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7.04 – 6.66 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.

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.

12.3 – 11.8 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 speciesof 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 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:

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.

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

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. 

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.

18.3 – 17.4 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).

There’s a dark side to being a primate. A few years back a review article summarized data on rates of lethal aggression in non-human animals. The figure below shows some of the results. Several clusters of especially violent species stand out in the figure, including primates (redder is more violent). Bats are pretty nice, though (too bad about all the viruses).

Much of the lethal aggression in primates involves infanticide. Sarah Hrdy demonstrated back in the 1970s that infanticide occurs regularly in Hanuman langurs, monkeys in India. A male who takes over a group of females will systematically kill offspring sired by the previous male. If you think evolution is about the survival of the species, this is hard to explain. But it makes sense given the logic of the selfish gene. Females who lose an infant return more quickly to breeding again, and the father of the next infant is likely to be the killer of the previous one.

Primates may be particularly vulnerable to this grim logic, because they spend a long time as infants. Among primates, commonly,

           L/G>1

That is to say that the time, L, that a female spends lactating for an infant (during which she is unlikely to conceive), is usually greater than the time, G, that she spends gestating an infant. This puts particular pressure on males to hurry things along by eliminating nursing infants fathered by other males.

Death of Astyanax

As a result, infanticide is relatively common among primates, and females under particularly strong pressure to find ways to avoid it. Hanuman langurs live in one-male units, where a female has little choice about who she mates with. In other species, by contrast (most baboons, chimpanzees), multiple males reside with multiple females. In these species females are often sexually promiscuous, sometimes actively soliciting multiple males for sex. This is probably mostly a matter of confusing paternity sufficiently to suppress the threat of infanticide. There’s a general lesson here: females are not always monogamously inclined, but female promiscuity generally has different evolutionary roots than male promiscuity.

30.3 – 28.8 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. (You’ll get this view earlier in the night. It will fall below the horizon later.) 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.

Short version: It looks like most mammals, at least most large mammals, have the brains they need, while primates, especially large primates, have the brains they can afford.

Longer version: One reason for being interested in monkeys is that they’re brainy mammals. Here’s the conventional graph illustrating that:

Larger mammals tend to have larger brains, but the relationship is non-linear. Multiplying body mass by x doesn’t multiply brain mass by x. Instead it multiplies brain mass by about x.75. In other words, Brain Mass is proportional to (Body Mass).75. Equivalently (taking the logarithm of both sides) Log[Brain Mass] is equal to .75 times Log[Body Mass], plus a constant. So Log[Brain Mass] plotted against Log[Body Mass] gives a straight line with a slope of .75. That means that if one mammal has 16 times the body mass of another, it’s expected to have 8 times the brain mass, 10,000 times the body mass means 1000 times the brain mass, and so on. The thing to note is that primates defy expectations. They have larger brains than would be expected based on their body sizes.

But we’ve recently learned that primates – especially big ones – are even more special than this graph suggests. Susan Herculano-Houzel has pioneered a technique that involves chopping up brains (or parts of brains), dissolving their cells to make a kind of brain soup, and counting cell nuclei. This allows her to estimate how many neurons there are in different brains.

Major findings: Among most mammals, the number of neurons increases more slowly than brain size. Increase brain size by x, and you increase number of neurons by about x.67. (H-H shows this flipped around. Increase number of neurons by x and you increase brain mass by x1.5.) But primates are exceptional; the relationship is nearly linear. An x-fold increase in primate brain size corresponds to about an x-fold increase in number of neurons. Humans follow the primate rule here. We have about the same density of neurons as other primates. When you combine the exceptionally large brain sizes of humans (exceptional even relative to our brainy primate relations) with a standard high primate neuron density, you get an animal with an enormous number of neurons. By contrast, a rodent with a human sized brain, if it followed rodent rules for how neuron numbers increase with brain size, would have only 1/7 as many neurons.

Neurons are expensive. Most large animals economize by cutting back on neuron density. A cubic centimeter of cow brain has fewer neurons, and consumes energy at a lower rate, than a cubic centimeter of mouse brain (although of course the cow’s brain is bigger). By contrast, large primates are extravagant, devoting exceptionally large energy budgets to running their brains. And human brains are exceptionally costly. An important question for the study of human evolution is how we paid the bill for such costly brains. That’s a story for later. But another part of the story starts back in the early Cenozoic, when monkeys committed to a different set of rules for building brains.

And here is a chart giving absolute numbers of  cortical neurons (cneurons) for a bunch of species. Scott Alexander has some thoughts about the moral implications. Short version: skip the pork for dinner (and skip the elephant, chimp, and manflesh. But you knew that). Beef might be okay. Better is lobster.

And Werner Herzog is probably okay with you eating chickens.

35.9 – 34.0 million years ago

The Oligocene (starting today on Logarithmic History) sees a major diversification of anthropoid primates (monkeys, apes, and humans). Among the anthropoids, the major evolutionary split is a geographic one, between platyrrhines (New World monkeys) and catarrhines (Old World monkeys, apes, and humans). Aegyptopithecus is one of the earliest primates that clearly falls on the catarrhine side of that split (although the split must go back earlier).

At Logarithmic History we traffic in Big Questions, and one of the biggest questions of all is the balance of natural law and accident in making our world. Thus physicists have long hoped to find that the laws governing our universe reduce to just a few fundamental equations, but we saw at the beginning of this blog that they are now confronting the possibility that our universe is just one among many, and that the laws of physics in our universe may incorporate a large dose of historical accident. With the discovery of extra-solar planets, we’re just beginning to get an idea of how typical or atypical our solar system is. And we’ll have a lot of opportunities to ask whether there are Laws of History (an old idea now undergoing a revival in the new field of cliodynamics*) when we move into the historical period later in the year.

The field of biogeography – the study of the geographic distribution of species – has seen some major pendulum swings in this regard. Darwin was intensely interested in questions of biogeography mainly because they could provide support for the theory of evolution. His approach could fairly be called eclectic. From sometime in the second half of the twentieth century however, a lot of biologists thought they could do better than just answering particularistic questions about how species A got to island Z (sometimes disparaged as “stamp collecting”). They wanted to find scientific laws.

Edward O. Wilson was an early pioneer in this area. Along with Robert MacArthur, he developed a theory of island biogeography which was supposed to get the field out of its natural history phase, and turn it into a predictive science. According to MacArthur and Wilson, the number of species on an island is set by a predictable equilibrium between extinction (smaller islands have higher extinction rates) and colonization (remote islands have lower colonization rates). Being a good scientist Wilson actually put this theory to the test by getting an exterminator to “defaunate” (it means what you think it means) some little mangrove islets, and showing that they returned to very close to their predicted equilibrium numbers of animal species after a while.

For the biogeography of continents (and larger islands once part of continents) the quest for scientific laws took a different turn. The discovery of continental drift and plate tectonics encouraged a school of “vicariance biogeography.” Vicariance biogeographers liked to trace current biogeographic distributions to the wanderings of continents. They were highly allergic to explanations involving accidental long-distance dispersal over big stretches of ocean.

Alan de Queiroz, in The Monkey’s Voyage: How Improbable Journeys Shaped the History of Life, provides a highly readable overview of the decline (if not quite the extinction) of the vicariance school in the face of mounting evidence for flukish dispersals as a major factor in biogeography. The dispersal of monkeys to the New World is a dramatic case in point. (Guinea pigs and their relatives are another.) About the only scenario that makes sense involves a raft of trees washing out to sea (most likely from the Congo basin) and eventually delivering a few parched, scared monkeys to the island continent of South America, where they eventually spawned the whole range of species – spider monkeys, squirrel monkeys, howler monkeys, tamarins, marmosets, capuchins – we know today. Sheer accident: change the weather a little, leave the monkeys stranded at sea a little longer, and the whole history of primates in the New World is erased.

* so new my spellchecker doesn’t recognize it.