Fossils recovered at ‘Cradle of Humankind’ site
Sterkfontein is a set of limestone caves near Krugersdorp, South Africa. It is one of the most important hominin fossil-bearing sites in the world and finds include the female australopithecine Mrs Ples, discovered in 1947 and recently voted No.95 in a list of 100 Great South Africans. Sterkfontein has yielded stone tools in addition to hominin fossils and it is now part of the Cradle of Humankind, a World Heritage Site named by UNESCO in 1999. Most of the finds have been made in Members 4 and 5 of the cave’s sedimentary sequence, but rather less well known is the large underground chamber known as Milner Hall.
It is from Milner Hall that the discovery is reported of a hominin adult upper right molar (M1) tooth and a proximal phalanx finger bone, probably from a left hand. The chamber has previously yielded only stone tools, and association with these suggests that the fossils are 2.18 million years old.
The tooth is broadly closer to Homo than to Australopithecus or Paranthropus. It most closely resembles the Olduvai OH 6 first molar assigned to Homo habilis and a first molar assigned to the recently-proposed Homo naledi. The shape and size of the tooth’s cusps align it to early Homo.
The finger bone is larger and more robust than that of any hominin so far discovered in South Africa. It resembles the Olduvai Homo habilis fossil OH 7, but is much larger. It is markedly curved, within the range of Australopithecus afarensis and suggesting adaptation for tree-climbing, but it lacks other features associated with arborealism, such as a strongly developed flexor apparatus and a mediolaterally expanded diaphysis; these features are present in A. afarensis, Homo habilis and present-day chimpanzees. The finger bone possesses an enigmatic mixture of primitive, derived and unique characteristics. It is not clear whether or not it belonged to the same individual and its taxonomic affinities are at this stage uncertain.
Stratford, D., Heaton, J., Pickering, T., Caruana, M. & Shadrach, K., First hominin fossils from Milner Hall, Sterkfontein, South Africa. Journal of Human Evolution 91, 167-173 (2016).
New skull with ‘enigmatic’ jawbone and differing tool technologies suggests that two different hominin groups are represented by Dmanisi remains.
The former Soviet republic of Georgia is located at the crossroads of Europe and Asia. Lying on the eastern shores of the Black Sea, it was the destination of Jason and the Argonauts in their quest for the Golden Fleece, but long before this it was a stopping point for the earliest-known hominin migration out of Africa. In 1984, stone tools were discovered at the small medieval town of Dmanisi in the southeast of the country, 93 km (58 miles) southwest of the capital, Tbilisi. Archaeologists broke through the foundations of a medieval building into an ancient river deposit, where simple stone tools resembling those made by the earliest humans were found with the bones of extinct mammals.
During the 1990s, the remains of early humans were recovered, including two partial skulls and a lower jawbone. The fossils were dated by palaeomagnetic, potassium-argon and argon-argon methods, giving an age for the remains of 1.77 million years old (Gabunia, et al., 2000). Subsequent dating of the stone tools indicated that the site was first occupied 1.85 million years ago, and that repeated occupations continued over a period of 80,000 years. There was evidently a long-term human presence in the Caucasus at around or even before the time of the earliest evidence for Homo erectus in Africa (Ferring, et al., 2011).
There have been a number of subsequent discoveries of human remains at the site. These include the skull, lower jawbone and partial skeleton of an adolescent (Vekua, et al., 2002; Lordkipanidze, et al., 2007); the skulls and lower jawbones of two adults (Lordkipanidze, et al., 2006; Lordkipanidze, et al., 2013); and postcranial bones from three other individuals, all adults (Lordkipanidze, et al., 2007). One of the skulls belonged to an elderly male who had lost all but one of his teeth some years prior to his death. He could not have survived unaided and must have been cared for by his companions throughout those last years of his life (Lordkipanidze, et al., 2005; Lordkipanidze, et al., 2006). The other skull, the fifth to be discovered at the site and hence known as Skull 5, is characterised by a large face and thick browridges. Skull 5 is complete and undeformed; it is the only known fully-preserved adult hominin skull from the early Pleistocene (Lordkipanidze, et al., 2013).
From the various remains, body size metrics have been estimated for the Dmanisi hominins. They were 1.45 to 1.66 m (4 ft. 9 in. to 5 ft. 5 in.) tall and weighed 40.0 to 50.0 kg (88 to 110 lb.). The cranial capacities of the five skulls range from 546 to 730 cc, about half that of a modern human. The encephalization quotient (a measure of brain size in relation to body size) lies in the range from 2.4 to 3.13; a figure that is at the lower end of the estimates for African Homo erectus, and is more comparable to that of Homo habilis or Australopithecus (Lordkipanidze, et al., 2007; Lordkipanidze, et al., 2013).
The Dmanisi hominins display a mosaic of primitive and derived (more modern) features. Their limb proportions were similar to those of a modern human. The lower limbs and feet were essentially modern, although the feet turned slightly inwards. On the other hand, the forearm lacked what is known as humeral torsion. In modern humans, the elbow joint is typically rotated relative to the shoulder joint, so that the forearm naturally hangs with the palms facing inwards; but the Dmanisi forearm lacked this rotation, so their palms were oriented more forwards. The inward-turning feet, lack of humeral torsion, small body size and small brain size may be seen as primitive traits, sharing more in common with Homo habilis than with Homo erectus (Lieberman, 2007; Lordkipanidze, et al., 2007).
Initially assigned to African Homo erectus (Vekua, et al., 2002), the Dmanisi hominins were later put forward as a new human species, Homo georgicus (Gabunia, et al., 2002); though this proposal has since been retracted (Lordkipanidze, et al., 2013), and it has been suggested that early African Homo erectus was not only quite widespread, but also unusually variable in both body and brain size, and also less modern than sometimes supposed (Lieberman, 2007).
Two more radical (and diametrically-opposed) possibilities have recently been put forward. The first is that the various species often proposed for early African Homo (Homo habilis, Homo rudolfensis, Homo ergaster and Homo erectus) were all actually variants of the same species, and that early Homo was a single lineage which evolved over time without differentiating into multiple species. This conclusion is based on a claim that shape variation between the five Dmanisi skulls is roughly the same as that seen among the various early Homo skulls from East Africa, even though the former represents a single species and the latter are generally thought to represent several (Lordkipanidze, et al., 2013).
The second proposal (Bermúdez de Castro, et al., 2014) is that Skull 5 represents a different group of early hominins to that of the other Dmanisi remains. The lower jawbone is larger than those of others, and is said to represent a ‘large and somewhat enigmatic individual’. Its shape differs, and the differences cannot be accounted for in terms of body size or sex. It possesses a mosaic of primitive and derived features that are absent from other Dmanisi specimens. Furthermore, patterns of dental wear suggest a higher intake of fibrous and abrasive foods. It has accordingly been suggested that the jawbone is adapted to a different ecological niche to the other Dmanisi hominins, and that it represents a different species.
Although tools document a long-term human presence at Dmanisi, all the actual human remains were found in the same geological layer. This makes the ‘two species’ scenario problematic, as it implies that both species lived at about the same time. However, the stratigraphy of Dmanisi is complex, and it is possible that the fossil remains were re-deposited in the same geological layer after initially occupying sediments of different ages. It has also been claimed that the tools found at Dmanisi are consistent with the existence of two different populations.
More evidence is needed to determine just where the Dmanisi hominins fit into the broader human evolutionary picture, but it is becoming clear that the first hominin dispersal out of Africa was a far more complex process than was at one time supposed.
1. Gabunia, L. et al., Earliest Pleistocene Hominid Cranial Remains from Dmanisi,Republic of Georgia: Taxonomy, Geological Setting, and Age. Science 228, 1019-1025 (2000).
2. Ferring, R. et al., Earliest human occupations at Dmanisi (Georgian Caucasus) dated to 1.85–1.78 Ma. PNAS 108 (26), 10432-10436 (2011).
3. Vekua, A. et al., A New Skull of Early Homo from Dmanisi, Georgia. Science 297, 85-89 (2002).
4. Lordkipanidze, D. et al., Postcranial evidence from early Homo from Dmanisi, Georgia. Nature 449, 305-310 (2007).
5. Lordkipanidze, D. et al., A Fourth Hominin Skull From Dmanisi, Georgia. The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology 288A, 1146–1157 (2006).
6. Lordkipanidze, D. et al., A Complete Skull from Dmanisi, Georgia, and the Evolutionary Biology of Early Homo. Science 342, 326-331 (2013).
7. Lordkipanidze, D. et al., The earliest toothless hominin skull. Nature 434, 717-718 (2005).
8. Lieberman, D., Homing in on early Homo. Nature 449, 291-292 (2007).
9. Gabunia, L., de Lumley, M.-A., Vekua, A., Lordkipanidze, D. & de Lumley, H., Découverte d’un nouvel hominidé à Dmanissi (Transcaucasie, Géorgie). C.R. Palévol. 1, 243–253 (2002).
10. Bermúdez de Castro, J., Martinón-Torres, M., Sier, M. & Martín-Francés, L., On the Variability of the Dmanisi Mandibles. PLoS One 9 (2), e88212 (2014).
2.0 to 2.6-million-year-old tooth is from australopithecine or early human.
A reassessment of a fossil tooth from an old archaeological collection suggests that early hominins had extended their range to the western branch of Africa’s Great Rift Valley by no later than two million years ago. Since the late 1950s, large numbers of early hominin fossils have been found in the eastern branch of the Great Rift Valley, which is often described as the Cradle of Humanity. However, up until now, none have been found in the western branch.
Ishango 11 is an archaeological site in the Democratic Republic of Congo; it is located alongside the Semliki River, in the western branch of the Great Rift Valley. In the 1950s, the site was excavated by the Belgian geologist Jean de Heinzelin, who recovered numerous fossil human and animal remains, together with stone and bone artefacts. The assemblage dates mainly to the early part of the African Late Stone Age, from 25,000 to 19,000 years ago. It is housed in the Department of Anthropology and Prehistory at the Royal Belgian Institute of Science, Brussels.
However, the finds also included an upper left first molar that did not appear to be from such a recent period. Known as #Ish25, doubts were cast on its affinities to modern humans as long ago as 1958. A recent study has shown that #Ish25 probably originated from an earlier geological layer than the other fossils and artefacts. Animal remains associated with this layer suggest that it dates to between 2.6 and 2.0 million years ago. These dates make #Ish25 the earliest fossil hominin find from the western branch of the Great Rift Valley (though not the earliest from Central Africa, as much earlier hominins are known from Chad).
Various statistical analyses of the shape and size of #Ish25 suggest closer affinities to hominins from the Late Pliocene/Early Pleistocene than those from the Middle Pleistocene to Recent epochs. The exact hominin species to which the tooth belongs cannot be determined with certainty; Australopithecus africanus, Paranthropus robustus and early Homo are all possibilities.
The western Great Rift Valley underwent episodes of climate change 3.0, 2.6 and 1.8 million years ago; these led to the partial replacement of Congo flora and fauna with those typical of the East Africa; the latter are adapted to more open grassland conditions. The #Ish25 findings suggest that these conditions led to a dispersal of hominins into the region from either East Africa or South Africa.
The study also demonstrates how valuable knowledge can often be gained by applying modern techniques to old anthropological collections.
1. Crevecoeur, I. et al., First Early Hominin from Central Africa (Ishango, Democratic Republic of Congo). PLoS One 9 (1), e84652 (2014).
Earliest unambiguous evidence for meat-eating by early hominins.
Modern humans are the only existent primates anatomically adapted for the regular consumption of significant quantities of meat. The human gut is reduced compared with that of other primates, a configuration more suited to a meat-eating diet than the predominantly vegetarian diet of other primates. Although crucial to many models of hominin evolution, however, the timing of and circumstances in which early hominins began to include significant quantities of meat in their diet remain poorly understood.
The earliest-known stone tools, from Gona, Ethiopia, are 2.6 million years old and are often taken to be early evidence for meat eating (Semaw, et al., 1997; Semaw, 2000). No hominin remains were recovered in association with the tools, but in 1999, anthropologists working at the nearby Bouri Formation reported the discovery of large mammal bones bearing cut-marks apparently made by stone tools, possibly as a result of dismembering and filleting carcasses. Animals appeared to have been defleshed, and their long bones broken open, presumably to extract marrow. The bones were found in association with 2.5 million-year-old australopithecine remains, thought to be of Australopithecus garhi (de Heinzelin, et al., 1999).
It has also been claimed that 3.39 million-year-old animal bones from Dikika, Ethiopia, show stone tool cut-marks for flesh removal, and signs of having been struck with hammerstones to extract bone marrow (McPherron, et al., 2010). In the absence of any associated tools, there is no way to tell whether the cut-marks were produced with specially-made tools or naturally-sharp pieces of stone. Some are sceptical and argue that as the bones were buried in coarse-grained, sandy deposits, it is likely that trampling by animals produced the marks (Domınguez-Rodrigo, et al., 2011).
Even if the above is accepted as evidence of carcass-processing by early hominins, it is too insubstantial to show whether these were one-off forays into meat-eating or part of a more substantial shift in hominin dietary adaptations. To demonstrate ‘persistent carnivory’ requires a geologically-stratified series of relatively large assemblages of animal remains, each showing extensive signs of persistent hominin activity. The sum of the assemblages must demonstrate that this activity persisted over the course of at least a thousand years (Ferraro, et al., 2013).
Although rather more recent than the above dates, such evidence has now been reported from Kanjera South, a small site located on the shores of Lake Victoria, southwestern Kenya (Ferraro, et al., 2013). Three excavations along 50 metres have yielded several thousand well-preserved animal remains, approximately 2 million years old, and associated with stone tools. There is a consistent record of hominin activities throughout the stratified sequence, which spans hundreds or possibly thousands of years.
The animal remains included gazelle and other small bovids, together with a smaller number of medium-sized bovids. The remains showed clear evidence of butchery by hominins in the form of cut-marks and damage caused by hammerstones. Patterns of tooth-marks made by carnivores such as lions and hyenas suggest that these animals only had access to the carcasses after the hominins had removed the bulk of the meat and bone marrow. Carnivores typically chew on the mid-shafts of long bones, but the percentage of bones that were so marked was low.
Small bovids are invariably wholly consumed by carnivores within hours of death, implying that the hominins acquired and butchered them very soon after death. A possible implication is that these animals were hunted rather than scavenged, and that Kanjera represents the earliest archaeological record of hunting activities by hominins.
The skeletal remains of the small bovids suggest that they were transported to the site for butchery more or less intact. However, in the case of the medium-sized bovids, head and limb parts predominate. These animals were too large to transport intact, so the hominins removed the limb parts, leaving the rest of the body behind. Although head contents are nutritious, they are difficult to exploit and would thus be ignored by other scavengers. They therefore represent a niche that tool-using hominins could exploit. It is therefore likely that hominins scavenged leftover head parts from carnivore kills and transported them to the site for processing.
The Kanjera data not only provides the required evidence of hominin meat-eating over a period of many centuries: it also provides clues about specific activities. Thus, it seems, the hominins obtained much of their meat by hunting small bovids, but they also scavenged medium-sized bovid heads as a separate by complimentary activity. The date of 2 million years ago is somewhere between 200,000 and 500,000 years earlier than the previous earliest evidence for persistent hominin carnivory.
1. Semaw, S. et al., 2.5-million-year-old stone tools from Gona, Ethiopia. Nature 385, 333-336 (1997).
2. Semaw, S., The World’s Oldest Stone Artefacts from Gona, Ethiopia: Their Implications for Understanding Stone Technology and Patterns of Human Evolution Between 2.6–1.5 Million Years Ago. Journal of Archaeological Science 27, 1197–1214 (2000).
3. de Heinzelin, J. et al., Environment and Behavior of 2.5-Million-Year-Old Bouri Hominids. Science 284, 625-629 (1999).
4. McPherron, S. et al., Evidence for stone-tool-assisted consumption of animal tissues before 3.39 million years ago at Dikika, Ethiopia. Nature 466, 857-860 (2010).
5. Domınguez-Rodrigo, M., Pickering, T. & Bunn, H., Reply to McPherron et al.: Doubting Dikika is about data, not paradigms. PNAS 108 (21), E117 (2011).
6. Ferraro, J. et al., Earliest Archaeological Evidence of Persistent Hominin Carnivory. PLoS One 8 (4) (2013).