Tag: Human evolution

Advances in hominin evolution

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For decades, most of the news concerning our deep ancestry emerged from discoveries in sub-Saharan Africa at sites in Zambia, Tanzania, Kenya, South Africa, Ethiopia. The first week of 2026 decisively shifted that focus northwards to Chad and Morocco in two separate publications.

In 2002 ago the world of palaeoanthropology was in turmoil following the first discovery of fragments of what was then thought to be a hominid, or great-ape, cranium in Chad dated at around 7 Ma ago (Brunet, M. and 37 others 2002. A new hominid from the Upper Miocene of Chad, central Africa. Nature, v. 4418, p. 145-151;DOI:10.1038/nature00879). When pieced together the cranium looked like a cross between that of a chimpanzee and an australopithecine. Some suggested that the creature may have been a ‘missing link’ between the hominids and hominins; perhaps the ultimate ancestor of humans. Sahelanthropus tchadensis (nicknamedToumaï­ or ‘hope of life’ in the local Goran language) was undoubtedly enigmatic. The ‘molecular-clock’ age estimate for the branching of hominins from a common ancestor with chimpanzees was, in 2002, judged to be two million years later the dating of Sahelanthropus, so controversy was inevitable. Another point of contention was the size of Sahelanthropus’s canine teeth: too large for australopithecines and humans, but more appropriate for a gorilla or chimp. Moreover, Toumaï­ showed no indisputable evidence for having been bipedal. The Chadian site subsequently yielded three lower jaw bones and a collection of teeth, a partial femur (leg bone) and three fragmentary ulnae (forearm bones). The finds suggested that as many as five individuals had been fossilised. The femur gave an unresolved hint of an upright gait, yet the ulnas suggested Toumaï­ might equally have been arboreal; as could also be said for the australopithecines.

Reconstructed skull of Sahelanthropus tchadensis. (Credit: Didier Descouens, University of Toulouse)

All the limb bones of Toumaï­have now been anatomically compared with those of hominins and apes (Williams S.A. et al. 2026. Earliest evidence of hominin bipedalism in Sahelanthropus tchadensis. Science Advances, v. 12, article eadv0130; DOI: 10.1126/sciadv.adv0130). Scott Williams of New York University and co-workers from other US institutions show that although the leg bones are much the same size as those of chimpanzees, their proportions were more like those of hominins. They also showed features around the knees and hips needed for bipedalism and an insertion point for a tendon for the gluteus maximus muscle (buttock) vital for sustained upright locomotion, similar to the femurs of Orrorin tugenensis (see: Orrorin walked the walk; May 2008) and Ardipithecus ramidus. Unfortunately, an intact Sahelanthropus cranium showing a foramen magnum – where the skull attaches to the spine – continues to elude field workers. Its position distinguishes upright gait definitively.

See also: This ancient fossil could rewrite the story of human origins.  Science Daily, January 3, 2026)

The second new advance concerns the joint ancestry of Neanderthals, Denisovans and anatomically modern humans (AMH), whose ancient genetics crudely suggest a last common ancestor living between 765 to 550 ka. This had previously been attributed to Homo antecessor found in the Gran Dolina cave at Atapuerca in northern Spain, roughly dated between 950 ka and 770 ka. (Incidentally, Gran Dolina has yielded plausible evidence of cannibalism). A novel possibility stems from hominin fossils excavated from a cave in raised-beach sediments near Casablanca in Morocco (Hublin, JJ. and 28 others, 2026  Early hominins from Morocco basal to the Homo sapiens lineage. Nature, v. 649 ; DOI: 10.1038/s41586-025-09914-y). The fossil-bearing sediments contain evidence for a shift in the Earth’s magnetic field (the Brunhes–Matuyama reversal) dated at 773 ka, much more precisely than the Atapuerca age span for H. antecessor. Jean-Jacques Hublin of CNRS in Paris and his multinational colleagues report that the fossils are similar in age to H. antecessor, yet are morphologically distinct, displaying a combination of primitive traits and of ‘derived features reminiscent of’ later Neanderthal, Denisovan and AMH fossils. The differences and shared features suggest that there may have been genetic exchanges between the Moroccan and Iberian population over a considerable period. The most obvious route would have been across the Straits of Gibraltar, but would have required some kind of water craft.  An important question is ‘which population gave rise to the other?’

Artistic reconstruction of a juvenile Homo antecessor, Based on skeletal remains from Gran Dolina Cave

Larger and more robust hominin remains in Algeria dated at 1,000 ka – H. heidelbergensis? – resemble those found near Casablanca. They may have evolved to the latter. Similar possible progenitors to Iberian Homo antecessor have yet to be found in Western Europe. Homo erectus appeared in Georgia and Romania between 2.0 and 1.9 Ma, but the intervening million years or more have yielded no credible European forebears of H. antecessor. For the moment, incursion of a North African population into Europe followed by sustained contact is Hublin et al’s favoured hypothesis, rather than a European origin for Homo antecessor. For Neanderthals and Denisovans to have originated from such an African group, as has been suggested, requires finds of African fossils with plausible resemblance to what are predominantly Eurasian groups. The Iberian population migrated far and wide in Western Europe, as witnessed by stone tools and footprints dating to between 950 to 850 ka in eastern England. So it is equally possible that the Iberian group were progenitors of Neanderthals and Denisovans in Eurasia itself. At least for the moment, ancient genomes of the two H. antecessor groups are unlikely to be found in either Iberian or African fossils of the same antiquity. But, as usual, that will not stifle debate: a resort to the adage ‘absence of evidence is not evidence of absence’ seems appropriate to several research teams!

The oldest anatomically modern human fossils dated at ~300 ka, were also discovered in Morocco (see: Origin of anatomically modern humans, June 2017). Their isolation in the NW corner of the African continent poses a similar conundrum, as since then such beings went on to occupy wide areas of sub-Saharan Africa and then the world.

A possible Chinese ancestor for Denisovans, Neanderthals and modern humans

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Assigning human fossils older than around 250 ka to different groups of the genus Homo depends entirely on their physical features. That is because ancient DNA has yet to be found and analysed from specimens older than that. The phylogeny of older human remains is also generally restricted to the bones that make up their heads; 21 that are fixed together in the skull and face, plus the moveable lower jaw or mandible. Far more teeth than crania have been discovered and considerable weight is given to differences in human dentition. Teeth are not bones, but they are much more durable, having no fibrous structure and vary a great deal. The main problem for palaeoanthropologists is that living humans are very diverse in their cranial characteristics, and so it is reasonable to infer that all ancient human groups were characterised by such polymorphism, and may have overlapped in their physical appearance. A measure of this is that assigning fossils to anatomically modern humans, i.e. Homo sapiens, relies to a large extent on whether or not their lower mandible juts out to define a chin. All earlier hominins and indeed all other living apes might be regarded as ‘chinless wonders’! This pejorative term suggests dim-wittedness to most people, and anthropologists have had to inure themselves to such crude cultural conjecture.

The extraction, sequencing and comparison of ancient DNA from human fossils since 2010 has revealed that three distinct human species coexisted and interbred in Eurasia. Several well preserved examples of ancient Neanderthals and anatomically modern humans (AMH) have had their DNA sequenced, but a Denisovan genome has only emerged from a few bone fragments from the Denisova Cave in western Siberia. Whereas Neanderthals have well-known robust physical characters, until 2025 palaeoanthropologists had little idea of what Denisovans may have looked like. Then proteins and, most importantly, mitochondrial DNA (mtDNA) were extracted from a very robust skull found around 1931 in Harbin, China, dated at 146 ka. Analysis of the mtDNA and proteins, from dental plaque and bone respectively, reveal that the Harbin skull is likely to be that of a Denisovan. Previously it had been referred to as Homo longi, or ‘Dragon Man’, along with several other very robust Chinese skulls of a variety of ages.

The distorted Yunxian cranium (right) and its reconstruction (middle) [Credit: Guanghui Zhao] compared with the Harbin Denisovan cranium (left) [Hebei Geo University]

The sparse genetic data have been used to suggest the times when the three different coexisting groups diverged. DNA in Y chromosomes from Denisovans and Neanderthals suggest that the two lineages split from a common ancestor around 700 ka ago, whereas Neanderthals and modern humans diverged genetically at about 370 ka. Yet the presence of sections of DNA from both archaic groups in living humans and the discovery that a female Neanderthal from Denisova cave had a Neanderthal mother and a Denisovan father reveals that all three were interfertile when they met and interacted. Such admixture events clearly have implications for earlier humans. There are signs of at least 6 coexisting groups as far back as the Middle Pleistocene (781 to 126 ka), referred to by some as the ‘muddle in the middle’ because such an association has increasingly mystified palaeoanthropologists. A million-year-old, cranium found near Yunxian in Hubei Province, China, distorted by the pressure of sediments in which it was buried, has been digitally reconstructed.

This reconstruction encouraged a team of Chinese scientists, together with Chris Stringer of the UK Museum of Natural History, to undertake a complex statistical study of the Yunxian cranium. Their method compares it with anatomical data for all members of the genus Homo from Eurasia and Africa, i.e. as far back as the 2.4 Ma old H. habilis (Xiabo Feng and 12 others 2025. The phylogenetic position of the Yunxian cranium elucidates the origin of Homo longi and the Denisovans. Science, v. 389, p. 1320-1324; DOI: 10.1126/science.ado9202). The study has produced a plausible framework that suggests that the five large-brained humans known from 800 ka ago – Homo erectus (Asian), H. heidelbergensis, H. longi (Denisovans), H. sapiens, and H. neanderthalensis – began diverging from one another more than a million years ago. The authors regard the Yuxian specimen as an early participant in that evolutionary process. The fact that at least some remained interfertile long after the divergence began suggests that it was part of the earlier human evolutionary process. It is also possible that the repeated morphological divergence may stem from genetic drift. That process involves small populations with limited genetic diversity that are separated from other groups, perhaps by near-extinction in a population bottleneck or as a result of the founder effect when a small group splits from a larger population during migration. The global population of early humans was inevitably very low, and migrations would dilute and fragment each group’s gene pool.

The earliest evidence for migration of humans out of Africa emerged from the discovery of five 1.8 Ma old crania of H. erectus at Dmanisi to the east of the Black Sea in Georgia. similar archaic crania have been found in eastern Eurasia, especially China, at various localities with Early- to Middle Pleistocene dates. The earliest European large-brained humans – 1.2 to 0.8 Ma old H. antecessor from northern Spain – must have migrated a huge distance from either Africa or from eastern Eurasia and may have been a product of the divergence-convergence evolutionary framework suggested by Xiabo Feng and colleagues. Such a framework implies that even earlier members of what became the longi, heidelbergensis, neanderthalensis, and sapiens lineages may await either recognition or discovery elsewhere. But the whole issue raises questions about the widely held view that Homo sapiens first appeared 300 ka ago in North Africa and then populated the rest of that continent. Was that specimen a migrant from Eurasia or from elsewhere in Africa? The model suggested by Xiabo Feng and colleagues is already attracting controversy, but that is nothing new among palaeoanthropologists. Yet it is based on cutting edge phylogeny derived from physical characteristics of hominin fossils: the traditional approach by all palaeobiologists. Such disputes cannot be resolved without ancient DNA or protein assemblages. But neither is a completely hopeless task, for Siberian mammoth teeth have yielded DNA as old as 1.2 Ma and the record is held by genetic material recovered from sediments in Greenland that are up to 2.1 Ma old. The chances of pushing ancient human DNA studies back to the ‘muddle’ in the Middle Pleistocene depend on finding human fossils at high latitudes in sediments of past glacial maxima or very old permafrost, for DNA degrades more rapidly as environmental temperature rises.

See also: Natural History Museum press release. Analysis of reconstructed ancient skull pushes back our origins by 400,000 years to more than one million years ago. 25 September 2025; Bower, B. 2025. An ancient Chinese skull might change how we see our human roots. ScienceNews, 25 September 2025; Ghosh, P. 2025. Million-year-old skull rewrites human evolution, scientists claim. The Guardian, 25 September 2025

How did African humans survive the 74 ka Toba volcanic supereruption?

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The largest volcanic eruption during the 2.5 million year evolution of the genius Homo, about 74 thousand years (ka) ago, formed a huge caldera in Sumatra, now filled by Lake Toba. A series of explosions lasting just 9 to 14 days was forceful enough to blast between 2,800 to 6,000 km3 of rocky debris from the crust. An estimated 800 km3 was in the form of fine volcanic ash that blanketed South Asia to a depth of 15 cm. Thin ash layers containing shards of glass from Toba occur in marine sediments beneath the Indian Ocean, the Arabian and South China Seas. Some occur as far off as sediments on the floor of Lake Malawi in southern Africa. A ‘spike’ of sulfates is present at around 74 ka in a Greenland ice core too. Stratospheric fine dust and sulfate aerosols from Toba probably caused global cooling of up to 3.5 °C over a modelled 5 years following the eruption. To make matters worse, this severe ‘volcanic winter’ occurred during a climatic transition from warm to cold caused by changes in ocean circulation and falling atmospheric CO2 concentration, known as a Dansgaard-Oeschger event.

There had been short-lived migrations of modern humans out of Africa into the Levant since about 185 ka. However, studies of the mitochondrial DNA (mtDNA) of living humans in Eurasia and Australasia suggest that permanent migration began about 60 ka ago. Another outcome of the mtDNA analysis is that the genetic diversity of living humans is surprisingly low. This suggests that human genetic diversity may have been sharply reduced globally roughly around the time of the  Toba eruption. This implies a population bottleneck with the number of humans alive at the time to the order of a few tens of thousands (see also: Toba ash and calibrating the Pleistocene record; December 2012). Could such a major genetic ‘pruning’ have happened in Africa? Over six field seasons, a large team of geoscientists and archaeologists drawn from the USA, Ethiopia, China, France and South Africa have excavated a rich Palaeolithic site in the valley of the Shinfa River, a tributary of the Blue Nile in western Ethiopia. Microscopic studies of the sediments enclosing the site yielded glass shards whose chemistry closely matches those in Toba ash, thereby providing an extremely precise date for the human occupation of the site: during the Toba eruption itself (Kappelman, Y. and 63 others 2024. Adaptive foraging behaviours in the Horn of Africa during Toba supereruption. Nature, v. 627; DOI: 10.1038/s41586-024-07208-3).

Selection of possible arrowheads from the Shinfa River site (Credit: Kappelman et al.; Blue Nile Survey Project)

The artifacts and bones of what these modern humans ate suggest a remarkable scenario for how they lived. Stone tools are finely worked from local basalt lava, quartz and flint-like chalcedony found in cavities in lava flows. Many of them are small, sharp triangular points, some of which show features consistent with their use as projectile tips that fractured on impact; they may be arrowheads, indeed the earliest known. Bones found at the site are key pointers to their diet. They are from a wide variety of animal, roughly similar to those living in the area at present: from monkeys to giraffe, guinea fowl to ostrich, and even frogs. There are remains of many fish and freshwater molluscs. Although there are no traces of plant foods, clearly those people who loved through the distant effects of Toba were well fed. Although a period of global cooling may have increased aridity at tropical latitudes in Africa, the campers were able to devise efficient strategies to obtain victuals. During wet seasons they lived off terrestrial prey animals, and during the driest times ate fish from pools in the river valley. These are hardly conditions likely to devastate their numbers, and the people seem to have been technologically flexible. Similar observations were made at the Pinnacle Point site in far-off South Africa in 2018, where Toba ash is also present. Both sites refute any retardation of human cultural progress 74 ka ago. Rather the opposite: people may have been spurred to innovation, and the new strategies may have allowed them to migrate more efficiently, perhaps along seasonal drainages. In this case that would have led them or their descendants to the Nile and a direct route to Eurasia; along ‘blue highway’ corridors as Kappelman et al. suggest.

Yet the population bottleneck implied by mtDNA analyses is only vaguely dated: it may have been well before or well after Toba. Moreover, there is a 10 ka gap between Toba and the earliest accurately dated migrants who left Africa – the first Australians at about 65 ka. However, note that there is inconclusive evidence that modern humans may have occupied Sumatra by the time of the eruption.  Much closer to the site of the eruption in southeast India, stone artifacts have been found below and above the 74 ka datum marked by the thick Toba Ash. Whether these were discarded by anatomically modern humans or earlier migrants such as Homo erectus remains unresolved. Either way, at that site there is no evidence for any mass die-off, even though conditions must have been pretty dreadful while the ash fell. But that probably only lasted for little more than a month. If the migrants did suffer very high losses to decrease the genetic diversity of the survivors, it seems just as likely to have been due to attrition on an extremely lengthy trek, with little likelihood of tangible evidence surviving. Alternatively, the out-of-Africa migrants may have been small in number and not fully representative of the genetic richness of the Africans who stayed put: a few tens of thousand migrants may not have been very diverse from the outset.

An evolutionary bottleneck and the emergence of Neanderthals, Denisovans and modern humans

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The genetic diversity of living humans, particularly among short, repetitive segments of DNA, is surprisingly low. As they are passed from generation to generation they have a high chance of mutation, which would be expected to create substantial differences between geographically separated populations. In the late 1990s and early 2000s some researchers attributed the absence of such gross differences to the human gene pool having been reduced to a small size in the past, thereby reducing earlier genetic variation as a result of increased interbreeding among survivors. They were able to assess roughly when such a population ‘bottleneck’ took place and the level to which the global population fell. Genetic analysis of living human populations seemed to suggest that around 74 ka ago the global human population fell to as little as 10 thousand individuals. A potential culprit was the catastrophic eruption of the Toba supervolcano in Sumatra around that time, which belched out 800 km3 of ash now found as far afield as the Greenland and Antarctic ice caps. Global surface temperature may have fallen by 10°C for several years to decades. Subsequent research has cast doubt on such a severe decline in numbers of living hummans; for instance archaeologists working in SE India found much the same numbers of stone tools above the Toba ash deposit as below it (see: Toba ash and calibrating the Pleistocene record: December 2012). Other, less catastrophic explanations for the low genetic diversity of modern humans have also been proposed. Nevertheless, environmental changes that placed huge stresses on our ancestors may repeatedly have led to such population bottlenecks, and indeed throughout the entire history of biological evolution.

An improved method of ‘back-tracking’ genetic relatedness among living populations, known as fast infinitesimal time coalescence or ‘FitCoal’, tracks genomes of individuals back to a last common ancestor. In simple language, it expresses relatedness along lineages to find branching points and, using an assumed mutation rate, estimates how long ago such coalescences probably occurred. The more lineages the further back in time FitCoal can reach and the greater the precision of the analysis. Moreover it can suggest the likely numbers of individuals, whose history is preserved in the genetics of modern people, who contributed to the gene pool at different branching points. Our genetics today are not restricted to our species for it is certain that traces of Neanderthal and Denisovan ancestry are present in populations outside of Africa. African genetics also host ‘ghosts’ of so-far unknown distant ancestors. So, the FitCoal approach may well be capable of teasing out events in human evolution beyond a million years ago, if sufficient data are fed into the algorithms. A team of geneticists based in China, Italy and the US has recently applied FitCoal to genomic sequences of 3154 individual alive today (Hu, W.and 8 others 2023. Genomic inference of a severe human bottleneck during the Early to Middle Pleistocene transition. Science, v. 381, p. 979-984; DOI I: 10.1126/science.abq7487). Their findings are startling and likely to launch controversy among their peers.

Their analyses suggest that between 930 and 813 ka ago human ancestors passed through a population bottleneck that involved only about 1300 breeding individuals. Moreover they remained at the very brink of extinction for a little under 120 thousand years. Interestingly, the genetic data are from people living on all continents, with no major differences between the analyses for geographically broad groups of people in Africa and Eurasia. Archaeological evidence, albeit sparse, suggests that ancient humans were widely spread across those two continental masses before the bottleneck event. The date range coincides with late stages of the Mid-Pleistocene climatic transition (1250 to 750 ka) during which glacial-interglacial cycles changed from 41 thousand-year periods to those that have an average duration of around 100 ka. The transition also brought with it roughly a doubling in the mean annual temperature range from the warmest parts of interglacials to the frigid glacial maxima: the world became a colder and drier place during the glacial parts of the cycles.

Genomes for Neanderthals and Denisovans suggest that they emerged as separate species between 500 and 700 ka ago. Their common ancestor, possibly Homo heidelbergensis, H. antecessor or other candidates (palaeoanthropologists habitually differ) may well have constituted the widespread population whose numbers shrank dramatically during the bottleneck. Perhaps several variants emerged because of it to become Denisovans, Neanderthals and, several hundred thousand years later, of anatomically modern humans. Yet it would require actual DNA from one or other candidate for the issue of last common ancestor for the three genetically known ‘late’ hominins to be resolved. But Hu et al. have shown a possible means of accelerated hominin evolution from which they may have emerged, at the very brink of extinction.

Oxygen-isotope record and global temperature changes over the last 5 million years, green lines showing the times dominated by 41 and 100 ka climatic cycles. The mid-Pleistocene climatic transition is shown in pink (Credit: Robert A Rohde)

There is a need for caution, however. H. erectus first appeared in the African fossil record about 1.8 Ma ago and subsequently spread across Eurasia to become the most ‘durable’ of all hominin species. Physiologically they seem not to have evolved much over at least a million years, nor even culturally – their biface Acheulean tools lasted as long as they did. They were present in Asia for even longer, and apparently did not dwindle during the mid-Pleistocene transition to the near catastrophic levels as did the ancestral species for living humans. The tiny global population suggested by Hu et al. for the latter also hints that their geographic distribution had to be very limited; otherwise widely separated small bands would surely have perished over the 120 ka of the bottleneck event. Yet, during the critical period from 930 to 813 ka even Britain was visited by a small band of archaic humans who left footprints in river sediments now exposed at Happisburgh in Norfolk. Hu et al. cite the scarcity of archaeological evidence from that period – perhaps unwisely – in support of their bottleneck hypothesis. There are plenty of other gaps in the comparatively tenuous fossil and archaeological records of hominins as a whole.

The discovery of genetic evidence for this population bottleneck is clearly exciting, as is the implication that it may have been the trigger for evolution of later human species and the stem event for modern humans. Hopefully Hu et al’s work will spur yet more genetic research along similar lines, but there is an even more pressing need for field research aimed at new human fossils from new archaeological sites.

See also: Ashton, N. & Stringer, C. 2023. Did our ancestors nearly die out? Science (Perspectives), v. 381, p. 947-948; DOI: 10.1126.science.adj9484.

Ikarashi, A. 2023. Human ancestors nearly went extinct 900,000 years ago. Nature, v. 621; DOI: 10.1038/d41586-023-02712-4

Di Vicenzo, F & Manzi, G. 2023. An evolutionary bottleneck and the emergence of Neanderthals, Denisovans and modern humans. Homo heidelbergensis as the Middle Pleistocene common ancestor of Denisovans, Neanderthals and modern humans. Journal of Mediterranean Earth Sciences, v, 15, p. 161-173; DOI: 10.13133/2280-6148/18074

A book on archaeology, radiocarbon dating, ancient DNA, and how modern humans evolved

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Since 2001 Tom Higham, now Professor of Scientific Archaeology at the University of Vienna, helped develop new ways of refining radiocarbon dating at Oxford University’s Research Lab for Archaeology and the History of Art. Specifically his lab learned how to remove contamination of ancient samples by recent carbon and to reduce the detection limit of their accelerator mass spectrometer for the 14C atoms that remained from when they were in living organisms. The Oxford Radiocarbon Accelerator Unit pushed sample dates to the absolute limit of the method: around 50 thousand years. Being among the very best, the ORAU had a path beaten to its doors by archaeologists from across the world keen to get the most believable dates for their samples. Equally, Higham engaged in the field work itself and in the interpretation of other data from sites, such as ancient DNA. An outcome of Higham’s energetic efforts over two decades is his book The World Before Us: How Science is Revealing a New Story of Our Human Origins (paperback edition 2022, Penguin Books,ISBN-10: ‎0241989051). One reviewer commented ‘The who, what, where, when and how of human evolution’.

The World Before Us is not only comprehensive and eminently clear for the lay-reader, but it is more exciting than any science book that I have read. For the moment, it is the latest ‘word’ on early, anatomically modern humans and on the closely related Neanderthals and Denisovans. Its core is about how these three key groups ‘rubbed along’ once they met  in the Late Pleistocene. As an amateur interested in palaeoanthropology, I have tried to keep pace with all the developments in the field since 2001 through Earth-logs, but Higham shows just how much I have missed that is important to the human story. If you have followed my many posts on human evolution and migrations with interest, read his book for a great deal more and a coherent story of how things stand.

The ancestry of our opposable thumbs

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Since the appearance of smart phones and the explosion of social media our thumbs have found a new niche; typing while holding a mobile. At a desktop keyboard, most of us don’t use thumbs very much, unless we have mastered fast touch typing, but for a huge variety of manual tasks thumbs are essential. The first makers of sophisticated stone tools must have been able to grip between fingers and thumb to manipulate the materials from which they were made and to perform the various stages in creating a razor sharp edge. To do that, as most of us are aware, the tip of the thumb must be capable of touching the tips of all four fingers; an opposable thumb is essential for the ‘precision grip’. Being able to tell when opposable thumbs evolve depends, of course, on finding hand-bone fossils. Being made of many bones disarticulated hands are a lot more fragile than long bones or those of the skull. Complete fossil hands are rare, as are feet, but a number have been found more or less complete. Whichever hominin had evolved opposable thumbs, their potential would have given them a considerable advantage over those that hadn’t.

The main muscles that control the movements of modern human fingers and thumb (Credit: Wikipedia)

Simply comparing the shapes of fossilised bones of fingers and thumbs with those of modern humans and other living primates has, so far, not proved capable of resolving with certainty which hominin groups either did or did not have opposable thumbs. The key lies in the muscles that operate them. It has become commonplace to reconstruct faces and even whole bodies from fairly complete skeletal remains by modelling musculature from the positioning and shape of the points of attachment of muscles to bone. But that become increasingly difficult for the small-scale and intricate attachments in hands. The critical muscle for opposable thumbs is known as the Opponens pollicis (the Latin for thumb is Digitus pollex); a small triangular muscle that operates in conjunction with three others (with pollicis in their Latin names).

Fotios Karakostis and six colleagues from German, Swiss and Greek universities have devised software that can model muscles in 3-D (F.A. Karakostis et al. 2021. Biomechanics of the human thumb and the evolution of dexterity. Current Biology, v.31,  online; DOI: 10.1016/j.cub.2020.12.041). Based on the anatomy of human and chimpanzee hand muscles and the positions of their attachment to individual bones, they have been able to establish a series of parameters that clearly distinguish the morphological and probably functional characteristics of the thumbs of these living primates. Complete sets of thumb bones from four Neanderthal skeletons show that they were significantly, but only slightly, different from anatomically modern humans. Those from three species of Australopithecus (africanus, sediba and afarensis) lie between ours and chimps’, with significantly closer affinity to chimpanzees. It seems that australopithecines of whatever age were not equipped with opposable thumbs and were possible tool producers and users with the very limited capabilities of modern chimps; holding, pounding and poking. A single set of hominin thumb bones from about two million years ago that were found in the famous Swartkrans Cave in South Africa show just as close affinity in thumb opposability to humans as do Neanderthals. So at 2 Ma there was a hominin species sufficiently dextrous to make and use sophisticated tools. The problem is, the bones are not directly associated with others and have been ascribed by different authors either to H. habilis or Paranthropus robustus. Interestingly, this paranthropoid has also been suggested (controversially) to have been the first known hominin to use fire, and it also used digging sticks. No one has ever suggested that the genus Homo descended from a paranthropoid ancestor or vice versa; these massively jawed beings did coexist with early humans in East Africa for over a million years. The other hominin who left hands in the geological record was Homo naledi; a controversial species because it was found in a barely accessible cave chamber, and took a while to date. This context gave rise to the notions that it was the direct ancestor of humans and that it buried its dead in a special place. However, it turned out to be relative recent, at about 280 ka (see: Homo naledi: an anti-climax; May 2017). Homo naledi does seem to have had opposable thumbs, but there is no associated evidence to suggest either tool making or use.

Fascinating as the methodology outlined by Karakostis et al. is, their findings do not take early human capabilities very much further than what is already known. Tools were made and used as far back as 3.3 Ma ago, and we know that H. habilis was doing this by about 2.6 Ma; i.e. long before the first evidence for opposable thumbs, and who had them first is uncertain. What is clear is that sophisticated tools, such as the bifacial Acheulian artifacts whose manufacture demands great dexterity, only appeared after the potential for nimble dexterity (about 1.8 Ma). The same goes for the first migration out of Africa, at about the same time, which demanded resourcefulness that may have sprung from the ability to manipulate natural materials effectively and carefully

See also: Handwerk B. 2012. How dexterous thumbs may have helped shape evolution two million years ago. (Smithsonian Magazine, 28 January 2021); Bower, B. 2021. Humanlike thumb dexterity may date back as far as 2 million years ago. (Science News, 28 January 2021)

A protein clue to H. antecessor’s role in human evolution

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Homo_antecessor child
Forensic reconstruction of the remains of a Homo antecessor child from Gran Dolina Cave in northern Spain (credit Élisabeth Daynès, Museo de la Evolución, Burgos, Spain)

The older a fossil, no matter how well preserved it is, the less chance it has to contain enough undegraded DNA for it to be extracted and sequenced using the most advanced techniques. At present the oldest fossil DNA not to have passed its ‘sell-by’date is that of a 560 to 780 thousand year-old horse’s legbone found in Canadian permafrost. For human remains the oldest mtDNA is that of a ~430 ka individual from the Sima de los Huesos in northern Spain (see: Mitochondrial DNA from 400 thousand year old humans; Earth-logs December 2013). But there is another route to establishing genetic relatedness from the amino-acid sequences of proteins recovered from ancient individuals (see: Ancient proteins: keys to early human evolution?). Fossil teeth have proved to be good repositories of ancient protein and are the most commonly found hominin fossils.

A key species for unravelling the origins of the three most recent human groups (ourselves, Neanderthals and Denisovans) is thought to be Homo antecessor who inhabited the Gran Dolina Cave in the Atapuerca Mountains in northern Spain between about 1.2 Ma and 800 ka ago (see: Human evolution: bush or basketwork? Earth-logs, January 2014). Palaeoanthropologists excavated 170 skeletal fragments from six individuals in the most productive layer at Gran Dolina. Incomplete facial bones suggest a ‘modern-like’ face, although the remains as a whole are insufficient to reconstruct the oldest Europeans with sufficient detail to place them in anatomical relation to the younger groups. But there are several teeth. One of them, a permanent molar, has yielded informative proteins (Welker, F. and 26 others 2020. The dental proteome of Homo antecessor. Nature, v. 580, p. 235-238; DOI: 10.1038/s41586-020-2153-8) and has been dated to between 772 to 949 ka.

Amino acids in the dental proteins, sequenced using mass spectrometry, were compared with those of other hominins. Because protein sequences are coded by an animal’s genome they are a ‘proxy’ for DNA. The outcome is that the Gran Dolina proteins are roughly equally related to Denisovans, Neanderthals and ourselves, suggesting that, although the younger three groups are closely related, H. antecessor is an ‘outlier’. Being significantly older, it is likely to be the common ancestor of all three. Another species with close anatomical affinities is H. heidelbergensis (700 to 300 ka) found in Africa as well as in Europe. Its mtDNA (see: Mitochondrial DNA from 400 thousand year old humans; Earth-logs December 2013) matches that of Denisovans better than it does Neanderthals, yet without protein and full-genome analysis all that can be concluded is that it may be an intermediary between H. antecessor and the well known interbreeding triad of more recent times.

We are getting closer to a documented web of interrelationships between humans in general whose time span from 2 Ma ago is now well established. The remaining genetic link to be documented is that to H. erectus, the longest lived and most travelled of all ancient humans. Frido Welker and co-workers also had a shot at the proteomics of one of the first humans known to have migrated from Africa, using an isolated, presumably H. erectus, molar found at the 1.77 Ma site at Dmanisi in the Caucasus foothills of Georgia. Although inconclusive in placing that precociously intrepid group firmly in the human story, the fact that dental proteins were discovered is cause for optimism.

See also: Campbell, M. 2020. Protein analysis of 800,000-year-old human fossil clarifies dispute over ancestors (Technology Networks, 1 April 2020)

Further back in the Eurasian human story

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About 800 to 950 thousand years (ka) ago the earliest human colonisers of northern Europe, both adults and children, left footprints and stone tools in sedimentary strata laid down by a river system that then drained central England and Wales. The fossil flora and fauna at the Happisburgh (pronounced ‘Haze-burra’) site in Norfolk suggest a climate that was somewhat warmer in summers than at present, with winter temperatures about 3°C lower than now: similar to the climate in today’s southern Norway. At that time the European landmass extended unbroken to the western UK, so any hunter-gatherers could easily follow migrating herds and take advantage of seasonal vegetation resources. These people don’t have a name because they left no body fossils. A group known from their fossils as Homo antecessor had occupied Spain, southern France and Italy in slightly earlier times (back to 1200 ka). Since the discovery of their unique mix of modern and primitive traits, they have been regarded as possible intermediaries between H. erectus and H. heidelbergensis – once supposed to be the predecessor of Neanderthals and possibly anatomically modern humans (AMH). Since the emergence about 10 years ago of ancient genomics as the prime tool in examining human ancestry the picture has been shown to be considerably more complex. Not only had AMH interbred with Neanderthals and Denisovans, those two groups were demonstrably interfertile too, and a complex web of such relationships had been pieced together by 2016. But there has been a new development.

700 ka Homo erectus from Java: a possible Eurasian ‘super-archaic’ human (credit: Gibbons 2020)

Population geneticists at the University of Utah, USA, have devised sophisticated means of making more of the detailed ATCG nucleotide sequences in ancient human DNA, despite there being very few full genomes of Neanderthals and Denisovans (Rogers, A.R. et al. 2020. Neanderthal-Denisovan ancestors interbred with a distantly related hominin. Science Advances, v. 6, article eaay5483; DOI: 10.1126/sciadv.aay5483). In Earth-logs you may already have come across the idea of the ancestral ‘ghosts’ that are represented by unusual sections of genomes from living West African people. Those sections seem likely to have resulted from interbreeding with an unknown archaic population – i.e. neither Neanderthal nor Denisovan. It now seems that both Neanderthal and Denisovan genomes also show traces of such introgression with ‘ghost’ populations during much earlier times. The ancestors of both these groups separated from the lineage that led to AMH perhaps 750 ka ago. Rogers et al. refer to the earliest as ‘neandersovans’ and consider that they split into the two groups after they entered Eurasia, at some time before 600 ka – perhaps around 740 ka. This division may well have occurred as a result of a population of ‘neandersovans’ having spread over the vastness of Eurasia and growing genetic isolation. The reanalysis of both sets of genomes show evidence of a ‘neandersovan’ population crash before the split. Thereafter, the early Neanderthal population may have risen to around 16 thousand then slowly declined to ~3400 individuals.

A ‘state-of-play’ view of human interbreeding in Eurasia since 2 Ma ago (credit: Gibbons 2020)

However, the ‘neandersovans’ did not enter a new continent devoid of hominins, for as long ago as 1.9 Ma archaic H. erectus had arrived from Africa.  Both Neanderthal and Denisovan genomes record the presence of sections of ‘super-archaic’ DNA, which reflect early  interbreeding with earlier Eurasian populations. Indeed, Denisovans seem to have repeated their ancestors’ sexual exploits, once they became a genetically distinct group.  From the ‘ghost’ DNA fragments Rogers et al. conclude that the ‘super-archaics’ separated from other humans about two million years ago. They were descended from the first ‘Out-of-Africa’ wave of humans, represented by the fossils humans from Dmanisi in Georgia (see First out of Africa, November 2003 and An iconic early human skull,  October 2013 in Earth-logs Human evolution and migrations). A measure of the potential of novel means of analysing available ancient human DNA is the authors’ ability even to estimate the approximate population size of the interbreeding ‘super-archaic’ group at 20 to 50 thousand. Long thought to be impossible, it now seems possible to penetrate back to the very earliest human genetics, and the more DNA that can be teased out of other Neanderthal and Denisovan fossils the more we will know of our origins.

See also: Gibbons, A. 2020. Strange bedfellows for human ancestors. Science, v. 367, p. 838–839; doi:10.1126/science.367.6480.838

Ancient proteins: keys to early human evolution?

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A jawbone discovered in a Tibetan cave turned out to be that of a Denisovan who had lived and died there about 160,000 years ago (see: Denisovan on top of the world; 6 May, 2019). That discovery owed nothing to ancient DNA, because the fossil proved to contain none that could be sequenced. But the dentine in one of two molar teeth embedded in the partial jaw did yield protein. The teeth are extremely large and have three roots, rather than the four more common in modern, non-Asian humans, as are Denisovan teeth from in the Siberian Denisova Cave. Fortunately, those teeth also yielded proteins. In an analogous way to the genomic sequencing of nucleotides (adenine, thymine, guanine and cytosine) in DNA, the sequence of amino acids from which proteins are built can also be analysed. Such a proteomic sequence can be compared with others in a similar manner to genetic sequences in DNA. The Tibetan and Siberian dentine proteins are statistically almost the same.

collagen
Triple helix structure of collagen, colour-coded to represent different amino acids (credit: Wikipedia)

At present the most ancient human DNA that has been recovered – from an early Neanderthal in the Sima de los Huesos in Spain – is 430,000 years old (see: Mitochondrial DNA from 400 thousand year old humans; December 2013). Yet it is proving difficult to go beyond that time, even in the cool climates that slow down the degradation of DNA. The oldest known genome of any animal is that of mtDNA from a 560–780 thousand year old horse, a leg bone of which was extracted from permafrost in the Yukon Territory, Canada. The technologies on which sequencing of ancient DNA depends may advance, but, until then, tracing the human evolutionary journey back beyond Neanderthals and Denisovans seems dependent on proteomic approaches (Warren, M. 2019. Move over, DNA: ancient proteins are starting to reveal humanity’s history. Nature, v. 570, p. 433-436; DOI: 10.1038/d41586-019-01986-x). Are the earlier Homo heidelbergensis and H. erectus within reach?

It seems that they may be, as might even earlier hominins. The 1.8 Ma Dmanisi site in Georgia, now famous for fossils of the earliest humans known to have left Africa, also yielded an extinct rhinoceros (Stephanorhinus). Proteins have been extracted from it, which show that Stephanorhinus was closely related to the later woolly rhinoceros (Coelodonta antiquitatis). Collagen protein sequences from a 3.4 Ma camel preserved in the Arctic and even from a Tanzanian 3.8 Ma ostrich egg shell show the huge potential of ancient proteomics. Most exciting is that last example, not only because it extends the potential age range to that of Australopithecus afarensis but into tropical regions where DNA is at its most fragile. Matthew Warren points out potential difficulties, such as the limit of a few thousand amino acids in protein sequences compared with 3 million variants in DNA, and the fact that the most commonly found fossil proteins – collagens –  may have evolved very little. On the positive side, proteins have been detected in a 195 Ma old fossil dinosaur. But some earlier reports of intact diosaur proteins have been questioned recently (Saitta, E.T. et al. 2019. Cretaceous dinosaur bone contains recent organic material and provides an environment conducive to microbial communities. eLife, 8:e46205; DOI: 10.7554/eLife.46205)

 

Multiregional human evolution in Africa

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Africa is not only a large continent, but is subdivided into many different climatic zones and ecosystems and these have changed drastically over the last 2 Ma. It is further subdivided by terrain features, such as the courses of major rivers, large plateaus, tectonic rift systems and the mountains that frequently define their flanks. Getting around Africa is not easy today, was more difficult before modern transport, and many geomorphic provinces may have been mutually inaccessible in the distant past. For instance, the Sahara Desert forms a major barrier to travellers on foot because access to surface water is non-existent except at widely spaced oases. Without boats or rafts the Nile and Congo cannot be crossed for a thousand miles or more. Migration was perhaps a very rare event outside of periods of widespread humid climates or when great environmental stress forced people either to move or perish. Despite these physical and ecological divisions and barriers palaeoanthropologists have, until recently, tended to regard the evolution of Homo sapiens and earlier human and hominin species as having occurred within single populations: a linear view forced on them by scanty fossil remains and limited methodologies. Logically, when human numbers were small Africa probably had several isolated population Physical isolation would have engendered genetic isolation in which our ancestors evolved for tens of thousand years.

Anatomically modern human (AMH) remains found at Jebel Irhoud in Morocco turned out to be 315 ka old, displacing those from Ethiopia (190 ka) as the earliest known examples of AMH. Several more archaic H. sapiens fossils have turned up in southern Africa and as far afield as the Middle East, suggesting that the early evolution of AMH was in an Africa-wide context rather than in one area – the rift system of Ethiopia and Kenya – from which a new species radiated outwards. This breadth of finds has encouraged Eleanor Scerri of Oxford University and her many international colleagues to resurrect what was once a widely discarded hypothesis; a multiregional model of modern human origins, originally proposed to have arisen from pre-sapiens groups in Eurasia by Milford Wolpoff but which was sunk once genetic connections among living humans turned out to be rooted in Africa. (Scerri, E.M.L. and 22 others 2018. Did our species evolve in subdivided populations across Africa, and why does it matter? Trends in Ecology & Evolution, v. 33, p. 582-594; (PDF) doi: 10.1016/j.tree.2018.05.005). Scerri et al’s model is sited in Africa and the paper’s authors include several leading palaeoanthropologists who once opposed multiregionalism and established the Recent African Origin hypothesis on the back of the early genetic data.

early homo
Different early AMH cranium shapes: left Jebel Irhoud, Morocco (315 ka), right Qafzeh, the Levant (85 ka) (credit: Scerri et al, 2018; Figure 1)

From region to region in Africa, the oldest AMH crania show significant differences from each other, but within a distinct combination of features that clearly distinguish us from our fossil relatives and ancestors, such as Homo heidelbergensis from Zimbabwe and the primitive-looking H. naledi found in a South African cave in 2015. Improved dating now shows that the Zimbabwean H. heidelbergensis and H.naledi remains are roughly the same age as the Jebel Irhoud AMH specimens. The first has long been held as the progenitor of AMH and descended from H. antecessor, perhaps the common ancestor for AMH, Neanderthals and Denisovans about 700 ka ago. The three human species cohabited Africa early in the evolutionary history of AMH. It is now abundantly clear from ancient and modern genomes that AMH, Neanderthals and Denisovans interbred in Eurasia. The proximity in time and space of earlier African AMH to two more ancient human species opens up a similar possibility earlier in the emergence of all living humans. There is evidence for that too: Yoruba people living in West Africa, whose genomes have been analysed, carry up to 8% of genetic ancestry that originated in an unidentified ancient population that was non-sapiens. At present, DNA analysis with the same high precision and information content from other living Africans has not been performed, and deterioration of ancient DNA in African climates has so far thwarted genomic studies of ancient African fossils.

The new view of our origins points to repeated hybridisation involving other coexisting human species, as well as evolution in isolation, from the outset. It continued through later times while Neanderthals and Denisovans survived. Even recent human genetic history is peppered with intermingling of a great variety of migrants passing through all the habitable continents. Another issue: In the earliest times, were cultures exchanged as well as genes? The first appearance of AMH coincides with that of a new stone technology (Levallois technique), moving away from the earlier dominance by handaxes towards more delicate, leaf-shaped points, that characterise the African Middle Stone Age. Similar techniques reached Europe with the Neanderthals. Was this an invention of the earliest AMH or a joint venture?

You can find an excellent review of these issues in the September 2018 issue of Scientific American (Wong, K. 2018. Last hominin standing.  Scientific American, v. 319(3), p. 56-61) along with several other articles on human evolution.

A revised and updated edition of Steve Drury’s book Stepping Stones: The Making of Our Home World can now be downloaded as a free eBook

Early modern humans in Sumatra before the Toba eruption

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In late July 2017 news emerged that modern humans first reached Australia at least 65 thousand years ago. Confirming that the date of departure from Africa to end up in SE Asia and Australasia was  considerable earlier than previously believed, deposits in Sumatra that contain remains of early Home sapiens have yielded even older ages (Westaway, K.E. and 22 others 2017. An early modern human presence in Sumatra 73,000–63,000 years ago. Nature v. 548 online; doi:10.1038/nature23452). This resulted from a re-examination of material from the Padang Caves first excavated more than a century ago by Eugène Dubois, famous for his discovery in Java of the first H. erectus remains. A richly fossiliferous breccia in the Lida Ajer cave yielded a fauna characteristic of a rainforest biome and included two teeth that Dubois considered to be human. Several later palaeontologists confirmed his identification as have hominin specialists in the present Australian-Indonesian-American-British-Dutch-German team. The fossil assemblage has long suggested great antiquity for the site, but only now has it been dated precisely. The dating employed three methods: optically stimulated luminescence dating of quartz grains from the breccia (85±25 to 62±5  ka); uranium-series dating of speleothem including fragments of hollow ‘soda-straw’ stalactites(84±1 to 71±7 ka); uranium-series dating of gibbon and orangutan teeth found together with the human teeth (86±13 to 76±7 ka). Statistical analysis of the age data suggests 73 to 63 ka for the fauna, with a maximum age for deposition of the breccia of 84±1 ka.

Satellite image of Lake Toba, the site of a VE...
Satellite image of Lake Toba in NW Sumatra (at centre), the site of the largest volcanic eruption during the history of human evolution ~71,600 years ago (credit: Wikipedia)

Stone tools which may have been carried by anatomically modern humans into the area have previously been used to suggest a minimum date of the arrival of migrants, though they may have been carried by ­H. erectus. Remarkably, such tools have been found beneath a thick bed of volcanic ash found throughout southern Asia and in Indian Ocean sediment cores. This has been dated at 71.6 ka and represents the explosive collapse of the caldera now containing Lake Toba in NW Sumatra that was the largest volcanic event in the entire history of the genus Homo. The new age data from Lida Ajer suggests that modern humans were present in its vicinty before the eruption, a view also supported by ‘molecular-clock’ dating of the range of mitochondrial DNA carried by living SE Asian people (79 to 75 ka). So, despite the stupendous magnitude of the Toba eruption is seems likely that some of the migrants survived.  Together with the dating of the earliest Australians the Sumatran evidence is at odds with the view, widely held by palaeoanthropologists, that the ‘Out of Africa’ exodus began by crossing the Straits of Bab el Mandab between 74 and 58 ka when global sea-level fell markedly during marine oxygen-isotope Stage 4 (MIS4). A problem with that hypothesis has been that climatic and ecological conditions in southern Asia during MIS4 were unfavourable. But is seems that modern humans were already there and capable of adapting to both the climate shift and to the devastation undoubtedly caused by Toba.

Stepping Stones eBook

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Title

A revised and updated edition of Stepping Stones: The Making of Our Home World by Steve Drury, first published in 1999, has been released as a free eBook on the book’s web site https://earthstep.wordpress.com/. The revision incorporates the hundreds of commentaries on geoscientific advances written since 2000 by Steve for earth-pages. It is a personal view of the evolution of the Earth System and the emergence of humanity from it. First published by Oxford University Press, Stepping Stones was widely acclaimed by  fellow Earth scientists and general readers.

Homo naledi: an anti-climax

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In September 2015 a barrage of publicity announced the remarkable unearthing of the remains of 15 diminutive hominins, dubbed Homo nadeli, from the floor sediments of an almost inaccessible South African cave, part of the equally hyped ‘Cradle of Humankind’ UNESCO World Heritage Site near Johannesburg. An international team of lithe women speleo-archaeologists was recruited for the excavation, for which the original discoverers were incapably burly. The remains included numerous examples of still articulated intricate bones, such as those of feet and hands, and none show signs of dismemberment by large scavengers. Indeed the discovery chamber was so far from the cave entrance that such animals probably were unaware of their presence. These features and the sheer complexity of the system strongly suggested that cadavers had been deliberately taken to the chamber; implying that the deep penetration had been accomplished using fire-brand illumination. What seized the headlines was the possibility of ritual burial, although sanitary disposal or panicked refuge from predators seem equally, if not more likely.


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Lee Burger and the reconstructed skull of Homo naledi

Now yet more fossils have been reported from a separate chamber at a crawling distance about 150 m away from the original but closer to the system’s main entrance (~85 m). These add at least other 3 individuals to the H. nadeli association, with sufficient similarity to indicate that all 18 belong to H. naledi. This wealth of detail enabled the team of authors (Hawks, J. and 37 others 2017. New fossil remains of Homo naledi from the Lesedi Chamber, South Africa. eLife, v. 6, online; http://dx.doi.org/10.7554/eLife.24232) to perform a detailed comparative anatomic analysis of the species. The results are a mosaic, showing some post-cranial affinities with australopithecines, H. habilis, H.floresiensis, H. erectus, Neanderthals and anatomically modern humans, and others, such as the hands and shoulders, that are not well matched with other hominins. Their crania show a similar broad spectrum of resemblances, and as regards dentition they are distinctly primitive. They are also on the small-brained side of the hominin clade. Despite the astonishing abundance of fossil material, not a single artifact was found in the cave system, despite the apparent similarity of its hands to those of ourselves and Neanderthals.

With plenty of scope for speculation, H. nadeli remains enigmatic. The big question looming over the 2015 announcement of the species was its age, the discovers suggesting about 2 Ma, and placing on the direct line of human descent. On the same day as the fossil description there appeared a multi-method dating analysis (Dirks, P.H.G.M. and 19 others 2017. eLife, v. 6, online; http://dx.doi.org/10.7554/eLife.24231.001), which showed that with little doubt that the H. nadeli association was deposited between 236 ka and 335 ka; around the time when anatomically modern humans first emerged and stone tools had undergone a >2 Ma technological evolution. To me, the only sensible conclusion at present is that H. nadeli is another addition to the 6 species living and in some cases coexisting across the late Pleistocene world, and that expansion of ideas beyond that must await DNA analysis; a definite possibility considering the age of the fossils, their seemingly good preservation in a relatively dry cave system and the new possibility of cave soils as well as bones yielding genetic materials. The leader of the research team, Lee Berger of the University of the Witwatersrand now maintains, together with four other members of the research team, that H. nadeli may be a coelacanth-like survivor of Homo’s earliest diversification and that ‘we cannot exclude that this lineage was responsible for the production of Acheulean or Middle Stone Age tool industries’.

Related articles

Barras, C. 2017. Homo naledi is only 250,000 years old – here’s why that matters. New Scientist, 6 May 2017 Issue

Sample, I. 2017. New haul of Homo naledi bones sheds surprising light on human evolution. The Guardian, 9 May 2017

Human penis bone lost through monogamy?

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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.

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Tree-climbing australopithecines

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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.


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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

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Climatic conditions for early hominin evolution

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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.

Our ancestors parted from other humans earlier than expected

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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

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The ‘star’ hominin of South Africa

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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.

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Stone tools go even further back

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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.

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Genus Homo pushed back nearly half a million years

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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!

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Convincing, indirect evidence for early toolmakers

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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.

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Are modern humans ‘domesticated’?

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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.

Related articles: Gibbons, A. 2014. How we tamed ourselves – and became modern. Science, v. 346, p. 405-406.

Human evolution news

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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

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Human evolution: bush or basketwork?

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Analysis of DNA from ancient humans has revealed its power decisively in the last few years, and especially at the beginning of 2014 with publication of the sixth full genome of an individual who was not an anatomically modern human (Prüfer, K. and 44 others 2014. The complete genome sequence of a Neanderthal from the Altai Mountains. Nature, v. 505, p. 43-49). The newly sequenced material came from a toe bone found in the Denisova Cave in the Altai Mountains of southern Siberia; the same location made famous in 2010 by genetic evidence for unknown late hominins, the Denisovans . The bone occurred in the same layer of cave sediment, dated at 50.3 ka, which yielded the Denisovan finger bone, but from a lower sublayer. So there is no firm evidence that both groups cohabited the cave.

The genome reveals that the individual was female and related to the three Neanderthals from Croatia and another infant Neanderthal from the Caucasus, also analysed previously by Svante Pääbo’s team at the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany (Note that the toe-bone team also includes co-workers from US, Chinese, Austrian, French and Russian institutions). The closest statistical link is to the Caucasian infant Neanderthal’s DNA. Interestingly, it proved possible to demonstrate that the Siberian Neanderthal woman was from a population that was clearly inbred, her parents having been related at the level of half siblings. Her mtDNA shows that she shared a common ancestor with all 6 Neanderthals from whom mtDNA has been analysed.

Comparing genomes from the single Denisovan, the 5 Neanderthals and living humans from sub-Saharan Africans gives an estimated 550 to 765 ka time of divergence of a population leading to anatomically modern humans from the progenitors of Neanderthals and the Denisovan. The Neanderthal-Denisovan split was roughly 380 ka ago. It was already known that non-African living humans contain genetic evidence for past interbreeding with Neanderthals and that some people in Asia, Australia, Melanesia and the Philippines had acquired genes from Denisovans. More refined comparisons now show Oceanians to have 3 to 6% Denisovan make-up with Asians in general sharing 0.2%. Neanderthal to modern non-African gene flow is now estimated at between 1.5 and 2.1%, with Asians and Native Americans being at the high end.  Neanderthals and Denisovans also interbred, but only at the level of about 0.5% inheritance. However, that genetic sharing involved DNA regions known to confer aspects of immunity and sperm function, that also made their way into living non-African humans.

Since the common ancestor of Neanderthals and Denisovans left Africa long before modern humans appeared on the scene it would be expected that living Africans’ genomes would show the same level of similarity with both the now extinct groups, if all three originally shared a common ancestor. A surprising outcome from comparison of Neanderthal and Denisovan genomes with those of living sub-Saharan Africans is that there is a significant bias towards Neanderthal rather than Denisovan comparability.  There are three possibilities for this bias. After the Neanderthal-Denisovan split the former group may have continued to interbreed with the group leading to modern Africans (and indeed to modern non-Africans): that would require Neanderthal genetics to have originated in Africa before they migrated to Eurasia. Secondly, the gene flow could have been from the ancestors of modern humans to Neanderthal progenitors, making descendant Neanderthals more like modern humans. Prüfer et al. suggest that the evidence is less supportive of both and weighs towards a third possibility; that the Denisovans interbred with an unknown contemporary hominin, whose genetic make-up was yet more different from that of all three known groups of the late Pleistocene and therefore their common ancestor . This may have been Homo antecessor or possibly H. erectus who survived until as late as 20 ka in SE Asia.

Family tree of the four groups of early humans living in Eurasia 50,000 years ago and the gene flow between the groups due to interbreeding. Image credit: Kay Prüfer et al.
Family tree of the four groups of early humans living in Eurasia 50,000 years ago and the gene flow between the groups due to interbreeding. Image credit: Kay Prüfer et al.

As other commentators  on the paper (Birney, E. & Pritchard J.K. 20113. Four makes a party. Nature, v. 505, p. 32-34)  have observed, ‘…Eurasia during the late Pleistocene was an interesting place to be a hominin, with individuals of at least four quite diverged groups living, meeting and occasionally having sex.’ All this arises quite convincingly from the genetics of only 7 ancient individuals, to show that it may no longer be appropriate to consider human evolution as a tree or a bush linking permanently separated species. Either it is the history of a single, polymorphic species – remains of 1.7 Ma old Homo georgicus show clear evidence of such polymorphism – or a better metaphor for human development is an interwoven basket or twine. Rumour has it that attempts are being made to sequence an H. antecessor dated at 900 ka from Gran Dolina Cave in the Atapuerca Mountains in Northern Spain: as they say, ‘Watch this space’!

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Mitochondrial DNA from 400 thousand year old humans

Published on by zooks7771 Comment

The Sima de los Huesos (‘pit of bones’) site in the cave complex of Atapuerca in northern Spain has yielded one of the greatest assemblages of hominin bones. Well-preserved remains of at least 28 individuals date to the Middle Pleistocene (>300 ka). Anatomically the individuals have many Neanderthal-like features but also show affinities with earlier Homo heidelbergensis, who is widely considered to be the common ancestor for anatomically modern humans and Neanderthals, and perhaps also for the mysterious Denisovans. Most palaeoanthropologists have previously considered this Atapuerca group to be early Neanderthals, divergent from African lineages because they migrated to and became isolated in Europe.

English: Cranium 5 is one of the most importan...
Human cranium from the Sima de los Huesos, Atapuerca mountains (Spain). (credit: Wikipedia)

The riches of the Sima de los Huesos ossuary made it inevitable that attempts would be made to extract DNA that survived in the bones, especially as bear bones from the area had shown that mtDNA can survive more than 4300 ka. There has been an air of expectancy in hominin-evolution circles, and indeed among the wider public, since rumours emerged that the famous Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany had initiated genetic sequencing under the direction of Svante Pääbo: perhaps another ‘scoop’ to add to their reconstructing the first Neanderthal and Denisovan genomes. The news came out in the 5 December 2013 issue of Nature, albeit published on-line (Meyer, M. and 10 others 2013. A mitochondrial genome sequence of a hominin from Sima de los Huesos, Nature, v. 504; doi:10.1038/nature12788) with a discussion by Ewan Callaway (Callaway, E. 2013. Hominin DNA baffles experts Nature, v. 504, p. 16-17).

The bafflement is because the mtDNA from a femur of a 400 ka  individual does not match existing Neanderthal data as well as it does that of the Denisovan from Siberia by such a degree that the individual is an early Denisovan not a Neanderthal. Northern Spain being thousands of kilometres further west than the Denisova cave heightens the surprise.  Indeed, it may be on a lineage from an earlier hominin that did not give rise to Neanderthals. The full Neanderthal and Denisovan genomes suggest that they shared a common ancestor up to 700 ka ago. So the Sima de los Huesos individual presents several possibilities. It could be a member of an original population of migrants from Africa that occupied wide tracts of Eurasia, eventually to give rise to both Neanderthals and Denisovans. That genetic split may have arisen by the female line carrying it not surviving into populations that became Neanderthals – mtDNA is only present in the eggs of mothers. Mind you, that begs the question of who the Neanderthal females were. Another view is that the Sima de los Huesos individual may be descended from even earlier H. antecessor, whose 800 ka remains occur in a nearby cave. Pääbo’s team have even suggested that Denisovans interbred with a mysterious group: perhaps relics of the earlier H. antecessor colonists.

Established ideas of how humans emerged, based on bones alone and very few individuals to boot, are set to totter and collapse like a house of cards. Interbreeding has been cited three times from DNA data: modern human-Neanderthal; modern human-Denisovan and Denisovan with an unknown population. Will opinion converge on what seems to be obvious, that one repeatedly errant species, albeit with distinct variants, has been involved from far back in the human evolutionary journey?  There seems only one avenue to follow for an answer, which is to look for well preserved H. heidelbergensis. H. antecessor and H. erectus remains and apply ever improving techniques of genetic retrieval. Yet there is a chance that stretches of ancient DNA can be teased out of younger fossils.

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