± 7.6 ka (MIS 5, Late Pleistocene). The Larache footprints are, therefore, the oldest attributed to Homo sapiens in Northern Africa and the Southern Mediterranean.

Biomechanical and taxonomic diversity in the Early Pleistocene in East Africa: Structural analysis of a recently discovered femur shaft from Olduvai Gorge (bed I)

Recent Plio-Pleistocene hominin findings have revealed the complexity of human evolutionary history and the difficulties involved in its interpretation. Moreover, the study of hominin long bone remains is particularly problematic, since it commonly depends on the analysis of fragmentary skeletal elements that in many cases are merely represented by small diaphyseal portions and appear in an isolated fashion in the fossil record. Nevertheless, the study of the postcranial skeleton is particularly important to ascertain locomotor patterns. Here we report on the discovery of a robust hominin femoral fragment (OH 84) at the site of Amin Mturi Korongo dated to 1.84 Ma (Olduvai Bed I). External anatomy and internal bone structure of OH 84 were analyzed and compared with previously published data for modern humans and chimpanzees, as well as for Australopithecus, Paranthropus and Homo specimens ranging from the Late Pliocene to Late Pleistocene. Biomechanical analyses based on transverse cross-sections and the comparison of OH 84 with another robust Olduvai specimen (OH 80) suggest that OH 84 might be tentatively allocated to Paranthropus boisei. More importantly, the identification of a unique combination of traits in OH 84 could indicate both terrestrial bipedalism and an arboreal component in the locomotor repertoire of this individual. If interpreted correctly, OH 84 could thus add to the already mounting evidence of substantial locomotor diversity among Early Pleistocene hominins. Likewise, our results also highlight the difficulties in accurately interpreting the link between form and function in the human fossil record based on fragmentary remains, and ultimately in distinguishing between coeval hominin groups due to the heterogeneous pattern of inter- and intraspecific morphological variability detected among fossil femora.

The Pleistocene grassland-dominated mammal fauna from Tham Kra Duk (Nakhon Si Thammarat, Peninsular Thailand)

ABSTRACT Controversy exists as to whether the Pleistocene vegetation in northern Sundaland was dominated by lowland tropical grasslands or rainforests, due to limited palaeoecological evidence recorded from the region. We describe a new Pleistocene large mammal fauna from Tham Kra Duk, a cave in the Tham Phedan mountain, Nakhon Si Thammarat Province in Peninsular Thailand, with emphasis on its palaeoecological and palaeoenvironmental investigations using the stable isotope analysis of mammalian tooth enamel. The fossil site has yielded at least nine mammalian taxa almost comparable to late middle to latest Pleistocene faunas in the mainland, thus suggesting the same biogeographic mammal elements with a range extension south of the Kra Isthmus. The stable isotope results indicate that mixed woodland to grassland ecosystems were dominated by C4 vegetation in the area. This supports the assumption that the expansion of Pleistocene tropical savanna ecosystems might have held the key to facilitating the southward distribution range of grazing mammals such as gaurs and Himalayan gorals into the Thai-Malay Peninsula. The presence of the Tham Kra Duk fauna was probably linked to some major biogeographic events of Pleistocene hominin and mammal migration through the land-bridge peninsula into the islands of Southeast Asia during a period of glaciation.

Beaver exploitation, 400,000 years ago, testifies to prey choice diversity of Middle Pleistocene hominins

Data regarding the subsistence base of early hominins are heavily biased in favor of the animal component of their diets, in particular the remains of large mammals, which are generally much better preserved at archaeological sites than the bones of smaller animals, let alone the remains of plant food. Exploitation of smaller game is very rarely documented before the latest phases of the Pleistocene, which is often taken to imply narrow diets of archaic Homo and interpreted as a striking economic difference between Late Pleistocene and earlier hominins. We present new data that contradict this view of Middle Pleistocene Lower Palaeolithic hominins: cut mark evidence demonstrating systematic exploitation of beavers, identified in the large faunal assemblage from the c. 400,000 years old hominin site Bilzingsleben, in central Germany. In combination with a prime-age dominated mortality profile, this cut mark record shows that the rich beaver assemblage resulted from repetitive human hunting activities, with a focus on young adult individuals. The Bilzingsleben beaver exploitation evidence demonstrates a greater diversity of prey choice by Middle Pleistocene hominins than commonly acknowledged, and a much deeper history of broad-spectrum subsistence than commonly assumed, already visible in prey choices 400,000 years ago.

“Dragon man” prompts rethinking of Middle Pleistocene hominin systematics in Asia

“Dragon man” prompts rethinking of Middle Pleistocene hominin systematics in Asia

Close encounters vs. missed connections? A critical review of the evidence for Late Pleistocene hominin interactions in western Eurasia

Recent advances in the study of ancient DNA recovered from fossils and cave sediments have profoundly changed our views on the biological and cultural interactions between populations and lineages of fossil Homo in the Later Pleistocene of Eurasia. A spatiotemporally complex picture emerges, with multiple population admixture and replacement events. Focusing on the evidence from Western Eurasia, we consider here how the mapping out of between-species interactions based on fossil and material cultural evidence is being replaced by a broader approach. Traditional narratives about human migrations and the biological and/or cultural advantages of our own species over the Neanderthals are now giving way to the study of the biological and cultural dynamics of past human populations and the nature of their interactions in time and space.

Late Acheulean occupations at Montagu Cave and the pattern of Middle Pleistocene behavioral change in Western Cape, southern Africa

Patterns of so-called modern human behavior are increasingly well documented in an abundance of Middle Stone Age archaeological sites across southern Africa. Contextualized archives directly preceding the southern African Middle Stone Age, however, remain scarce. Current understanding of the terminal Acheulean in southern Africa derives from a small number of localities that are predominantly in the central and northern interior. Many of these localities are surface and deflated contexts, others were excavated prior to the availability of modern field documentation techniques, and yet other relevant assemblages contain low numbers of characteristic artifacts relative to volume of excavated deposit. The site of Montagu Cave, situated in the diverse ecosystem of the Cape Floral Region, South Africa, contains the rare combination of archaeologically rich, laminated and deeply stratified Acheulean layers followed by a younger Middle Stone Age occupation. Yet little is known about the site owing largely to a lack of contextual information associated with the early excavations. Here we present renewed excavation of Levels 21–22 at Montagu Cave, located in the basal Acheulean sequence, including new data on site formation and ecological context, geochronology, and technological variability. We document intensive occupation of the cave by Acheulean tool-producing hominins, likely at the onset of interglacial conditions in MIS 7. New excavations at Montagu Cave suggest that, while Middle Stone Age technologies were practiced by 300 ka in several other regions of Africa, the classic Acheulean persisted later in the Fynbos Biome of the southwestern Cape. We discuss the implications of this regionalized persistence for the biogeography of African later Middle Pleistocene hominin populations, for the ecological drivers of their technological systems, and for the pattern and pace of behavioral change just prior to the proliferation of the southern African later Middle Stone Age.

Structural properties of the Late Pleistocene Liujiang femoral diaphyses from southern China

The characterization of the femoral diaphysis in Pleistocene hominins with chronoecogeographical diversity plays a crucial role in evaluating evolutionary shifts in locomotor behavior and body shape. However, Pleistocene hominin fossil remains in East Asia are scarce and are widely dispersed temporally and spatially, impeding our comprehension of the nature and polarity of morphological trends. Here, we present qualitative and quantitative analyses of the cross-sectional properties and structural organization of diaphyses in two Late Pleistocene hominin femora (Liujiang PA91 and PA92) from southern China, comparing them to other Eurasian and African Pleistocene hominins. By integrating surface features and internal structure, our findings reveal that the Liujiang femora exhibit modern human-like characteristics, including a developed pilaster, a gluteal buttress, and minimum mediolateral breadth located at the midshaft. The presence of a femoral pilaster may relate to posterior cortical reinforcement and an increased anteroposterior bending rigidity along the mid-proximal to mid-distal portion of the diaphysis. Compared to archaic Homo, Liujiang and other Late Pleistocene modern human femora show a thinner mediolateral cortex and lower bending rigidity than the anteroposterior axis, and a lack of medial buttress, potentially indicating functionally related alterations in a range of pelvic and proximal femoral features throughout the Pleistocene. The femoral robusticity of the Liujiang individual resembles that of other Pleistocene hunter–gatherers from East Asia, implying comparable overall mobility or activity levels. The investigation of Liujiang femoral diaphyseal morphology contributes to a more comprehensive understanding of early modern human postcranial structural morphology in East Asia.

Did Early Pleistocene hominins control hammer strike angles when making stone tools?

In the study of Early Pleistocene stone artifacts, researchers have made considerable progress in reconstructing the technical decisions of hominins by examining various aspects of lithic technology, such as reduction sequences, hammer selection, platform preparation, core management, and raw material selection. By comparison, our understanding of the ways in which Early Pleistocene hominins controlled the delivery and application of percussive force during flaking remains limited. In this study, we focus on a key aspect of force delivery in stone knapping, namely the hammerstone striking angle (or the angle of blow), which has been shown to play a significant role in determining the knapping outcome. Using a dataset consists of 12 Early Pleistocene flake assemblages dated from 1.95 Ma to 1.4 Ma, we examined temporal patterns of the hammer striking angle by quantifying the bulb angle, a property of the flake's Hertzian cone that reflects the hammer striking angle used in flake production. We further included a Middle Paleolithic flake assemblage as a point of comparison from a later time period. In the Early Pleistocene dataset, we observed an increased association between the bulb angle and other flake variables related to flake size over time, a pattern similarly found in the Middle Paleolithic assemblage. These findings suggest that, towards the Oldowan–Acheulean transition, hominins began to systematically adjust the hammer striking angle in accordance with platform variables to detach flakes of different sizes more effectively, implying the development of a more comprehensive understanding of the role of the angle of blow in flake formation by ∼1.5 Ma.

A new assemblage of late Neanderthal remains from Cova Simanya (NE Iberia)

This study presents an exceptional collection of 54 Late Pleistocene human remains that correspond to at least three Neanderthal individuals from Simanya Gran, the main gallery of Cova Simanya, located in the northeastern Iberian Peninsula. The collection comprised 53 unpublished remains that were unearthed during the 1970s and an additional tooth discovered during 2021 excavations. The specimens represent an adult with a small stature, a periadolescent aged approximately 11.5 years, and an immature individual aged approximately 7.7 years, thus offering a more complete demographic perspective. The collection encompasses diverse anatomical parts including upper and lower dentition, mandible, vertebrae, and limb bones from both the upper and lower extremities. Attempts to extract aDNA were unsuccessful. Renewed archaeological investigations at Cova Simanya have facilitated the reevaluation of the original stratigraphic context of these remains, leading to the discovery of the additional tooth, aligning with the periadolescent individual. This assemblage is currently the most extensive Neanderthal collection from the northeastern Mediterranean Iberia, offering invaluable insights into the morphology and evolutionary trajectory of Late Pleistocene hominins. Hence, Simanya Neanderthals will enhance our understanding of Neanderthal demographics and evolution, paving the way for an in-depth examination of the morphological diversity and evolutionary context of Iberian Neanderthals.

Identifying Palaeolithic birch tar production techniques: challenges from an experimental biomolecular approach

The intentional production of birch bark tar by European Neanderthals as early as 190,000 years ago plays an important role in discussions about the technological and behavioural complexity of Pleistocene hominins. However, research is hampered because it is currently unknown how Neanderthals were producing birch tar. There are several different techniques that could have been employed, but these differ in their apparent production complexity, time and resource efficiency. Identifying production processes in the archaeological record is therefore paramount for furthering research on the technical behavioural repertoire. Organic biomarkers, identified with Gas Chromatograph–Mass Spectrometry (GC–MS), have been used to identify possible production processes during the Neolithic. Here we test whether these biomarkers can also distinguish Palaeolithic (aceramic) tar production methods. We produced tar using five different methods and analysed their biomolecular composition with GC–MS. Our results show that the biomarkers used to distinguish Neolithic tar production strategies using ceramic technology cannot be reliably used to identify tar production processes using aceramic Palaeolithic techniques. More experimentation is required to produce a larger reference library of different tars for future comparisons. To achieve this, complete GC–MS datasets must also be made publicly available, as we have done with our data.