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:

Apes, on the road to great

14.1 – 13.4 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, but there have been interesting discoveries from Europe that we’ll cover in days to come. 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 thoats

There’s a great expansion in the diversity of horses in the mid-Miocene, especially horses that are adapted to grazing rather than browsing. The shift to grazing is going 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.)

Thoatherium reconstruction

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.

Planet of the horses

16.7 – 15.8 million years

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 generate 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.

Bunches of monkeys

Our descent, then, is the origin of our evil passions!! The devil under form of Baboon is our grandfather.

Charles Darwin, Noteboook M

Maimoun angushti shaitan ast.

(A monkey is the devil’s fingers.)

Tajik proverb

Monkeys and apes are not only exceptionally brainy, but also distinctively social. Most mammals are solitary (apart from mothers and their juvenile offspring of course). Among a minority of mammals, adult males and females set up pairbonds. And some mammals form larger groups. In most cases, however, these are relatively unstructured aggregations: a herd of buffalo is more like a crowd of people than a human community. A handful of mammals – elephants, cetaceans, and the majority of monkeys and apes – form more structured groups, enduring and internally differentiated.

Social evolution is path dependent: primate social organization is affected by ecology, but also has a strong phylogenetic component.  This makes it possible to offer a tentative reconstruction of the stepwise evolution of stable sociality in primates. Here’s an evolutionary tree, showing inferred transitions between solitary living, and multimale/multifemale, unimale/multifemale, and pairbonded groups:

A diagram of the possible evolutionary dynamics looks like this:

And the accompanying story goes like this: about 52 million years ago, the solitary nocturnal ancestor of monkeys and apes switched to being diurnal. This allowed for the exploitation of a whole range of new foods, but it also exposed the ancestor to new forms of predation. The first step in the evolution of monkey and ape sociality, then, was aggregation in multimale/multifemale groups to cut predation risk. At first these groups would have been loosely structured and unstable, but eventually they would have evolved into something like what we see today among most Old World monkeys: stable (sometimes lifelong) networks of relatives and friends, dominants and subordinates, nested within enduring communities.

A later development, going back to 20 million years ago or less, was a shift, among some of these social primates, to unimale/multifemale groups, or pair-living family groups.

The inferred development of structured social groups in primates bears a remote similarity to the evolution of eusociality among social insects. According to current theories, the starting point for eusociality is the development of a defended nest site where females lay eggs and raise offspring. Sometimes an established nest site is so valuable that it’s adaptive for the next generation of offspring to stay on when they mature rather trying to found new nests. The eventual result may be the evolution of a highly structured society, with strong reproductive skew: some nest members specialize in reproduction, others in foraging or defending the nest. The latter may evolve into an obligately sterile caste.

Primates too have developed an intensified, structured sociality as a response to obligate group living. But the parallels with eusocial insects go only so far. The great majority of primates give birth to one offspring at a time. There are no queen bee baboons whelping one vast litter after another and pushing subaltern kin into caring for them. This goes for humans as well. Our species matches the social insects in the scale of cooperation, but we manage this through a complicated dance of coalition-building and reputation management. For a primate, building a honey-bee style superorganism has to be an aspiration rather than a reality.

We were never chimps

It’s natural to turn to our closest living relatives, the great apes (chimpanzees and bonobos, gorillas, orangutans), for insights into what our remote ancestors were like. But the fossil evidence suggests that current great apes aren’t a good guide to our past. Below is a figure from a recent article reconstructing diet and habitat for Morotopithecus and some relatives, from just over 20 million years ago, and later. It looks like all these guys inhabited relatively open woodland – trees interspersed with grass – rather than the closed canopy tropical forest that is the modal habitat for all the living great apes. Also, they may have specialized in consuming more young leaves, and less fruit than, say, chimpanzees. On current evidence, then, our closest living relatives have all evolved away from our common ancestors, to become (not terribly successful) tropical forest specialists.

Also, more on this later: gorilla and chimpanzee knuckle walking may be a poor model for locomotion in our common ancestor. Asking how we evolved from a chimp-like ancestor is probably asking the wrong question. 

Pace Jared Diamond: a good book and a snappy title, but we are not The Third Chimpanzee.

Planet of the apes

17.6 – 16.7 million years ago

We are now covering history at the rate of one million years a day

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).

Gould’s Belt

29.2 – 27.7 million years ago

Logarithmic History has had a lot of geology and biology lately, not so much astronomy. But all is not peaceful in the heavens.

Benjamin Gould is a nineteenth century astronomer who noted that a lot of bright stars in the sky — especially the bright blue stars that we know are very young — seem to fall along a ring tilted at a 20 degree angle to the Milky Way. This ring has come to be called Gould’s Belt (or the Gould Belt). The Belt is an ellipse about 2400 by 1500 light years across where there has been a recent wave of star formation. Our Sun lies within the belt, somewhat off center; the center lies in the direction of the Pleiades.

The Belt began forming maybe thirty million years ago. We’re not sure what happened. A supernova may have set off a wave of star formation, but it would have to have been a huge one. Or it may be that a gas cloud or a clump of dark matter passed at an angle through our part of the Milky Way, and started stars forming with its shock wave. There are features resembling Gould’s Belt in other galaxies. In any case, the Belt is one of the really striking features of our part of the Milky Way.

Whatever its cause, no one disputes its magnificence. Gould’s belt is the most prominent starry feature in the Sun’s neighborhood, contributing most of the bright young stars nearby. Nearly two thirds of the massive stars within 2,000 light-years of the Sun belong to Gould’s belt. If I were kidnapped by an alien spaceship and taken to some remote corner of the Galaxy, Gould’s belt is what I’d look for to find my way back home.

Ken Crosswell. Gould’s Belt.

If you’re in the Northern hemisphere you can look at the sky tonight and see the Milky Way in an arc in the Western sky, stretching from North to South. West of the Milky Way you’ll see some of Gould’s belt, an arc of bright stars running north to south from the Pleiades, through Taurus and the bright stars of Orion, and Canis Major. So tonight look at the stars, and drink a toast if you want, to your ape ancestors, who were just on the cusp of splitting off from monkeys thirty million years ago.