Tag Archives: bipedalism

Homo erectus

Evolutionary theory implies that the transition from one species to another takes many generations. There’s never going to be a point at which a non-human animal gives birth to a human offspring. But on the scale we use to measure things at Logarithmic History, the time 1.8 million years ago has a good claim to be the time when human beings began. Genus Homo has been around for a while, but there are major evolutionary changes around now in the human direction. We can start with geography. It’s now that we find the first hominins outside of Africa, at least as far as Georgia in the Caucasus. The Dmanisi fossils from Georgia can probably be assigned to the new species Homo erectus, albeit somewhat shorter and smaller-brained than later erectus. Homo erectus also appears around this time in Africa.

It’s still possible that hominins got out of Africa even before erectus. Just this April, an article came out suggesting that Homo floresiensis, the famous diminutive “hobbit” from Flores island, Indonesia, has its closest affinities with African Homo habilis, raising the possibility of an earlier exit. It’s even possible that H. erectus evolved outside Africa from this earlier emigrant (so far undiscovered), and then some migrated back to Africa.

H. erectus has a bigger brain than earlier forms, and reduced jaws and teeth. And there are dramatic changes below the neck. Erectus has a body shape and size quite similar to ours. Strikingly, the changes in body form seem to be systematically related to distance running. Tendons in the feet and calves turn into springs that put a bounce in our running stride (but also rule out serious tree-climbing). Our neck gets longer and shoulders and head get more independent so we can swing arms for balance without twisting our heads from side to side. And the gluteus maximus becomes the largest muscle in the body, to prevent our bodies from toppling forward with each step. Homo erectus is the first hominin with a serious butt.

Moving from what we know to what we guess, it looks likely that Homo erectus had shifted to a new diet and a new mode of acquiring food. David Carrier argues that H. erectus was a persistence hunter, running after prey until they were exhausted. Human beings, although pretty poor sprinters, have a big advantage in distance running, in that our breathing is uncoupled from our running. This lets us run efficiently at whatever speed we choose. Most mammals, by contrast, have to breathe and run in synch, and pay a heavy price – wasting energy and overheating – for running at non-optimal speed. Bipedal dinosaurs enjoyed a similar advantage.

Like anything else in paleoanthropology, there are arguments about this. For example, fire may or may not have played a significant role at this early stage. We’ll cover some of these arguments in posts to come.

Four legs good, three legs better

Having grasping hands (and having them coordinate with the eyes) is one of the important things that distinguishes primates from other mammals. And a special version of bipedalism, which allows hands to specialize for manipulation and feet for locomotion, is one of the first things that distinguishes hominins from other primates, even before hominin brains get big.

You find the same arrangement — a pair of arms with hands and a pair of legs with feet – with most science fiction aliens. (For TV and movie science fiction this just reflects the fact that aliens, pre-CGI, were mostly played by actors made up with pointy ears or fur suits or whatever.) But there are wilder possibilities, with no Earth analog. One of the most imaginative is the Pierson’s Puppeteers invented by Larry Niven:

“…. I was fed up with humanoids. Chad Oliver in particular, an anthropologist, wrote story after story claiming that this is the only workable shape for an intelligent being. The puppeteers were my first attempt to show him a shape that could evolve to intelligence. …”


The Puppeteers’ brains are safely tucked away inside their bodies, but they have two “necks” ending in “heads” each including one eye, one mouth, and a set of “fingers” around the lips. And the body has three legs. Decapitation is bad news for a Puppeteer,  like having a limb amputated, but not a death sentence.

Even more exotic are Vernor Vinge’s “Tines.” These are dog-like aliens who have evolved a short-range ultrasonic communication system that transmits information at such a high baud rate that a pack of half a dozen separate organisms is integrated into an enduring single individual with a shared consciousness. Losing one member of the pack is more like losing a limb, or having a stroke, than like the death of an individual. The mouths of the pack act together, as coordinated as the fingers on a hand, allowing the Tines to build up a medieval level civilization. (Vinge is a computer scientist, not an evolutionary biologist, however, and he glosses over some potential problems in Tine sociobiology: “all for one and one for all” is all very well, but which member of the pack actually gets to pass on their genes when it’s time to mate?)

But we don’t have to travel to other planets to find alternatives to two hands / two feet. Elephant trunks, for example, let elephants browse while avoiding the need for a giraffe/diplodocus-style long neck. The trunks even have “fingers” (2 for African elephants, 1 for Asian elephants) that are sensitive enough to pick up a single piece of straw.

We’ll spend a lot of time on Logarithmic History asking how human beings got to be such an extraordinary species. Hands are an important part of the story, although the elephant case suggests that hands (or their near-equivalent) are merely unusual, not absolutely unique to humans and near relations.

Four legs good, two legs better

4.75-4.49 million years ago

There’s been a lot of hullabaloo in the last few days over claims that a jaw assigned to the 7.2 million year old Graecopithecus freygbergi represents the earliest known human relative after the hominin/chimp split. The jaw was found in Greece, which suggests that the split happened around the Mediterranean, rather than in Africa. (This doesn’t take anything way from the claim that Africa is the main center of later human evolution, up to 2 million years ago, which would have taken place when Graecopithecus’ descendants migrated to Africa).

All this needs to be taken fairly skeptically: a mandible with one tooth isn’t overwhelming evidence.

Here I cover some of what we know about the evolution of bipedalism. This is mostly in the context of Ardipithecus ramidus, but I have some suggestions at the end of the post about how the Graecopithecus find might be relevant.

ardipithecusWith Ardipithecus ramidus (4.5 million years ago) we have the strongest evidence so far that hominins have adopted bipedalism. Earlier fossils, including the earlier Ardipithecus kadabba, are too fragmentary to be very sure. Even “Ardi” was not bipedal quite the way we are. She had a somewhat diverging big toe, and arms and hands well-adapted for suspension, suggesting she was bipedal on the ground, but still spent a lot of time in trees.

We’ve seen bipedalism before on Logarithmic History. Bipedalism allowed ancestral dinosaurs to overcome the tight coupling of locomotion and respiration that prevents sprawling lizards from breathing while they run. But human bipedalism, with no counterbalancing tail, is different. As far as we know it evolved only once in the history of life (or maybe twice if Oreopithecus was bipedal).

In part human bipedalism is related to the general primate phenomenon of having grasping hands. Both humans and macaques, for example, devote separate areas of the brain (within the somato-sensory cortex, specifically) to each finger on each hand. Brain areas for the toes, by contrast, are more smooshed together.

Human bipedalism is more specifically related to tradeoffs in locomotion in  great apes. Other great apes pay a big price for being the largest animals well-adapted for moving around under and among branches: great ape locomotion on the ground is particularly inefficient. Chimpanzees spend several times as much energy knuckle-walking on all fours as you would expect based on comparisons to similar sized quadrupedal mammals. Remarkably, chimpanzees don’t take any more energy walking on two legs than they do walking on all fours, even though they aren’t at all well-adapted to bipedalism. Humans by contrast take a little less energy to walk around than a same-size four-legged mammal, and way less than a chimp.

That said, efficiency isn’t everything. Human beings are lousy at sprinting – try outsprinting your dog, or a squirrel for that matter. Our top speed is less than half that of a chimpanzee.

So there’s a tradeoff between the efficiency advantages of bipedalism (at least compared to knuckle walking), and the loss of speed. It may be that bipedalism evolved initially in an environment where predation pressure wasn’t very intense, and the need for speed was not as great. This argument has been made for Oreopithecus, living on an island in the Mediterranean. Perhaps Graecopithecus initially enjoyed a similar isolation, and freedom from predation, associated in some way with the drying and flooding of the Mediterranean.


7.04-6.65 million years ago

Sahelanthropus is a 7-6 million year old species whose remains have been found in Chad. “Toumaï” (“hope of life” in the Daza language) is the nickname for one individual, represented by a fairly complete skull. Otherwise Sahelanthropus is known from some jaws and teeth.


One of the things that distinguishes hominins (the human line) from great apes is that the front teeth – canines and incisors – are reduced. (Back teeth are another story. They stay big, or even get bigger, for a long time.) By this standard, Sahelanthropus looks like an early hominin. It’s got reduced incisors and canines and a short mid-face. And depending on who you talk to, it might or might not have been bipedal, although the foramen magnum (where the spine enters the skull) was maybe not positioned to balance the skull on top of the spine. Not that there was much brain inside the skull: the cranial capacity (maybe 360 cc) is at the low end for a chimp.

So Sahelanthropus could be one of the very first species related to us after the chimp/human split. Chad, where Sahelanthropus was found, is a long way from East Africa, where most other hominins have been found, which suggests there may have been a profusion of hominins across Africa, waiting to be discovered.

The face in the Logarithmic History banner for the month of May is a Sahelanthropus.

Stories of O

There were several interesting apes around 9 million years ago.

Ouranopithecus (sometimes called Graecopithecus) could fit almost anywhere on the great ape tree. Some people think it looks like an Asian great ape. Others think it looks more like the African great apes, maybe gorillas especially. This would be consistent with African great apes evolving outside Africa, then moving back. But maybe it only looks gorilla-like because it’s pretty big. In any case, we should expect that at this point different lineages of great ape will be hard to tell apart; they have only recently split.

But the award for weird goes to Oreopithecus. (If you think that sounds like a good species name for the Cookie Monster – you’re not the first person to have that thought.) From 9 to 6.5 million years ago, Tuscany and Sardinia were part of an island chain. Oreopithecus evolved there in relative isolation. It may be important that big predators weren’t abundant. Oreopithecus spent significant time arm-hanging. It’s when it was on the ground that things get strange. O’s big toe stuck out sideways at an extreme angle, so its foot was tripod-like, with a triangle formed by heel, little toes, and big toe. It’s possible that O was a biped, walking around on its two tripod feet when it was down on the ground. (Although measurements on the lower spine published in 2013 cast doubt on the biped theory.)

oreopithecus foot

There’s no reason to think Oreopithecus was close to the human line. If it’s true that it was a biped, this suggests that several versions of bipedalism evolved independently as solutions to the problem of how does an arm-hanger get around on the ground – knuckle-walking being another solution.

Biped or not, Oreopithecus was probably pretty awkward on the ground. When a land bridge reconnected O’s island chain with the mainland, predators arrived and Oreopithecus went extinct.

Oak ape

13.8-13 million years ago

We’ve known about Dryopithecus (“Oak ape”) for a while. The first specimen was found in France in 1856. They’ve since been found all over Europe, from Spain to Hungary. There are about 4 species of Dryopithecus, roughly chimp-sized.

The various Dryopithecuses are interesting because they look like good candidates for being somewhere in the ancestry of the great apes, Asian and/or African. (They could just as easily be on a side branch though. It’s easier to tell whether something is a close or distant relative than to figure out whether it’s an ancestor or a collateral.) Dryopithecus had made the move to suspensory brachiation – hanging from branches – and had the freely-rotating shoulders, long arms, and strong hands you need for that. But it wasn’t specialized for knuckle walking like a gorilla or a chimpanzee. This could mean it spent almost all its time in trees. Later on (10 Mya) at Rudabanya, Hungary, we find Dryopithecus living in a moist subtropical forest, among fauna including Miocene versions of pigs, horses, rhinos, and elephants. The fauna also included predators: the lynx-like Sansanosmilus, weighing about 170 lbs, and “bear-dogs” up to five feet long. So maybe up in the trees all day was the safest place to be.

The evolutionary position of Dryopithecus matters for one of the big unsettled questions in human evolution: did bipedal human ancestors evolved directly from a tree-dweller like Dryopithecus, or were human ancestors chimp-like semi-terrestrial knuckle walkers before they started standing upright? Many scenarios for human evolution start with something that looked like a chimp and maybe lived in chimp-style social groups (dominated by gangs of males ready to rumble with neighboring gangs), but there’s a lot of guesswork in this.

David Begum has recently written a book, The Real Planet of the Apes, covering this period in the evolution of human ancestors and collaterals. Begum argues that Dryopithecus was not just a great ape (now generally accepted) but close to the ancestry of present-day African great apes (i.e. gorillas, chimps (genus Pan), and humans, as opposed to Asian great apes – orangutans (genus Pongo)). This implies that African great apes may have originally evolved in Eurasia, and migrated back to Africa. Here’s one possible evolutionary tree, from Begum’s book:

dryopithecus tree

Apes, on the road to great

16.3-15.5 million years ago

Teeth are tough, and survive better than most bones. We can recognize apes by their teeth: ape and human molars have 5 cusps that form a distinctive Y pattern. Early Miocene apes like Proconsul already had this pattern. They had also already lost their tails. But in other respects they were more like monkeys than living great apes. They walked on their palms like monkeys, meaning they mostly walked on top of branches, instead of hanging underneath them.

How we get to modern great apes is somewhat mysterious. Apes may have left Africa for Europe and Asia as early as 16 million years ago, or maybe more like 14 Mya. A variety of great apes develop in Asia, although orangutans are now the only survivors. But we’re not sure whether the ancestors of African great apes are apes that stayed in Africa, or whether they’re apes that developed more modern features in Eurasia and then migrated back to Africa.

The various genera of great apes all make some kind of compromise between walking and hanging from branches. When orangutans are on the ground (which is not very often), they walk on the edges of their hands. Chimpanzees and gorillas walk on the knuckles of their hands. And of course humans walk on their hind legs. These are all pretty unusual ways to get around.

It would be nice to know whether human ancestors went through a knuckle walking phase. African fossils are skimpy for this period, and likely to remain so. Maybe genetics will have something to tell us about whether chimp ancestors took to knuckle walking before or after they spit from human ancestors.