Bottom-up apes

According to the latest research, chimpanzees recognize other chimps not just by their faces but by their butts. 

Which raises the further question: Why are chimps down on all fours, while we’re not? This and related matters are subjects of a major recent review, “Fossil apes and human evolution,” which contrasts “top-down” and “bottom-up” approaches to understanding comparative ape and human ancestry. The authors write “top down approaches have relied on living apes (especially chimpanzees) to reconstruct [human] origins.” By contrast, “bottom-up” approaches pay more attention to the fossil record. The review brings to the fore something that’s been brewing for a while: fossil apes from the mid to late Miocene, leading up to the time that gorillas, chimps, and humans go their separate ways, are a varied bunch, and the last common ancestors of gorillas, chimpanzees, and humans, and of chimpanzees and humans, may be creatures that didn’t look all that much like any of the three. More specifically, the common ancestors may have been “orthograde,” standing upright and using both feet and hands to clamber vertically through trees. From ancestors like this, gorillas and chimps may have evolved similar innovations in parallel, getting bigger, evolving longer arms, larger palms, and shorter backs – all of which helped them as big animals to get around in trees, but also led them to take a more crouching posture, and to adopt the expedient of knuckle-walking on the ground (and hence to spend a lot of time looking at one another’s butts). The ancestors of humans, on this account, followed a different path, adopting bipedalism – not such a big step, already a big part of their postural repertoire – when walking on the ground. 

In some ways, gorillas and chimps seem to be caught in a “specialization trap.” This seems to show up in the energetics of locomotion. For a human being, it takes about 50 kilocalories to walk a mile (or about 30 kcal to walk a kilometer). This is about the same as you’d expect for a standard mammalian quadruped of our size. But for chimpanzees, committed to knuckle-walking, the energy cost is about double.

Stories of O

9.00 – 8.51 million years ago

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 is just one find showing that apes early in the Late Miocene, well before our ancestors parted ways with chimpanzees, were experimenting with a lot of different types of locomotion, possibly including versions of bipedalism. Many of these experiments were taking place in Europe. (A few more examples: Danuvius guggnemosi and  Rudapithecus hungaricus.)

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, perhaps in consequence, Oreopithecus went extinct.

Oak ape

12.6 – 12.0 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. As any good cladist will tell you, it’s easier to say 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 million years ago) 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) consistent with reconstructions of ancestral multi-male/multi-female groups among monkeys and apes. But there’s a lot of guesswork in this; probably we were never chimps.

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:

It’s a small world after all

475 – 450 thousand years ago

The story of human origins is partly a story of Big Things like The Taming of Fire and  The Dawn of Speech. But it’s also the story of some odd byways and quiddities. A nice introduction to some of these is Chip Walter’s book Thumbs, Toes, and Tears: And Other Traits That Make Us Human. (His more recent Last Ape Standing is good too.) Walters considers funny bits of anatomy like our unique big toes and thumbs, and funny bits of behavior like our habits of laughing, weeping, and kissing. Toes and thumbs fossilize, but behaviors can be hard to date, evolutionarily. Presumably these behaviors appeared sometime before modern humans evolved and spread, so let’s pick today’s date. It’s also hard to figure out the exact evolutionary rationale for some of these behaviors. Humor, for example, is not a simple phenomenon: intellectually appreciating a joke, actually finding it funny and enjoying it, and finally laughing, each involve separate areas of the brain.

Laughter, specifically, is a minor human oddity that sheds an interesting light on some big events in human evolution. Robert Provine, a leading laughter researcher, spells out the argument in “Curious Behavior: Yawning, Laughing, Hiccuping, and Beyond.” Chimpanzees have a kind of laugh, a modified vocalized panting synchronized with inhalation and exhalation. Presumably laughter first resulted when panting-during-play evolved into a play signal. But the short bursts of human laughter go further, having freed themselves from synchrony with the inhalation/exhalation cycle. Laughter, in other words, is just one instance of the more general phenomenon of humans having separate controls for vocalization and for respiration. Interestingly, the most prominent examples of complex vocalization – songbirds and some other birds, whales, bats, and humans – are all found in non-quadrupeds. In quadrupeds, breathing is tightly coupled with locomotion: lungs need to be full to stiffen the thorax when the forelimbs hit the ground. Giving up quadrupedalism seems to have allowed for an “adaptive release” in the evolution of vocal abilities in a number of unrelated lineages. So the study of laughter (and other vocalizations) suggests that two key human adaptations – bipedalism and spoken language – are more closely linked than one might have expected.

Another and overlapping set of human particularities involves facial expressions of the emotions. Darwin got a whole book out of this. He concluded (admittedly based on somewhat anecdotal evidence) that different emotional expressions are largely innate. It’s an interesting illustration of his ability to reason from small facts to large conclusions that he also drew a big conclusion about human evolution from this. In Darwin’s day, there were scientists who believed that different human races had evolved from very different prehuman progenitors: one prehuman species giving rise to Europeans, another to Africans, and so on. But Darwin reasoned that the very close similarity in facial expressions (and he had traveled a lot, and witnessed a lot of expressions in a lot of places) and the very similar emotional makeup of humans around the world was evidence that human populations shared a fairly recent common ancestry. Here as in several other cases, a mixture of close reasoning and sheer luck led Darwin to the correct conclusion about evolution long before there was much solid evidence.

Darwin’s work on emotions was neglected for most of the twentieth century by anthropologists favoring a blank slate view of human behavior, but was eventually largely vindicated by a number of researchers, notably Paul Ekman. There is now good evidence for six basic facially expressed emotions: Fear, Disgust, Joy, Anger, Sadness, and Surprise.

If you’re a movie watcher, this list may seem familiar. These emotions (all except for Surprise) are all depicted as little homunculi living inside the head of an 11 year old girl in the animated feature “Inside Out.” Somebody at Pixar Studios knew their Ekman.

So the sappy song is right: There is just one moon and one golden sun, and a smile means friendship to everyone.

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 (or maybe a bit earlier in South Africa, according to recent findings.)

It’s still possible that hominins got out of Africa even before erectus.  A recent article came out suggests 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.

Lucy

AL (Afar locality)-288-1 is better known as Lucy: probably the most famous individual hominin fossil. Her skeleton is particularly complete, and demonstrated the existence of a previously unknown species, Australopithecus afarensis.

When she was discovered in 1974, she pushed back the hominin fossil record by over a million years. A. afarensis looked like a plausible ancestor to all the later hominins. (Things have gotten complicated since then.) “The first family” (AL 333) was discovered just a year later: bones of at least 13 individuals, a mixture of adults and juveniles, maybe all victims of a flash flood. This helped to fill in knowledge of individual variation among A. afarensis. And subsequent findings have documented the species over a million year span from 4-3 million years ago. At the same time, later discoveries have also suggested that there were likely multiple hominid species around in Lucy’s day.

Also: Her pelvis is basin-shaped, and she stood with her legs under her body, not rocking from side to side. All this is like a human, not a chimpanzee. So she was certainly bipedal, but there are arguments about just how bipedal. The initial view was that she was an efficient walker (although not much of a runner: that comes later). But another school of thought points out that she’s got very long arms, and curved finger and toe bones, suggesting she spent a lot of time in trees. Her long toes might have made her an inefficient biped (a view derided by members of the Lucy-the-proficient-walker school as the clown shoe hypothesis).

Lucy got her name from this song, played on a camp loudspeaker the day she was discovered. John Lennon, who wrote the lyrics, always denied that Lucy in the Sky with Diamonds, in spite of the initials, had anything to do with LSD. He said that it was based on picture that his son Julian (then four years old) made in school, with some Lewis Carroll thrown in.

‘I could tell you my adventures—beginning from this morning,’ said Alice a little timidly; ‘but it’s no use going back to yesterday, because I was a different person then.’

Alice in Wonderland Lewis Carroll

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.

Even more exotic are octopuses (octopi, octopuses) – easily the smartest invertebrates, solitary creatures with little social life, but very handy with their tentacles. Peter Godfrey-Smith, philosopher and scuba diver gives his take here.

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 and octopus cases suggest that hands (or their near-equivalent) are merely unusual, not absolutely unique to humans and near relations.

Ardi

4.4 Mya

We’ve got earlier hominins – Sahelanthropus, Orrorin tugensis, even an earlier species of Ardipithecus – but Ardi (her nickname, her species is Ardipithecus ramidus) stands out because she left us an exceptionally complete skeleton. 

And she helped to upend a lot of theories based on drawing a straight line between modern chimpanzees and later australopithecines. She was a biped. Her legs tucked under her pelvis, with a short lower spine, and a broad pelvis (all closer to modern humans than to chimps). Her outer foot was adapted for walking on the ground. But her toes were long-ish. And her big toe stood way off to the side: well-adapted for grasping branches. So when she was walking on the ground, she had to use her second toe to push off, instead of – like you and me – her big toe.

Ardi really shifted people to realizing that gorillas and chimps are specialized beasts, specialized in being big apes adept at climbing trees, grasping branches, swinging around, and pulling themselves up by their arms, but paying a price in inefficient knuckle walking when down on the ground. Ardi was taking up a different specialization, still a tree climber (her habitat was woodland, more than savannah), but ambling around (not quite striding yet) two-legged on the ground. 

So the common ancestor of chimps and humans, back before Ardi, apparently didn’t look all that much like either. In some respects she might have been somewhat closer to humans. She might even have been a sometime biped on the ground.

For a great recent summary, including lots of information about Ardi, and also about the politics, academic and otherwise, of digging fossils, check out Fossil Men: The Quest for the Oldest Skeleton and the Origins of Humankind.

Toumaï

7.04 – 6.67 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) would be 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.

Bottom-up apes

According to the latest research, chimpanzees recognize other chimps not just by their faces but by their butts. 

Which raises the further question: Why are chimps down on all fours, while we’re not? This and related matters are subjects of a major recent review, “Fossil apes and human evolution,” which contrasts “top-down” and “bottom-up” approaches to understanding comparative ape and human ancestry. The authors write “top down approaches have relied on living apes (especially chimpanzees) to reconstruct [human] origins.” By contrast, “bottom-up” approaches pay more attention to the fossil record. The review brings to the fore something that’s been brewing for a while: fossil apes from the mid to late Miocene, leading up to the time that gorillas, chimps, and humans go their separate ways, are a varied bunch, and the last common ancestors of gorillas, chimpanzees, and humans, and of chimpanzees and humans, may be creatures that didn’t look all that much like any of the three. More specifically, the common ancestors may have been “orthograde,” standing upright and using both feet and hands to clamber vertically through trees. From ancestors like this, gorillas and chimps may have evolved similar innovations in parallel, getting bigger, evolving longer arms, larger palms, and shorter backs – all of which helped them as big animals to get around in trees, but also led them to take a more crouching posture, and to adopt the expedient of knuckle-walking on the ground (and hence to spend a lot of time looking at one another’s butts). The ancestors of humans, on this account, followed a different path, adopting bipedalism – not such a big step, already a big part of their postural repertoire – when walking on the ground. 

In some ways, gorillas and chimps seem to be caught in a “specialization trap.” This seems to show up in the energetics of locomotion. For a human being, it takes about 50 kilocalories to walk a mile (or about 30 kcal to walk a kilometer). This is about the same as you’d expect for a standard mammalian quadruped of our size. But for chimpanzees, committed to knuckle-walking, the energy cost is about double.