Showing posts with label Palaeoarchaeology. Show all posts
Showing posts with label Palaeoarchaeology. Show all posts

Monday, 29 December 2025

A new reconstruction of the Yunxian 2 skull, and its implications for the relationship between the 'Archaic Homo sapiens' of China and Modern Humans.

The Middle Pleistocene, or Chibanian (roughly 774 100 to 129 000 years ago), has produced a range of Hominin fossils, with a surprising range of morphological diversity. Recent finds from Callao Cave on Luzon Island in 2007 and Rising Star Chamber in South Africa in 2013, have led to the description of two new Human species, Homo luzonensis and Homo naledi respectively, from this age. 

China has produced a diverse assemblage of Chibanian and older Hominin fossils, often labelled as 'Archaic Homo sapiens', although their relationship to Modern Humans has been the subject of debate for a long time. The Yunxian 1 & 2 crania were discovered on on a terrace of the Hanjiang River in the Yunyang District (formerly Yunxian) of Shiyan City, in Hubei Province, China, in 1989 and 1990. These skulls have been dated to between 1.1 million and 940 000 years ago, and are considered important for the understanding of the genus Homo in China, and to a certain extent the world, although both are heavily distorted, hampering attempts to reconstruct their original morphology and therefore phylogenetic relationships. 

The Yunxian 2 skull. Although the least distorted of the two crania from Yunxian, it is still partially crushed, and therefore difficult to interpret. Garry Todd/Hubei Provincial Museum/Wikimedia Commons.

In a paper published in the journal Science on 25 September 2025, a team of scientists led by Xiaobo Feng of the School of History and Culture at Shanxi University, Hubei Polytechnic University, and the Institute of Yunxian Man Site and School of History, Culture and Tourism, at Hanjiang Normal University, Qiyu Yin of the Institute of Vertebrate Paleontology and Paleoanthropology of the Chinese Academy of Sciences, and the College of Earth and Planetary Sciences of the University of the Chinese Academy of Sciences, Feng Gao of the Yunnan Institute of Cultural Relics and Archaeology, and Dan Lu, also of the Institute of Vertebrate Paleontology and Paleoanthropology of the Chinese Academy of Sciences, and the College of Earth and Planetary Sciences of the University of the Chinese Academy of Sciences, present a new reconstruction of the Yunxian 2 skull, and discuss the implications of that reconstruction for the relationship between the 'Archaic Homo sapiens' of China and Modern Humans.

Our understanding of the relationship between Modern Humans and our extinct Hominin relatives is based almost entirely on the reconstruction of the anatomy of extinct species from fossil specimens, many of which are quite distorted. Of the three fossil crania discovered at Yunxian, one (Yunxian 3) is still under preparation, having been found in 2022. Both of the previously discovered specimens are distorted, with Yunxian 1 being quite badly crushed as well. Computed tomography scans of Yunxian 2 have suggested that most of the deformation is due to the fragmentation and displacement of parts of the skull, rather than actual warping of the material. Feng et al. built a digital reconstruction of this fossil, using software to move parts back into their original position, and filling in some gaps with data from Yunxian 1. 

The reconstructed cranium is large and long, with a braincase which appears flattened in lateral view. It is smaller than the Harbin Xuchang crania, approximately the same size as the as Kabwe, Petralona, Bodo, Jinniushan, and Sangiran 17 crania, and larger than the Jebel Irhoud 1, Dali, and Maba crania. The reconstruction still lacks small parts of the zygomatic arches and the central incisors, but is otherwise intact. It contains a number of traits associated with earlier members of the genus Homo, including a thick supraorbital torus, a broad basicranium and palate, a long and low vault in lateral view, a receding frontal con￾tour, and a rather flat parietal contour, and a high, anteriorly projecting face. However, it lacks other traits associated with early members of the genus, such as a strongly angulated occipital with a prominent transverse torus. The specimen lacks the occipital bun, forward projecting midface, and general rounded 'en bombe' shape of Neanderthal skulls, and the upper parietal expansion of Modern Humans. The zygomaxillary region is transversely flat and faces anteriorly, similar to the Harbin, Dali, Jinniushan, and Hualongdong crania from Asia, as well as specimens assigned to Homo antecessor from Europe, as wellas Modern Humans. Its cheek￾bones are large and high.

Reconstruction of the Yunxian 2 cranium in standard views. (A) to (F) Anterior, posterior, inferior, superior, left, and right views, respectively. Brown colour indicates the fossil bone. The zygomatic bone and the tip of the left maxilla, as indicated with dark brown, were grafted and reconstructed by incorporating elements of Yunxian 1. White colour indicates the reconstructed parts inferred from the fracture edge and Yunxian 1. Neutral grey indicates the bones crushed and covered by other bones and matrix. Scale bar is 5 cm. Feng et al. (2025).

The matrix filling the skull is dense, and does not produce a good contrast with the bone material in computed tomography scans, preventing analysis of the features of the interior of the skull. However, Feng et al. were able to calculate an endocranial capacity of 1143 cm². However, the frontal lobe appears low and narrow, little expanded from the situation in early Homo, and unlike the expanded frontal lobes of Neanderthals and Modern Humans. 

Reconstruction of the endocranial cast of Yunxian 2. (A to F) Anterior, posterior, inferior, superior, left, and right views, respectively. Scale bar is 5 cm. Feng et al. (2025).

The traits which can be measured suggest that morphologically, Yunxian 2 appears to be intermediate between early Homo species and specimens such as Home erectus, Homo ergaster, and the Kabwe and Petralona crania, and later Asian skulls such as Harbin, Dali, and Jinniushan. 

A phylogenetic analysis based upon the new reconstruction of Yunxian 2, found it grouped with a group of Asian specimens including Dali, Jinniushan, Xujiayao, and Hualongdong and the Xiahe and Penghu mandibles, which have together been referred to as the Homo longi clade, based upon the suggestion that the Xujiayao, Xuchang, Xiahe, and Penghu specimens, as well as the Denisovans should be grouped together as a new species, Homo longi (although Feng et al. did not recover the Xuchang specimens as part of this group). Feng et al.'s reconstruction found this clade to be the sister to Homo antecessor, with the Homo longi clade plus Homo antecessor being the sister clade to Homo sapiens. Yunxian 2 is the oldest member of the Homo longi clade as recovered by Feng et al., but not the earliest branching. 

Phylogeny and divergence time of 57 selected fossil operational taxonomic units from the genus Homo. The topology of the tree was the majority consensus of the most parsimonious trees from the parsimony analysis in TNT. The divergence time was inferred from the Bayesian tip-dating analysis in MrBayes 3.2. Branch lengths are proportional to the division age in thousands of years (Ka). Numbers at the internal nodes are the median ages, and the blue bars indicate the 95% highest posterior density interval of the node ages. The red half-brackets on the right indicate the ranges of the Neanderthal, Homo longi, and Homo sapiens clades. The numbers in red highlight the ages of division of the three clades. Yunxian is also highlighted in red. Feng et al (2025).

The term 'Denisovan' was coined for a group of highly fragmentary fossils from the Denisova Cave in the Altai Mountains of Russia. Because of the fragmentary nature of these fossils, it has been possible to determine little about their original morphology, but DNA has been recovered from the specimens, revealing a great deal of genetic information. Studies of mitochondrial DNA from these specimens has suggested that this group branched off from the lineage that led to modern Humans before the Neanderthal line. However, studies based upon nuclear DNA have suggested that Denisovans and Neanderthals were sister groups, with a common ancestor more recent than their last common ancestor with Modern Humans. While the limited amount of data available makes this hard to resolve, Feng et al. suggest that the Denisovans were probably members of the Homo longi clade, with those morphological traits known all consistent with membership of this group. 

The divergence between the ancestors of Neanderthals and those of Modern Humans was for a long time considered to have happened between 700 000 and 500 000 years ago. However, more recent studies incorporating DNA recovered from Neanderthal specimens have suggested a much earlier split. Feng et al.'s analysis suggests that the Homo longi clade emerged as a distinct lineage about 1.2 million years ago, with Homo sapiens first appearing about 1.02 million years ago, around the time when Yunxian 2 was alive. The Homo longi and Homo sapiens clades are recovered as having diverged about 1.32 million years ago, while the Neanderthal clade diverged from the ancestors of these two groups about 1.38 million years ago.

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Sunday, 24 August 2025

Hominin remains from the Late Pliocene of Ledi-Geraru, Ethiopia.

The genera Homo and Paranthropus are common in the fossil record from about 2.0 million years ago. Both are thought to have derived from an earlier Australopithecus ancestor, with the most likely ancestor for the genus Homo generally thought to be Australopithecus afarensis. However, Australopithecus afarensis is not known after 2.95 million years ago, with Hominid fossils being rare over the intervening interval, spanning the latest Pliocene and earliest Pleistocene. Examples of Paranthropus have recently been described from 2.7 million-year-old deposits in the Omo-Turkana Basin of Ethiopia and Nyayanga in Kenya, and the 2.66 million-year-old deposits at Laetoli in Tanzania, while a jawbone attributed to the genus Homo has been found at Ledi-Geraru in Ethiopia which has been dated to 2.78 million years ago, pushing the presence of both these 'Pleistocene' genera back into the latest Pliocene, while a new species of Australopithecus, Australopithecus garhi, has recently been described from 2.5 million-year-old (earliest Pleistocene) deposits in the Afar region of Ethiopia.

In a paper published in the journal Nature on 13 August 2025, a team of scientists led by Brian Villmoare of the Department of Anthropology at the University of Nevada Las Vegas describe a series of recent Hominin finds made by the Ledi-Geraru Research Project in the Afar Basin of Ethiopia.

The Ledi-Geraru sites are located at the northern end of the palaeoanthropological sites of the Afar Basin, and has produced the only known evidence of the genus Australopithecus surviving after 2.95 million years ago, as well as the earliest evidence for the appearance of the genus Homo. Paranthropus has not been found in this area, but it is unclear whether this represents a genuine absence. The two sites of Ledi-Geraru, Lee Adoyta and Asboli are to the west of the Awash River, in an area cut through by the Mille and Geraru rivers and their tributaries. The deposits here are between 2.5 and 3.0 million years old, and have been dated by Argon-Argon radioisotope stratigraphy of volcanic layers, as well as magnitostratigraphy.

Map of the Ledi-Geraru Research Project area. (a) The  Ledi-Geraru Research Project area (yellow star) is located towards the northern  extent of palaeontological sites (red circles) in the Afar depression, Afar Region,  Ethiopia. (b) Within the Ledi-Geraru project area, the Lee Adoyta and Asboli fossil  sites are located approximately 12.5 km apart. Villmoare et al. (2025).

Argon-Argon dating relies on determining the ratio of radioactive Argon⁴⁰ to non-radioactive Argon³⁹ within minerals from igneous or metamorphic rock (in this case volcanic ash) to determine how long ago the mineral cooled sufficiently to crystallise. The ratio of Argon⁴⁰ to Argon³⁹ is constant in the atmosphere, and this ratio will be preserved in a mineral at the time of crystallisation. No further Argon³⁹ will enter the mineral from this point, but Argon⁴⁰ is produced by the decay of radioactive Potassium⁴⁰, and increases in the mineral at a steady rate, providing a clock which can be used to date the mineral.

Magnitostratigraphy uses traces of ancient magnetic fields preserved in iron minerals in rocks to trace ancient pole reversals; the poles only have two possible orientations (north pole in the north/south pole in the south or south pole in the north/north pole in the south) and these occasionally flip, with the poles exchanging positions. Pole reversals happen more-or-less at random, with periods between reversals occurring at intervals ranching from tens of thousands to millions of years, and reflected across the globe. This creates a pattern of magnetic reversals in sedimentary rocks that can be matched in different rocks across the globe.

The first specimen described by Villmoare et al. comes from the Gurumaha Sedimentary Package, which outcrops in narrow fault-bounded exposures in the central Lee Adoyta basin and in drag-faulted blocks adjacent to basalt ridges  bounding the basin to the east. This sedimentary package is cut through by the Gurumaha Tuff, which has been dated to 2.782 million years before the present. This is the unit which previously produced specimen LD 350, a 2.78 million-year-old mandible which is the oldest fossil assigned to the genus Homo

The specimen derived from this unit, LD 302-23, is a third right lower premolar found 22 m to the southwest and 7 m bellow specimen LD 350, but still above the Gurumaha Tuff layer. This tooth measures 11.5 mm in length and 10.5 mm in width, and has a fragment of enamel missing from its lingual corner, being otherwise well-preserved. The shape of the premolar is consistent with that seen in some examples from Australopithecus afarensis, but the pattern of cusps is quite different to anything seen in any member of the genus, making it unlikely that this tooth came from an Australopithecus. It also falls within the size range of both species of Paranthropus, but is quite different in shape. Third premolars from early members of the genus Homo are quite variable, but clearly differ from both those of Australopithecus and Paranthropus. Since this tooth falls within the size and shape variation found in these early Homo specimens, Villmoare et al. assign it to the genus Homo

New Hominin dentition from the Ledi-Geraru Research project. Right, from top: LD 302-23 P₃,  LD 750 P₄, AS 100 M¹ and AS 100 M².  Left, images show the LD 760 assemblage  (top, from left: maxillary molar, I², I¹, maxillary canine; bottom, mandibular molars). Villmoare et al. (2025).

The second specimen described, LD 750-115670, is an isolated lower fourth premolar, found at the base of an 8 m exposure of fossiliferous mudstones and sandstones at site LD 750. This site is located stratigraphically between the 2.63 million-year-old Lee Adoyta Tuffs and the 2.59 million-year-old Giddi Sands Tuff. 

The tooth crown is unworn, with all cusps preserved, although the root is broken off, giving a maximum root height of about 2 mm. The lack of wear may imply that the tooth was unerupted at the time of death. The tooth is 12.4 mm long and 11.4 mm wide, placing it at the upper end of the size range for Australopithecus afarensis or Australopithecus africanus, and too large for Australopithecus anamensis. No lower jaw or teeth are known for Australopithecus garhi, but the specimen is within a plausible size range for the species. It also falls within the size range of both Paranthropus species, but again is quite different in shape. It does resemble several fourth premolars attributed to early Homo, though Villmoare et al. note that these attributions are provisional, and that it is difficult to distinguish between early Homo and Australopithecus fourth premolars. Since this tooth lacks any distinctively Homo features, Villmoare et al. provisionally assign it to aff. Australopithecus sp..

A set of five associated lower molars were discovered at a site identified as LD 760, a flat sandy area approximately below the 2.63 million-year-old Lee Adoyta Tuffs. These are worn, with dentine exposed on their outer cusps, and wide for their lengths, giving them a squarish profile. Notably, the third molar is larger than the second molar, and the second molar is larger than the first, the first and second molars a relatively square, and the first and second molars lack a seventh cusp, all traits compatible with Australopithecus afarensis, but not early Homo. However, these teeth also differ from those of Australopithecus afarensis in several ways; they do not taper towards the rear, and lack a distinctive bilobate buccal contour. 

LD 760 molars compared to Australopithecus afarensis. Left molars from Ledi-Geraru specimen LD 760 (left) and Hadar specimen A.L. 400-1 (right). Measurements in mm of the LD 760 molars (length × width): LM1: 13.3 × 13.4, LM2: 14.5 × 14.6, LM3: 14.0 (estimated) × 15.7, RM1: 13.2 × 13.1, RM2: 14.8 × 15.2. Specimens are oriented with their buccal surfaces to the left and mesial surfaces up. Villmoare et al. (2025).

A partial upper molar was also recovered from this locality. This preserves the lingual grove of the tooth, which appears to be quite distinct. In the upper molars of Australopithecus garhi this groove is indistinct. Furthermore, the upper molars of Australopithecus garhi has a greatly reduced hypercone cusp, which leads the protocone cusp to take on a triangular shape. The hypercone is absent in the LD 760 specimen, but the protocone is present, and shows no sign of modification due to a reduced hypercone. However, the sample size for Australopithecus gahri is small, so this cannot be ruled out as a natural variation within the species.

Also found at the LD760 site were a right maxillary (upper) canine, a complete left maxillary lateral incisor and a left fragmentary maxillary central incisor. Thecanine (LD 760-115979) is well  preserved, lacking only the tip of the root. This has mesial and distal interproximal contact facets (wear marks caused by contact between teeth), with the interproximal contact facet having a matching distal interproximal contact facet on the second incisor. This means that the second incisor and canine were in contact, something typical of the the genus Homo, and unlike the situation in many Australopithecus specimens (including the Australopithecus gahri maxilla BOU-VP-12/130) where these two teeth are separated by a diastema (this trait is variable in Australopithecus afarensis, which may-or-may not have a diastema, so this could conceivably also be the case in Australopithecus gahri). The canine is also notably large, towards the upper end of the size range seen in Australopithecus (and much larger than anything seen in Paranthropus).

Comparative maxillary canine morphology. (a) Lingual (left) and  labial (right) views of the Ledi-Geraru LD 760-115979 canine (left) with Hadar Australopithecus afarensis specimens A.L. 763-1 (middle) and A.L. 333x-3 (right). Note that the  LD 760 canine is a right canine, whereas the Australopithecus afarensis canines are from the  left and are mirrored in these images. (b)–(d) LD 760-115979 ((b) shown in lingual view) contrasted with Hadar Australopithecus afarensis specimen A.L. 199-1 ((c) right canine  shown; distal to the upper right) and Bouri Australopithecus garhi specimen BOU-VP-12/130  (d) left canine, mirrored; distal is to the right). Note the simple mesial–distal  chisel-like wear pattern on the LD 760 canine (b) in contrast to the complex  multi-faceted wear pattern of Australopithecus afarensis (c) and the broad curved basin on the  distal side of the Australopithecus garhi upper canine ((d) this is seen on both left and right  canines). Images are oriented differently to emphasise the distinctive relevant morphology. Images in (b)–(d) are not to scale. Vallmoare et al. (2025).

The canine of Australopithecus gahri has a unique structure, with a shallow distal basin reminiscent of a talon which is contiguous with a wide wear furrow which runs along the entire post-canine dental row, something not seen in any other Hominin, and absent in the LD 760 canine. Morphologically, this tooth resembles those of Australopithecus afarensis, however, it has different wear patterns. In Australopithecus afarensis wear is mostly seen on the distal crest, whereas in the the LD 760 canine it is predominantly on the apex, suggesting a difference in diet and/or lifestyle.

The LD 760 individual clearly does not belong to Paranthropus, and is not morphologically consistent with any described species of Australopithecus. However, since it resembles Australopithecus afarensis more closely than anything else, Vallmoare et al. refer it to Australopithecus sp. indet.

The final specimens discussed come from the Giddi Sands unit in the Asboli region. These were found immediately below the 2.593 million-year-old Giddi Sands Tuff, and comprise a partial upper left first molar, and two fragments of an upper left second molar, which can be assembled to form a whole crown. They show little wear, and are close in shape to those of Australopithecus afarensis, although they lack the pronounced lingual occlusocervical sloping and general 'puffy' appearance of the molars of that species. They closely resemble the molars of early Homo specimens such as the 2.3 million-year-old specimen from the Busidima Formation at Hadar, or the 2.5-2.4 million-year-old specimen from Mille-Logya. It is quite different in form from the molars of Australopithecus gahri, and is small compared to the molars of either Australopithecus gahri or Paranthropus

Vallmoare et al. believe that although the Ledi-Geraru material is very limited, it provides clear evidence that both Australopithecus and Homo were present during the 3.0-2.5 million years ago interval, suggesting that multiple non-robust Hominin lineages were present in East Africa before 2.5 million years ago. 

The molar fragments from Asboli sufficiently resemble the Homo specimens from Hadar and Mille-Logya that Vallmoare et al. are confident that they represent the same species. However, they predate the older of these specimens (Mille-Logya) by at least 150 000 years. This adds to the evidence for an early appearance of Homo as Ledi-Geraru previously established by the LD 350-1 mandible, as does the LD 302-23 premolar which Vallmoare et al. describe from the Gurumaha sedimentary package.

The LD 750 and LD 760 material both come from the Lee Adoyta sedimentary package, although they are separated by 24 m of strata and the 2.63 million-year-old Lee Adoyta Tuffs. Nevertheless, both appear to represent a single species of Australopithecus (an assumption based in part upon the unlikelyhood of two similar species of Austrlopithecus coexisting in the same area). 

Vallmoare et al. consider four potential explanations for this material. Firstly, they might represent a late surviving population of Australopithecus afarensis (approximately 350 000 younger than the current youngest member of the species, from the Kada Hadar 2 Submember at Hadar). Secondly, they may represent an unknown species of Australopithecus ancestral to Paranthropus; the presence of Homo in this region implies that the Homo and Paranthropus lineages had diversified by this time, but Paranthropus itself appears to be absent. However, the oldest currently accepted member of the genus coming from about 2.7 million years ago from Nyayanga in Kenya and 2.66 million years ago from the Upper Ndolanya Beds at Laetoli in Tanzania, which makes this scenario less likely. Thirdly, they may represent earlier examples of Australopithecus gahri, which have not yet developed the distinctive features of the 2.5 million-year-old specimen BOU-VP-12/130, although the lack of similarities makes this unlikely. Finally, these specimens represent a new, as yet undescribed, species of Australopithecus.

Villmoare et al. conclude that were at least three species of Hominins present in the Afar Region between 3.0 and 2.5 million years ago, Australopithecus gahri, an unknown species of Homo, and an unknown species of Australopithecus. At the same time, Australopithecus africanus was still present in South Africa, and Paranthropus had already appeared in Kenya, Tanzania, and southern Ethiopia. The Ledi-Geraru environment was drier and more open than was typical for Australopithecus, very much the sort of environment associated with appearance and proliferation of the genus Homo, suggesting that, at least locally, Australopithecus may have been able to adjust to these more open environments, at least for a while.

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Tuesday, 6 August 2024

Using a multivariate analysis of the teeth of Sahelanthropus tchadensis to assess its status as a Hominin.

During the twentieth century palaeoanthropologists searching for the origin of the Hominins became convinced that the group has an East African origin. However, in 2001 scientists from the working on Upper Miocene deposits in the Toros Menalla region of Chad uncovered a group of possible Hominin fossils which they assigned to a new species, Sahelanthropus tchadensis, and which included a nearly complete, is distorted, cranium as well as a mandible and some isolated teeth.

Cranium TM 266-01-060-1, nicknamed Toumai, the holotype of Sahelanthropus tchadensis. Smithsonian Human Origins Program.

If correctly interpreted as a Hominin, then Sahelanthropus tchadensis would be the oldest known member of the group. As such the discovery has been subjected to a considerable amount of scrutiny, particularly as the original specimens had been subject to considerable taphonomic (post-mortem) alteration. One of the problems with the original material was that it was had to interpret the position of the foramen magnum (the whole in the skull through which the brain is connected to the central nervous system) because of distortion of the skull. This is important, because in (upright) Hominins it is located roughly in the centre of the bottom of the skull, while in Apes it is typically towards the back of the skull. A virtual reconstruction of the cranium of Sahelanthropus tchadensis has suggested that the foramen magnum would have been on the base. 

Sahelanthropus tchadensis also has a shorter, more vertical face, reduced canines and a related lack of a honing complex (a gap between the lower canine and the first premolar, into which the upper canine fits, allowing the two canines to rub together and sharpen, or 'hone' one-another), and a downward facing lip on the nuchal crest at the rear of the skull, all of which are Hominin traits. However, it also has a small, Ape-like neurocranium, as well as having a size and number of tooth-roots consistent with an Ape, presenting a mosaic of features consistant with an very early Hominin, or possibly an Ape closely related to the earliest Hominins.

Different analyses of a femur and two ulnae found at the same location as the cranium have suggested both that Sahelanthropus tchadensis was either habitually bipedal, or not habitually bipedal, and that it probably spent at least some of its time in the trees. Since a cladistic analysis has recovered Sahelanthropus tchadensis as a Hominin, this raises at least the possibility that Hominins were habitually walking on two legs 7 million years ago.

One aspect of Sahelanthropus tchadensis which has not been studied extensively is its dentition, despite a number of teeth and tooth fragments being available. In a paper published in the South African Journal of Science on 31 July 2024, Walter Neves, Leticia Valota, and Clovis Monteiro of the Institute of Advanced Studies at the University of São Paulo, present the results of a morphometric analysis which compared the upper posterior dentition of Sahelanthropus tchadensis to that of living Apes and fossil Plio-Pleistocene Hominins.

The upper posterior teeth were chosen as these were the teeth for which the mesiodistal and buccolingual diameters could be determined in Sahelanthropus tchadensis. These were compared to a selection of teeth from Pan troglodytes (Chimpanzees), as well as the Hominins Orrorin tugenensisArdipithecus ramidusAustralopithecus afarensisAustralopithecus africanusParanthropus boiseiParanthropus robustusHomo habilis, and Homo erectus.

For each tooth the ratio between the mesiodistal and buccolingual diameters (length and width) was calculated, and these were plotted on a distribution map. The teeth fell into approximately three groups on this map, one comprising modern Chimpanzees, one comprising the 'Robust Autralopithecines' Paranthropus boisei and Paranthropus robustus, and one containing all other Hominins.

Distribution of the species and specimens included in the study along the morphospace defined by principal component 1 (PC1) x principal component 2 (PC2). Neves et al. (2024).

The teeth of Sahelanthropus tchadensis fell within the non-Robust Hominin cluster, plotting closest to Ardipithecus ramidus, a Miocene-Pliocene species with a large number of known specimens, which again has some Ape-like characteristics, but which is generally accepted as a Hominin by palaeoanthropologists today. Neves et al. interpret this as supporting the hypothesis that Sahelanthropus tchadensis is a Hominin rather than an Ape.

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Sunday, 5 November 2023

Evidence for shoes in the Middle Stone Age of the Cape Coast, South Africa.

The appearance of clothing is considered to be one of the key stages in the development of Modern Humans, and the development of footwear, a complex form of clothing, is in itself an important technological leap. However, the exact time when footwear first appeared is unknown, with the first foot-coverings presumed to have been made of perishable materials unlikely to have survived. In the absence of preserved footwear, ichnology (the study of tracks and traces) provides the most plausible technique for detecting the first use of footwear among Hominins.

The oldest known shoes in the archaeological record are sandals made of woven Sagebrush bark from Oregon in the United States, the oldest of which have been dated to between 10 500 and 9200 years old. Also from the United States, a variety of sandals, moccasins, and slip-on footwear have been uncovered in Missouri, dating from between 8000 and 1000 years ago. The oldest shoe from Eurasia is a leather wrap-around shoe from southeastern Armenia, dated to between 5600 and 5400 years ago. Also from Eurasia, ‘Ötzi the Iceman' a frozen natural mummy found in the High Alps on the border between Austria and Italy, dated to more than 5000 years old, had a complete set of clothing, including complex footwear made from Bear-skin, Deer hide, and tree bark. A pair of Cow-hide sandals from Israel has also been dated to more than 5000 years ago.

Hominin trackways are known from many sites around the world, and can be identified by features such as the alignment and shape of the hallux, relative digit lengths, and the presence of a prominent medial longitudinal arch. However, these identifying features are based upon the assumption that the track-maker was unshod, and it is unlikely that all would be present were they to have been wearing some form of foot-covering. Thus, unsurprisingly, the majority of known Hominin tracks are considered to have been made by unshod tracemakers, although a few exceptions are known.

The oldest apparently shod Human tracks in the Americas are in Jaguar Cave, Tennessee, which are thought to be about 5000 years old. In Europe such traces include traces associated with a Magdalenian (Upper Palaeolithic) tool assemblage in Fontanet Cave, France, which appear to have been made by an individual wearing a soft shoe or sock. Also in France, Cussac Cave, footprints associated with a Gravettian assemblage and dated to between 38 000 and 31 000 years ago lack any digit impressions, despite these being present on both Human handprints and Bear footprints within the same cave, leading to the possibility that the trace-makers may have been shod.

Footprints without digit impressions are also known from a third site in France, Le Rozel, although these are about 80 000 years old, implying that the trace-makers were Neanderthals rather than Modern Humans. Possible evidence for footwear use by Neanderthals comes from Theopetra Cave in Greece, where a mixture of shod and unshod traces appears to have been left by a group of children around 130 000 years ago.

In South Africa, a number of possible footwear traces have been found on the Cape Coast, an area noted for its excellent Pleistocene tracks, including one trace which appears to show a clear sandal-imprint.

The use of clothing is thought to have been developed by Hominins inhabiting cool environments during Pleistocene cold spells, although the perishable nature of the material from which such cloths are likely to have been made, makes it unlikely that any preserved material will ever be found. Instead, tools used for the processing of skins, leather, and textiles are used as proxies for the materials themselves. This begins with the appearance of scrapers presumed to have been used for the preparation of skins, shortly followed by bone awls, needles and eyed needles, which imply the manufacture of increasingly complex clothing. Based upon this, it has been calculated that the first simple clothing probably appeared around 800 000 years ago, long before the emergence of Modern Humans, with several different Hominin species probably manufacturing and wearing cloths, with the possibility that footwear appeared at the same time. Molecular clock analysis suggests that Body Lice probably diverged from Head Lice between 170 000 and 80 000 years ago, by which time is is assumed that Humans were habitually wearing cloths all the time.

The Contrebandiers Cave site in Morocco, which has been dated to between 120 000 and 90 000 years ago, has yielded a variety of bone tools thought to have been used for the processing of leather and furs, but for the most part Pleistocene sites with evidence for tools likely to have been used in clothing manufacture and tracks made by Hominins are restricted to two areas, Western Europe and the Cape Coast of South Africa.

Such tools in Europe are known from the Middle Palaeolithic of southwestern France, with the oldest dating to about 51 400 years ago, and other examples between 48 000 and 41 000 years ago. These tools were presumably made by Neanderthals, and include lissoirs, tools which can be used to process hides, giving a smooth, tough, and reasonably impermeable finish (although not the same as modern commercially produced leathers).

The oldest sites in South Africa yielding tools which could have been made for making cloths are at Klasies River and Blombos Cave on the Cape Coast, with the Klasies River site dated to about 100 000 years ago and Blombos Cave mostly to about 80 000 years ago, although one awl, made from the bone of a Bird, was dated to at least 125 000 years ago.

Although these tools are taken as the earliest tools which are clearly associated with clothing manufacture, in both Europe and South Africa earlier stone tools could have been used to cut or pierce hides during the manufacture of simple garments, as could sharpened shell tools known from the Cape Coast. However, the fact that the oldest surviving footwear was made from woven Sagebrush rather than leather does suggest that caution should be applied when judging what ancient populations would have seen as suitable materials for making footwear.

It has also been suggested that wearing shoes might affect the way in which the foot develops, something which has been implied for a set of Human remains from Tianyuan in China dated to about 50 000 years ago, although the phalanges of the foot are seldom well preserved, limiting the number of instances in which this method can be applied.

Another Upper Palaeolithic site at Sunghir, in northern Russia, yielded a buried individual with remnant body decoration implying leggings or boots, as well as having extremely gracile lateral phalanges, something thought to be associated with habitual shoe wearing. This was particularly surprising as all Upper Palaeolithic Hominin tracks from Eurasia appear to have been made by barefoot trace-makers.

The relationship between footwear and foot morphology is complicated. Individuals who minimise their use of footwear are thought to have stronger foot muscles, and fewer pathologies of the feet, but this would be unlikely to result in any detectable skeletal difference between someone who never wore shoes and someone who occasionally did so, or even somebody who habitually wore very lightweight shoes.

In a paper published in the journal Ichnos on 28 August 2023, Charles Helm of the African Centre for Coastal Palaeoscience at Nelson Mandela University, Martin Lockley, also of the African Centre for Coastal Palaeoscience at Nelson Mandela University, and of the Dinosaur Trackers Research Group at the University of Colorado Denver, Hayley Cawthra, again of the African Centre for Coastal Palaeoscience at Nelson Mandela University, and of the Minerals and Energy Unit at the South African Council for Geoscience, Jan De Vynck, again of the African Centre for Coastal Palaeoscience at Nelson Mandela University, and of the Evolutionary Studies Institute at the University of the Witwatersrand, Mark Dixon, again of the African Centre for Coastal Palaeoscience at Nelson Mandela University, Renée Rust, again of the African Centre for Coastal Palaeoscience at Nelson Mandela University, and of the School of Geography, Archaeology and Environmental Studies at the University of the Witwatersrand, Willo Stear and Monique Van Tonder, once again of the African Centre for Coastal Palaeoscience at Nelson Mandela University, and Bernhard Zipfel, also of the Evolutionary Studies Institute at the University of the Witwatersrand, describe three Middle Stone Age sites on South Africa’s Cape Coast where tracks appear to have been made by Hominins wearing shoes, and discuss ways in which future studies of this topic could be approached.

Map of the Cape Coast of South Africa, showing Cenozoic deposits and places mentioned in the text. Helm et al. (2023).

Scientists from the African Centre for Coastal Palaeoscience have been studying the trace fossils of the Cape Coast since 2007. Here, a 350 km section of coast has frequent coastal aeolianites (wind-blown sand deposits) which have been buried by subsequent similar deposits than set with a carbonate cement, preserving a record of the people and animals which moved over them during the Middle Stone Age. As well as trackways left by Hominins, these studies revealed imprints left by Crocodiles, Giraffe, breeding Sea Turtles and very large Tortoises, none of which have left skeletal remains in the area.

The Cape Coast region has a remarkably rich archaeological record, tracing the appearance and development of numerous stages considered key steps on the way to Modern Human behaviour, including personal adornments, jewellery, the heat treatment of stone tools, art, and the use of abstract symbols. Palaeoclimatic studies of the same region indicate that the coastal plains upon which these deposits were laid down went through a cycle of exposure and inundation throughout the Pleistocene, which probably helped to maintain an ideal Hominin environment in the region. 

Prior to this project, Hominin tracks had previously been discovered at Langebaan on the west coast of South Africa, and Nahoon on the east coast of the country. Extensive study of the Cape Coast has produced four new track localities, and while another probable set of tracks has recently been identified at Langebaan. Combined with a set of tracks on an ancient lakebed in the Nefud Desert of Saudi Arabia, this represents the entire global inventory of tracks more than 46 000 years old which are attributed to Modern Humans.

All of the Hominin tracks on the Cape Coast are within the aeolianites of the Waenhuiskrans Formation, which together with the cemented foreshore, shoreface and lagoon deposits of the Klein Brak Formation, comprise the Bredasdorp Group. These deposits have been dated to between 400 000 and 36 000 years old, although the majority belong to Marine Isotope Stage 5, making them between 130 000 and 80 000 years old. To the east of Robberg, the correlate of the Waenhuiskrans Formation is the Nahoon Formation, which forms part of the Algoa Group. These Pleistocene aeolianites split readily along their bedding planes, so that tracks are often exposed on fallen blocks on the coast. Thes blocks are ephemeral, quickly being eroded away by the action of the waves.

Due to this ephemeral nature, tracks needed to be recorded quickly when exposed. This was done by recording their locality with a GPS unit and taking numerous photographs, which were then used to build photogrammetric models. Exposed trackways and footprints were also measured for track length, track width, track depth, pace length, stride length, and thickness of foresets, and individual footprints were examined for evidence of strap attachment points. 

In addition, studies were made of newly created tracks on sandy surfaces on the Cape Coast, with varying levels of moisture, slope, and firmness, and a variety of shoe types, including shoes with an open soft sole, an open hard sole, a closed soft sole and a closed hard sole. The best match for the fossil footprints was made by using an open hard soled shoe on soft, moist sand.

Shoe designs for this study were based upon shoes made by examples of sandals made by San peoples of southern Africa from the collections of the Blombos Museum of Archaeology in South Africa and Zambezi Heritage Museum in Namibia. The shoe which gave the closest results to the preserves footprints was based upon two such sandals, and was made from two layers of Cow-hide glued together, and laced by piercing three holes through this sole, one between the big toe and the adjacent digit, and two about half way along the length of the shoe, one at each side. The laces were made from the same Cow-hide as the upper part of the shoe, and were threaded through the holes and knotted underneath, protruding below, although in the museum examples prolonged use had levelled out the knots with the base of the sole. These sandals proved easy to attach to the foot.

Helm et al. present details of three footprint-bearing sites on the Cape Coast, these being, from west to east, Kleinkrantz, Goukamma, and Woody Cape.

At Kleinkrantz a slab measing 55 cm x 55 cm and 20 cm thick with apparent footprints on its upper surface, was found lying on a modern coast dune beneath a vegetated slope, which in turn lies beneath a cliff with an exposure of the Waenhuiskrans Formation. There was no sign of a recent rockfall, and samples taken from the Waenhuiskrans Formation in this area have been dated to between 148 000 and 79 000 years old.

(A) The Kleinkrantz site in the Garden Route National Park; scale bar is 10 cm. (B) Photogrammetry colour mesh of the Kleinkrantz site, using 48 images. Photos were taken average 36.4 cm from the surface. The reprojection error is 0.39 pix. Vertical and horizontal scales are in metres. Arrows indicate possible strap attachment point impressions. Helm et al. (2023).

This slab has two similar depressions, one in front of the other, each with crisp three crisp and well-defined margins, two straight parallel, the third near semicircular. One of these appears to have been a double impression, with a slightly narrower impression overlying an slightly wider one. The wider of these is 9 cm wide, the narrower 7 cm, while the single impression is 8 cm wide. Both features are at least 16 cm long. The narrowest impression also has three depressions within it consistent with the position of the strap knots on the experimental sandal.

Two other possible track prints are also present, although these are more amorphous, and were apparently made by a person travelling in a different direction, though again possible strap-attachment knot impressions are present. 

The second site lies within the Goukamma Nature Reserve, between Sedgefield and Knysna, where the Waenhuiskrans Formation has been dated to between 136 000 and 79 000 years old, although recent dates obtained suggest parts of the formation may be as young as 73 000 years old. Here a fallen slab has three footprints with crisp margins and no signs of digits, one of which particularly resembles the imprint of a shod foot. This print measures 11.5 cm in length and 6 cm in width.

(A) The Goukamma surface; scale bar = 10 cm. (B) Angled view of the Goukamma site; scale bar is 10 cm. (C) Photogrammetry colour mesh of the Goukamma site, using 59 images. Photos were taken average 29.9 cm from the surface. The reprojection error is 0.39 pix. Vertical and horizontal scales are in metres. Helm et al. (2023).

The third site, Woody Cape, is situated in the coastal portion of the Addo Elephant National Park. Here a detached slab from the Nahoon Formation shows a trackway comprising four footprints, one of which is partial. The prints are 10-12 cm long and 5-6 cm wide, with a pace length of 19 cm. Raised areas to the left of each print imply some downslope movement.

Photogrammetry colour mesh of the Woody Cape site, using 37 images. Photos were taken average 29.2 cm from the surface. The reprojection error is 0.57 pix. Vertical and horizontal scales are in metres. Helm et al. (2023).

Helm et al. also attempted to recreate a trackway similar to the Pleistocene examples, using a hard-soled sandal based upon museum specimens, on a modern, moist, soft, level dune surface. This produced footprints with slight out-toeing about 26 cm in length, with a width of 11.5 cm and a pace length of 63 cm. The front and back margins of the prints were semicircular, the sides parallel. Strap attachment marks can be seen.

Photogrammetry colour mesh of the neoichnological trackway (level surface, hard-soled sandal, soft substrate) using 63 images. Photos were taken average 37 cm from the surface. The reprojection error is 2.07 pix. Vertical and horizontal scales are in metres. Helm et al. (2023).

Helm et al. accept that the evidence they provide cannot be seen as irrefutable, however, the presence of similar tracks at three separate sites, combined with the recreation of similar prints with a pair of modern sandals based upon a historic design, does strongly suggest that similar sandals were being worn by Humans in the area more than 70 000 years ago. All three sets of footprints appear to have been made by trackmakers smaller than modern adult Humans, suggesting that they were either adults of smaller stature, or children.

A pair of sandals on exhibit in the Blombos Museum of Archaeology in Still Bay, viewed from above (A), and below (B), showing strap attachment points; scale bars are 10 cm. Helm et al. (2023).

The tracks of bare feet are distinctive, due to the presence of digits and the raised arch of the foot. The impression likely to have been made by a shoe of unknown design is less easy to predict, but it is likely to have followed the general plan of a Hominin footprint, i.e. twice as long as wide, and soft foot-coverings may still preserve the presence of an arch, and rounded front and back margins. It is also quite likely that the front portion of the print will be wider than the rear portion, and that the curve of the front and back parts of the foot will be different. Trackways will reflect the fact that Hominins are bipeds with a narrow stride. In some cases, distinctive traces may be made by the footwear, such as attachment marks from straps.

A pair of sandals on exhibit in the Zambezi Heritage Museum in Katima Mulilo, Namibia, viewed from above (A), and below (B), showing strap attachment points; scale bars are 10 cm. Helm et al. (2023).

Previous attempts to recreate impressions likely to have been made by ancient footwear mostly concentrated on the use of soft leather shoes by persons walking over clay substrates. These studies found that shod tracks were typically longer and narrower than unshod tracks, and that this became more exaggerated as the substrate got moister and the footprints deeper. Shod footprints were also typically simpler than unshod ones, most obviously in the lack of toe-prints. 

These findings may have some implications for studies being carried out on the Cape Coast of South Africa, where some footprints have been considered questionable due to their apparently elongate shape.

Historic sandals in the collections of museums shed light on the appearance of shoes within the region in the recent past, and therefore possibly those that might have been used by people in the remote past armed with similar resources and environmental challenges. These sandals are remarkably symmetrical, with little difference between the width of the front and rear portions of the shoe. The pair from the Zambezi Heritage Museum in Namibia are believed to have been made from Buffalo hide in the 1920s.

The San people who made these sandals are known to have used a variety of Animal skins, with different groups using specific hides, including Eland, Hartebeest, and Wildebeest, while other Animals, such as Gemsbok, were avoided by all groups. Furthermore, different types of sandals were made for different purposes, including special shoes for seniors, shoes for uphill travel, and running sandals with a hook under the toe giving extra purchase when running over soft sand (possibly in imitation of the foot of an Ostrich, the fastest bipedal Animal present in the region).

A pair of ‘running sandals’ in the Fourie Collection in Museum Africa, accession number MM40-69-2416; scale bar is 10 cm. Justine Wintjes in Helm et al. (2023).

Rock art in Southern Africa also provides a record of the use of footwear in the region, albeit one that only stretches back a few thousand years. The artwork at Baviaanskloof in Eastern Cape Province is thought to be about 2000 years old, and includes a 21 cm high figure interpreted as a shaman, naked apart of a pair of laced sandals and a kaross (cloak made from Animal skin). The fact that a largely naked figure is wearing elaborate sandals suggests that these were important items to the people of the region.

(A) A male figure, apparently with footwear and shoelaces (magnified in inset), in a San pictograph from Baviaanskloof.  (B) Rock art of footwear that appears to depict shoelaces, Baviaanskloof. Helm et al. (2023).

In total, four figures with footwear are present within at Baviaanskloof, two of which have identifiable laces. Sandals can also be seen in the rock art of the western US, where Pueblo petroglyphs made between 1000 and 1200 AD also depict footwear.

Helm et al. note that the identification of tracks made by Hominins with footwear can be difficult, and suggest a set of criteria which may make this easier for future researchers. Firstly, any trackway examined should be of sufficient length for the study to be useful. Secondly, Hominins are bipedal, with a narrow gait, and any track made by them should reflect this. Thirdly, Hominin footprints are roughly twice as long as wide, regardless of footwear. Fourthly, Hominin tracks tend to have rounded fronts and backs. Fifthly, any trackway analysed for the presence of footwear should be of sufficient quality, with crisp margins. Sixthly, shod footprints will lack the diagnostic features of unshod footprints, such as digit marks. Finally, the footprints may have diagnostic features associated with the footwear, such as strap attachment points.

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