Denisovan(?) remains in the garden

On the edge of the small town of Lingjing near Xuchang City in Henan Province, China, local people have long practiced intensive vegetable gardening because the local soil is naturally irrigated by the water table beneath the flood plain deposits of the Yinghe River. In the mid 1960s, around a small spring, they began to find dozens of small stone tools together with animal bones. Only in 2005, after the spring had stopped flowing, did systematic excavation begin (Li, Z.-Y. et al. 2017. Late Pleistocene archaic human crania from Xuchang, China. Science, v. 355, p. 969-972; doi: 10.1126/science.aal2482) About 3.5 m below the surface tools and bone fragments, including one with a carved representation of a bird, occurred just above the base of the modern soil profile. Radiocarbon dating of charcoal from the layer clustered around 13 500 years ago, just before the start of the Younger Dryas cooling episode; probably products of modern humans, although no human remains were found in the layer. Continued excavation penetrated sediments free of fossils and tools down to a depth of 8 m, when stone tools and bone fragments began to turn up again through the lowest 2 m of sediment. Optically stimulated luminescence (OSL) dating of mineral grains, which shows the last time that sediments were exposed to sunlight, produced much older dates between 78 to 123 ka. The thousands of stone flakes and cores, and cut marks on the animal bones found through the fossil-rich layer suggests that this was a site long used for tool making and food preparation, that had begun in the last interglacial period. Among the bones were fragments of the crania of as many as five individual humans.

Who were they? Their age range is tens of thousands of years before anatomically modern humans began to migrate into east Asia, so they are likely to have been an earlier human group. Homo erectus is known to have inhabited China since as early as 1.6 Ma ago and may be a possibility. The other possible group are the Denisovans, known only from their DNA in a small finger bone from a cave in eastern Siberia. Fragments of Denisovan DNA are famously present in that of many living indigenous people from eastern Asia, Melanesia and the Americas, but hardly at all in west Asians and Europeans. They also interbred with Neanderthals and may share a common ancestor with us and them, who lived about 700 ka ago.

Map showing the proportion of the genome inferred to be Denisovan in ancestry in diverse non-Africans. The color scale is not linear to allow saturation of the high Denisova proportions in Oceania (bright red) and better visualization of the peak of Denisova proportion in South Asia. (Credit: Sankararaman et al./Current Biology 2016;  http://dx.doi.org/10.1016/j.cub.2016.03.037)
Map showing the proportion of the genome inferred to be Denisovan in ancestry in non-Africans. The color scale ranges from black – 0, through greens – present to red – highest . (Credit: Sankararaman et al./Current Biology 2016; http://dx.doi.org/10.1016/j.cub.2016.03.037)

Unfortunately the human bones are completely fragmented and lack any teeth, jaw bones or elements of the face. However, the Chinese-US team used sophisticated computer refitting of CT-scanned fragments to reconstruct two of the crania, revealing one individual with prominent brow ridges and a flat-topped skull extended towards the back, similar to that of Neanderthals but with a much larger brain than H. erectus. The semi-circular canals associated with the ears, but used in balancing, are well preserved and also resemble those of Neanderthals. Yet east Asia has yielded not a single Neanderthal fossil. Could these be the elusive Denisovans? Even if more diagnostic bones turn up, especially teeth, such is the state of late hominin taxonomy that only DNA will provide definitive results: the Denisovans are defined entirely by DNA. The authors, perhaps wisely, do not speculate, but others may not be able to resist the temptation.

For more information on recent human evolution see here.

Gibbons, A. 2017. Close relative of Neandertals unearthed in China. Science, v. 355, p. 899; doi: 10.1126/science.355.6328.899

Human penis bone lost through monogamy?

The baculum or penis bone is arguably the most variable of mammalian bones, present in some species but not others. Among those in which it does occur the baculum varies enormously in shape, length and breadth relative to body size. This makes it likely to have been subject to the most divergent evolution among mammals. Yet its evolution has remained somewhat puzzling until recently. Observation has shown that the width of the baculum in male house mice is positively correlated with reproductive success. So one factor in the bone’s evolution may be postcopulatory sexual selection: female mice seem to favour males well endowed in this department once they have mated with them, a notion supported by careful laboratory experimentation. The physical role of the penis bone is to support and protect the penis during sexual intercourse. Sturdy dimensions are increasingly efficacious the longer the duration and the greater the frequency of copulation, particularly among polygamous and seasonally breeding species. They also tend to delay or inhibit a female mating with another male after copulation.

Walrus baculum, approximately 22 inches long
Walrus baculum, approximately 0.6 metres long (credit: Wikipedia)

Matilda Brindle and Christopher Opie of University College London have applied advanced phylogenetic statistical analysis to data on the dimensions of penis bones among 2000 mammal species (Brindle, M. & Opie, C. 2016. Postcopulatory sexual selection influences baculum evolution in primates and carnivores. Proceedings of the Royal Society, B, v. 283, doi: 10.1098/rspb.2016.1736) and suggest that the baculum first evolved in mammals between 145 to 95 Ma ago, earlier mammals likely having no penis bone. Ancestral primates and carnivorous mammals, however, were so endowed. Yet some mammalian species have lost the baculum.  Among the primates human males do not have one whereas male chimpanzees and bonobos, with which we share a last common ancestor, do: both are boisterously promiscuous whereas humans are pair-bonded to a large degree.

The issue of polygamy versus monogamy among human ancestors, and when the latter emerged, continues to exercise palaeoanthropologists. The former in other living primates is often associated with a marked contrast in size between males and females – sexual dimorphism. The earliest hominins, such as species of Australopithecus, did exhibit such dimorphism whereas species of Homo show significantly less size contrast, which some have taken to mark the emergence of pair-bonding amongst members of the earliest human species to be passed on to their successors. Another indicator of competitiveness among primate males for females, and their dominance over the latter, is the near universal possession of large canine teeth among males of polygamous primates; an odd feature for species whose diet is dominantly and often exclusively vegetarian. Not only do living humans not have prominent canines, neither do any known fossil hominins. Despite the views of a small minority of anthropologists who demand that modern human females won social parity with males only in the last 100 thousand years, only to lose it following the Neolithic ‘revolution’, the physical evidence suggests that a trend towards that emerged with other distinct characteristics of hominins and concretised in early Homo. An assiduous search for fossil hominin penis bones may yet reveal the moment of monogamy.

Tree-climbing australopithecines

We know that Lucy, the famous Australopithecus afarensis, could climb trees because her many bone fractures show that she fell out of a tree to her death. But that does not mean her species was an habitual tree-climber: plenty of modern humans fall to their deaths from trees, cliffs and the like. But the issue seems to have been resolved by using X-ray tomography of Lucy’s limb bones (Ruff, C.B. et al. 2016. Limb bone structural proportions and locomotor behaviour in A.L. 288-1 (“Lucy”).  PLOS ONE v. 11, e0166095. doi:10.1371/journal.pone.0166095) during the skeleton’s triumphal series of exhibits in the US.


Embed from Getty Images

The authors, including two of those who showed that Lucy died after a fall using similar data, compared the digital 3-D models of her surviving arm- and leg bones with those of other hominins and living primates, estimating their relative strengths at different positions. Lucy was probably stronger in the arm than in the leg, but not to the same degree as chimpanzees. This is a feature that would significantlyassist climbing , but her bipedal locomotion on the ground would have been only slightly different from that of later Homo species. If anything, her strength relative to size would have been greater than ours, perhaps reflecting less reliance on tools for getting food and defending herself. But almost certainly Australopithecus afarensis habitually spend more time in trees, perhaps foraging and as a defence against predation, especially at night.

The new data for Lucy allows palaeoanthropologists to better judge the capabilities of other hominins. Interestingly Homo habilis, the earliest of our genus, may have had similar habits. But later species, beginning with H. erectus/ergaster, were as Earth-bound as we are. This suggests a shift in hominin ecology from an early and probably long history of semi-arboreal behavior until humans became masters of their terrain about 1.9 Ma ago, probably through their invention of better tools and the controlled use of fire.

Read more about human evolution here and here

A challenge to the concept of species

Comparison of DNA from ancient hominin fossils with that obtained from a broad spectrum of living people showed that on the road out of Africa in the last 130 thousand years some anatomically modern humans successfully interbred with Neanderthals and Denisovans to produce fertile offspring. All non-African people contain a trace of those liaisons and include fertile hybrids in their ancestry, whereas Africans do not. Using quick, low-cost and sensitive genomic analyses these discoveries made similar searches for hybridisation among other supposedly distinct species a popular and fruitful line of research (Pennisi, E. 2016. Shaking up the tree of life. Science, v. 354, p. 817-820; doi: 10.1126/science.354.6314.817). They also challenge the long-held view that individual species are incapable of fertile interbreeding with others. Yet fertile hybrids have long been known among plants and butterflies without recourse to genomics. Now that it is a basic tool, it has been shown that up to 10% of known plant species have arisen from hybrids and examples are quickly being found among birds, insects, fish and mammals, including the famous Galapagos finches. Hybridisation introduces genetic variation more quickly than does mutation, potentially a major advantage in adaptive radiation.

Page from Darwin's notebooks around July 1837 ...
Page from Darwin’s notebooks around July 1837 showing his first sketch of an evolutionary tree. (credit: Wikipedia)

Of course, the concept of ‘species’ is arbitrarily based on biological ‘form and function’, and in the same fashion as discoveries about epigenetics have shown genetic determinism to have an air of dogma, so hybridisation suggests that a ‘web’ is more apt as shorthand for the progress of evolution than is Darwin’s ‘tree’ or even a tangled ‘bush’. Another welcome outcome spurred by the pioneers of hominin comparative genetics is a powerful challenge to the dominant philosophies of reductionism and dualism among scientists; legacies of René Descartes bound up with the ‘scientific method’ – especially among physical scientists – and ideas such as ‘nature versus nurture’. A major revolution is in progress, from which the seekers for a Theory of Everything, from quantum mechanists through particle physicists to cosmologists need to draw some sharp and perhaps embarrassing lessons.

It is appropriate that the driving agency lies within anthropology, and thrilling too, for everyone can quickly learn a new way of approaching the world by contemplating their own origins. They would be hard-pressed to do that by pondering on the early nanoseconds of the cosmos …

Neanderthal culture confirmed

The Châtelperronian material culture represents the earliest sign of the Upper Palaeolithic in Europe and its products span a period from about 45 to 40 ka. It includes stone tools, such as points and long, thin blades with a single cutting edge and a blunt back, reminiscent of a modern knife, and others with notched, or denticulate edges that resemble saw blades. A great many of the tools, including ivory and bone ones, are probably designed for working and stitching skins. But the most revealing worked objects are animal teeth, shells and fossils that are either bored or grooved to be strung together. The best have been found in the Grotte du Renne in eastern France. The most controversial aspect of the Châtelperronian is that its artefacts are sometimes found with the fossil remains of Neanderthals who had previously produced less sophisticated, Mousterian tools since around 160 ka. The controversy centres on whether or not Neanderthals created the Châtelperronian culture, and if so, did they develop them independently or through cultural exchange with or copying from the newly arrived anatomically modern humans (AMH).

Science Magazine
Châtelperronian ornaments from the Grotte du Renne eastern France, probably parts of a necklace. (Credit: ©Marian Vanhaeren, CNRS, University of Bordeaux)

The Grotte du Renne material is especially rich in ornaments, but insufficient fossil material is present to tell from anatomical characteristics whether or not they were made by AMH or Neanderthals. It has now become possible using traces of bone proteins to detect hominin bone fragments and DNA to assess which group is implicated (Welker, F. and 127 others, 2016. Palaeoproteomic evidence identifies archaic hominins associated with the Châtelperronian at the Grotte du Renne. Proceedings of the National Academy of Science, www.pnas.org/cgi/doi/10.1073/pnas.1605834113). Analyses of mtDNA and radiometric dating of the bones that yielded it show that the Grotte du Renne tools and ornaments link with Neanderthals who lived there about 37 ka ago. Interestingly, the stratigraphic horizon beneath the definite Neanderthal occupation level contains their earlier, Mousterian artefacts. So it seems that they developed new manufacturing techniques and material culture. Yet, the findings do not resolve the issue of independent invention or copying AMH methodology.

Importantly, Grotte du Renne shows that Neanderthals, even if they copied AMH techniques, were capable of appreciating, producing and using personal ornamentation: they could learn and transmit ideas. In that respect, here is support for the notion that, apart from significant anatomical differences from AMH they were not that different intellectually.

More on Neanderthals, Denisovans and anatomically modern humans

Wade, L. 2016. Neandertals made jewelry, proteins confirm. Science, v. 353, p. 1350.

Out of Africa: a little less blurred?

DNA from the mitochondria of humans who live on all the habitable continents shows such a small variability that all of us must have had a common maternal ancestor, and she lived in Africa about 160 ka ago. Since this was first suggested by Rebecca Cann, Mark Stoneking and Allan Wilson of the University of California, Berkeley in 1987 there has been a stream of data and publications – subsequently using Y-chromosome DNA and even whole genomes – that both confirm an African origin for Homo sapiens and illuminate it. Analyses of the small differences in global human genetics also chart the routes and – using a ‘molecular clock’ technique – the timings of geographic and population branchings during migration out of Africa. As more and better quality data emerges so the patterns change and become more intricate: an illustration of the view that ‘the past is always a work in progress’. The journal Nature published four papers online in the week ending 25 September 2016 that demonstrate the ‘state of the art’.

Three of these papers add almost 800 new, high-quality genomes to the 1000 Genomes Project that saw completion in 2015. The new data cover 270 populations from around the world including those of regions that have previously been understudied for a variety of reasons: Africa, Australia and Papua-New Guinea. All three genomic contributions are critically summarized by a Nature News and Views article (Tucci, S & Akey, J.L. 2016. A map of human wanderlust. http://dx.doi.org/10.1038/nature19472). The fourth paper pieces together accurately dated fossil and archaeological findings with data on climate and sea-level changes derived mainly from isotopic analyses of marine sediments and samples from polar ice sheets (Timmermann, A & Friedrich, T. 2016. Late Pleistocene climate drivers of early human migration. Nature, doi:10.1038/nature19365). Axel Timmermann and Tobias Friedrich of the University of Hawaii have attempted to simulate the overall dispersal of humans during the last 125 ka according to how they adapted to environmental conditions; mainly the changing vegetation cover as aridity varied geographically, together with the opening of potential routes out of Africa via the Straits of Bab el Mandab and through what is now termed the Middle East or Levant. They present their results as a remarkable series of global maps that suggest both the geographic spread of human migrants and how population density may have changed geographically through the last glacial cycle. Added to this are maps of the times of arrival of human populations across the world, according to a variety of migration scenarios. Note: the figure below estimates when AMH may have arrived in different areas and the population densities that environmental conditions at different times could have supported had they done so. Europe is shown as being possibly settled at around 70-75 ka, and perhaps having moderately high densities for AMH populations. Yet no physical evidence of European AMH is known before about 40 ka. Anatomically modern humans could have been in Europe before that time but failed to diffuse towards it, or were either repelled by or assimilated completely into its earlier Neanderthal population: perhaps the most controversial aspect of the paper.

timmermann
Estimated arrival time since the last continuous settlement of anatomically modern human migrants from Africa (top); estimated population densities around 60 thousand years ago. (Credit: Axel Timmermann University of Hawaii)

The role of climate change and even major volcanic activity – the 74 ka explosion of Toba in Indonesia – in both allowing or forcing an exodus from African homelands and channelling the human ‘line of march’ across Eurasia has been speculated on repeatedly. Now Timmermann and Friedrich have added a sophisticated case for episodic waves of migration across Arabia and the Levant at 106-94, 89-73, 59-47 and 45-29 ka. These implicate the role of Milankovich’s 21 ka cycle of Earth’s axial precession in opening windows of opportunity for both the exodus and movement through Eurasia; effectively like opening and closing valves for the flow of human movement. The paper is critically summarised by a Nature News and Views article (de Menocal, P.B. & Stringer, C. 2016. Climate and peopling of the world. Nature, doi:10.1038/nature19471.

This multiple-dispersal model for the spread of anatomically modern humans (AMH) finds some support from one of the genome papers (Pangani, L. and 98 others 2016. Genomic analyses inform on migration events during the peopling of Eurasia. Nature (online). http://dx.doi.org/10.1038/nature19792). A genetic signature in present-day Papuans suggests that at least 2% of their genome originates from an early and largely extinct expansion of AMH from Africa about 120 ka ago, compared with a split of all mainland Eurasians from African at around 75 ka. It appears from Pangani and co-workers’ analyses that later dispersals out of Africa contributed only a small amount of ancestry to Papuan individuals. The other two genome analyses (Mallick, S. and 79 others 2016. The Simons Genome Diversity Project: 300 genomes from 142 diverse populations. Nature (online) http://dx.doi.org/10.1038/nature18964; Malaspinas, A.-S. and 74 others 2016. A genomic history of Aboriginal Australia. Nature (online). http://dx.doi.org/10.1038/nature18299) suggest a slightly different scenario, that all present-day non-Africans branched from a single ancestral population. In the case of Malaspinas et al. an immediate separation of two waves of AMH migrants led to settlement of Australasia in one case and to the rest of Mainland Eurasia. Yet their data suggest that Australasians diverged into Papuan and Australian population between 25-40 ka ago. Now that is a surprise, because during the lead-up to the last glacial maximum at around 20 ka, sea level dropped to levels that unified the exposed surfaces of Papua and Australia, making it possible to walk from one to the other. These authors appeal to a vast hypersaline lake in the emergent plains, which may have deterred crossing the land bridge. Mallick et al. see an early separation between migrants from Africa who separately populated the west and east of Eurasia, with possible separation of Papuans and Australians from the second group.  These authors also show that the rate at which Eurasians accumulated mutations was about 5% faster than happened among Africans. Interestingly, Mallick et al. addressed the vexed issue of the origin of the spurt in cultural, particularly artistic, creativity after 50 ka that characterizes Eurasian archaeology. Although their results do not rule out genetic changes outside Africa linked to cultural change, they commented as follows:

‘… however, genetics is not a creative force, and instead responds to selection pressures imposed by novel environmental conditions or lifestyles. Thus, our results provide evidence against a model in which one or a few mutations were responsible for the rapid developments in human behaviour in the last 50,000 years. Instead, changes in lifestyles due to cultural innovation or exposure to new environments are likely to have been driving forces behind the rapid transformations in human behaviour …’.

Variations in interpretation among the four papers undoubtedly stem from the very different analytical approaches to climate and genomic data sets, and variations within the individual sets of DNA samples. So it will probably be some time before theoretical studies of the drivers of migration and work on global human genomics and cultural development find themselves unified. And we await with interest the pooling of results from all the different genetics labs and agreement on a common data-mining approach.

Lucy: the australopithecine who fell to Earth?

The specimen of Australopithecus afarensis known far beyond the confines of palaeoanthropology as Lucy remains the iconic figure of hominin evolution, 42 years after her discovery by Donald Johanson and Tom Gray near Hadar in the Awash valley of the Afar Depression, Ethiopia. Her skeletal remains were not complete, but sufficient to recognize that they were from the oldest known upright hominin and that they were female, the pelvis having affinities to that of human women rather than other extant apes. Yet her skull was more akin to apes with a brain volume about the same as a modern chimpanzee’s. Part of the reason for her fame stems from being named after a character in a somewhat mystical song by the British pop group, the Beatles, which was played over and over in the palaeontologist’s camp – good job the find wasn’t during the 1990’s acme of gangsta rapper Apache.

Subsequently, the entombing strata were radiometrically dated at around 3.2 Ma. Lucy, in common with most fossils roughly in the human line of descent, has from the outset been the subject of controversy, even at one time being said to be misnamed because of alleged male characteristics; a view swiftly discarded. Like the treasures of Tutankhamun, Lucy’s actual remains have been exhibited far and wide, including a 6-year stay in the US. Fears of damage in transit led the Ethiopian authorities to produce casts for distribution, and Lucy is now restricted to the National Museum in Addis Ababa. As a further precaution, all the actual bones were rendered in digital 3-dimensions using a high-resolution CT scanner during her US sojourn. It is these scans that have led to a surprising development as regards her original fate. Apart from signs of a single carnivore tooth mark, her remains were not devoured by scavengers, nor did early anatomical examinations suggest any sign of disease and she was estimated to have been a young mature female when she died – the cause of death was unknown.

Model of the australopithecus Lucy in the muse...
Model of the Lucy (Australopithecus afarensis) in the museum of Barcelona (credit: Wikipedia)

Detailed forensic analysis of the CT scans (Kappelman, J. and 8 others 2016. Perimortem fractures in Lucy suggest mortality from fall out of tall tree. Nature, v. 537, published online 29 August 2016, doi:10.1038/nature19332) revealed far more than did the original bones, including evidence for numerous fractures in Lucy’s limbs, ribs and cranium, many of which are of the compressive or ‘greenstick’ kind. Those in the left ankle and leg bones (talus, tibia, fibula and femur) are compressive and suggest a severe vertical impact of the heel with enough force to smash the strongest bones in the body, driving the hip into the pelvis. Damage to the ribs, pelvis and lower spine (sacrum) is commensurate with a further horizontal, frontal impact of the torso. Arm (humerus), wrist (radius)  shoulder blade (scapula) and collar (clavical) bone fractures are typical of injuries sustained when a falling person tries to break a fall by stretching out the arms. Damage to the cranium and lower jaw (mandible) suggest this instinctive defence posture was futile. None of the fractures show signs of healing, so the multiple traumas were immediately fatal.

Forensic reconstruction of how Lucy fell to meet her end. (credit: John Kappelman et al, doi: 10.1038/nature19332)
Forensic reconstruction of how Lucy fell to meet her end. (credit: John Kappelman et al, doi: 10.1038/nature19332)

The traumatic pattern is reminiscent of someone falling onto hard ground from great height; perhaps equivalent to a four- or five-storey building (see animated reconstruction here). In Lucy’s case, the most likely scenario is from a large tree, perhaps while foraging or sleeping in a safe refuge from ground predators. Forensic analysis of newly dead victims of severe falls generally show massive soft tissue damage by penetration of bone fragments or a ‘hydraulic ram effect’ in which abdominal organs are thrust upwards to produce cardiac damage. That Lucy was found almost intact rules out dismemberment by scavengers or transport by flood water. Indeed, the preservation of even tiny slivers of fractured bone seems to suggest her burial in flood plain sediments before decomposition had set in. A question that the authors do not address is whether or not she may have been deliberately interred, which to me seems a possibility that could be drawn from the detailed evidence. I wonder what a modern coroner might conclude: probably misadventure.

More on hominin evolution can be found here.

Climatic conditions for early hominin evolution

Until about 1.8 Ma ago, when early Homo erectus and perhaps other archaic hominins strode into Eurasia, our forerunners lived and evolved on only one continent – Africa. The physical and environmental conditions underlying the steps from a common ancestor with modern chimpanzees through a growing number of upright species are not well charted by the Pliocene and early Pleistocene terrestrial evidence. All that is know of this formative period is that global climate cooled in an oscillating fashion that culminated in the onset of Northern Hemisphere glaciations in the late Pliocene (~3 Ma) and a shift to drier conditions in East Africa around 2.8 Ma suggested by pollen records off the east coast. Marine sediments of the Indian Ocean, Red Sea and Gulf of Aden still offer the most convenient means of charting environmental change in detail for this crucial episode in human history. As well as oxygen-isotope and pollen-type variations, modern core analysis offers a growing number of wind-blown proxies for onshore vegetation. These include organic geochemistry plus carbon and hydrogen isotopes from trace amounts of leaf waxes. During the May to September East African Monsoon, high speed winds in the upper atmosphere drag dusty continental air from the East African Rift System over the Gulf of Aden, making sea-floor sediment an important target for tracking variations in the proxies (Liddy, H.M. et al. 2016.  Cooling and drying in northeast Africa across the Pliocene.  Earth and Planetary Science Letters, v. 449, p. 430-438. doi:10.1016/j.epsl.2016.05.005). Hannah Liddy and colleagues from the Universities of Southern California and Arizona, USA, applied these techniques to a Gulf of Aden core from offshore Somaliland to open a window on this crucial period.

Early history of hominin evolution and evidence for climate change in East Africa. Based on a diagram at the handprint.com website
Early history of hominin evolution and evidence for climate change in East Africa. Based on a diagram at the handprint.com website and in Stepping Stones Chapter 22

Early Pliocene East Africa (5.3 to 4.3 Ma), the time of Ardepithecus ramidus, was characterized by evidence for a climate wet enough to sustain grasses and riverine woodlands. Yet around 4.3 Ma conditions had shifted to ecosystems more dominated by shrubby plants able to thrive in more arid conditions. At about that time the earliest australopithecines appear in the fossil record, with A. anamensis. Yet the later Pliocene was not devoid of grasses or herbivores. There is ample carbon-isotope evidence from the teeth of hominins that shows that after 3.4 Ma the diet of A. afarensis and A. africanus included increasing amounts whose carbon derived from grasses, when. This apparent paradox can be explained by a major turn to eating meat from herbivores as vegetable foods declined with increasing aridity. This is all very interesting, especially the detailed record of δ13C in plant waxes, but there is little to indicate that steps in hominin speciation or extinction had much direct connection with fluctuations in climatic conditions. Environmental change may have formed a background to other influences that may have been wholly down to early hominin’s social and technological behaviour.

Hobbit time

A few months after the diminutive hominin fossil Homo floresiensis, which because of its relatively large feet was quickly dubbed the ‘Hobbit’, turned out to be considerably older than previously thought it hit has the headlines again because its ancestors may have colonized the Indonesian island of Flores far earlier still. A pair of articles in the 9 June 2016 issue of Nature consider evidence from another site on the island where fluvial sediments offer more easily interpreted stratigraphy than the complex Liang Bua cave assemblage where the original skeletal remains were unearthed. The site in the So’a Basin became an important target for excavation following the discovery there in the 1950’s of stone artefacts, east of Wallace’s Line – a fundamental faunal and floral divide once thought to be due to the difficulty of crossing a deep, current-plagued channel in the Indonesian archipelago. The unexpected presence of artefacts drew palaeoanthropologists from far afield, but it was almost 50 years later before their exploration yielded hominin remains.

English: homo from flores
Homo floresiensis (credit: Wikipedia)

One of the papers reports sparse new finds of hominin material from the So’a Basin, a fragment of mandible and 6 isolated teeth thought to be from at least three individuals (van den Bergh, G.D. et al., 2016. Homo floresiensis-like fossils from the early Middle Pleistocene of Flores. Nature, v.  534, p. 245-248). The other covers newly discovered artefacts, the stratigraphic and palaeoecological setting, and radiometric dates of the finds (Brumm, A. and 22 others, 2016. Age and context of the oldest known hominin fossils from Flores. Nature, v.  534, p. 249-253). The jaw fragment shows signs of having once held a wisdom tooth, showing that it belonged to an adult. Yet although it resembles the dentition of the younger Liang Bua specimens, it seems more primitive and is even smaller. The other dental finds are most likely to be deciduous teeth of juveniles. Fission-track, uranium-series and 40Ar/39Ar dating indicates that the fossils entered the sediments about 700 ka ago. But tools and remains of prey animals in deeper sedimentary layers here and at other Flores sites indicate the presence of hominins back as far as about 1 Ma, before which there are no such signs.

So, at least a million years ago Flores was colonised by hominins. Either the original immigrants were uniquely small compared with other hominins of that vintage in Asia and Africa, or within 300 ka they had decreased in size through the evolutionary influence of limited resources on Flores and the process of island dwarfism. The second may also have been influenced by an initially small population of migrants or a later population ‘bottleneck’ that added a loss of genetic variability – a founder effect.   These two alternatives may point respectively to either the even earlier migration out of Africa and across most of Asia of perhaps H. habilis, or the dwarfing of a limited population of H. erectus who made their way there from their known occupation of Java. The authors painstaking analysis of the meagre remains suggest a closer dental resemblance to Asian Homo erectus than to earlier African hominins, so the second alternative seems more likely. However, even that scenario poses palaeoanthropology with a major problem; yet another evolutionary process that helps cryptify the links among our earlier relatives. (See also: Gomez-Robles, A., 2016. The dawn of Homo floresiensis. Nature, v.  534, p. 188-189.)

Breaking news: Cave structures made by Neanderthals

Neanderthals were well equipped and undoubtedly wore clothing, made shelters, hunted, used fire and famously lived in caves. Deliberate burial of their dead, in some cases arguably with remains of flowers, indicates some form of ritual and belief system. Those in Spain wore necklaces and pendants of bivalve shells, some of which retain evidence of having been painted. Excavators there even found a paint container and painting tools made of small bones from a horse’s foot. The container and tools retain traces of the common iron colorants goethite, jarosite and hematite. One large, perforated scallop shell, perhaps used as a pectoral pendant, shows that its white interior was painted to match its reddish exterior. Given the evidence for adornment by earlier hominins, to find that Neanderthals created art should not be surprising. In May 2016 it emerged that about 177 thousand years ago and earlier, they had broken stalagmites off the cave roof to create curious semi-circular structures in Bruniquel Cave near Montauban in southern France (Jaubert, J. and 19 others, 2016. Early Neanderthal constructions deep in Bruniquel Cave in southwestern France. Nature, v. 533,  online publication, doi:10.1038/nature18291). Each of the structures contains incontrovertible evidence that fires were made within them. Rather than being near the well-lit cave entrance the structures are more than 300 m deep within the cave system surrounded by spectacular stalagmites and stalactites that are still in place. Were the structures younger than 42 ka they would probably have been attributed to the earliest anatomically modern Europeans and to some ritual function. Instead they were made during the climatic decline to the last but one glacial maximum.

Related article

Neanderthals built mystery underground circles 175,000 years ago

 

Homo floresiensis, aka the ‘Hobbit’, is somewhat older

In 2004 a newly discovered hominin fossil from the Indonesian island of Flores made headlines worldwide. Although an adult, it was tiny – about a metre tall, had a commensurately small brain (the size of a grapefruit), had made tools and hunted small elephants and giant rats. Dates from the cave floor sediments that had entombed it gave ages as young as 13 to 11 thousand years and as far back as 850 ka. So H. floresiensis was regarded as being the last human to share the Earth with us; that is, if it was a different species rather than a product of evolutionary shrinkage of anatomically modern humans stranded and isolated on the island for a very long time. Then there was talk among locals of the legendary Ebo Go-Go, with whom their ancestors had shared the island – they had arrived between 35 to 55 thousand years ago.

Homo floresiensis (the "Hobbit")
Homo floresiensis (the “Hobbit”) ( credit: Wikipedia)

Unsurprisingly, a major controversy raged in palaeoanthropology circles, between those who demanded either island dwarfism or congenital deformity of modern humans, and the other camp focused on many anatomical differences that pointed to a bona fide companion to later immigrants who perhaps survived into modern times. The ‘Hobbit’ became a cause celebre, but many of the original protagonists are now left with the proverbial egg on their faces. The cave sediments turn out to have a much more complex stratigraphy than previously thought, following further excavations led by the original discoverer Thomas Sutikna of the Pusat Penelitian Arkeologi Nasional in Jakarta Indonesia (Sutikna, T. and 19 others 2016. Revised stratigraphy and chronology for Homo floresiensis at Liang Bua in Indonesia. Nature, v. 532, p. 366-369.

English: Cave where the remainings of ' where ...
Liang Bua cave on Flores island, Indonesia, where the remains of Homo floresiensis were discovered in 2003. (credit: Wikipedia)

The delayed appearance of the revision is hardly surprising, given the lengthy political squabbles surrounding access to the site. And neither are the outcomes, for cave sediments are notoriously tricky because of their episodic reworking by cave floods and roof falls, together with the difficulty in finding materials suited to dating in tropical settings. The original charcoal used in radiocarbon dating and sand grains subject to the thermoluminescence method were in fact from a  unit that lies unconformably against the stratum that hosted the fossils. More sophisticated luminescence dating of the actual fossil-hosting sediments yield ages between 100 to 60 ka, tool-bearing units range from 190 to 50 ka. The origins of H. floresiensis are thus pushed back beyond the date of supposed colonisation by H. sapiens, and remain an open question.

Neanderthals and Denisovans at it more often

Palaeogeneticists certainly have the bit between their teeth as DNA sequencing methods become faster and more productive and statistical methods of sequence analysis and comparison are made more powerful. Only last month I reported on the two-way breeding unearthed from the data on single-chromosome DNA extracted from Croatian and Spanish Neanderthals, as well as some of the tangible inheritance from Neanderthals found in living non-African people. Now a team of statisticians, anthropologists and genetic sequencers have applied the new approaches to the genomes of over 1500 non-Africans, including 35 living Melanesian people from Papua-New Guinea (Vernot, B. and 16 others 2016.  Excavating Neandertal and Denisovan DNA from the genomes of Melanesian individuals. Science, v. 351 doi:10.1126/science.aad9416).  Melanesians had previously shown evidence of hybridization with both Neanderthals and Denisovans. The most interesting outcome is that the analyses pointed towards yet more instances of interbreeding between ancestors of modern non-Africans and Neanderthals. Many East Asians have 3 Neanderthals in their family trees, for Europeans and South Asians the score is 2, while Melanesians show descent from one Neanderthal and one Denisovan. Moreover, it emerges that interbreeding episodes were at different times among different populations since anatomically modern humans migrated from Africa, beginning perhaps as long ago as 130 ka and recurring later, after different regional groups of AMH had proceeded on their separate ways.

English: Melanesia, a cultural and geographica...
Melanesia, a cultural and geographical area in the Pacific. (credit: Wikipedia)

A second study (Sankararaman, S. et al. 2016. The combined landscape of Denisovan and Neanderthal ancestry in modern humans. Current Biology, v. 26, p. 1-7) has teased out evidence for Denisovan ancestry among South Asians, their admixture with Melanesians after that group acquired Neanderthal forebears, and significant signs  of dwindling fertility among hybrid males.

Early 2016 has been very fertile as regards palaeoanthropology. Katherine Zink and Daniel Lieberman of Harvard University focus on the small teeth of Homo erectus and later humans, wondering if they arose following a major shift in culinary practices (Zink, K.D. & Lieberman, D.E. 2016. Impact of meat and lower Palaeolithic food processing techniques on chewing in humans. Nature, v. 531, p. 500-503). Their work is based  on experiments to discover how much chewing is needed to make it possible to swallow different uncooked foodstuffs (assuming that cooking did not arise until after 500 ka). It seems that simply introducing meat to the diet would have reduced mastication by around 13% (2 million chews) per year, with a 15% reduction in applied chewing force. Simply slicing and pounding takes out another 750 thousand annual chews and gives a 12% fall in average biting force. So, here’s a link between tools and human gnashers as well as with development of the hand. Fascinating, perhaps, but every hominin species since 7 Ma old Sahelanthropus tchadensis had far smaller canine teeth than are the norm among non-hominin living and fossil apes. Something else was going on with dentition during our evolution, which may have been a loss of the need for threatening teeth. From ‘Do that again and I’ll bite you’, to ‘Let’s chew this over’…

More on Neanderthals, Denisovans and anatomically modern humans

Neanderthal news

Note: Earth-Pages will be closing as of early July, but will continue in another form at Earth-logs

Increasingly sophisticated analysis of existing genomes from Neanderthal and Denisovan fossil bone, together with new data on single-chromosome DNA extracted from Croatian and Spanish Neanderthals continues to break new ground.

Artistic reconstruction of Neanderthal woman (credit: Natural History Museum, http://www.nhm.ac.uk/natureplus/blogs/tags/human_evolution)
Artistic reconstruction of Neanderthal woman (credit: Natural History Museum, http://www.nhm.ac.uk/natureplus/blogs/tags/human_evolution)

According to genome comparison between a Siberian specimen and modern humans, a population from which Neanderthals emerged separated from that which led to anatomically modern humans (AMH) sometime between 550 and 765 ka, although the fossil record can only confirm that divergence was before 430 ka. The comparison famously showed that Neanderthals contributed to modern, non-African humans between 47 and 75 ka, that is after the exodus of AMH from Africa that spread our species throughout all continents except Antarctica. This genetic exchange is thought to have taken place somewhere in the Middle East, which seems to have been a major staging post for our spread further east and also westward to Europe. A similar indication of liaison between Denisovans and AMH migrants is restricted to modern Melanesians, and probably took place in eastern Asia before 45 ka, when modern people began crossing from Eurasia to New Guinea and Australia. Neanderthal-Denisovan comparison suggests that those distinct groups separated between 380 and 470 ka ago (recently revised from an earlier estimate).

In both cases the gene flow was from the older groups to humans. Further examination of Siberian Neanderthal genomes now indicates that a reverse exchange occurred more than 100 ka ago (Kuhlwilm, M. and 21 others 2016. Ancient gene flow from early modern humans into Eastern Neanderthals. Nature, v. 530, p. 429-433). But the single-chromosome DNA from Croatian and Spanish Neanderthals shows no such sign This instance of two-way exchange is significant in another way: it took place before direct evidence of the generally accepted departure of African migrants to populate the rest of the world. At about 100 ka there is fossil evidence of possible AMH-Neanderthal cohabitation of the Levant, followed by a period with fossil evidence for Neanderthal presence there but not modern humans. Because stone tools from northern Arabia are dated as far back as 125 ka and closely resemble those associated with archaic modern humans, there is a possibility that AMH migration was far earlier than previously thought and passed through the Levant en route to points east.

Another tantalizing aspect of Neanderthal-modern human genetics is the tangible legacy of interbreeding with non-African humans. The first sign was that the gene (mc1r) that confers red hair on those of us blessed, or otherwise, with it may have Neanderthal origins, thus making us extremely proud of that heritage. The same gene is implicated in northern modern humans having developed pale skin, which might embarrass ‘white supremacists’! Similar studies in Svante Paabo’s lab at the Max Planck Institute for Evolutionary Anthropology in Leipzig also suggested 15 genome regions that include those involved in energy metabolism, possibly associated with type 2 diabetes; cranial shape and cognitive abilities, perhaps linked to Down’s syndrome, autism and schizophrenia; wound healing; skin, sweat glands, hair follicles and skin pigmentation; and barrel chests. There is more…

Joshua Akey of the University of Washington, Seattle, and evolutionary genomicist Tony Capra of Vanderbilt University in Nashville hit on the idea of ‘mining’ archived genetic information from more than 28 thousand living people for traces of 6000 Neanderthal DNA variants and comparing the results with physical traits and diseases logged in the human database (reported by Gibbons, A. 2016. Neanderthal genes linked to modern diseases. Science, v. 351, p. 648-9). On the plus side, Neanderthal ancestry may help boost immune responses to fungi, parasites and bacteria. Inheritance of enhanced blood coagulation, although greatly assisting recovery from wounds and hemorrhage when giving birth, confers a proclivity to heart attacks and strokes. Neanderthals also passed on ‘weak bladders’, solar keratoses that confer skin cancer risk, a tendency to malnutrition from modern diets low on meat and nuts, depression triggered by jet lag(!) and even a tendency to nicotine addiction. But a ‘pure’ line of modern human descent, shared by most Africans, also has its positive and negative heritable traits.

More on Neanderthals, Denisovans and anatomically modern humans

A rational view of the start of human influences on Life and Geology

Regular readers will know that I have strong views on attempts to burden stratigraphy with a new Epoch: the Anthropocene. The central one is that the lead-in to a putsch has as much to do with the creation of a bandwagon, to whose wheels all future geologists will be shackled, as it does to any scientific need for such a novelty. Bound up as it is with the fear that Earth may be experiencing its sixth mass extinction, the mooted Anthropocene will likely become a mere boundary marked by future stratigraphers as a Global Boundary Stratotype Section and Point or GSSP between the existing Holocene Epoch and that sequence of sedimentary strata and their fossil record that will be laid down on top of it. Or not, if humanity becomes extinct should the economically induced, dangerous modifications of our homeworld of the last few decades or centuries not be halted. Either way, it defies the stratigraphic ‘rule book’.

No one can deny that humanity’s activities are now immensely disruptive to surface geological processes. Nor is it possible to rule out such disruptive change to the biosphere in the near-future that a latter-day equivalent of the K/Pg or end-Permian events is on the cards: such confidence does not spring from the interminable succession of grand words and global inaction reiterated in December 2015 by the UN Paris Agreement on economically-induced climate change. Still, it was a bit of a relief to find that palaeontological evidence, or rather statistics derived from the fossil record in North American sedimentary rocks since the Carboniferous, emphasises that there is no need for the adoption of Anthropocene as an acceptable geological adjective.

To ecologists, extinctions are not the be all and end all of disruption of the biosphere. Major shifts in life’s richness are also recorded by the way entire ecosystems become disrupted. A classic, if small-scale, example is that way in which the ecosystem of the US Yellowstone National Park changed since the eradication by 1926 of the few hundred grey wolves that formerly preyed mainly on elk. In the 20 years since wolf reintroduction to the Park in 1995 the hugely complex but fragile Yellowstone ecosystem has showed clear signs of recovery of its pre-extirpation structure and diversity.

A consortium of mainly US ecologists, led by Kathleen Lyons of the National Museum of Natural History at the Smithsonian Institution in Washington DC, has assessed linkages between species of fossil animal and plants since the Carboniferous (S.K. Lyons and 28 others, 2015. Holocene shifts in the assembly of plant and animal communities implicate human impacts. Nature, published on-line 16 December 2015 doi:10.1038/nature16447). They found that of the 350 thousand pairs of species that occurred together at different times throughout the late Palaeozoic to the last Epoch of the Cenozoic, the Holocene, some pairs appeared or clustered together more often than might be expected from random chance. Such non-random association suggests to ecologists that the two members of such a pair somehow shared ecological resources persistently, hinting at relationships that helped stabilise their shared ecosystem. For most of post-300 Ma time an average of 64% of non-random pairs prevailed, but after 11.7 ka ago – the start of the Holocene – that dropped to 37%, suggesting a general destabilisation of many of the ecosystems being considered. This closely correlates with the first human colonisation of the Americas, the last of the habitable continents to which humans migrated. This matches the empirical evidence of early Holocene extinctions of large mammals in the Americas, which itself is analogous to the decimation of large fauna in Australasia during the late Pleistocene following human arrival from about 50 to 60 ka ago. Significant human-induced ecological impact seems to have accompanied their initial appearance everywhere. The ecological effects of animal domestication and agriculture in Eurasia and the Americas mark the Holocene particularly. In fact, in Europe the presence of Mesolithic hunter gatherers is generally inferred, in the face of very rare finds of artefacts and dwellings, from changes in pollen records from Holocene lake and wetland sediments, which show periods of tree clearance that can not be accounted for by climate change.

There is no need for Anthropocene, other than as a political device.

Surprising modern-human migrations into China and Africa

Caves figure highly in discoveries of hominin remains, fossil riches from those near Johannesburg in South Africa and at Atapuerca in northern Spain having set the world of palaeoanthropology reeling in the last few months. As often as not the caves chosen by hominins for day-to-day living, refuge or ritual, places where carnivores dragged some of our early relatives, or into which they fell accidentally, formed in limestones. There are few places so well endowed with karst features than southern China, a fair number of caves in them having rich deposits of bat guano to which farmers have beaten well-trodden paths to dig it out for fertiliser. One such is Fuyan Cave in Daoxian County, Hunan. Manure mining there had done a great deal of the heavy work faced by archaeologists, having stopped when it reached a hard layer of calcite speleothem or flowstone that underpaves more or less the entire cave floor. Initial trial investigations found three clearly human teeth at the surface, encouraging further work. Digging through the flowstone revealed sediments rich in fossils, mainly teeth which preserve better than other remains in humid conditions. As well as teeth from a variety of mammals, large and small, 47 human teeth emerged. Close study revealed dental features that are irrefutably those of anatomically modern humans (Liu, W. and 13 others 2015. The earliest unequivocally modern humans in southern China. Nature, doi:10.1038/nature15696). Remarkably, many of the teeth are in far better condition than my own, and those of many living people with access to dental expertise.

Some of the Daoxian human teeth. (Credit: Song Xing and Xiu-jie Wu of the 1Key Laboratory of Vertebrate Evolution and Human Origins at the Chinese Academy of Sciences
Some of the Daoxian human teeth. (Credit: Song Xing and Xiu-jie Wu of the 1Key Laboratory of Vertebrate Evolution and Human Origins at the Chinese Academy of Sciences)

The true significance of the excavation emerged only when 230Th dating revealed the age of the flowstone cap to the old cave sediments. A small stalagmite protruding from its surface yielded a minimum age of ~80 ka: by far the oldest date for anatomically modern human remains outside of Africa and the eastern Mediterranean. The dating produced older ages around 120 ka with equally good precision. Before this discovery the date of migration of Africans to populate Eurasia was thought to be about 60 ka from imprecise dating based on genetics of a range of living Eurasians and Africans – a ‘molecular clock’ – and the earliest sign of humans found in Australia. Consequently, finds in South India of artefacts beneath 74 ka ash from the super-eruption of the Mount Toba caldera have been regarded by many, other than the finders, as having been made by Homo erectus. Dates of 100 ka for modern human occupation of the Levant were thought to represent a failed attempt at migration out of Africa by a northern route. Both these important findings now take on renewed significance. Yet a 30 to 40 ka time gap between the Fuyan people and the previous dates for the earliest signs of migration into China, Borneo and Australia (40-50 ka) begs the question, ‘Did this early group of far-travelled migrants survive to become ancestors of modern Chinese people?’ There are many possible scenarios that only future discoveries might validate: simply goiung extinct; failure to survive the encounter with earlier migrants, such as H. erectus or the Denisovans; assimilation into those older populations.

Mitochondrial DNA-based chart of large human m...
Mitochondrial DNA-based chart of large human migrations: the consensus before these new data. Numbers are millennia before present. ( credit: Wikipedia)

As if to counter this, a multinational group of collaborators have sequenced and analysed the genome from a 4500 year-old male skeleton discovered in the Mota Cave of the Gamo highlands of southern Ethiopia (Llorente, M.G. and 18 others 2015. Ancient Ethiopian genome reveals extensive Eurasian admixture throughout the African continent. Science, DOI: 10.1126/science.aad2879). Comparison with what is now a virtual library of living human genomes showed that this man’s genetic make-up most closely matched that of the Ari, a tribe living in the area today. What was most interesting is that part of the modern Ari genome – between 4 to 7% – is not present in the 4500 year-old sequence. Instead, it matches those of modern Sardinians and a prehistoric German farmer. Yet it occurs in people living not only in Ethiopia, but also in central, western and southern Africa to varying degrees. There seems to have been a ‘backflow’ of people into the whole of Africa from Eurasia, estimated to have occurred some 3500-4000 years ago and probably involving a large influx. By that time farming was already established in Africa, so the migrants may have had some advantage, either culturally or physically, to encourage their wide spread through the continent.

In tropical climates, DNA is likely to break down quickly and little if any fossil DNA has been recovered from prehistoric Africans. In this case, burial in a cave at high elevation may have helped preserve it, but also the target for extraction was the petrous bone from the inner ear whose density seems to allow DNA a better change of long-term survival. With continually improving DNA analysis and sequencing techniques more news is surely going to emerge from past African populations.

More on human migrations

Related articles

Gibbons, A. 2015. Prehistoric Eurasians streamed into Africa, genome shows. Science, v. 350 (9 October 2015), p. 149.

Our ancestors parted from other humans earlier than expected

Despite the excitement raised by the discovery of remnants of 15 individuals of Homo naledi in a South African Cave the richest trove of hominin fossils remains that of Sima de los Huesos (‘pit of bones’) in northern Spain. In 2013 bone found in that cave from one of 28 or more individuals of what previous had been regarded as H. heidelbergensis, dated at around 400 ka, yielded mitochondrial DNA. It turned out to have affinities with mtDNA of both Neanderthals and Denisovans, especially the second. The data served to further complicate the issue of our origins, but were insufficient to do more than throw some doubt on the significance of H. heidelbergensis as a distinct species: nuclear DNA would do better, it was hoped by the palaeo-geneticists of the Max Planck Institute for Evolutionary Anthropology in Leipzig. Now a small fragment of those data (about 1 tro 2 million base pairs) have been presented to a London meeting of the European Society for the Study of Human Evolution – though not yet in a peer-reviewed journal. Anne Gibbons summarised the formal presentation in the 18 September 2015 issue of Science (Gibbons, Ann 2015. Humanity’s long, lonely road. Science, v. 349, p. 1270).

English: Cranium 5 is one of the most importan...
One of the best preserved discoveries in the Sima de los Huesos, Atapuerca (Spain). (credit: Wikipedia)

The partial nuclear DNA is a great deal more like that of Neanderthals from much more recent times than it is of either Denisovans and modern humans. It seems most likely that the Sima de los Huesos individuals are early Neanderthals, which implies that the Neanderthal-Denisovan split was earlier than 400 ka. That might seem to be just fine, except for one thing: Neanderthal and Denisovan DNA are much more closely related to each other than to that of ourselves. That implies that the last common ancestor of the two archaic human species must have split from the ancestral line leading to modern humans even further back in time: maybe 550 to 765 ka ago and 100 to 400 ka earlier than previously surmised. This opens up several interesting possibilities for our long and separate development. Since Neanderthals and perhaps Denisovans emigrated from Africa to Eurasia several glacial cycles ago, maybe people genetically en route to anatomically modern humans did so too. The Neanderthal and Denisovan genomes suggest that they interbred with each other and that could have been at any time after the genetic split between them. Famously, they also interbred with direct ancestors of living Eurasians, but there is no genetic sign of that among living Africans. The evidence suggests that the insertion of archaic genetic material was into new migrants from Africa around 100 to 60 ka ago at different points along their routes to Europe and East Asia. But, obviously, it is by no means clear cut what passed between all three long-lived groups nor when. It is now just as possible that surviving, earlier Eurasians on the road to modern humans passed on their own inheritance from relationships with Neanderthal and Denisovan to newcomers from Africa. But none of these three genetic groups ever made their way back to Africa, until historic times.

More on Neanderthals, Denisovans and anatomically modern humans

The ‘star’ hominin of South Africa

The week of 7 to 11 September 2015 was one of the most news-rich of the year. To name but two issues: the plight of tens of thousands of refugees fleeing Africa and the Middle East to Europe was made worse by total confusion, little action and downright obstruction by some of the most privileged governments on Earth ; in Britain one of the most exciting political dramas in decades – the leadership elections of the Labour Party – were reaching a climax of press and political skulduggery because of the unexpected direction both had taken. Something else burst onto the media scene that was, if anything, even more out-of-the-blue to the majority of people on Thursday 10 September: the remains of at least 15 individuals of a new hominin species found in a near-inaccessible cave were announced by a multinational team of geologists and anthropologists. The feature that ensured its wide publicity in competition with some pretty serious political and humanitarian developments was the suggestion that the corpses had been ritually laid to rest by beings that lived maybe 2 million years ago. This major scientific stir arose from the publication of two lengthy papers by the open-access, electronic journal eLife (Berger, L. R. and 46 others 2015. Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa. eLife DOI: 10.7554/eLife.09560. Dirks, P.H.G.M. and 23 others 2015. Geological and taphonomic context for the new hominin species Homo naledi from the Dinaledi Chamber, South Africa. eLife, DOI: 10.7554/eLife.09560).

Artist’s reconstruction of the face of Homo naledi (credit: John Gurche artist, Mark Thiessen photographer, National Geographic)

Homo naledi (naledi means ‘star’ in the Sotho language: the find was in the Rising Star cave system near Johannesburg) is known in more anatomical detail than any early hominin, and most closely resembles H. habilis and H. rudolphensis discovered 3 to 4 thousand miles away in Tanzania and Kenya. The Dinaledi deposit remains undated but likely to come out at around 2 Ma or older. The sheer wealth of anatomical detail, including complete foot- and hand-bone remains from individuals, evidence for a range of ages at death, and plenty of dental and cranial information, actually poses a taxonomic problem of comparison with remains of other early hominins. Most of them are fragmentary, and it seems likely that once a precise date is obtained H. naledi will assume greater importance in comparative anatomy. Comparison with australopithecines is easier because of their abundant remains, and H. naledi is clearly distinct from that clade as regards gait, chewing, overall physiognomy (see reconstruction video) and cranial dimensions, but does have some australopithecine affinities. They were certainly different from their near geographic neighbour Au. sediba, also found in a cave deposit within the great swath of Palaeoproterozoic limestones near Johannesburg, where the Cradle of Humankind UNESCO World Heritage Site is situated. The brain of Homo naledi was on a par with those of australopithecines as regards volume, yet larger than that of H. floresiensis: it does seem that brain size is not necessarily related to the uses to which it is put.

The route into the Dinaledi Chamber where bones of at least 15 individual members of Homo naledi were found (credit: National Geographic magazine http://news.nationalgeographic.com/2015/09/150910-human-evolution-change/)

Interestingly, it is reported that only the most diminutive members of the research team were able to enter the chamber where the remains were found because of the narrowness of the connecting passage. Also, access from the main cave system involved an upward ‘U-bend’, so that although water could – and did from time to time – enter the chamber in the past, it is unlikely that coarse material such as large bones could simply have been washed in, the more so as the chamber is on a minor spur from the main system and its outlet is through small floor drains that could not sustain torrential flow. Nor is there any direct access from the ground surface to this part of the system. Some of the more fragile body parts, such as a hand, are still articulated, which suggests a non-violent movement to the chamber. There are no signs of physical trauma to any of the bones, ruling out action by carnivores or transport by violent floods, nor any indicative of de-fleshing as by cannibalism. However, before fossilisation, many of the bones had been gnawed by beetles and snails. This combination of features leads to the possibility that corpses may have been deliberately placed in the chamber. If they had been, then to get to deepest recess of the cave system and find the Denalidi Chamber required illumination: fire brands.  That the chamber was actually a living space is highly unlikely because of its remoteness from the surface. One big question that cannot be answered is whether or not such possible disposal was by ritual or simply for sanitary arrangements. Another possibility, not considered by the authors is seeking refuge from predators and becoming trapped in the desperately constricted space.

The possibility of ritual burial is clearly what has seized headlines. Yet few palaeoanthropologists will accept that: only Neanderthals and anatomically modern humans are definitely considered to have adopted such a practice, in the last hundred thousand years. The association of a bifacial stone tool with 350 ka old H. heidelbergensis remains at Atapuerca in northern Spain has been suggested to be the earliest evidence for ritual burial, but is not widely accepted. There are no reports of artefacts in the Dinaledi Chamber.

Stone tools go even further back

Shortly after it seemed that the maker of the earliest stone tools (2.6 Ma) may have been Australopithecus africanus, thanks to a novel means of analyzing what hominin hands may have been capable of, some actual tools have turned up from even earlier times (Harmand, S. and 20 others 2015. 3.3-million-year-old stone tools from Lomekwi 3, West Turkana, Kenya. Nature, v. 521, p. 310-315). Their age is comparable with that (3.4 Ma) of animal bones from Dikika, Ethiopia showing cut marks and signs of deliberate breaking, which had previously been controversial as they suggested that local Australopithecus afarensis of a similar age had made them. What the authors claim to be ‘a new beginning to the known archaeological record’ almost a million years earlier than the first appearance of Homo fossils in the Lake Turkana area seems to point in that direction. But A. afarensis has not been found in that area, although a hominin known as Kenyanthropus platyops with roughly the same age as the tools has.

Australopithecus afarensis reconstruction
Reconstruction of Australopithecus afarensis (Photo credit: Wikipedia)

Almost 150 fine-grained basaltic artefacts turned up at the Lomekwi site, which may have been where knappers habitually worked as many of them were fragments or debitage. The cores from which flakes had been struck are large, weighing on average 3.1 kg. It seems that the tool makers may have been forcefully pounding out edged tools for a variety of uses, unlike the single-use hammer stones used by chimpanzees today. Compared with the well known Oldowan tools, however, these are cruder and made by a different knapping technique that seems not to have focused on exploiting the conchoidal fracturing that produces the sharpest tools and is a feature of the later Oldowan tools.

English: Chopper: one of the earliest examples...
Oldowan ‘chopper’ from Melka Kunture, Ethiopia. (credit: Wikipedia)

Frederick Engels, whose 1876 essay The Part played by Labour in the Transition from Ape to Man was among the first works to take Darwin’s ideas on human origins forward, would have had a field day with the new evidence. For him the vital step was freeing of the hands by a habitual bipedal gait and their manipulation of objects – together with changes to the hands that would arise by such a habit. What the first tool maker looked like, doesn’t really matter: the potential that act conferred was paramount. Nevertheless, there is a big step between early hominins and humans, from relatively small brains to those of H. erectus that were on the way to modern human capacity. The Lomekwi tools and the improved Oldwan artefacts spanned 1.7 Ma at least before H. erectus revolutionised manufacture to produce the bi-facial Acheulian hand ‘axe’, and going beyond that took almost a million years of little change in both tools and anatomy until the emergence of archaic modern humans.

Note added 28 May 2015: Within a week palaeoanthropologists’ focus shifted to the Afar Depression in Ethiopia where a new species of hominin has emerged from Pliocene sediments dated to between 3.3 and 3.5 Ma (Haile-Selassie, Y et al. 2015. New species from Ethiopia expands Middle Pliocene hominin diversity. Nature, v. 521, p. 483-488. doi:1038/nature14448). Australopithecus deyiremeda is represented by fragments of two lower- and one upper jaw plus several other lower facial specimens. So the species is differentiated from other hominins by dentition alone, but that is unmistakably distinct from extensive data on Au. afarensis which lived within a few kilometres over the same period. Until the last 15 to 20 years it was thought that Au. afarensis was the sole hominin around in the Middle Pliocene of East and Central Africa, but now it seems there may have been as many as five, the three mentioned above, plus Au. bahrelghazali from Chad and an as yet undesignated fossilised foot from Afar. For possibly three closely related species to coexist in Afar is difficult to understand: possibly they occupied different niches in the local food web or employed different strategies (Spoor, F. 2015. The middle Pliocene gets crowded. Nature, v. 521, p. 432-433). Another question is: did they all make and use tools? For the Lomekwi tools K. platyops is a candidate, but for the cut marks on bones at Dikika in Afar there are at least two: Au. afarensis and Au. deyiremeda. So multiple tool makers living at the same time suggests some earlier originator of the ‘tradition’.

Note added 4 June 2015: Add southern Africa into the equation and there is yet more breaking news about coeval hominin diversity. US, Canadian, South African and French collaborators have finally started to resolve the achingly complex stratigraphy of the fossil-rich Sterkfontein cave deposits in South Africa by using a novel approach to estimating ages of materials’ last exposure to cosmic rays (Granger, D.E. et al. 2015. New Cosmogenic burial ages for Sterkfontein member 2 Australopithecus and Member 5 Oldowan. Nature, v. 522, p. 85-88). Specifically, they managed to date the tumbling into a deep sinkhole of a recently found, almost complete skeleton of an australopithecine. It still resembles no other some 70 years after a less complete specimen was found by Raymond Dart in the mid 1940s. It was first informally dubbed ‘Little Foot’ and then Au. prometheus and up to now has been regarded as an odd contemporary of 2.2 Ma old Au. africanus. The new dating gives an age of about 3.7 Ma: so at least 6 hominids occupied Africa in the Middle Pliocene. It is beginning to look like a previously unsuspected time of sudden diversification.

Genus Homo pushed back nearly half a million years

Bill Deller, a friend whose Sunday is partly spent reading the Observer and Sunday Times from cover to cover, alerted me to a lengthy article by Britain’s doyen of paleoanthropologists Chris Stringer of the Natural History Museum. (Stringer, C. 2015. First human? The jawbone that makes us question where we’re from. Observer, 8 March 2015, p. 36). His piece sprang from two Reports published online in Science that describe about 1/3 of a hominin lower jaw unearthed – where else? – in the Afar Depression of Ethiopia. The discovery site of Ledi-Geraru is a mere 30 km from the most hominin-productive ground in Africa: Hadar and Dikika for Australopithecus afarensis (‘Lucy’ at 3.2 Ma and ‘Selam’ at 3.3 Ma, respectively); Gona for the earliest-known stone tools (2.6 Ma); and the previously earliest member of the genus Homo, also close to Hadar.

On some small objects mighty tales are hung, and the Ledi-Geraru jawbone and 6 teeth is one of them. It has features intermediate between Australopithecus and Homo, but more important is its age: Pliocene, around 2.8 to 2.75 Ma (Villmoare, B. And 8 others. Early Homo at 2.8 Ma from Ledi Geraru, Afar, Ethiopia. Science Express doi: 10.1126/science.aaa1343). The sediments from which Ethiopian geologist Chalachew Seyoum, studying at Arizona State University, extracted the jawbone formed in a river floodplain. Other fossils suggest open grassland rich with game, similar to that of the Serengeti in Tanzania, with tree-lined river courses. These were laid down at a time of climatic transition from humid to more arid conditions, that several authors have suggested to have provided the environmental stresses that drove evolutionary change, including that of hominins (DiMaggio, E.N. and 10 others 2015. Late Pliocene fossiliferous sedimentary record and the environmental context of early Homo from Afar, Ethiopia. Science Express doi: 10.1126/science.aaa1415).

Designating the jawbone as evidence for the earliest known member of our genus rests almost entirely on the teeth, and so is at best tentative awaiting further fossil material. The greatest complicating factor is that the earliest supposed fossils of Homo (i.e. H. habilis, H rudolfensis and others yet to be assigned a species identity) are a morphologically more mixed bunch than those younger than 2 Ma, such as H. ergaster and H. erectus. Indeed, every one of them has some significant peculiarity. That diversity even extends to the earliest humans to have left Africa, found in 1.8 Ma old sediments at Dmanisi in Georgia (Homo georgicus), where each of the 5 well-preserved skulls is unique.  The Dmanisi hominins have been likened to the type specimen of H. habilis, but such is the diversity of both that is probably a shot in the dark.

English: Cast replica of OH 7, the type specim...
Replica of OH 7, the deformed type specimen of Homo habilis. (credit: Wikipedia)

Coinciding with the new Ethiopian hominin papers a study was published in Nature the same week that describes how the type specimen of H. habilis (found, in close association with crude stone tools and cut bones, by Mary and Lewis Leakey at Olduvai Gorge, Tanzania in 1960) has been digitally restored from its somewhat deformed state when found (Spoor, F. et al. 2015. Reconstructed Homo habilis type OH 7 suggests deep-rooted species diversity in early Homo. Nature, v. 519, p. 83-86, doi:10.1038/nature14224). The restored lower jaw and teeth, and part of its cranium, deepened the mysterious diversity of the group of fossils for which it is the type specimen, but boosts its standing as regards probable brain size from one within the range of australopithecines to significantly larger –~750 ml compared with <600 ml – about half that of modern humans. The habilis diversity is largely to do with jaws and teeth: it is the estimated brain size as well as the type specimen’s association with tools and their use that elevates them all to human status. Yet, the reconstruction is said by some to raise the issue of a mosaic of early human species. The alternative is an unusual degree of shape diversity (polymorphism) among a single emerging species, which is not much favoured these days. An issue to consider is: what constitutes a species? For living organisms morphological similarity has to be set against the ability for fertile interbreeding. Small, geographically isolated populations of a single species often diverge markedly in terms of what they look like yet continue to be interfertile, the opposite being convergence in form by organisms that are completely unrelated.

Palaeontologists tend to go largely with division on grounds of form, so that when a specimen falls outside some agreed morphological statistics, it crosses a species boundary. Set against that the incontrovertible evidence that at least 3 recent human species interbred successfully to leave the mark in all non-African living humans. What if the first humans emerging from, probably, a well-defined population of australopithecines continued to interbreed with them, right up to the point when they became extinct about 2 Ma ago?

On a more concrete note, the Ledi Geraru hominin is a good candidate for the maker of the first stone tools found ‘just down the road’ at Gona!

Wet spells in Arabia and human migration

In September 2014, Earth Pages  reported how remote sensing had revealed clear signs of extensive fossil drainage systems and lakes at the heart of the Arabian Peninsula, now the hyper-arid Empty Quarter (Rub al Khali). Their association with human stone artifacts dated as far back as 211 ka, those with affinities to collections from East Africa clustering between 74-90 ka, supported the sub-continent possibly having been an early staging post for fully modern human migrants from Africa. Member of the same archaeological team based at Oxford University have now published late Pleistocene palaeoclimatic records from alluvial-fan sediments in the eastern United Arab Emirates that add detail to this hypothesis (Parton, A. ­et al. 2015. Alluvial fan records from southeast Arabia reveal multiple windows for human dispersal. Geology, advance online publication doi:10.1130/G36401.1).

The eastern part of the Empty Quarter is a vast bajada formed from coalesced alluvial fans deposited by floods rising in the Oman Mountains and flowing westwards to disappear in the great sand sea of dunes. Nowadays floods during the Arabian Sea monsoons are few and far between, and restricted to the west-facing mountain front. Yet, older alluvial fans extend far out into the Empty Quarter, some being worked for aggregate used in the frantic building boom in the UAE. In one of the quarries, about 100 km south of the Jebel Faya Upper Palaeolithic tool site , the alluvial deposit contains clear signs of cyclical deposition in the form of 13 repeated gradations from coarse to fine waterlain sediment, each capped by fossil soils and dune sands. The soils contain plant remains that suggest they formed when the area was colonized by extensive grasslands formed under humid conditions.

Dating the sequence reveals that 6 of the cycles formed over a 10 thousand-year period between 158 to 147 ka, which coincides with a peak in monsoon intensity roughly between 160 and 150 ka during the glacial period that preceded the last one. Three later cycles formed at times of monsoon maxima during the last interglacial and in the climatic decline leading to the last glacial maximum, at ~128 to 115 ka, 105 to 95 ka, 85 to 74 ka. So, contrary to the long-held notion that the Arabian Peninsula formed a hostile barrier to migration, from time to time it was a well watered area that probably had abundant game. Between times, though, it was a vast, inhospitably dry place.

English: SeaWiFS collected this view of the Ar...
Satellite view of the Arabian Peninsula. The Oman mountains sweep in a dark arc south eastwards from the Staits of Hormuz at the mouth of the Persian Gulf. The brownish grey area to the south of the arc is the bajada that borders the bright orange Empty Quarter (credit: NOAA)

The authors suggest that the climatic cyclicity was dominated by a 23 ka period. As regards the southern potential migration route out of Africa, via the Straits of Bab el Mandab, which has been highly favoured by palaeoanthropologists lately, opportunities for migration in the absence of boats would have depended on sea-level lows. They do not necessarily coincide with wet windows of opportunity for crossing the cyclically arid Arabian peninsula that would allow both survival and proceeding onwards to south and east Asia. So far as I can judge, the newly published work seems to favour a northward then eastward means of migration, independent of fluctuations in land-ice volume and sea level, whenever the driest areas received sufficient water to support vegetation and game. In fact most of NE Africa is subject to the Arabian Sea monsoons, and when they were at their least productive crossing much of Ethiopia’s Afar depression and the coastal areas of Eritrea, Sudan and Egypt would have been almost as difficult as the challenge of the Empty Quarter.

Human-Neanderthal cohabitation of the Levant

The earliest known remains of anatomically modern humans outside of Africa were found unearthed from the Skhul and Qafzeh caves in what is now northern Israel. Their context was that of deliberate burial at a time when climate was cooling from the last interglacial, between 90 to 120 ka. The Levant was also the repository for a number of well-preserved Neanderthal skeletons, most dating to between 35-65 ka, including ten individuals at Shanidar in today’s northern Iraq, some of whom were also deliberately buried including one whose grave reputedly contained evidence for a floral tribute. The 25 ka gap between the two populations has previous been regarded as evidence for lack of contact between them. However, the Tabun Cave in modern Israel has yielded tools attributed to Neanderthal Mousterian culture that may indicate their intermittent presence from 200 to 45 ka, and fossils of two individuals dated at ~122 and ~90 ka. The remains at Skhul and Qafzeh are significantly more rugged or robust than African contemporaries and have been considered possible candidates for Neanderthal-modern human hybrids. But whatever their parentage, it seems they became extinct as the climate of the Levant dried to desert conditions around 80 ka.

View of the exterior of Shanidar Cave, taken d...
Entrance to the Shanidar Cave, northern Iraq, occupied by Neanderthals between 35-65 ka (credit: Wikipedia)

A more promising overlap between modern human and Neanderthal occupation comes with the discovery by a group of Israeli, US, Canadian, German and Austrian scientists of a much younger anatomically modern human cranium from the Manot Cave, also in northern Israel (Herschkovitz, I. and 23 others 2015. Levantine cranium from Manot Cave (Israel) foreshadows the first European modern humans. Nature (online) doi:10.1038/nature14134). The cranium has a U-Th radiometric age of ~55 ka, well within the time span of Neanderthal occupation. Moreover, Manot Cave is one of a cluster of occupied sites in northern Israel, with separations of only a few tens of kilometres: undoubtedly, this individual and companions more than likely met Neanderthals. The big question, of course, is did the neighbours interbreed? If so the Levant would be the confirmed as the probable source of hybridisation to which the DNA of non-African living humans points. There may be a insuperable difficulty in taking this further: it is thought that the high temperatures of the region, despite its dryness, may have destroyed any chance of reconstructing ancient genomes. Yet one of the first Neanderthal bones to yield useful genetic material was from Croatia, which is not a great deal cooler in summer.

Convincing, indirect evidence for early toolmakers

A surprising number of animals pick up items from their surroundings and use them, mainly to get at otherwise inaccessible foodstuffs. What sets humans apart from such tool users is that we make them and for a long time part of our repertoire has been tools used to make other tools; so-called ‘machine tools’. An example is a piece of antler used to pressure-flake flint to give a stone blade a better edge, a more recent one is the increasing use of robots on assembly lines. Making a tool is impossible for a bird with only its beak and ill-adapted feet, while even a chimpanzee lacks various forms of grip needed for precisely directed force and manipulation. It was Frederick Engels who first focussed on the importance of the hand being freed to evolve the capacity for manual labour by the permanent adoption of an upright posture and gait, in his essay The Part Played by Labour in the Transition from Ape to Man written in 1876.

The earliest tools known turned up in 2.6 Ma old sediments at Gona in NE Ethiopia, while evidence for tool use is well accepted from cracked and sliced bones found in sediments dated at 2.5 Ma from Bouri in the same region. In neither case can the finds be tied to fossil remains of the makers and users, the earliest direct link emerging from famous Olduvai Gorge in western Tanzania, where crude Oldowan tools and worked bones occur with incomplete remains of a hominin, dubbed Homo habilis (‘handy man’) because of this association. Somewhat more controversial are bones that show cuts and scrape marks plus signs of having been cracked open that were found in a 3.4 Ma context at Dikika, also in Ethiopia, within the same sedimentary horizon as the young Australopithecus afarensis known as Selam (‘Hello’). The Dikika material is little different from 0.9 to 1.2 Ma younger bones at Bouri and Olduvai: the controversy seems to stem more from its much greater age and association with hominins deemed by some to have been incapable of creating tools.

English: Main division on the (right) human hand.
Bone structure of the (right) human hand. (credit: Wikipedia)

An entirely novel approach to the issue of the first tools and their makers, which with little doubt would have tickled Engels no end, is a careful anatomical and physiological examination of fossil hominin hand bones in comparison with those of chimps and living humans (Skinner, M.M. et al. Human-like hand use in Australopithecus africanus. Science, v. 347, p. 395-399). The bones being scrutinized are the five metacarpals that form the links in the palms from muscles of the forearm to finger and thumb movements and thus to various kinds of grip. In humans there are a host of ways of gripping objects from the precision of opposed thumb and finger pinching, especially that using the forefinger, to the squeezing power grip that wraps thumb and all fingers around an object and makes a fist. The best a chimp can do is grabbing a branch, to which its knuckle-walking hands are well adapted. The tips of the metacarpals are mechanically loaded according to the types of grip used repeatedly in life and that works to modify the physical density of the tips’ spongy bone tissue in patterns that vary according to habitual usage of the hand and its digits. This new approach is reputedly far more diagnostic than the actual shape of metacarpal bones, and requires high-resolution CT scanning.

Known early human and Neanderthal tool-makers show very similar patterns: in fact they suggest far more heavy loading through various kinds of grip than the metacarpals of humans from the modern period. In 1.8 to 3.0 Ma old A. africanus and Paranthropus robustus (a gorilla-like but bipedal australopithecine) from South Africa metacarpals suggest that both were habitually using a tree-climbing grip, much as chimpanzees do, but more closely resembled modern human and Neanderthal committed tool users. Both were certainly capable of using forceful precision grips to make and use tools up to 0.5 Ma earlier than the date of the earliest known tools. So far the technique has not been applied to the palm bones of earlier hominins such as A. afarensis (2.9-3.9 Ma) and Orrorin tugenensis (~6 Ma). Despite the suggestion of tool-making capability­, agreeing that it did take place in non-Homo hominins must await finds of tools, as well as signs of their use, in close association with fossil remains of their makers. The Dikika association is simply not enough. Yet, some bipedal being must have made tools before the date of the earliest ones (~2.6 Ma) discovered at Gona. Look at it this way: it is a lucky archaeologist who discovers every piece of evidence for a fundamental social change at one site. The fact that, by definition, the vast bulk of Pliocene and Pleistocene sediments that may contain the key evidence is either buried by younger material or was a victim of erosion, means that the chance of resolving the origin of the fundamental feature of human behaviour is tiny. The chance that scientists will continue looking is astronomically higher.

Art from half a million years ago

original fossils of Pithecanthropus erectus (n...
Original fossils of Pithecanthropus erectus (now Homo erectus) found By Eugene Dubois in Java in 1891 (credit: Wikipedia)

Eugene Dubois, an anatomist at the University of Amsterdam in the late 19th century, became enthralled by an idea that humans had evolved in what is now Indonesia, contrary to Charles Darwin’s suggestion of an African origin. So much so that Dubois took the extraordinary step of joining the Dutch army and scrounging a posting to the Dutch East Indies to facilitate his search for a ‘missing link’, accompanied by his wife and newborn daughter. After a four-year quest, in 1891 he discovered the upper cranium and brow of a being that was obviously related to us, but also quite distinct as regards its beetling brow ridges. Pithecanthropus erectus (now Homo erectus) raised a storm of controversy, sadly only resolved in Dubois’s favour after his death in 1940. Yet, as well as mounting the first deliberate search for human ancestors, Dubois collected everything possible in the sediments at Trinil, Java, so in a sense he was also an early palaeoecologist. The collection gathered dust in Leiden for the best part of a century, until archaeologist Josephine Joordens of the University of Leiden took on the task of reviewing its contents in 2007 (Joordens, J.C.A. and 20 others 2014. Homo erectus at Trinil on Java used shells for tool production and engraving. Nature (on-line): doi:10.1038/nature13962).

Progressively enlarged views of freshwater clam from Eugene Dubois's collecti9on from Trimil, showing clear evidence of deliberate engraving. (credit: Joordens et al., 2014 in Nature; photos by Wim Lustenhouwer, VU University Amsterdam
Progressively enlarged views of freshwater clam from Eugene Dubois’s collection from Trinil, showing clear evidence of deliberate engraving. (credit: Joordens et al., 2014 in Nature; photos by Wim Lustenhouwer, VU University Amsterdam)

Homo erectus clearly had a taste for freshwater clams and lots of their shells figure in the Trinil collection: all are of similar large size rather than showing a wide variation according to age, suggesting a shell midden rather than a natural assemblage. A piece of serendipity revealed what may prove to be the anthropological find of the year. High-quality photos of the shells taken by a visiting mollusc specialist showed up evidence that one of them had been meticulously engraved. Its surface had a near-perfectly geometric, zig-zag pattern deeply gouged by someone with a steady hand, who probably used an associated shark’s tooth as a scribing tool. Since the molluscs in life bear a dark, chitinous veneer the etching would have been more striking when freshly made. Another of these sturdy shells also show signs of having had its edge sharpened, suggesting that they were used for tools such as scrapers or graters.

The stratigraphy at Trinil suggested that the engraved shell and tools were coeval with Homo erectus, but that needed proof. Using sediment grains trapped in the shells and a combination of 40Ar/39Ar and thermoluminescence dating, the team have shown that they and the human fossils from Trinil date to between 430 and 540 thousand years ago: at least 350 ka older than the very similar engravings made by an anatomically modern human on ochre that was found at Blombos Cave in South Africa. The next-oldest putative artwork is the controversial ‘Venus’ found at Berekhat Ram on the Israel-Syria border, dated between 250 and 280 ka.

Engraved ochre from Blombos Cave, South Africa. (credit Chris Henshilwood)
Engraved ochre from Blombos Cave, South Africa. (credit: Chris Henshilwood)

Probably the majority of palaeoanthropologists have dismissed humans other than ­H. sapiens as being cognitively incapable of either abstract or figurative art. The general view is that the mental capacity to create art or design began with the creation of the Blombos engraving, was restricted to anatomically modern humans and only exploded in Europe after they had migrated there by about 40 ka. A few argue that portable art, such as the Trinil and Blombos engravings, is bound by its very nature to be rare and easily overlooked. Whether having some use – counting? – merely being the making of an idle ‘doodle’ or expressing some unknowable ritual significance, the Trinil etching is a result of creativity and controlled skill that could only be the product of the H. erectus mind. Moreover, the very close comparison with the 0.35 Ma younger Blombos engraving suggests the product of a consciousness little different from that of our direct ancestors of 75 ka ago.

Are modern humans ‘domesticated’?

While animals, especially dogs, underwent domestication the deliberate or unconscious human choice of favoured physiological and behavioural traits produced distinct differences between the ancestral species and the ‘breeds’ with which we are now familiar. In general domestication has resulted in dogs with reduced jaws and flatter faces, lower aggression, especially in the case of males, and reduced stressfulness in the company of humans and other tame dogs compared with their wolf ancestors. It is widely accepted that cats have ‘tamed themselves’ through the adoption of lifestyles associated with the benefits of close association with human communities, which have resulted in similar adaptations to those in more deliberately domesticated dogs. It is beginning to dawn on anthropologists that human social evolution may unwittingly have affected the course of our own evolution. Tighter social bonding among growing sizes of communities as brain capacity increased and the behavioural and cognitive attributes needed for that have been summarised recently by a group associated with the Social Brain hypothesis of Robin Dunbar of Oxford University, UK (Gamble, C., Gowlett, J. & Dunbar, R.I.M. 2014. Thinking Big: How the Evolution of Social Life Shaped the Human Mind. ISBN-13: 978-0500051801;Thames and Hudson: London).

It was Charles Darwin who first speculated that ‘Man in many respects may be compared with those animals which have long been domesticated’. But to what extent does the hominin fossil record support such a view? Collaborators from Duke University and the University of Iowa, USA, have set out to analyse physiological changes over the last 200 ka that may be explained in this way (Cieri, R.L. et al. 2014. Craniofacial feminization,social tolerance and the origins of behavioural modernity. Current Anthropology, v. 55, p. 419-443. Includes discussion and responses). They used the degree of projection of brow ridges, facial shape and cranial volume from 3 groups of Homo sapiens remains: skulls older than 80 ka (13 specimens); spanning 38 to 80 ka (41) and from recent humans (1367). They found that brow ridges shrank significantly over the last 80 thousand years, faces shortened and cranial capacity decreased, especially among males. This resulted in a convergence in appearance between males and females, which the authors attributed to general lowering of testosterone and stress hormone levels through selection for greater social tolerance: akin to similar physiognomic changes in domesticated dogs which DNA analyses have shown to be been linked with modification of genes associated with aggression regulation. The finding among dogs suggests that their domestication is accomplished by slower development; i.e. young animals are naturally less fearful and have a greater tendency to taming. This delayed development from foetus to adulthood, with retention in mature individuals of juvenile characteristics, is known as neoteny, and affects all manner of adult characteristics, including coloration, snout length and the adrenal glands: as adult dogs now more resemble wolf pups, so human adults are more like young chimps than elders. At a conference where Cieri et al.’s results were presented, it was observed that hunter gatherer bands are intolerant, to the point of capital punishment, of wife stealers, murderers and seriously dishonest men, whereas such reactions fall off significantly among members of larger social groups involved in agriculture and urban life. Such modern behavioural patterns tally with brow ridge, face length and cranial capacity, perhaps linked with selection for personalities more attuned to cooperation.

English: comparison of Neanderthal and Modern ...
Comparison of Neanderthal and Modern human skulls from the Cleveland Museum of Natural History (credit: Wikipedia)

Although earlier human species, such as H. neanderthalensis, heidelbergensis and erectus had significantly different skull anatomy, each had prominent brow ridges that, on this account, may signify both greater exposure to testosterone and less social tolerance, and smaller group sizes. But, so far, analysis of the Neanderthal genome has not led to publication of any comments about testosterone or stress-hormone related genes. However, a clear strand of discussion is developing around evidence rather than mere speculation about psychological/cognitive aspects of human evolution that challenges the old-style ‘what-you-see-is-what-there-was’ (WYSWTW) archaeological dogma: a dialectic of social and biological relationships.

Human evolution news

Since discovery of its fossilised remains in Liang Bua cave on the Indonesian island of Flores was discovered in 2004 the diminutive Homo floresienesis, dubbed the ‘hobbit’ by the media, has remained a popular news item each time controversies surrounding it have flared. To mark the tenth anniversary  of its publication of a paper describing the remains Nature has summarised the recollections of many of those involved in trying to understand the significance of H. floresiensis (Callaway, E. 2014. Tales of the hobbit. Nature, v. 514, p. 422-426). Two main schools of thought continue in dispute, one holding that it is anatomically so different from anatomically modern humans and earlier members of the genus Homo that it constitutes a new species, despite its youngest member dating back only 18 ka, the other that it is H. sapiens, its tiny size having resulted from some kind of genetic disorder, such as microcephaly or Down’s syndrome. There have been so many attempts to expunge the idea of such an odd fossil cohabiting an island with fully modern humans yet being a different and perhaps extremely archaic species that such an outlook itself seems somewhat pathological.

English: Homo floresiensis, replica Deutsch: H...
Replica of the Homo floresiensis skull from Liang Bua cave, Flores, Indonesia (credit: Wikipedia)

The evidence presented to force H. floresiensis into a deformed human mould has never been convincing, and the best way of combating that view is to document from a ‘non-combatant ‘standpoint the many ways in which its anatomy differs from ours and how it might have arisen; a job to which Chris Stringer of the Museum of Natural History in London is amply qualified (Stringer, S. 2014. Small remains still pose big problems. Nature, v. 514, p. 427-429). He, like the original discoverers, feels this is a case of evolution of small stature due to a limited population being isolated for a long time on a relatively small island, which is just what happened to elephants that colonised Flores to become the pigmy Stegodon that H. floresiensis seemingly hunted. These tiny Flores dwellers (adults were about 1 m tall) used fire and made tools, similar ones dating as far back as ~1 Ma. Stringer mentions the possibility of first human colonisation about that time by Asian H. erectus but also the view that if it happened once there may have been several waves of immigration to Flores. The unusual ‘hobbit’ anatomy is not restricted to tiny size and a small skull and brain cavity (400 cm3), but includes odd hips, wrist bones, shoulder joint and collar bone. In fact the remains bear as much or more resemblance to australopithecines like ‘Lucy’ (3.2 Ma) than to other members of our genus, even H. erectus that has been proposed as its possible ancestor. Could they be far-travelled descendants of the 1.8 Ma old H. georgicus from Dmanisi in Georgia? More fossils clearly need to be found, and Stringer raises the possibility of the search being widened to other islands east of Java, such as Sulawesi, the Philippines and Timor. He hints that in such a tectonically active region tsunamis may have led to animals and humans saving themselves and then being current dispersed on rafts of broken vegetation, rather like some survivors of the 2004 Indian Ocean tsunami who ended up 150 miles from their homes by such a means.

Another story that is set to ‘run and run’ is that of ‘alien’ DNA in the human genome and productive relations between early out-of-Africa migrants with Neanderthals, Denisovans and perhaps yet a mysterious, earlier human species. The oldest (45 ka) anatomically modern human genome sequence so far charted is from a leg bone found by a mammoth-ivory prospector in Siberian permafrost (Fu, Q. and 27 others 2014. Genome sequence of a 45,000-year-old modern human from western Siberia. Nature, v. 514, p. 445-449). Like a great many living non-Africans this individual carried about 2 % Neanderthal DNA, but unlike living people the 45 ka genome has it in significantly longer segments. That allowed the authors to re-estimate the timing of the genetic flow from Neanderthals into the individual’s ancestors. Previous estimates from living DNA geve the possibility of that being between 37-86 ka, but this closer data suggests that it happened between 7 to 13 ka before the date of the fossil femur, i.e. narrowing it down to between 52 and 58 ka closer to the widely suggested time of African exodus around 60 ka (but see an Earth Pages item from September 2014)

More on human evolution here and here