Rama’s ape

12.3-11.7 million years ago

Ramapithecus (Rama’s ape) is no more. Another Hindu god has taken over the franchise; Ramapithecus is now subsumed under Sivapithecus, an earlier discovery, and is no longer a valid taxon name.

The story is interesting from a history-of-science point of view. Ramapithecus used to be presented as the very first ape on the human line, postdating the split between humans and great apes, maybe even a biped. This was given in textbooks not so long ago as established fact. Then geneticists (Sarich and Wilson) came along, and declared that the genetic divergence between chimps and humans is so low that the split had to be way later than Ramapithecus. There was a lot of fuss over this. Paleoanthropologists didn’t like geneticists telling them their job. Eventually, though, the paleoanthropologists found some new fossils. These showed in particular that the line of Ramapithecus‘s jaw was not arch-shaped, like a human’s, but more U-shaped, like a non-human ape’s. So after thinking it over a while, paleoanthropologists decided that Ramapithecus (now part of Sivapithecus) looked more like an orangutan relative: likely ancestor of a great radiation of orangutan kin that left just one genus, Pongo, in the present.

rama jaw

There are plenty of examples of experts in different fields coming up with different answers. For example, paleontologists didn’t like physicists telling them why dinosaurs went extinct. And we’ll see other examples in days to come: geneticists, physical anthropologists, and archeologists arguing over modern human origins. And very recently geneticists coming in on the side of old-fashioned historical linguists, and against recent generations of archeologists, in the matter of Indo-European origins.

It would be nice if there were a simple rule of thumb to decide who’s right in these cases. Maybe experts know what they’re talking about (except that experts were telling us recently that low fat diets were the key to losing weight and eggs would kill us with cholesterol*). Or maybe harder science experts know better than softer science experts (except that physicists like Kelvin were telling geologists that the Sun couldn’t possibly have produced enough energy to support life on Earth for hundreds of millions of years – then along came Einstein and E=mc2). So the best we can do maybe is realize people, scientists included, are prone to overconfidence and group think – and not just those other people, either, but you and me.

* Memories of the fallout from 1990s nutrition expert wisdom

  • SnackWell’s cookies, low fat, loads of sugar
  • Jelly beans prominently labelled “No fat.”
  • A fellow grad student laughing about an older relative who said bread and pasta make you fat, and confidently declaring “Fat makes you fat.”
  • Me in Brazil talking with my landlord, who was planning to lose weight by sticking with sausage, and cutting starch. I told him modern science had shown this was precisely the wrong approach. He politely (Brazilians can be pretty good-natured) disagreed, stuck with his plans, and lost weight.

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. 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), 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.

Land of the thoats

17.2-16.4 million years ago

There’s a great expansion in the diversity of horses in the mid-Miocene, especially horses adapted to grazing rather than browsing. The shift to grazing goes on world wide among many different groups. In South America the big grazers are the liptoterns, ungulates not closely related to horses that evolve to look a lot like them, with high-crowned grazing teeth, single-toed hoofed feet and legs built for speed. (Edgar Rice Burroughs took the name thoat – what his characters rode around on on Barsoom/Mars — from one genus of liptotern, Thoatherium.)


We often think of evolution as a matter of organisms adapting to their environments, but when the environment is other organisms, each side may be chasing a moving target. Or sometimes the sides may reach an equilibrium. In the case of grazing animals, there’s a process of coevolution that goes on between grazers and grasses. Where grazers are active, the plants that survive are grasses, which keep leaves above the ground but grow from underground. And this works in the other direction: in moderately dry climates, grasses are more productive than taller brushy plants, so it’s when grasses take over that there’s enough food around for grazers – a mutually reinforcing cycle. With drier climates from the mid-Miocene on, grasslands and grazers get to be more and more important.

So a lot of the story of life on Earth is not just the appearance of this or that cool animal, but also the evolution of ecosystems. At the same time grasslands were spreading on land, for example, kelp forests were spreading in coastal oceans. We’ll see how important grasslands are in human evolution and history. And kelp forests, with their rich fish populations, might have been important too, as the highway that the earliest Americans followed along the Pacific coast to the New World.

High fidelity

Arms races have been a big engine of evolutionary progress, both in biological evolution and in the evolution of human societies. Another big driver has been improvements in the fidelity of inheritance. We see this in the evolution of genetic systems, including the evolution of life itself, and of the eukaryotic chromosome. And we’ll see it in human social evolution, including the evolution of language, of writing, of the alphabet, and printing.

Both arms races and improved information transmission may have been factors in the evolution of braininess.

jerison brain race

The figure above is from the classic work of Harry Jerison, one of the pioneers in studying the evolution of brain size. It’s several steps away from the raw data, but what it shows is how mammalian Encephalization Quotients (EQs), a measure of brain size relative to body size, evolved over the Cenozoic. The figure might be read as the record of a brainy arms race between prey and predators, leading to increased variance in the EQ bell curve for both.

Primates of course are particularly brainy mammals. One popular explanation for this is a series of arms races within species, with bright monkeys and apes outwitting dimmer ones. This has been called the Machiavellian Intelligence hypothesis (or, in the case of macaques, macachiavellian intelligence).


This hypothesis may not hold up too well, however. One complication is that, contrary to what a lot of evolutionary psychology might suggest, social intelligence in primates is not separate from other sorts of intelligence. The same primate species that are good at solving social problem (e.g. tricking other group members) are also clever about things like tool use and other complex foraging skills. Variation in intelligence across primate species mostly boils down to a single general factor, rather than a bunch of domain-specific aptitudes.

Also, the latest research suggests that variation in diet and ecology, like the distinction between fruit eaters (brainy) and leaf eaters (not-so-much), accounts a lot of variation in brain size, while differences in social complexity (measured by group size) don’t seem to matter.

An alternative to the Machiavellian Intelligence hypothesis is the cultural intelligence hypothesis, with brainier animals more likely to innovate and more likely to learn others’ innovations. The first part pf this equation holds up: across various groups of organisms, including birds and primates, brainy animals are more flexible in their behavior, more likely to discover new adaptive behaviors, and more successful in colonizing novel environments. The second part is trickier. In recent years we’ve learned that learning useful information by observing others (go ahead, call it culture, if you want to annoy anthropologists) is extremely widespread, and found in organisms like guppies and honeybees that no one thinks are terribly bright. So learning from others doesn’t take special smarts.

Where bigger brained animals may excel is not in how much social learning they do, but in how accurately they do it – in copying fidelity. Theoretical models of the evolution of copying suggest that accurate copying makes a big difference. Small changes in copying fidelity can lead to large changes in the persistence of cultural traits. Of course this will crucially important for human evolution: more on this in days to come.

copying fidelity

The expected lifetime, measured in generations, of a cultural trait as a function of the efficiency of social learning (p). Each learning trial uses a new cultural parent drawn from the parent population (see text). Parameter value: n 1⁄4 2.

For a wide-ranging introduction to this rapidly advancing area of research, written by a leader in the field, try Darwin’s Unfinished Symphony: How Culture Made the Human Mind.

Planet of the horses

18.29-17.29 million years ago.

We’re now running through Big History at the rate of 1 million years per day.

Horses have probably been the single most important domesticated animal in human history. Also, more than with other livestock, people get attached to horses as individuals. I’m guessing that in history and literature there are more horses with individual names than any other animal. (Alexander the Great’s horse was Bucephalus, “Ox-head”; Muhammed’s was al-Buraq*; Charlemagne’s was Tencendur; Don Quixote’s was Rocinante; Gandalf’s was Shadowfax.) We’ll be hearing a lot more about horses and horse folk on Logarithmic History once we get to human history.

Being so charismatic, horses have featured in a big way in arguments over evolution. Thomas Henry Huxley (1825-1895), “Darwin’s bulldog,” knew he needed to find good evidence for evolution. When he visited the United States in 1876, he was ready to give a lecture based on horse fossils from Europe. But visiting Yale, he was so impressed with O. C. Marsh’s collection of horse fossils from the western United States, that he rewrote his lecture around it.

Henry Fairfield Osborn (1857-1935) was director of the American Museum of Natural History and a huge presence in American paleontology. He was active at a time when most scientists accepted evolution, but many weren’t so keen on Darwin’s theory of natural selection. He thought horses were a fine example of “orthogenesis,” the tendency of species to follow a fixed line of evolution, reflecting internal forces, maybe related to willpower. He thought that humans shared a migratory spirit with horses, so that anywhere horse fossils were found would be a good place to look for human fossils. This theory didn’t pan out too well. A massive AMNH expedition to Central Asia led by Ray Chapman Andrews found all sorts of wonders – dinosaur eggs, baluchitheres – but no fossil “pro-men.” Orthogenesis leant itself naturally to diagrams showing evolution from early to modern horses going in a straight line.


George Gaylord Simpson (1902-1984), paleontologist, was one of the great figures in the evolutionary Modern Synthesis that brought together Darwin’s theory of natural selection and Mendel’s genetics. There was no room for orthogenesis in the Modern Synthesis, and Simpson emphasized that the evolution of horses was a matter of adaptation to a changing environment – especially the spread of grasslands. Also that horse evolution looked more like a bush than a ladder.


Stephen Jay Gould (1941-2002) was the most widely recognized American evolutionary biologist of recent times. (For example had a spot on The Simpson’s — “Lisa The Skeptic,” Season 9.) Gould had his own take on the modern synthesis, taking the “bushes not ladders” theme for horses and other animals (including human ancestors), and pushing it a step further. According to the theory of “punctuated equilibrium” (formulated in collaboration with Niles Eldredge), species mostly change relatively little during the time they exist (evolutionary stasis). Most evolutionary change happens when a small population buds off to form a new species and reproductive isolation allows it to conserve any evolutionary novelties it has developed. This opens up the possibility of “species selection.” Applied to horses, for example, this could mean that horses were evolutionarily successful for some time not so much because individual horses were well-adapted, but because something about horses collectively (their harem mating system, maybe) made one horse species especially likely to produce new species. Both horses and primates seem to be especially prone to bud off new species:

Speciation and chromosomal evolution seem fastest in those genera with species organized into clans or harems (e.g., some primates and horses) or with limited adult vagility and juvenile dispersal, patchy distribution, and strong individual territoriality (e.g., some rodents). This is consistent with the … hypothesis … that population subdivision into small demes promotes both rapid speciation and evolutionary changes in gene arrangement by inbreeding and drift.

 * Richard Dawkins doesn’t believe that Muhammed’s horse, al-Buraq, carried him (i.e. Muhammed) to heaven and back.

Planet of the apes

22.9-21.6 million years ago

The Miocene (23 – 5 million years ago) is a period of extraordinary success for our closest relatives, the apes. Overall there may have been as many as a hundred ape species during the epoch. Proconsul (actually several species) is one of the earliest. We will meet just a few of the others over the course of the Miocene, as some leave Africa for Asia, and some (we think) migrate back.

Sometimes evolution is a story of progress – not necessarily moral progress, but at least progress in the sense of more effective animals replacing less effective. For example, monkeys and apes largely replace other primates (prosimians, relatives of lemurs and lorises) over most of the world after the Eocene, with lemurs flourishing only on isolated Madagascar. This replacement is probably a story of more effective forms outcompeting less effective. And the expansion of brain size that we see among many mammalian lineages throughout the Cenozoic may be another example of progress resulting from evolutionary arms races.

But measured by the yardstick of evolutionary success, (non-human) apes — some of the brainiest animals on the planet — will turn out not to be all that effective after the Miocene. In our day, we’re down to just about four species of great ape (chimpanzees, bonobos, gorillas, and orangutans), none of them very successful. Monkeys, with smaller body sizes and more rapid reproductive rates, are doing better. For that matter, the closest living relatives of primates (apart from colugos and tree shrews) are rodents, who are doing better still, mostly by reproducing faster than predators can eat them.

So big brains aren’t quite the ticket to evolutionary success that, say, flight has been for birds. One issue for apes may be that with primate rules for brain growth – double the brain size means double the neurons means double the energy cost – a large-bodied, large brained primate (i.e. an ape) is going to face a serious challenge finding enough food to keep its brain running. It’s not until a later evolutionary period that one lineage of apes really overcomes this problem, with a combination of better physical technology (stone tools, fire) and better social technology (enlisting others to provision mothers and their dependent offspring).