Increasing evidence for both taxonomic diversity and early stone manufacture during the Pliocene highlights the importance of the hominin fossil record from this epoch in eastern Africa. Here, we describe dental remains from Lomekwi (West Turkana, Kenya), which date from between 3.2 and 3.5 Ma. The sample was collected between 1982 and 2009 and includes five gnathic specimens and a total of 67 teeth (mostly isolated permanent postcanine teeth). Standard linear dimensions indicate that, although the Lomekwi teeth are relatively small, there is broad overlap in size with contemporary Australopithecus afarensis and Australopithecus deyiremeda specimens at most tooth positions. However, some dental characters differentiate this sample from these species, including a relatively large P4 and M3 compared with the M1, a high incidence of well-developed protostylids, and specific accessory molar cuspules. Owing to a lack of well-preserved tooth crowns (and the complete absence of mandibular teeth) in the holotype and paratype of Kenyanthropus platyops, and limited comparable gnathic morphology in the new specimens, it cannot be determined whether these Lomekwi specimens should be attributed to this species. Attribution of these specimens is further complicated by a lack of certainty about position along the tooth row of many of the molar specimens. More comprehensive shape analyses of the external and internal morphology of these specimens, and additional fossil finds, would facilitate the taxonomic attribution of specimens in this taxonomically diverse period of human evolution.
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Sexual dimorphism is an important component of the total variation seen in populations and plays a key role in taxonomic debates. In this study, microtomographic (microcomputed tomography) techniques were applied to a sample of hominin teeth from the Sima de los Huesos site (Spain). Dental tissue proportions of the permanent canines were assessed to characterise the pattern and degree of sexual dimorphism within this population. In addition, the possible similarities and differences with the Homo neanderthalensis remains from Krapina (Croatia) and with a recent modern human sample were evaluated. A combination of classical statistical approaches with more novel techniques allowed us not only to ratify the sex allocation of the individuals previously assigned in the literature but also to estimate the sex of the youngest individuals, which were not assessed in previous studies. Likewise, the sexes of certain extensively worn canines and isolated pieces were estimated. As a result, the sex ratio observed in our dental sample from the Sima de los Huesos population is 5:9 (Nm:Nf). In general terms, both Sima de los Huesos and Krapina dental samples have a degree of sexual dimorphism in their permanent canine tissue proportions that does not surpass that of modern humans. The marked dimorphic root volume of Sima de los Huesos mandibular canines is the exception, which surpasses the modern human mean, although it falls within the 95% confidence interval. Therefore, our results do not support that dental tissue proportions of the European Middle Pleistocene populations were more dimorphic than in modern humans. However, the differences in canine tissue proportions are great enough to allow sex estimation with a high degree of confidence.
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The data set used within the present study includes hominin trackways that have been attributed to six distinct hominin species within two genera, spanning from the Pliocene to the Holocene. Even across such a broad sample of time and space, some aspects of track morphology are found to be remarkably consistent. However, between-sample differences were identified in three morphological aspects of the tracks. These differences are related to the prominence and position of the medial midfoot impression, the abduction angle of the hallux impression, and the length of the forefoot relative to the rest of the track. Generally, comparing sites across time from the Pliocene to the Holocene, the MLA is more prominent, the hallux is less abducted (this variable achieved the greatest discrimination between assumed species), and the forefoot is relatively shorter in more recent track samples. The linear dimensions classified the potential H. antecessor tracks from Happisburgh (pronounced Haysbra) as being most similar to the H. erectus prints from Ileret, suggesting the dimensions and shape of Pleistocene tracks were likely similar. Importantly, this is the first study to specifically examine the morphology of the Happisburgh tracks within such a broad comparative context. The Happisburgh tracks are found to be morphologically similar to other Early Pleistocene and Holocene hominin tracks consistent with the geological age of the site, yet distinct from the Pliocene tracks from Laetoli.
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Sediment cores retrieved from the Pleistocene Olduvai Basin by the Olduvai Gorge Coring Project (OGCP) provide a high resolution record of tuffs and other volcaniclastic deposits, together with a lacustrine sedimentary record full of paleoenvironmental indicators. Correlating tuffs between the cores and outcrops at Olduvai, where these tuffs are identified at paleoanthropologically important sites, is critical for applying the new paleoenvironmental data to the conditions under which hominins lived. Tuffs and other volcaniclastic deposits from three cores were analyzed for mineral assemblages and glass and mineral major element compositions (feldspar, augite, hornblende, titanomagnetite, and glass where possible) to compare to published geochemical fingerprint data, based on marker tuffs from outcrop equivalents at Olduvai Gorge. In combination with stratigraphic position, these mineralogical and geochemical data were used to correlate between the cores and outcrops, providing direct temporal tie-lines between the cores and sites of paleoanthropological interest. Direct correlations are most certain for Olduvai Bed I, where all major tuff markers from outcrop are identified for one or more of the three core sites, and for the upper part of the underlying Ngorongoro Formation, which includes the Coarse Feldspar Crystal Tuff (CFCT) and Naabi ignimbrites exposed in the oldest Pleistocene exposures of the Western Gorge. Also characterized were the mineral and glass compositions of tuffs and ignimbrites pre-dating the oldest exposed outcrop units, extending our record of explosive events from the Ngorongoro Volcano. While no specific correlations can be confirmed between individual Bed II tuffs in the cores and in outcrops, correlations are possible between the cores themselves (using newly identified tuff compositions), and some potential correlations (non-unique, based on individual mineral phases) between core and outcrop can be used in conjunction with other stratigraphic tools to help constrain the intervals in question.
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The cranium from Broken Hill (Kabwe) was recovered from cave deposits in 1921, during metal ore mining in what is now Zambia. It is one of the best-preserved skulls of a fossil hominin, and was initially designated as the type specimen of Homo rhodesiensis, but recently it has often been included in the taxon Homo heidelbergensis. However, the original site has since been completely quarried away, and—although the cranium is often estimated to be around 500 thousand years old—its unsystematic recovery impedes its accurate dating and placement in human evolution. Here we carried out analyses directly on the skull and found a best age estimate of 299 ± 25 thousand years (mean ± 2σ). The result suggests that later Middle Pleistocene Africa contained multiple contemporaneous hominin lineages (that is, Homo sapiens, H. heidelbergensis/H. rhodesiensis and Homo naledi), similar to Eurasia, where Homo neanderthalensis, the Denisovans, Homo floresiensis, Homo luzonensis and perhaps also Homo heidelbergensis and Homo erectus were found contemporaneously. The age estimate also raises further questions about the mode of evolution of H. sapiens in Africa and whether H. heidelbergensis/H. rhodesiensis was a direct ancestor of our species
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Figure 3. Stratigraphic section (left) shown with the image (right) of the outcrop highlighting the positions of the three major tuffs. All hominin fossils were found on the surface or in secondarily deposited sediment below the Ileret Tuff (1.52 ± 0.01 Ma). Some of the remains were found above the Lower Ileret Tuff (1.53 ± 0.01 Ma) indicating that the bones must have been buried above it. The large excavation area is visible on the right side of the outcrop and gully; the lower footprint level (Bennett et al., 2009, Dingwall et al., 2013, Hatala et al., 2017) is exposed on the left side of the image. All fragments of KNM-ER 47000 were located on the surface of the lower portion of the outcrop or secondarily buried in sediment eroded from the drainage that extends up the slope from the excavation site toward the right margin of the picture.
Paranthropus boisei was first described in 1959 based on fossils from the Olduvai Gorge and now includes many fossils from Ethiopia to Malawi. Knowledge about its postcranial anatomy has remained elusive because, until recently, no postcranial remains could be reliably attributed to this taxon. Here, we report the first associated hand and upper limb skeleton (KNM-ER 47000) of P. boisei from 1.51 to 1.53 Ma sediments at Ileret, Kenya. While the fossils show a combination of primitive and derived traits, the overall anatomy is characterised by primitive traits that resemble those found in Australopithecus, including an oblique scapular spine, relatively long and curved ulna, lack of third metacarpal styloid process, gracile thumb metacarpal, and curved manual phalanges. Very thick cortical bone throughout the upper limb shows that P. boisei had great upper limb strength, supporting hypotheses that this species spent time climbing trees, although probably to a lesser extent than earlier australopiths. Hand anatomy shows that P. boisei, like earlier australopiths, was capable of the manual dexterity needed to create and use stone tools, but lacked the robust thumb of Homo erectus, which arguably reflects adaptations to the intensification of precision grips and tool use. KNM-ER 47000 provides conclusive evidence that early Pleistocene hominins diverged in postcranial and craniodental anatomy, supporting hypotheses of competitive displacement among these contemporaneous hominins.
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Figure 1. Right upper limbs of a modern human (left), chimpanzee (center), and KNM-ER 47000 (right), which preserves lateral portions of the scapula, the distal portion of the humerus, most of the ulna, and most of metacarpals (MCs) 1-3 and proximal phalanges 2-4. KNM-ER 47000 has primitive traits including a gracile thumb MC, lack of MC 3 styloid process, curved phalanges with prominent flexor sheaths, a long and curved ulna, a humerus with thick cortical bone and a prominent brachioradialis flange, and obliquely oriented scapular spine. Derived traits include a relatively long thumb, short manual phalanges, and a lateral scapular glenoid orientation. Scale bar at right is 10 cm.
Here we present the descriptive and comparative study of two immature scapulae recovered from the TD6.2 level of the Gran Dolina cave site (Sierra de Atapuerca, Spain) and assigned to Homo antecessor. This is the first time that data on the morphology and dimensions of the scapulae of a European late Early Pleistocene hominin population are provided. Considering the state of development and the linear dimensions, the scapula ATD6-116 could belong to a child of about 2–4 years. The morphology of ATD6-116 clearly departs from that of the Australopithecus afarensis juvenile specimen DIK-1-1, pointing to functional differences in locomotor behavior between Australopithecus and the late Early Pleistocene hominins. The immature scapula ATD6-118 belonged to an immature individual with a development of the scapula equivalent to that of adolescents of recent human populations. The scapulae ATD6-118 and KNM-WT 15000 present a similar state of development. Although the scapula KNM-WT 15000 is clearly larger than ATD6-118, these two specimens share some characteristics such as their relative narrowness and the value of the axilloglenoid and spinoglenoid angles. The glenoid fossa of ATD6-116 show a lateral orientation, whereas in ATD6-118 the glenoid fossa is slightly cranially oriented, but still within the range of variation of modern humans. The glenoid index of both ATD6-116 and ATD6-118 is low in accordance to the values usually observed in other early hominins, thus showing the primitive condition for this feature. Both scapulae show a ventrally placed axillary sulcus. The presence of this primitive feature in ATD-116 confirms that the shape of the axillary border has a genetic basis and it is not related to physical activity.
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Palaeolithic wooden spears provide rare but unique insights into early hunting technology. Examples from Schöningen, Germany indicate that spear tips were sometimes asymmetrical. This asymmetry has previously been interpreted as evidence for planning depth. A more parsimonious explanation, however, is that asymmetrical tips could be more efficiently produced (i.e., took less time to make) than symmetrical ones. Here, we experimentally investigated two different manufacturing processes, producing asymmetrical and symmetrical spear tips, while also testing the influence of biometric factors on spear-tip manufacturing efficiency (measured by time). One group of experimental participants sliced the wooden stave into an asymmetrical shape (slicing on one side of the stave with a steel blade), while the other group created a symmetrical shape (slicing on multiple sides of the stave). Based on time taken, results demonstrated no significant difference in efficiency between symmetrical and asymmetrical spear-tip manufacture. Conversely, some biometric characteristics (specifically pinch strength) present a more dominant influence in explaining time variation. These results demonstrate that the asymmetrical tips at Schöningen were not merely a byproduct of maximizing efficiency during the manufacturing process, but rather are evidence of planning and the associated cognitive capacities of these later Middle Pleistocene hominins.
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Along the Angolan coast, the Early Paleolithic sites of Dungo IV and V (Baia Farta, Benguela) have delivered a rich pre-Acheulean lithic industry testifying the antiquity of the hominin settlement in western Africa despite the current absence of any hominin fossil in the area. In Dungo IV, the Paleolithic level is located on a conglomeratic paleo-beach (104 m a.s.l.) buried under an at least 3 m thick sandy layer. In Dungo V, two unearthed large whale fossils are associated with numerous lithic tools intimately mixed with the whale bones. This is the oldest evidence of stranded marine mammal scavenging by hominins in this part of Africa. The lack of volcanism and fossils makes chronological constrain difficult. Considering its configuration, the Dungo IV site may be relevant for a dating based on both the 10Be and 26Al cosmogenic nuclides. For this purpose, a depth profile all along the sandy layer overlying the archeological layer has been sampled. Statistical treatments performed on the 26Al/10Be ratios obtained for the depth profile demonstrate that they all belong to the same population. If we consider that the samples have always been at or close to their sampling depth, the regression modeling allows computing that the surface sedimentary layer has been emplaced at least 614 ka ago and less than 662 ka ago. On the other hand, if we consider that the surface deposit has been truncated, burial durations ranging from 585 ka to 786 ka and truncations lower than 4 m result from the modeling of the evolution of the 10Be and 26Al concentrations as a function of depth.
The analyses of four pre-Acheulean artefacts lead to a minimum burial duration of 730 ka and a maximum burial duration of 2.11 Ma.
The low pre-burial denudation rates modeled from the data acquired for the stone tools as well as for the overlying layer (1–16 m.Ma−1) imply large inherited 26Al and 10Be concentrations. The post-depositional maximum denudation rate of 71 m.Ma−1associated with both the lithic artefacts and the surface sedimentary layer (considering that the samples have always been at or close to their sampling depth) as well as the deduced maximum uplift rate of ∼170 m.Ma−1 are in agreement with the known tectonic evolution and the climatic variability of this area.
This study confirms the antiquity of the hominin presence in western Africa more than 2000 km away from the closest old hominin fossil sites.