Are modern humans ‘domesticated’?

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

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

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

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

Human evolution news

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

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

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

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

More on human evolution here and here

Human evolution: bush or basketwork?

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

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.

Early humans could probably kill at a distance

It is always refreshing when physical anthropologists perform experiments as well as pondering on bones. It turns out that examining the bio-mechanics of college baseball players can provide useful clues about where in fossil anatomy to look for signs of potential big-game hunters. Anyone who can hurl a baseball, or one of the smaller but much harder red ones preferred by non-Americans, at speeds exceeding 100 kph could in all likelihood bring down a substantial prey animal with a rock and even more so with a spear. At the heart of an important examination of what our forebears might have done to get a meaty meal (Roach, N.T. et al. 2013. Elastic energy storage in the shoulder and the evolution of high-speed throwing in Homo. Nature, v. 498, p. 483-486) is a US-Indian team’s sophisticated study of college baseball players’ throwing action using high-speed video, radar and precise timing techniques.

Matt Kata throwing
Matt Kata throwing for the Houston Astros (Photo credit: Wikipedia)

It seems that there are several physiological phases in demon ball throwing: rotation of the torso; rotation flexion and extension of the shoulder; flexion and extension of the elbow; and wrist extension. All of these contribute to acceleration of the ball before release. While the thrower steps forward the arm is cocked so that ligaments, tendons and muscles crossing the shoulder become stretched, thereby storing energy. During the acceleration phase the bend in the elbow is snapped straight adding yet more power. Readers should note the difference between this action and that of a bowler in cricket, where the elbow snap is banned on pain of severe penalty and public humiliation of the bowler who ‘chucks’. Since a fast bowler also adds energy by running into the crease, this is a humanitarian aspect of the Rules of cricket, although several legal West Indian bowlers of the past 40 years are still remembered with terror by their batsmen contemporaries. No such stricture is placed on the baseball pitcher who has no run-up.

These observations focus attention on the structure of shoulder and elbow, yielding a robust means of predicting how fast throwers with different configurations may have thrown objects. Chimpanzees make poor players of ball games, although they will throw the odd stick, but just for aggressive show. The same goes for the earliest hominins for which we have suitable fossil material: australopithecines may occasionally have eaten carrion but they couldn’t throw rocks or spears with enough force to bring down anything and their throwing range would have been pathetic. Not so Homo erectus! They were well equipped in the hurling department and could, were they so inclined, have hunted equally as well as modern humans. Interestingly, earlier hominins had some of the physiological necessities of decent throwing, but not all of them. So it seems that the full combination emerged in the evolution of our own genus around 2 Ma ago,

This is in contrast to a view held by some anthropologists, such as Christopher Boehm of the University of Southern California, that big game hunting using projectile weapons emerged only with anatomically modern humans after 250 ka, and most likely only reached its acme 45 ka ago. That assumption, at least by Boehm, is central to notions of how social activities centred on meat sharing may have helped evolve morals, such as altruism and shame (see Boehm, C. 2012. Moral Origins: The Evolution of Virtue, Altruism and Shame. Basic Books, New York). That H.erectus would have been able to harness sufficient energy to kill at a distance casts doubt on such assertions. Mere foraging does not require throwing-capable physiology, so how it evolved in early humans with neither the inclination nor bodies to at least begin throwing projectiles at potential prey is something that school might consider.

 

Could the Toba eruption have affected migrating humans?

Around 73 thousand years ago a supervolcano in Sumatra erupted on a scale unprecedented in the last 2 million years. It left a 100 by 30 km elliptical caldera now occupied by Lake Toba, and explosively ejected 2800 of magma, about 800  km3 falling as ash as far afield as the Greenland ice cap. Although ice-core records show little if any sign of associated climate change in polar regions, the vast amount of ash and sulfate aerosols blasted into the stratosphere must have had some ‘global winter’ effect. Large areas of South Asia were blanketed by thick beds of ash. Human migration from Africa into Eurasia was probably underway at the time, indeed stone tools are found directly beneath and above the Toba ash in southern India and Malaysia. Some palaeoanthropologists have seen the stresses imposed by the Toba eruption as possible means of reducing the entire human population to a mere few thousand: a genetic ‘bottleneck’ that could have led to rapid evolution among surviving generations that may have shaped changes in human behaviour and culture.

Landsat image of Lake Toba, the largest volcan...
Landsat image of Lake Toba, the largest volcanic crater lake in the world. (credit: Wikipedia)

There is a widening range of views on the climate changes that may have followed Toba. It has even been suggested that global mean surface temperature fell by as much as 10°C (Robock, A. et al. 2009. Did the Toba volcanic eruption of ∼74 ka B.P. produce widespread glaciation? Journal of Geophysical Research: Atmospheres, v. 114, DOI: 10.1029/2008JD011652), although not so far as to produce a worldwide glacial surge but sufficient to devastate vegetation. This bleak look back to a critical point in human affairs resulted from modeling of the effects of a global reflective cloud of ash and sulfate. A later modeling study factored in particle and aerosol sizes (Timmreck, C. et al. 2010. Aerosol size confines climate response to volcanic super-eruptions. Geophysical Research Letters, v. 37, doi:10.1029/2010GL045464) to give a less dramatic, but still severe maximum global cooling due to Toba of ~3.5°C.

The focus has now shifted from modelling to a more direct look at the environmental effects of the Toba super-eruption, preserved in sediments beneath Lake Malawi in southern Africa (Lane, C.S. et al. 2013. Ash from the Toba supereruption in Lake Malawi shows no volcanic winter in East Africa at 75 ka. Proceedings of the National Academy of Science, v. 110, doi/10.1073/pnas.1301474110). The sediments contain a thin ash layer that is very different from those produced by East African Rift volcanism but chemically and texturally similar to the Toba ash from the Indian Ocean and India. The sediments, diatom fossils and chemical biomarkers immediately above the ash show little sign of a significant temperature fall. At most it records a 1.5°C fall, and the authors conclude little chance of a human genetic bottleneck among Africans living at the time.

There is clearly a conflict between results of modeling and real-world climatic data, which is interesting in its own right. But the Malawi findings do not rule out ‘bottlenecks’ resulting from severe stress in South Asia where the ash itself would have severely affected game and vegetation for long enough to face migrating human bands with the prospect of starvation. Obviously, some survived to move on and to leave their tools behind on top of the Toba Ash.

Hybridisation in human evolution

A press release from the Max Planck Institute for Evolutionary Anthropology in Leipzig, Germany, announces the completion of the most precise genome from a third Neanderthal individual. For the first time it is possible to distinguish copies of the genes inherited by the individual from both parents. The data release coincided with a review of genetic evidence for interbreeding between early Homo sapiens and other species.

The full item can be read at Earth-logs in the Human evolution and migrations archive for 2013

https://upload.wikimedia.org/wikipedia/commons/thumb/4/41/Spreading_homo_sapiens.svg/640px-Spreading_homo_sapiens.svg.png

Modern human migration out of and within Africa relative to the domains of coeval archaic humans 1 = modern humans 2 = Neanderthals 3 = other archaic humans (credit: Wikipedia)

Further support for Homo floresiensis (the ‘hobbit’)

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

When they were first discovered in Liang Bua cave on the Indonesian island of Flores diminutive hominin remains sparked off a heated debate. Part of the reason for dispute was the age of the deposit in which they were found (18 to 850 ka), so young that it indicated possible cohabitation on the island with anatomically modern humans. On the one hand, the finders claimed that they represented a previously unknown hominin species. Other specialists considered that the tiny size (adults no taller than about a metre with brain capacity around that of australopithecines) indicated some congenital  dwarfism.

Homo floresiensis (the "Hobbit")
Homo floresiensis skull (credit: Wikipedia)

In the 9 years since the remains came to light, several anatomically features have been cited to support the view of a distinct hominin species: their lack of a chin and different arm and shoulder anatomy, which H. floresiensis shares with H. erectus and H. georgicus. The fossils are associated with simple stone tools and bones of a variety of prey animals that show cut marks and charring, suggesting that cooking was part of these hominins’ lifestyle; despite having small brains they were not unintelligent.

Substantial remains of nine or more individuals have been unearthed so that anatomical detail is almost complete. In 2007 details were published of three well-preserved wrist bones from the original find. They too were sufficiently different from modern and Neanderthal humans to warrant confirmation that H. floresiensis is indeed a distinct hominin species. Further work on wrist bones from other individuals has now more or less put the seal on this identity (Orr, C.M. et al. 2013. New wrist bones of Homo floresiensis from Liang Bua (Flores, Indonesia). Journal of Human Evolution, v. 64, p. 109-129), the authors  concluding that ‘The pattern of morphology … supports H. floresiensis as a valid taxon and refutes the hypothesis that these specimens represent modern humans with some kind of pathology or growth disturbance’. They take matters further by suggesting that their lineage was established before divergence of modern humans and Neanderthals. As with the shoulder morphology that of their wrists would have somewhat hindered tool-making dexterity, but nonetheless they did make tools.

Hominin evolution becoming a thicket

Scientific American is renowned for its eminently readable reviews of both emerging and perennial topics. Its February 2013 issue takes on one that is guaranteed to run and run; the evolutionary course that produced us (Harman, K. 2013. Shattered ancestry. Scientific American, v. 308 (February 2013), p. 36-43). Since its launch Earth Pages has covered much of the new science in the field but did not anticipate the depth of the stir towards which it has led.

Australopithecus afarensis reconstruction
Australopithecus afarensis reconstruction (credit: Wikipedia)

For a decade it has become increasingly clear that anatomically modern humans are unique in one respect: they are the first species in perhaps 4 million years to be the sole extant member of the cladistic tribe Hominini. As recently as 30 ka Homo sapiens shared the planet with Neanderthals, Denisovans, H. erectus and H. floresiensis. At the time the genus Homo emerged around 2.0-2.5 Ma ago there were at least four other fossil groups that shared the major characteristic of upright gait, all australopithecines in ‘robust’ and ‘gracile’ guises.

As time goes by there will likely be more fossil discoveries that show important anatomical signs of other novel evolutionary divergence, which therefore warrant new species. Pliocene-Pleistocene time is becoming crowded, and the more diversity in its fossil record the less likely it is that some clear evolutionary pathways can be devised to explain just what was going on. Katherine Harmon of Scientific American’s editorial team touches on the thorny issues of upright walking and gait, tree climbing, precise use of the fingers and thumb, and brain size that are raised by 22 species; 2 living and 20 extinct.

Genetics clearly indicates that our nearest living relatives belong to two species in the genus Pan(chimpanzees and bonobos). It has been generally assumed that the common ancestor of this extant kinship some 8 Ma back was chimp-like, and that evolutionary divergence from its habits and anatomy produced the growing ‘bramble patch’ of hominin evolution. That assumption is based on the principle of parsimony, i.e. the simplest view of the evidence – what there is now and fragments from the past eight million years. The trouble is there is a dearth of fossils that can be said to be en route to chimps in some way.

In fact today’s chimps and bonobos are more or less restricted to clambering in tropical forest habitats, for which they are well-adapted. Maybe they are the survivors of evolutionary vagaries just as complex as those leading to us. For one thing, almost embarrassingly, their brain size is substantially larger than those of quite a few fossil hominins; and why not? How they behave socially may possibly have arisen as part of their specialisation too, of which more shortly. Our big difference from them is being supreme generalists, as well as consciousness.

All the fossils classed as hominins show some signs of being able to walk upright, classically the forward position of the foramen magnum at the base of the skull where it joins to the backbone, but in some cases merely the geometry of the hip joint to the pelvis for that is all that has been found. Yet that anatomical likelihood glosses over the vital detail of the actual gait – heel-to-toe like us (Australopithecus afarensis),  on the outside edge of the foot akin to chimps (Ardepithecus ramidus) or differently again but possible as efficient as us (Au. sediba). Then there is the matter of arboreal abilities: chimps are masters despite their bulk, but every hominin whose foot bones have been found shows some evidence of grasping with the big toe. Indeed humans are pretty nimble climbers but do not brachiate from branch to branch.

As regards the hands, an interesting point is that while chimpish knuckle walking is not seen in fossils, Ardipithecus probably could walk on all fours with hands flat on the ground but had fingers quite capable of precise manipulation, an ability shown spectacularly well by 2 Ma old Au. sediba. Upright walking may have evolved more than once, and it is even possible that chimps evolved specifically for climbing in forestlands, their highly adapted grasping hands only capable of knuckle walking on the ground.

English: Fossil of Oreopithecus bambolii, an e...
Oreopithecus bambolii from the Upper Miocene of northern Italy(credit: Wikipedia)

The late-Miocene of Africa – the likely time range for the Pan-Homo common ancestor – is a fossil desert as regards primates. Yet its Italian equivalent has yielded a fascinating and well-preserved creature; Oreopithecus bambolii has skeletal features compatible with an upright posture and bipedal locomotion. Until the African Miocene yields something more appropriate, Oreopithecus is a candidate for a common ancestor, and interesting in another respect. Its dentition does not include prominent canine teeth that in the predominantly vegetarian, though occasionally carnivorous, Pan species serve well in their aggression-based, hierarchical social systems, as they do in the even more spectacular baboons.

Christopher Boehm, primate behaviouralist cum anthropologist, in his recent book Moral Origins (2012 Basic Books, ISBN-13: 978-0465020485) uses the principle of parsimony to reconstruct the social system of the Miocene Pan-Homo common ancestor from those of chimps and surviving human hunter-gatherers. His thesis is that it was centred on the hierarchical dominance of ‘alpha’ males, as is that of chimps. Prolonged social selection in hominin evolution largely tempered such a ‘Big Man’ tendency through a variety of strategies directed by majorities. Social punishments, including capital punishment, evolved to combat free-loading, theft and individual dominance in favour of cooperative egalitarianism. Such measures developed increasingly conscious self-suppression of such traits that eventually manifested themselves as what we now regard as human morals. Boehm considers that this psychological trend in evolution accelerated once Homo sapiens began hunting of large prey animals that added substantially to diet.

Aggressive male chimpanzee (Credit: Daily Mail)
Aggressive male chimpanzee (Credit: Daily Mail)

There is a major problem for this view: like Oreopithecus every well-preserved hominin species, even the earliest Sahelanthropus tchadensis, do not have prominent canines irrespective of whether they show evidence of at least partial meat-eating or pure vegetarianism. For some species with many fossil members, such as Au. afarensis, there are signs of sexual dimorphism – larger males than females – but that does not necessarily signify hierarchical social behaviour. With the appearance of H. erectus that difference wanes to the present slight differences between modern male and female humans.

Scrum
Agressive male humans, note gumshields (credit: John_Scone via Flickr)

If it is valid – and who knows? – for morphology to give clues to social behaviour, then it is equally likely that the beginnings of the hominin evolutionary thicket may well have involved a trend in social behaviour towards cooperative action; 8 million years ago. For generally small, gracile creatures with habits no more threatening to the big predators of the African savannahs that that of the porcupine, there would have been a powerful selection pressure towards a united front. Of course, in the last ten thousand years since the shift to economic strategies based on storable surpluses and their expropriation, hierarchical social systems with violence at their heart emerged among modern humans. Judging by the body shapes and dentition of extant ‘alphas’, as in capital’s boardrooms and among the frontbenchers at Westminster, anthropology clearly is in need of some refinement…

Toba ash and calibrating the Pleistocene record

Landsat image of Lake Toba, the largest volcan...
Landsat image of the Lake Toba caldera, Sumatra (credit: Wikipedia)

The largest volcanic catastrophe during the evolution of humans formed the huge caldera at Lake Toba near the Equator in Sumatra about 70 thousand years ago. Explosive action erupted 2800 cubic kilometres of magma, of which 800 km3 was deposited as thick ash across most of South Asia and the northern Indian Ocean. Sulfates derived from the gas emissions by Toba form clear ‘spikes’ in ice cores from both Greenland and Antarctica. Its effects were global through the mixing of sulfate aerosols in the stratosphere of both hemispheres, encouraged by its position close to the Equator. By reflecting incoming solar energy the aerosols resulted in a century-long 10°C fall in temperature over the Greenland ice cap. Such global cooling almost certainly affected anatomically modern humans, but it is possible that in South Asia Toba had an even more devastating effect.

Jwalapuram
The Toba ash at the Jwalapuram excavations in South India(Photo credit: Sanjay P. K. via Flickr)

At several sites in the Indian state of Tamil Nadu and in Malaysia Toba ash has buried artifacts that arguably may have been made by the earliest modern emigrants from Africa. Immediately above the ash are yet more tools that suggest humans did survive the eruption. Palaeoanthropologists have argued that the stress of Toba’s environmental effects on all hominins living at the time may have resulted in population crashes from which only the fittest individuals emerged. Major evolutionary changes have been ascribed to ‘bottlenecks’ of that kind to result in changes in human behaviour detectable from the archaeological record, such as the creation of completely new kinds of tools, art and language.  However, recent finds in Africa suggest that many such shifts are much older than Toba.

Perhaps Toba’s greatest contribution to palaeoanthropology is that it is an easily recognised event in the geological record, but compared with its sulfate spike in the Greenland ice core at ~71 ka the existing radiometric dates have uncertainties of several thousand years. Using the latest 40Ar/39Ar dating methods on fresh crystals of sanidine (volcanic K-feldspar) from new excavations in Malaysia these uncertainties have been reduced significantly (Storey, M. et al. 2012. Astronomically calibrated 40Ar/39Ar age for the Toba supereruption and global synchronization of late Quaternary records. Proceedings of the National Academy of Sciences, v. 109, p. 19684-18688 ). The sulfate peak and the ash can now be attributed to an age of 73.88 ± 0.32 ka; better than a golden spike in Late Pleistocene stratigraphy. The ice-cores have a check on chronology just beyond the limit of counting annual layering, as do ocean sediment cores for a time older than 14C can ever achieve. Toba now links too with events recorded by the precise U-Th series dating of cave deposits