Middle Pleistocene Humans Research Articles

A double-pointed wooden throwing stick from Schöningen, Germany: Results and new insights from a multianalytical study.

The site of Schöningen (Germany), dated to ca. 300,000 years ago, yielded the earliest large-scale record of humanly-made wooden tools. These include wooden spears and shorter double-pointed sticks, discovered in association with herbivores that were hunted and butchered along a lakeshore. Wooden tools have not been systematically analysed to the same standard as other Palaeolithic technologies, such as lithic or bone tools. Our multianalytical study includes micro-CT scanning, 3-dimensional microscopy, and Fourier transform infrared spectroscopy, supporting a systematic technological and taphonomic analysis, thus setting a new standard for wooden tool analysis. In illustrating the biography of one of Schöningen's double-pointed sticks, we demonstrate new human behaviours for this time period, including sophisticated woodworking techniques. The hominins selected a spruce branch which they then debarked and shaped into an aerodynamic and ergonomic tool. They likely seasoned the wood to avoid cracking and warping. After a long period of use, it was probably lost while hunting, and was then rapidly buried in mud. Taphonomic alterations include damage from trampling, fungal attack, root damage and compression. Through our detailed analysis we show that Middle Pleistocene humans had a rich awareness of raw material properties, and possessed sophisticated woodworking skills. Alongside new detailed morphometrics of the object, an ethnographic review supports a primary function as a throwing stick for hunting, indicating potential hunting strategies and social contexts including for communal hunts involving children. The Schöningen throwing sticks may have been used to strategically disadvantage larger ungulates, potentially from distances of up to 30 metres. They also demonstrate that the hominins were technologically capable of capturing smaller fast prey and avian fauna, a behaviour evidenced at contemporaneous Middle Pleistocene archaeological sites.

Three-dimensional geometric morphometric study of the Xuchang 2 cranium

Results of traditional metric and nonmetric assessments suggest that the Xuchang hominin shares features with Neanderthals. To comprehensively compare the nuchal morphology of XC 2 to those of the genus Homo, we conduct a three-dimensional geometric morphometric study with 35 cranial landmarks and surface semilandmarks of XC 2, Homo erectus, Middle Pleistocene humans, Neanderthals, and early and recent modern humans. Results reveal that the centroid size of XC 2 is larger than that of early and recent modern humans and can only be compared to that of Middle Pleistocene humans and H. erectus. Early and recent modern humans share a nuchal morphology distinct from archaic hominins (Ngandong H. erectus, Middle Pleistocene humans, and Neanderthals), except for SM 3, Sangiran 17, and Asian and African H. erectus. Although Ngandong specimens differ from the other H. erectus, it is unclear whether this represents a temporal or spatial trend in the process of evolution of this species. The nuchal morphological resemblance between Middle Pleistocene humans and Neanderthals may be attributed to similar cranial architecture and cerebellar shape. The great nuchal morphological variation shared by recent modern humans may indicate a particular developmental pattern. In conclusion, the nuchal morphology of different human groups is highly variable and may be caused by different factors including brain globularization and developmental plasticity. XC 2 shares similar nuchal morphology with Middle Pleistocene humans and Neanderthals, but these results are insufficient to fully resolve the taxonomic status of XC 2.

Shaped stone balls were used for bone marrow extraction at Lower Paleolithic Qesem Cave, Israel.

The presence of shaped stone balls at early Paleolithic sites has attracted scholarly attention since the pioneering work of the Leakeys in Olduvai, Tanzania. Despite the persistent presence of these items in the archaeological record over a period of two million years, their function is still debated. We present new results from Middle Pleistocene Qesem Cave on the use of these implements as percussion tools. Use-wear and abundant bone and fat residues found on ten shaped stone balls indicate crushing of fresh bones by thrusting percussion and provide direct evidence for the use of these items to access bone marrow of animal prey at this site. Two experiments conducted to investigate and verify functional aspects proved Qesem Cave shaped stone balls are efficient for bone processing and provide a comfortable grip and useful active areas for repeated use. Notably, the patina observed on the analyzed items precedes their use at the cave, indicating that they were collected by Qesem inhabitants, most probably from older Lower Paleolithic Acheulian sites. Thus, our results refer only to the final phases of the life of the items, and we cannot attest to their original function. As bone marrow played a central role in human nutrition in the Lower Paleolithic, and our experimental results show that the morphology and characteristics of shaped stone ball replicas are well-suited for the extraction of bone marrow, we suggest that these features might have been the reason for their collection and use at Qesem Cave. These results shed light on the function of shaped stone balls and are consistent with the significance of animal fat in the caloric intake of Middle Pleistocene humans as shown by the archeozoological evidence at Qesem Cave and possibly beyond.

External auditory exostoses among western Eurasian late Middle and Late Pleistocene humans.

External auditory exostoses (EAE) have been noted among the Neandertals and a few other Pleistocene humans, but until recently they have been discussed primary as minor pathological lesions with possible auditory consequences. An assessment of available western Eurasian late Middle and Late Pleistocene human temporal bones with sufficiently preserved auditory canals (n = 77) provides modest levels of EAE among late Middle Pleistocene archaic humans (≈20%) and early modern humans (Middle Paleolithic: ≈25%; Early/Mid Upper Paleolithic: 20.8%; Late Upper Paleolithic: 9.5%). The Neandertals, however, exhibit an exceptionally high level of EAE (56.5%; 47.8% if two anomalous cases are considered normal). The levels of EAE for the early modern humans are well within recent human ranges of variation, frequencies which are low for equatorial inland and high latitude samples but occasionally higher elsewhere. The Early/Mid Upper Paleolithic frequency is nonetheless high for a high latitude sample under interpleniglacial conditions. Given the strong etiological and environmental associations of EAE development with exposure to cold water and/or damp wind chill, the high frequency of EAE among the Neandertals implies frequent aquatic resource exploitation, more frequent than the archeological and stable isotopic evidence for Middle Paleolithic/Neandertal littoral and freshwater resource foraging implies. As such, the Neandertal data parallel a similar pattern evident in eastern Eurasian archaic humans. Yet, factors in addition to cold water/wind exposure may well have contributed to their high EAE frequencies.

Cross-sectional properties of the lower limb long bones in the Middle Pleistocene Sima de los Huesos sample (Sierra de Atapuerca, Spain)

The recovery to date of three complete and five partial femora, seven complete tibiae, and four complete fibulae from the Atapuerca Sima de los Huesos site provides an opportunity to analyze the biomechanical cross-sectional properties in this Middle Pleistocene population and to compare them with those of other fossil hominins and recent modern humans.We have performed direct comparisons of the cross-sectional geometric parameters and reduced major axis (RMA) regression lines among different samples. We have determined that Atapuerca Sima de los Huesos (SH) fossils have significantly thicker cortices than those of recent modern humans for the three leg bones at all diaphyseal levels, except that of the femur at 35% of biomechanical length. The SH bones are similar to those of Neandertals and Middle Pleistocene humans and different from Homo sapiens in their diaphyseal cross-sectional shape and strength parameters.When standardized by estimated body size, both the SH and Neandertal leg bones have in general greater strength than those of H. sapiens from the early modern (EMH), Upper Paleolithic (UP), and recent populations (RH). The Sima de los Huesos human leg bones have, in general terms, an ancestral pattern similar to that of Pleistocene humans and differing from H. sapiens.

Inner ear morphology of the cioclovina early modern European calvaria from Romania.

The morphology of the human bony labyrinth is thought to preserve a strong phylogenetic signal and to be minimally, if at all, affected by postnatal processes. The form of the semicircular canals is considered a derived feature of Neanderthals and different from the modern human anatomy. Among other hominins, European Middle Pleistocene humans have been found to be most similar to Neanderthals. Early modern humans have been proposed to show a pattern that is distinct, but most similar to that of Holocene people. Here we examine the inner ear structures of the Cioclovina calvaria, one of the earliest reliably dated and relatively complete modern human crania from Europe, in the context of recent and fossil human variation. Bony labyrinths were virtually extracted from CT scans of recent Europeans and Cioclovina. Using univariate and multivariate methods, measurements of the semicircular canals were compared with published measurements of other fossil specimens. Our results show that Cioclovina's inner ear morphology falls within the range of modern variation, with affinities to both Late Pleistocene modern humans and recent Europeans. Using discriminant functions, the sex of the Cioclovina specimen is estimated as male. Results agree with previous work showing that Cioclovina exhibits fully modern cranial morphology.

The oral pathological conditions of the Broken Hill (Kabwe) 1 cranium

The Broken Hill (Kabwe) 1 cranium exhibits dental caries on ten teeth, multiple periapical lesions, periodontal disease, severe anterior dental wear, and hypercementosis, conditions all little-studied in Middle Pleistocene humans and making Broken Hill 1 of great value to discussions of the antiquity of oral pathological conditions. These individual pathological conditions, however, have never previously been described in detail; the focus has been on diagnosing an overarching syndrome connecting the temporal and oral lesions. This case study examines the individual dentoalveolar pathological conditions to provide data for cross-comparison using ordinal scores (e.g., carious lesions, interdental septum condition), descriptions (e.g., hypercementosis, periapical lesions), and continuous measurements (cemento-enamel junction to alveolar crest distances). Differential diagnoses are explored including age-related hyposalivation, dental wear, lead poisoning, and diet, and are discussed within the context of past published attempts. Studies on recent humans have demonstrated correlations between oral disease, systemic health, and diet, suggesting Broken Hill 1 likely suffered from larger systemic inflammation related to its age, health, and perhaps subsistence.

Evaluating developmental shape changes in Homo antecessor subadult facial morphology

The fossil ATD6-69 from Atapuerca, Spain, dated to ca. 900 ka (thousands of years ago) has been suggested to mark the earliest appearance of modern human facial features. However, this specimen is a subadult and the interpretation of its morphology remains controversial, because it is unclear how developmental shape changes would affect the features that link ATD6-69 to modern humans. Here we analyze ATD6-69 in an evolutionary and developmental context. Our modern human sample comprises cross-sectional growth series from four populations. The fossil sample covers human specimens from the Pleistocene to the Upper Paleolithic, and includes several subadult Early Pleistocene humans and Neanderthals. We digitized landmarks and semilandmarks on surface and CT scans and analyzed the Procrustes shape coordinates using multivariate statistics. Ontogenetic allometric trajectories and developmental simulations were employed in order to identify growth patterns and to visualize potential adult shapes of ATD6-69. We show that facial differences between modern and archaic humans are not exclusively allometric. We find that while postnatal growth further accentuates the differences in facial features between Neanderthals and modern humans, those features that have been suggested to link ATD6-69's morphology to modern humans would not have been significantly altered in the course of subsequent development. In particular, the infraorbital depression on this specimen would have persisted into adulthood. However, many of the facial features that ATD6-69 shares with modern humans can be considered to be part of a generalized pattern of facial architecture. Our results present a complex picture regarding the polarity of facial features and demonstrate that some modern human-like facial morphology is intermittently present in Middle Pleistocene humans. We suggest that some of the facial features that characterize recent modern humans may have developed multiple times in human evolution.

The Cognitive Implications of Controlled Fire Use by Early Humans

This article proposes a framework for investigating the cognitive implications of controlled fire use by Middle Pleistocene humans. By identifying the simplest strategies they could have used to control fire, given the constraints individuals had to overcome, we can establish a behavioural basis for making inferences about cognition. Accessing, maintaining and benefiting from fire involved a range of behaviours that imply future-directed planning, response inhibition and group-level cooperation. I argue that we can infer human cognitive abilities such as an extended working memory, episodic memories, collective intentionality and intersubjective communication from these fire-related behaviours.

Middle Pleistocene human facial morphology in an evolutionary and developmental context

Neanderthals and modern humans exhibit distinct facial architectures. The patterning of facial morphology of their predecessors, the Middle Pleistocene humans, is more mosaic showing a mix of archaic and modern morphologies. Significant changes in facial size and robusticity occurred throughout Pleistocene human evolution, resulting in temporal trends in both facial reduction and enlargement. However, the allometric patterning in facial morphology in archaic humans is not well understood. This study explores temporal trends in facial morphology in order to gain a clearer understanding of the polarity of features, and describes the allometric patterning of facial shape.The modern human sample comprises cross-sectional growth series of four morphologically distinct human populations. The fossil sample covers specimens from the Middle Pleistocene to the Upper Paleolithic. We digitized landmarks and semilandmarks on surface and computed tomography scans and analyzed the Procrustes shape coordinates. Principal component analyses were performed, and Procrustes distances were used to identify phenetic similarities between fossil hominins. In order to explore the influence of size on facial features, allometric trajectories were calculated for fossil and modern human groups, and developmental simulations were performed.We show that facial features can be used to separate Pleistocene humans into temporal clusters. The distinctly modern human pattern of facial morphology is already present around 170 ka. Species- and population-specific facial features develop before two years of age, and several of the large-scale facial differences between Neanderthals and Middle Pleistocene humans are due to scaling along a shared allometric trajectory. These features include aspects of the frontal bone, browridge morphology, nasal aperture size and facial prognathism. Infraorbital surface topography and orientation of the midface in the European Middle Pleistocene hominins is intermediate between the African Middle Pleistocene and Neanderthal condition. This could suggest that the European Middle Pleistocene hominins display incipient Neanderthal features.

Communicative capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain

The present study presents new data on the abilities of Homo heidelbergensis to produce and perceive the sounds emitted during modern human spoken language. The pattern of sound power transmission was studied through the outer and middle ears in five individuals from the Sima de los Huesos, four chimpanzees and four modern humans. The results were then used to calculate the occupied bandwidth of the outer and middle ears, an important variable related with communicative capacities. The results demonstrate that the Atapuerca SH hominins were similar to modern humans in this aspect, falling within the lower half of the range of variation, and clearly distinct from chimpanzees. Specifically, the Atapuerca SH hominins show a bandwidth that is slightly displaced and considerably extended to encompass the frequencies that contain relevant acoustic information in human speech, permitting the transmission of a larger amount of information with fewer errors. At the same time, the presence of a complete cervical segment of the spinal column associated with Cranium 5 from the Sima de los Huesos Middle Pleistocene site (Sierra de Atapuerca, Spain) makes it possible to estimate the vocal tract proportions in H. heidelbergensis for the first time. The results demonstrate that it is similar to the reconstructed vocal tract in the La Ferrassie 1 Neandertal individual, which has been suggested to have been capable of producing the full range of sounds emitted during modern human spoken language. These results in the Atapuerca (SH) hominins are consistent with other recent suggestions for an ancient origin for human speech capacity.

The postcranial dimensions of the La Chapelle‐aux‐saints 1 Neandertal

The La Chapelle-aux-Saints 1 Neandertal has figured prominently in considerations of Neandertal body size and proportions. In this context, a reassessment of its major long bones and a reassembly of its principal pelvic elements (sacrum and right ilium) was undertaken. There are secure measurements for its humeral and radial lengths and its femoral head diameter, but the femoral and tibial lengths were almost certainly greater than previous values. The resultant humeral, femoral and tibial lengths are similar to those of other male Neandertals, its femoral head diameter is among the largest known for Middle and Late Pleistocene humans, but its radial length is relatively short. The pelvic assembly provides modest bi-iliac and inlet transverse diameters compared with the few sufficiently complete and undistorted Middle and Late Pleistocene archaic human pelves, but its dimensions are similar to those of large male early modern humans.

Craniosynostosis in the Middle Pleistocene human Cranium 14 from the Sima de los Huesos, Atapuerca, Spain

We report here a previously undescribed human Middle Pleistocene immature specimen, Cranium 14, recovered at the Sima de los Huesos (SH) site (Atapuerca, Spain), that constitutes the oldest evidence in human evolution of a very rare pathology in our own species, lambdoid single suture craniosynostosis (SSC). Both the ecto- and endo-cranial deformities observed in this specimen are severe. All of the evidence points out that this severity implies that the SSC occurred before birth, and that facial asymmetries, as well as motor/cognitive disorders, were likely to be associated with this condition. The analysis of the present etiological data of this specimen lead us to consider that Cranium 14 is a case of isolated SSC, probably of traumatic origin. The existence of this pathological individual among the SH sample represents also a fact to take into account when referring to sociobiological behavior in Middle Pleistocene humans.

Paleoneurology of an “early” Neandertal: endocranial size, shape, and features of Saccopastore 1

The Saccopastore 1 cranium was found near Rome in 1929, and its most probable age is about 120 ka (OIS 5e). The Neandertal morphology of the specimen was recognized just after the discovery by the Italian anthropologist S. Sergi, and subsequently confirmed by several authors. The present paper provides a complete description and analysis of the endocranial shape and features of this specimen, considering anatomical traits, metrics, and landmark data. The main endocranial diameters and the vascular traces resemble the morphology displayed by Middle Pleistocene humans, although lacking some traits described in the European samples referred to as ante-Neandertals. Nevertheless, proportions and endocranial shape support a definite Neandertal morphology, mostly taking into account the lateral development of the frontal lobes and the shape of the parietal areas. Therefore, it may be hypothesized that the Neandertal neurocranial architecture was present since at least OIS 5, as already suggested on the basis of ectocranial morphology.

Body size, body proportions, and encephalization in a Middle Pleistocene archaic human from northern China.

The unusual discovery of associated cranial and postcranial elements from a single Middle Pleistocene fossil human allows us to calculate body proportions and relative cranial capacity (encephalization quotient) for that individual rather than rely on estimates based on sample means from unassociated specimens. The individual analyzed here (Jinniushan) from northeastern China at 260,000 years ago is the largest female specimen yet known in the human fossil record and has body proportions (body height relative to body breadth and relative limb length) typical of cold-adapted populations elsewhere in the world. Her encephalization quotient of 4.15 is similar to estimates for late Middle Pleistocene humans that are based on mean body size and mean brain size from unassociated specimens.

Reciprocal evolution of the cerebellum and neocortex in fossil humans.

Human brain evolution involved both neurological reorganization and an increase in overall brain volume relative to body mass. It is generally difficult to draw functional inferences about the timing and nature of brain reorganization, given that superficial brain morphology recorded on fossil endocasts is functionally ambiguous. However, the cerebellum, housed in the clearly delineated posterior cranial fossa, is functionally and ontologically discrete. The cerebellum is reciprocally connected to each of 14 neocortical regions important to human cognitive evolution. Cerebellar volume varies significantly relative to overall brain volume among mammalian orders, as well as within the primate order. There is also significant diachronic variation among fossil human taxa. In the australopithecines and early members of the genus Homo, the cerebral hemispheres were large in proportion to the cerebellum, compared with other hominoids. This trend continued in Middle and Late Pleistocene humans, including Neandertals and Cro-Magnon 1, who have the largest cerebral hemispheres relative to cerebellum volume of any primates, including earlier and Holocene humans. In recent humans, however, the pattern is reversed; the cerebellum is larger with respect to the rest of the brain (and, conversely, the cerebral hemispheres are smaller with respect to the cerebellum) than in Late Pleistocene humans. The cerebellum and cerebral hemispheres appear to have evolved reciprocally. Cerebellar development in Holocene humans may have provided greater computational efficiency for coping with an increasingly complex cultural and conceptual environment.

Auditory capacities in Middle Pleistocene humans from the Sierra de Atapuerca in Spain.

Human hearing differs from that of chimpanzees and most other anthropoids in maintaining a relatively high sensitivity from 2 kHz up to 4 kHz, a region that contains relevant acoustic information in spoken language. Knowledge of the auditory capacities in human fossil ancestors could greatly enhance the understanding of when this human pattern emerged during the course of our evolutionary history. Here we use a comprehensive physical model to analyze the influence of skeletal structures on the acoustic filtering of the outer and middle ears in five fossil human specimens from the Middle Pleistocene site of the Sima de los Huesos in the Sierra de Atapuerca of Spain. Our results show that the skeletal anatomy in these hominids is compatible with a human-like pattern of sound power transmission through the outer and middle ear at frequencies up to 5 kHz, suggesting that they already had auditory capacities similar to those of living humans in this frequency range.

Later Middle Pleistocene human remains from the Almonda Karstic system, Torres Novas, Portugal

Later Middle Pleistocene archeological deposits of the Galeria Pesada (Gruta da Aroeira), Almonda Karstic System, Torres Novas, Portugal, yielded two archaic human teeth, a mandibular canine and a maxillary third molar. The C 1presents moderate and asymmetrical shoveling with a stout root. The slightly worn M 3exhibits at least four cusps with a large hypocone, three roots with large radicular plates, and an absence of taurodontism. They are moderately large for later Middle Pleistocene humans in their buccolingual crown diameters, although the M 3mesiodistal diameter is modest. The C 1exhibits labial calculus and multiple linear hypoplastic defects, but the M 3is lesion free. Both teeth are morphologically similar to those of other Middle Pleistocene European humans and reinforce a pattern of dental hypertrophy among these archaic Homo.

Cranial discrete traits in the Middle Pleistocene humans from Sima de los Huesos (Sierra de Atapuerca, Spain). Does hypostosis represent any increase in “ontogenetic stress” along the Neanderthal lineage?

Cranial discrete traits may be regarded as markers of dynamic responses to general and local perturbations of the morphogenetic pattern, particularly when they are viewed and examined in terms of hypostosis vs. hyperostosis. There are indications, in fact, that the variation between these two opposite conditions relates to mechanical stress suffered by the bony structures during early stages of growth and development. In a previous comparison between Neanderthals and modern humans, variable degrees and contrasting distribution patterns of hypostosis were found [Manzi et al. (1996),JHE30 : 511–527]. In the present paper, the occurrence, expression and cranial distribution of 20 hypo-hyperostotic traits are examined in the Middle Pleistocene sample from Atapuerca - Sima de los Huesos (Spain), with the principal aim being to test whether or not the degree of cranial hypostosis increases during the evolution of the Neanderthals. Other Middle Pleistocene representatives of the genus Homo (Kabwe and Petralona), the Italian Neanderthals, and a large recent European sample are also considered. A general consistency between the gradual appearance and stabilization of the Neanderthal cranial features and the results of the present analysis is found and is interpreted as an indication that hypostosis does mark the occurrence of “ontogenetic stress”. As suggested more than half a century ago by S. Sergi, an increase in “ontogenetic stress” in the Neanderthal lineage could result from the relationship between intracranial pressures and other (heterochronic) effects produced by the growth of a large brain (encephalization) and the ossification of an archaic (platycephalic) cranial vault.

Intrapopulational body size variation and cranial capacity variation in middle pleistocene humans: The Sima de los Huesos sample (Sierra de Atapuerca, Spain)

A sexual dimorphism more marked than in living humans has been claimed for European Middle Pleistocene humans, Neandertals and prehistoric modern humans. In this paper, body size and cranial capacity variation are studied in the Sima de los Huesos Middle Pleistocene sample. This is the largest sample of non-modern humans found to date from one single site, and with all skeletal elements represented. Since the techniques available to estimate the degree of sexual dimorphism in small palaeontological samples are all unsatisfactory, we have used the bootstraping method to asses the magnitude of the variation in the Sima de los Huesos sample compared to modern human intrapopulational variation. We analyze size variation without attempting to sex the specimens a priori. Anatomical regions investigated are scapular glenoid fossa; acetabulum; humeral proximal and distal epiphyses; ulnar proximal epiphysis; radial neck; proximal femur; humeral, femoral, ulnar and tibial shaft; lumbosacral joint; patella; calcaneum; and talar trochlea. In the Sima de los Huesos sample only the humeral midshaft perimeter shows an unusual high variation (only when it is expressed by the maximum ratio, not by the coefficient of variation). In spite of that the cranial capacity range at Sima de los Huesos almost spans the rest of the European and African Middle Pleistocene range. The maximum ratio is in the central part of the distribution of modern human samples. Thus, the hypothesis of a greater sexual dimorphism in Middle Pleistocene populations than in modern populations is not supported by either cranial or postcranial evidence from Sima de los Huesos. Am J Phys Anthropol 106:19–33, 1998. © 1998 Wiley-Liss, Inc.