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).

Strange relations and island continents

54.0 – 51.1 million years ago

We’ve seen a great many catastrophes in the history of life, and been reminded of the role of sheer chance in evolution. But the Cenozoic also sees a dramatic adaptive radiation and the steady progress of arms races among survivors of the great dinosaur die-off. Four large scale groupings of placental mammals have already appeared: Afrotheres (aardvarks, hyraxes, elephants, and sea cows), Xenarthrans (anteaters, armadillos, and sloths), Laurasiatheres (shrews, hedgehogs, pangolins, bats, whales, hoofed animals, and carnivores), and Supraprimates (aka Euarchontoglires, including rodents, tree shrews, and primates). This grouping of mammals is anything but obvious – it’s only with DNA sequencing that it has emerged. What’s noticeable is the association with different continents: Afrotheres with Africa, Xenarthrans with South America, and the others with the monster content of Laurasia (Eurasia and North America). Looking beyond placental mammals we see other continental associations: marsupials flourish in South America and Australia, and giant flightless “terror birds” carry on rather like predatory dinosaurs in South America.

There is a pattern here. Evolutionary arms races are most intense in the supercontinent of Laurasia (eventually joined by India and Africa). The island continents of South America and Australia stand apart, and they fare poorly when they start exchanging fauna with the rest of the world. We’ll see a similar pattern – large areas stimulate more competition, and more intense evolution, isolated areas are at a disadvantage – when we look at modern history, with ocean voyages effectively reuniting Pangaea. (This is a major theme of Alfred Crosby’s Ecological Imperialism and Jared Diamond’s Guns, Germs, and Steel.)

On the Origin of Seafood by Means of Natural Selection,

Or the Preparation of Favorite Dishes in the Struggle for Dinner.

567 – 537 million years ago

The Cambrian explosion — shells and skeletons, and all the major animal phyla of today — is one of the major events in the history of life. It’s hard to miss – Darwin was well aware of it – because for the first time you have abundant well-preserved fossils of animals with hard parts. From now on, if I miss a tweet one day or another, it’s because I didn’t get to it, not because the evidence isn’t there.

Genetic evidence seemingly clashes with the fossil evidence. A “molecular clock” based on rates of gene divergence suggests that major animal phyla had begun diverging from one another long before the Cambrian explosion. But maybe the genetic evidence is wrong, and the “molecular clock” was running faster in the past than more recently. Or maybe complex organisms evolved long before the Cambrian, but left little or no fossil evidence. Or maybe the ancestors of today’s animals really did diverge early, but didn’t get complex until the Cambrian.

And why the explosion happened when it did is unresolved. Here’s a recent review. The early Snowball Earth episodes probably contributed in some way, and the rise of oxygen in the atmosphere must have been important. Or maybe there was some dramatic biological event that triggered the explosion: the origin of eyes, and/or the beginning of predation, setting off an arms race between predators and prey that’s been going on ever since.

If this last possibility is correct, then the transition from an Edenic, predator-free Ediacaran world to the Cambrian is a form of “symmetry breaking.” There is maybe an analogy here in human social evolution to the transition from an egalitarian society to a world of inequality, of rulers and ruled  (which also followed a – much milder – glacial episode).

Speaking of predation: the Logarithmic History blog is partly about commemorating great events in the past. It seems fitting to celebrate the Cambrian as the origin of seafood. If you take your time machine back to any time before the Cambrian, pickings will be slim – algae mostly, although we don’t really know what the Ediacara would have tasted like. The time-traveller’s menu gets a lot better with the Cambrian (although wood for a fire is still a problem). Nowadays you can’t hope to dine on trilobite, alas. (Check out March 13 last year for more of this sad story.) But sometime in the next few days why not have some mussels for dinner? (The recipe below has some non-Cambrian ingredients. It will be a few more days, incidentally, before the evolution of anything kosher.)

Steamed mussels, 4 servings

Wash and debeard:
4 to 6 pounds mussels
Place them in a large pot and add:
½ cup dry white wine
½ cup minced fresh parsley or other herbs
2 tablespoons chopped garlic
Cover the pot, place it over high heat, and cook, shaking the pot occasionally, until most of the mussels are opened, about 10 minutes. Use a slotted spoon to remove mussels to a serving bowl, then strain the cooking liquid over them. Drizzle over the mussels:
1 tablespoon extra-virgin olive oil
Juice of 1 lemon
Serve with:
Plenty of crusty bread (invented< 15,000 years ago, but who’s counting?)

Your cuisinart, a prehistory

A famous movie cut, from 2001: A Space Odyssey, transitions from a bone club, hurled aloft by an australopithecine 2.5 million years ago, to a spacecraft in the year 2001.

Arthur Clarke and Stanley Kubrick, coming up with the plot for the movie/book, were influenced by the popular author Robert Ardrey. In his book African Genesis, Ardrey casts human evolution as a version of the story of Cain and Abel, except in his version the peaceful vegetarians (robust australopithecines) get clobbered by the club-wielding meat-eaters (gracile australopithecines).

We were born of risen apes, not fallen angels, and the apes were armed killers besides. And so what shall we wonder at? Our murders and massacres and missiles, and our irreconcilable regiments?  Or our treaties whatever they may be worth; our symphonies however seldom they may be played; our peaceful acres, however frequently they may be converted into battlefields; our dreams however rarely they may be accomplished. The miracle of man is not how far he has sunk but how magnificently he has risen. We are known among the stars by our poems, not our corpses.

Ardrey, along with Konrad Lorenz and Desmond Morris, was much in vogue in the 1960s: Sam Peckinpah was another movie director influenced by him. The man could turn a phrase. Unfortunately his speculations on evolution and human behavior are probably not of enduring value: he had the misfortune to take up the topic too early to take on board the sociobiological revolution pioneered by William Hamilton, Robert Trivers, and George Williams, and popularized by E. O. Wilson and Richard Dawkins.

Ardrey may not have been off-base in thinking that weaponry and warfare have been an important motive force in human biological and social evolution (more on this later). But where early stone tools are concerned, a different segue, from Oldowan chopper to Cuisnart may be more appropriate.

Recent research argues that early hominins could have dramatically increased available food energy by pounding vegetables and chopping up meat into more digestible pieces. Tool use may have been an early step in our ancestors’ move to high energy diets. Meat-eating began to be important in human evolution around 2.6 million years ago. Somewhat later we see evidence that some hominins have lighter jaws and aren’t chewing as much. So to celebrate this early dietary revolution, here’s a recipe:

Steak Tartare

Place in a food processor fitted with the metal chopping blade:

1 ½ pounds lean beef (tenderloin, top round, or sirloin) cut into ½ inch cubes

Pulse until meat is coarsely ground, 7-10 seconds. Do not over-process. Remove meat to a chilled platter or individual plates and gently form into 6 individual mounds.

[Optional: Make a spoon shaped indentation on top of each mound and crack into each

1 egg yolk.]

Divide and arrange in small piles around each serving:

½ cup minced onions
½ cup minced shallots
½ cup minced fresh parsley
¼ cup minced drained capers
8-12 anchovies (optional)

Serve immediately and pass separately:

Fresh lemon juice
Worcestershire sauce
Dijon mustard
Hot pepper sauce
Freshly ground black pepper
Salt

From The Joy of Cooking 1997

Planet of the apes

20.5 – 19.4 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).

Strange relations and island continents

53.1 – 50.3 million years ago

We’ve seen a great many catastrophes in the history of life, and been reminded of the role of sheer chance in evolution. But the Cenozoic also sees a dramatic adaptive radiation and the steady progress of arms races among survivors of the great dinosaur die-off. Four large scale groupings of placental mammals have already appeared: Afrotheres (aardvarks, hyraxes, elephants, and sea cows), Xenarthrans (anteaters, armadillos, and sloths), Laurasiatheres (shrews, hedgehogs, pangolins, bats, whales, hoofed animals, and carnivores), and Supraprimates (aka Euarchontoglires, including rodents, tree shrews, and primates). This grouping of mammals is anything but obvious – it’s only with DNA sequencing that it has emerged. What’s noticeable is the association with different continents: Afrotheres with Africa, Xenarthrans with South America, and the others with the monster content of Laurasia (Eurasia and North America). Looking beyond placental mammals we see other continental associations: marsupials flourish in South America and Australia, and giant flightless “terror birds” carry on rather like predatory dinosaurs in South America.

There is a pattern here. Evolutionary arms races are most intense in the supercontinent of Laurasia (eventually joined by India and Africa). The island continents of South America and Australia stand apart, and they fare poorly when they start exchanging fauna with the rest of the world. We’ll see a similar pattern – large areas stimulate more competition, and more intense evolution, isolated areas are at a disadvantage – when we look at modern history, with ocean voyages effectively reuniting Pangaea. (This is a major theme of Alfred Crosby’s Ecological Imperialism and Jared Diamond’s Guns, Germs, and Steel.)

Enemies

339- 322 million years ago

You’re trying to live without enemies. That’s all you think about, not having enemies.

Isaac Babel, Red Cavalry

Enemies are the most important agencies of selection.

Geerat Vermeij, Evolution and Escalation

Much of what we’ve been seeing since the onset of the Cambrian, Monday, February 27, is the outcome of evolutionary arms races, leading to steady improvements in teeth, claws, armor, and mobility. It may well be that the onset of predation is what triggered the Cambrian explosion in the first place. The paleontologist Geerat Vermeij argues that arms races and escalation – not adaptation to the physical environment – are the greatest cause of progressive evolution.

We’ll see when we start getting into human evolution, biological and social, that enemies – other people especially – and arms races go on being a major motor of change. But arms races and escalation are going to look different in human evolution than they do in most non-human evolution. People are super-cooperators, and violent competition in humans tends to involve more group-against-group competition, with rival groups monopolizing and competing over territory. And in the human analog of predation – the formation of stratified societies, where elites live off the mass of the population – the human “predators” commonly band together under the aegis of the state to regulate their competition. At their best, human elites are less like wolves and more like sheepdogs.

Arms races operate with greater intensity in some environments than others. Races are more intense on large landmasses than small. Hence the common pattern in both biological evolution and human social evolution that isolated small continents and islands are especially vulnerable to invasion when their isolation ends. And arms races may be more intense, and the pace of evolution correspondingly greater, in the (more or less) 2-D terrestrial environment compared to the 3-D oceans.

Yet there may be something else involved in the initial move onto land – it’s sometimes among refugees from arms races that the greatest evolutionary advances arise. Fish moving onto land may have been doing it partly to get to someplace where enemies were weak or scarce. Human analogs might be the early Ionian Greeks fleeing the Dorian invasions, the settlers of Polynesia lighting out for the territories to escape a lowly position in a social order of ranked lineages, or the New England Pilgrims fleeing an un-Godly England. Or Vermeij himself – he is competitively handicapped, having lost his sight at three years old, but has made a distinguished career studying shelled invertebrates by touch.

On the Origin of Seafood by Means of Natural Selection

Or the Preparation of Favorite Dishes in the Struggle for Dinner.

562-532 million years ago

The Cambrian explosion — shells and skeletons, and all the major animal phyla of today — is one of the major events in the history of life. It’s hard to miss – Darwin was well aware of it – because for the first time you have abundant well-preserved fossils of animals with hard parts. From now on, if I miss a tweet one day or another, it’s because I didn’t get to it, not because the evidence isn’t there.

Genetic evidence seemingly clashes with the fossil evidence. A “molecular clock” based on rates of gene divergence suggests that major animal phyla had begun diverging from one another long before the Cambrian explosion. But maybe the genetic evidence is wrong, and the “molecular clock” was running faster in the past than more recently. Or maybe complex organisms evolved long before the Cambrian, but left little or no fossil evidence. Or maybe the ancestors of today’s animals really did diverge early, but didn’t get complex until the Cambrian.

And why the explosion happened when it did is unresolved. Here’s a recent review. The early Snowball Earth episodes probably contributed in some way, and the rise of oxygen in the atmosphere must have been important. Or maybe there was some dramatic biological event that triggered the explosion: the origin of eyes, and/or the beginning of predation, setting off an arms race between predators and prey that’s been going on ever since.

If this last possibility is correct, then the transition from an Edenic, predator-free Ediacaran world to the Cambrian is a form of “symmetry breaking.” There is maybe an analogy here in human social evolution to the transition from an egalitarian society to a world of inequality, of rulers and ruled  (which also followed a – much milder – glacial episode).

Speaking of predation: the Logarithmic History blog is partly about commemorating great events in the past. It seems fitting to celebrate the Cambrian as the origin of seafood. If you take your time machine back to any time before the Cambrian, pickings will be slim – algae mostly, although we don’t really know what the Ediacara would have tasted like. The time-traveller’s menu gets a lot better with the Cambrian (although wood for a fire is still a problem). Nowadays you can’t hope to dine on trilobite, alas. (Check out March 13 last year for more of this sad story.) But sometime in the next few days why not have some mussels for dinner? (The recipe below has some non-Cambrian ingredients. It will be a few more days, incidentally, before the evolution of anything kosher.)

Steamed mussels, 4 servings

Wash and debeard:
4 to 6 pounds mussels
Place them in a large pot and add:
½ cup dry white wine
½ cup minced fresh parsley or other herbs
2 tablespoons chopped garlic
Cover the pot, place it over high heat, and cook, shaking the pot occasionally, until most of the mussels are opened, about 10 minutes. Use a slotted spoon to remove mussels to a serving bowl, then strain the cooking liquid over them. Drizzle over the mussels:
1 tablespoon extra-virgin olive oil
Juice of 1 lemon
Serve with:
Plenty of crusty bread (invented< 15,000 years ago, but who’s counting?)

A famous movie cut, from 2001: A Space Odyssey, transitions from a bone club, hurled aloft by an australopithecine 2.5 million years ago, to a spacecraft in the year 2001.

Arthur Clarke and Stanley Kubrick, coming up with the plot for the movie/book, were influenced by the popular author Robert Ardrey. In his book African Genesis, Ardrey casts human evolution as a version of the story of Cain and Abel, except in his version the peaceful vegetarians (robust australopithecines) get clobbered by the club-wielding meat-eaters (gracile australopithecines).

We were born of risen apes, not fallen angels, and the apes were armed killers besides. And so what shall we wonder at? Our murders and massacres and missiles, and our irreconcilable regiments?  Or our treaties whatever they may be worth; our symphonies however seldom they may be played; our peaceful acres, however frequently they may be converted into battlefields; our dreams however rarely they may be accomplished. The miracle of man is not how far he has sunk but how magnificently he has risen. We are known among the stars by our poems, not our corpses.

Ardrey, along with Konrad Lorenz and Desmond Morris, was much in vogue in the 1960s: Sam Peckinpah was another movie director influenced by him. The man could turn a phrase. Unfortunately his speculations on evolution and human behavior are probably not of enduring value: he had the misfortune to take up the topic too early to take on board the sociobiological revolution pioneered by William Hamilton, Robert Trivers, and George Williams, and popularized by E. O. Wilson and Richard Dawkins.

Ardrey may not have been off-base in thinking that weaponry and warfare have been an important motive force in human biological and social evolution (more on this later). But where early stone tools are concerned, a different segue, from Oldowan chopper to Cuisnart may be more appropriate.

Recent research argues that early hominins could have dramatically increased available food energy by pounding vegetables and chopping up meat into more digestible pieces. Tool use may have been an early step in our ancestors’ move to high energy diets. Meat-eating began to be important in human evolution around 2.6 million years ago. Somewhat later we see evidence that some hominins have lighter jaws and aren’t chewing as much. So to celebrate this early dietary revolution, here’s a recipe:

Steak Tartare

Place in a food processor fitted with the metal chopping blade:

1 ½ pounds lean beef (tenderloin, top round, or sirloin) cut into ½ inch cubes

Pulse until meat is coarsely ground, 7-10 seconds. Do not over-process. Remove meat to a chilled platter or individual plates and gently form into 6 individual mounds.

[Optional: Make a spoon shaped indentation on top of each mound and crack into each

1 egg yolk.]

Divide and arrange in small piles around each serving:

½ cup minced onions
½ cup minced shallots
½ cup minced fresh parsley
¼ cup minced drained capers
8-12 anchovies (optional)

Serve immediately and pass separately:

Fresh lemon juice
Worcestershire sauce
Dijon mustard
Hot pepper sauce
Freshly ground black pepper
Salt

From The Joy of Cooking 1997

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.

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 modularist 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, such as 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 cultural 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.

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.