Lucy: the australopithecine who fell to Earth?

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

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

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

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

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

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

More on hominin evolution can be found here.

Stone tools go even further back

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

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

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

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

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

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

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

Genus Homo pushed back nearly half a million years

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

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

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

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

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

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

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

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.

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…

Feet of the ancients

Cast of Footprints, Laetoli Museum
Cast of footprints, probably of Au. afrensis, from the famous trackway of Laetoli in Tanzania (Photo credit: GIRLintheCAFE)

Much of what palaeoanthropologists have surmised about the evolution of humans and their hominin forebears has come from fossils of their heads. Crania, jaws and teeth can reveal a lot about human ancestors and related species, and inevitably smart modern humans would dearly like to know how brainy and clever they were and when possible intellectual changes, such as the acquisition of language, might have taken place. But only the rest of the body gives us clues about what they did and potentially might have done. If, like Darwin, and following his lead Frederick Engels (, we believe that the single most important development was adopting an upright gait and thereby freeing the hands to manipulate the world, then fossil hands and feet are of very high importance. Yet they are among the most fragile appendages consisting of a great many separate bones, each being small enough to be transported by flowing water once soft tissues decay and a corpse falls apart. And they are easily bitten off by scavengers.  Heads are a lot bigger, heavier and robust, and being round and smooth, quite difficult for, say, a hyena or porcupine to gnaw. Moreover, disaggregated hominin foot and hand bones are not easy to recognise in fossiliferous sediments, especially if they have been scattered far and wide: the big prize being heads jaws and teeth, professional hominin hunters become expert at spotting them, but not necessarily the other 80% of skeletons.

Ardi (Ardipithecus ramidus)
Artists reconstruction of female Ardipithecus ramidus (Photo credit: Mike Licht,

So, the discovery of hominin hands or feet is a rare cause for celebration. A new partial foot has turned up in the hominin ‘bran-tub’ that is the Afar depression of NE Ethiopia (Haile-Selassie, Y. et al. 2012. A new hominin foot from Ethiopia shows multiple Pliocene bipedal adaptations. Nature, v. 483, p. 565-569) and has caused quite a stir. It is significantly different from the few other feet known from the hominin record. Moreover, it adds a sixth design to those already know, leaving out those of chimps, taken as likely to be similar to those of our shared common ancestor, Homo sapien, Neanderthals and H. erectus whose feet are much the same. While being easily distinguished from the feet of Homo species, those of australopithecines are sufficiently like them in basic morphology to suggest that Au. africanus and sediba both walked the savannas as upright as we do. But one of the earlier hominins, Ardipithecus ramidus, also from Afar but dated at more than 4 Ma, has provided an almost complete foot whose geometry , including a spayed-out, short big toe capable of grasping, almost certainly indicates that the creature was equally at home in trees as it was on the ground. Ardipithecus walked upright, but probably could not run as its gait placed the side of the foot on the ground, much like a chimpanzee, instead of proceeding heel-to-toe as we do (Lieberman, D.E. 2012. Those feet in ancient times. Nature, v. 483, p. 550-551). The new find seems similar, although better adapted for upright walking. Yet no other body parts have been found so it has not been assigned to a species, though it almost certainly represents a new one. The excitement concerns its age, which at 3.4 Ma is within the time range of Australopithecus afarensis, a family of which left the famous trackway at Laetoli in Tanzania whose foot prints strongly suggest full adaptation to human-like gait: walking, running and abandonment of partially habitual life in the trees.

It seems therefore that the multiplicity of co-existing hominins from 2 million years ago to very recently existed much further back in their evolutionary history. That raises several possibilities, among which is the possibility of repeated evolution of bipedality, hinted at by some similarities to the feet of modern gorillas in that of the newly found foot. Another implication is that simply being able to walk upright did not lead quickly to a tool-making ability because the earliest stone tools capable of cutting through meat, skin and sinew did not arise until 2.6 Ma. Like fossils of feet, those of hominin hands are extremely rare. The first crucial evidence of a hand with potential to manipulate objects delicately and with purpose is around 2 Ma, with the astonishingly well preserved hand of a young Au. sediba unearthed in South Africa ( Frustratingly, the 2.6 Ma tools are not associated with fossil hominins, and the Au. sediba skeletons had no tools.