How the wolf became the dog

The Russian geneticist Dmitry Belyaev was interested in the inheritance of fur colour in Silver Foxes.  When he started his long experiment, Stalin was dead, but the atavistic simplicities that seem to underlie the flowering of dictatorships lingered.  Stalin had favoured the theories of Trofim Lysenko, who believed that characteristics acquired during a lifetime could be inherited; that weeds are spontaneously transmuted into food grains, and that genetics was a fake science.  Thus, anybody who advanced Darwinian or geneticist ideas was an enemy of Stalin and of the Soviet state.  Dmitry’s much older brother had done just that, and was executed without trial in 1937.  Stalin died in 1953, but Mendelian genetics was not permitted in the Soviet Union until 1964.

By 1959 Belyaev had observed that some silver foxes in captivity, in fur farms, were naturally tamer than others.  Darwin, who in 1859 did not know that genes existed, and referred in his theory to the loci of heritability and variation as “unknown factors”, had written in the On the Origin of Species that he suspected that the evolution of some characteristics caused by some of these “unknown factors” would be chained to other characteristics through the same “unknown factors”; or in genetics terms, that some genes operated in concert with other genes, so if you get the effect of one you get the effect of the whole suite.  Belyaev applied this to silver foxes, and surmised that the genetic substructure of tameness, and thus domestication, might correlate with certain physical characteristics.  In 1959 he began his series of experiments in which he selected for breeding, from a pool of thirty male foxes and a hundred vixens, only those who in very brief and sparse windows of contact were tolerant of, or even attracted to, human presence.  This experiment lasted the final twenty six years of his life, but that was a ridiculously short time in which to imagine that a process of evolution, replication, variation, selection, could take place.

Take place it did, nonetheless.  Darwin had noted that domestic animals tended to have characteristics such as as floppy ears, curly tails and spotted coats that were absent in their wild forebears, and by the twelfth generation of Belyaev’s selective breeding the tame silver foxes were not only tamer—the corticosteroid level in the their plasma had fallen to half the level in a control group, and after 30 generations the level had halved again — but, as Darwin had seen already, their coats became patchy, their ears floppy, their tails more curled in a ways that “tend to make animals appear appealingly juvenile to humans”.

Thus it seems likely that some animals have become tame or domesticated because long before they did so such developmental potentialities for variation have been established in their genetic and epigenetic structure.  If we are to avoid teleology —that these animals had these potentialities so that they could be domesticated and serve mankind— then we must suppose that this range of developmental potentialities, from calm, accommodating and deferential to aggressive, stressed and unpredictable, had some function that allowed for a stable complexity of social structure in the wolf pack.  Within the pack the tractable and cooperative, and the aggressive and dominant, are kept in a balance by selection, and this optimises predatory and reproductive success.

It has recently been observed that the same species of bird will, in times of plenty, feed the offspring that squawks the most and therefore is most in need, but in times of famine will feed the biggest. There is a general rule of reproductive success and maximum proliferation of genes here.  If there’s a lot to eat, feed the whole brood.  If there’s a dearth, feed the one most likely to live.

The size of a wolf pack will depend on the availability of prey.  The availability of prey will depend on absolute numbers of prey animals, and on competition from other predators, both conspecifics in other packs[1], and other predatory species.

With excessive expression of dominant and aggressive characteristics, alpha wolves of both sexes spend too much time strutting their stuff and fighting.  Hunting and rearing young become secondary, and such a pack might well fail entirely and vanish.  Too much mutual tenderness, care and sympathy would be fine as long as prey was abundant, all young fed, the weaker cherished (as in the bird in time of plenty).  But even in the best of times such a pack would expand until it started to deplete its resources, at which point the diminution of resources, because they too have to reproduce, becomes exponential.  The TLC-driven pack would try to spread the rapidly diminishing nutrition evenly among the young, and they’d all die.

In this case the constant state is a balance between the size of the wolf pack and its nutrition resources.  The pack size increases when food is plentiful, and decreases in time of dearth.  The selection takes place at the level of the genes and epigenetic processes that allow variation between TLC and aggression.  In the optimum circumstances the pack increases with TLC, this causes dearth, the residual aggressive individuals bully the softy TLC individuals into subservient cooperation; that is they are allowed to hunt, which is to their own advantage, because lone wolves are less successful hunters, but they are not allowed to breed within the pack, and if they do, their offspring are killed.

This leads to an excess of aggressive-dominant individuals.  There is no way the TLC lot can survive in this environment.  Food is scarce, the alphas use the TLCs’ hunting skills but do no allow them to feed adequately, and because the aggression of the alphas prevents them from hunting efficiently anyway, the level of nutrition is ever decreasing.  The desperate TLCs are driven off the kill.  They either fight, and die, because they’re not so good at fighting, or they flee to the wolf-wilderness, the debatable lands of the territory.  Where they will hear each other, smell each other, meet each other.  And because they are naturally cooperative, they will form a hunting group pair, threesome, and in time a pack, that is much more cooperative and efficient than the relict alpha population, staggering around on their last legs, famished but still snarling at each other rather than getting on with useful work.  But within the new TLC population will still be genes of dominance, ready for selection when food grows scarce again

The central point is that within individual wolves, and incidentally foxes, there is a developmental factor, the potential for genetic and epigenetic variability, that can be, and apparently has been, in the last 40,000 years, and probably much less, expressed as a bias towards co-existence with Homo sapiens.  Cooperation with humans was a potential present in the wolf organism.

Wolves that were proto-dogs were so because they were tolerant of, and over generations even attracted by, human beings.  But it takes two to tango, and it’s not so clear what humans saw in the proto-dog.  Maybe it is the human, particularly the juvenile human, tendency to neophilia, to explore new things.xplained so glibly.  Function must play a part.

The problem of explanation is one common in evolution.  It’s the transitional stage.  Once a complex system with two initially discrete components, as it might two hunting parties, one composed of dogs and one of humans; or indeed the system of a large-brained species such as humans co-existing in mutual dependency with their clothes, homes, tools, utensils and weapons; once these complex systems are up and running, it is easy to see how they work .  The intraspecific bit of the taming between dog and human, the combination of two mutually accommodating species that each gain advantage from the other, most certainly happened, since here we are together.  How the competition and mutual hostility between wolf and human made the transition to cooperation is harder to work out. The speculation has a limited development space.  It must not be teleological. There must be a functional advantage to both.   The possible time frame for the domestication of the dog coincides with the major wave of Homo sapiens spreading through Asia and Europe.  The transition took place in a cold landscape where the principal prey animals of both wolf and human, by biomass, were caribou.

The behaviour of the modern wolf is the nearest evidence we have of how the wolf ancestor to dogs might have behaved.  At the mythical level a sharp distinction between dog and wolf has been drawn over the evolutionary history of the taming; dog is archetypal friend to man, wolf archetypal foe.  This is marked in Eurasian folk literature and is with us to this day.

In the climate of the Palaeolithic when dogs emerged, the principle prey by biomass, and by use (meat, fur, antler), the caribou, migrated long distances between winter and summer feeding grounds, and humans as a result did the same.

Both wolves and humans are social hunters.  That is to say there is a complex of interdependent roles.  And many features of their total hunting repertoire would overlap.  They seem to be in direct, stable opposition and competition, as the man wolf archetype suggests.  In order that cooperation should emerge from this close competition, there must be an overall energy saving, which might be smooth or might be lumpy, that is it might bring in absolutely more energy for less effort, or it might raise the level of dangerous minima, times of potential extinction of the group through famine.  Either way , the taming must move towards a least energetic state compatible with homeostasis for the human/proto-dog combination, which is lower than the equivalent for each group on its own.

Wolves optimally hunt large mammals.  When the caribou are on the move, which they are most of the time, they follow them and watch for signs of weakness or panic.  They then run down chosen victims, which may change according to the course of the hunt.  They take various roles, chasing, heading off, distracting, bringing down, and killing.  They will utilise contingent bits of landscape, rocks, trees, cliffs, small round boulders over which the caribou are ill-equipped to run and become easily exhausted, gullies and uneven ice to check and turn and the prey with as little energy expenditure as possible.  The hunt economy, as well as minimising energy expended in running, can have bigger deficits.  The killing of a large mammal is a dangerous moment and wolves can easily be fatally wounded or killed by hooves or antlers which, if the pack is at an optimum size, is itself a serious energy depletion.

Upper Palaeolithic humans hunted, I guess, in much the same way.  There would have been a big overlap in technique.  So one must look at what specialisms each species brought to the deal.

Wolves had superior speed, wider field of vision, faster vision processing and better topographical navigation systems based on this speed, and also, possibly, better and faster-acting protocols for cooperation across short time durations.  (Their visual processing was not nearly as versatile as that of humans, who at that time could carve an image of the hunt on a sliver of prepared bone).

What wolves lacked was efficient killing ability, which was a danger area for them.  Otherwise they seem, as a species, well set-up; the present grey wold has survived to this day.  The present grey wolf survived those thirty or so thousand years.

And what do humans bring to the deal?

Wolves and caribou have around the same to speed.  Humans are the slowest runners.  Caribou protect themselves by keeping close to other caribou.  If threatened by the presence of a wolf pack their first  tendency is to move away, but they do so en masse, they are much bigger than wolves, in summer they have sharp hooves and they have big antlers.  A caribou can kill a wolf (our own red deer in our national parks can kill a human being, and do).  A wolf getting too close to a fleeing herd is liable to get a broken jaw or leg, soon fatal.  They have to wait for exhaustion, panic, and spatial vulnerability to make things easier.

What all three species have in common is the ability to cover ground fast, around the same speed, over long distances.  Performance wise they are all in the same sort of envelope, except that the human top speed, and I’m talking Usain Bolt here, is ten miles an hour slower than wolf or caribou.  A human being is not likely to catch a caribou with its bare hands.

What humans bring to the deal is their extended phenotype.  There are two parts to this that are relevant, technology and cosmology.

By the time of the mutual taming, our species had a lot of evolved extended phenotype.  It had enough art, painting, ceramic and stone and bone sculpture and relief work, music from bone flutes, even landscapes carved on bone that look like large scale maps, to provide an exhibition at the British Museum in 2013.   And they had weapons that could kill at a distance, the kind of thing that would be very useful to a wolf because it minimised the danger of the kill.  First, humans had an evolved physiology that could whip the hand up to speeds approaching ninety miles an hour.  Then they had projectile points stuck on sticks hooked to throwing sticks, which amplified the speed of missile launch.  You didn’t need to get within the reach of antlers and hooves in order to kill.

In addition to this humans had their cosmology.  This, the evidence is wide-ranging, was not just a bed-time story they told to the little ones.  As with the Australian songlines, cosmology was an evolved historical record, composed not of kings and battles but of, at the finest grain, of rocks, swamps and clouds, rain and stars, animals, insects, plants and rivers, fire, human beings, spirits, weapons, paintings and work in fibre, sculpture and aerial maps, canoes and kangaroos, each in and out of human brains in huge numbers of combinations and alliances, modulations of the total meaning space that has found homeostasis in a multi-media narrative.  It is an epistemological account, if lacking in detail, an account of how they knew things, not as lone things, as a kangaroo or a caribou or a wolf or a projectile point, but in huge intermodulating complexes.

This cosmology is not just an evolved historical record.  It is the ancient and durable account of how things are and how they work, and therefore it is an analytical and predicative instrument.  It has survived because it is a record of how to survive, not for an hour or a day but over tens of millennia, in an environment, rich in nutrition it is true, but with hostile maxima and minima of weather and nutrition which will kill, in days or sometimes in hours, a group without the knowledge of the Songlines.

And those competernces are what the humans will have brought to the deal with the wolves.  Wolves are opportunist hunters, following the herds, approaching them, making them anxious, pursuing them to panic, exhaustion and chaos, isolating and killing.  Man 30,000 years ago did much the same, but the evolved cosmology gave him a space-time chart that was with him always, a map of the land and the seasons.  If, at that time, you asked a wolf when the caribou would start to move north, or how long they would move for, or where they would stop and turn again, a wolf wouldn’t know what you were talking about.  A Palaeolithic human would not answer you in a way which was familiar.  They would regard the wolf with an entirely different intellectual apparatus.  The animal’s ontology would come across as ambivalent.  It would be in some senses an enemy, but in others a figure of respect.  One can infer from the modern wolf that the relationship of human and wolf was dual.  Honour, respect, correct manners and close cooperation were features of both societies.  And both were hunter-killers.  The old dog-ancestor wolf and Eurasian upper Palaeolithic humans would have existed in the same tundra landscape, among the same fauna; not just caribou but hare, fox, elk, bison and mammoth.  In winter the humans and caribou are clad in the same skins and two-layer fur, a fine thick undercoat next to the skin (for the humans this would be a second skin), and an upper layer of long hollow fibres, like quills.  The wolf hair is of the same two part type, the outer guard hairs particularly good at shedding ice, which is why the human hunters may have wolf pelt trims around the borders of their hoods to avoid their breath freezing out on the fibres.  Thus, more or less uniformly insulated in fur from the freezing air, this triarchy moves across the space time-continuum, wolf, human, reindeer, a stereotypical Christmas card or, since that particular tripartite deity was not to appear for another twenty or so millennia, a boreal universal White Moon Amanita card.

Which is all well and good, but it doesn’t account for how the dog came to be.

So: the function of the big human brain is to accumulate for yourself and your descendants as much extended human phenotype as possible.  The human extended phenotype thirty thousand years ago was large, but hardly the minutest fraction of what it is today.  Yet it is a feature of the architecture of the human brain in a healthy ecology that it registers and processes everything available.  We may feel, because we are more informed by quasi-infinite feeds of news, opinion and information generally, that our brains have and use massively greater capacity than the hunter gatherers’, but I mix with modern humans on a daily basis and I’m pretty sure this is a delusion.  It is true that the content of my ideoverse and the content of the ideoverse of a Palaeolithic caribou/reindeer hunter would be very different.  Their insentient material culture, stone, bone, antler, wood and hide and amanita, meat and fire and spears and berries dried, masks and other objective loci of rites, would take up a lot of meaning space, but so would the landscape and its inhabitants.  Animals would play a big part, particularly those they hunted and those they hunted alongside.  Wolves were a cosmological  conformation of tooth and fur and speed and ferocity, but alongside that also a confirmation of well ordered life in lupine camp and in the hunt.  They would  already have had a considerable presence in the paleo-meaning-space long before some of them became dogs.

Still, how?

There are various notions.  The first is that wolves kind of hung around camp fires and humans tolerated them so long as they didn’t come too close because to continually chase them off was a waste of energy.  Wolves have long canine teeth evolved to rip the perineum out of large mammals so that they bled to death, but they also have molars and jaws powerful enough to crack their main bones and get out the marrow.  But humans used stone tools for the same purpose, so the likelihood of humans tossing the wolves whose eyes glowed with the reflected gleam beyond the firelight the odd bone to keep them happy is slight.  I can see the picture, but I can’t see how we move on from there.

There is the neophiliac plasticity of young of both species which might attract them to each other, on the same lines as the tapir became a sporadic part of village life with the anaconda people.  It is plausible.  It has a lot of anecdotal stuff going for it. Except that I don’t think their parents would have allowed it.  I think both sets of offspring would be severely discouraged from playing with alien and dangerous neighbours, however much you might respect them for certain qualities you have in common.  Wolf pups and children, however much freedom they were given to play, however much cheek they were permitted to adults that would not be tolerated in low status adults themselves, would have been guided into superordinate protocols upon with the homeostasis and survival of the group depended.  These would have prioritised not getting killed or injured, and watching and learning so as to become an actively and appropriately useful co-operator in the optimally functioning group.  It would not have included hobnobbing with trans-specific bad influences.  I guess the caste prejudice might have been stronger on the wolf side than on the human.

Other-species familiarisation with the human organism is not unknown.  There is sequence on You-Tube.  Two elderly men, tourists not scientists by the look of them,  are sitting on the edge of a camp somewhere in the central African rain forest.  Two mature male gorillas shamble up with three adolescents.  They approach the men.  The adults stop about three metres from the men, and squat.  They allow the adolescents to go forward, touch the men, feel their clothes and skin, take off a hat to look underneath.  After a couple of minutes the adults grunt and gesture.  The adolescents go back to them and make as if to walk off down the path but one of the adults, with the polite arm gesture of an official who points out that the exit to an art gallery is not the way you came in, diverts them off the path and into the forest.  The End.  It’s lovely to watch, and makes you proud to be a hominid.  What it means scientifically, on You-Tube, is less certain, but as I’ve already suggested, while anthropomorphising other species can be overdone, theriomorphising humans is probably corrective.

Adventitious correlation of hunting practice is not unknown in nature. With wolves and humans, both species hunt, and in a manner that is concordant.  What is more, young wolves, or at least young grey wolves, when they are first taken on a hunt, just watch from a distance, and only gradually physically explore the roles of fast runner, balker, distractor, topography exploiter, perineum ripper and, if particularly large and powerful, muzzle clamper and suffocator.  We can assume that the same pattern applied to humans, though anthropological evidence would suggest that, unlike with wolves, hunting was a male only occupation.  Extended phenotype was to blame, perhaps.  Once the by then obligate clothing had emerged (unnecessary in the African cradle of Homo sapiens, indispensable in the ice-age tundra), and cooking, and fire, somebody had to manage the complex technology of domestic life.  There is an argument that early man, as a sex, traded meat for copulation.  The evidence of my grandparents’ generation suggests rather that they traded meat for a share in the outcomes of the technology of home, even when home lacked a roof, and may well have had to earn sex in much the way we do, by charm and status, good grooming and good humour, and excellent manual skills.  A big penis, valuable as a display item where the mean temperature is 25οC, is less so in the icy wastes under a couple of inches of hide and fur.  Of course nowadays the technology of the home is, hopefully, as much the burden of men as women, at least in northern climes.

So there in the tundra, still, we have the caribou and, following them at a short distance, the wolves and the humans.  If increasing anxiety, then fear, has accelerated the caribou herd to a headlong rush, and two predators in its path check this charge slightly, causing an energy draining eddy in the caribou ranks, this small loss for the collective prey, gain for the collective predators, will be one small factor in the economy which, when finally summed, will lead to the outcome, a kill or not a kill.  It doesn’t much matter in the heat of the moment whether the diverters are wolves who have cut across the rocky shorter distance between two points to get in front of the herd, or humans who have used the larger spatio-temporal metaversal map at their disposal to take a short cut at dawn in the expectation that that was the way the caribou would go.  It might even be a man and a wolf who had got there at the same time by their different journeys and could see advantage in each other’s presence as long as each kept a seemly distance.  If we can imagine a proliferation of instances like this, where certain roles are confined to one species, and others distributed among both, we get a pattern of the hunt with men and dogs which emerged millennia later.

There is still the problem of the kill.  If one of the advantages for wolves hunting with men was that men with projectiles could kill at a distance and this saved the wolves the risk of physical damage by hoof or antler, then for cooperation to be of any use the men would have to share the kill with the wolves, and this seems unlikely.  It would only be in the pre-kill stages of the hunt that the interspecific fluidity of certain roles would benefit both, and each would make their separate kills.  But there might have been occasions when, with a group of caribou cut off from the main herd and corralled not only by relatively static men with spears but behind them fast moving wolves casting a virtual net across lines of escape— a map capable of realisation in the human metaverse— the kill would be enhanced, a slaughter, a pile of dead; hecatombs for the sacrifice.  And evolved in the cosmology of the men was already a totemic wolf worthy of respect, probably one with a mythically shared phylogeny with men, though not perceived as such in the Darwinian sense, more in the sense of the Amazonian hunters who were descended from the ancestral anaconda or early North Americans descended from the Salmon People.

In this situation, some sort of deposition of the surplus kill, that benefited the wolves, might seem politic.  It would be better determined in a cosmological context, for several reasons.  First, it would probably be unpopular at a demotic level: “You’re not giving our meat to them fucking vermin, mate.”  The average shaman probably wasn’t possessed of  the dullest intellect of the group by any means, and their ideoverses would have developed along a phylogenetic continuum that was at variance with that of the, let us happily stereotype, the gorilla-muscled spear chucker, the one who if he was teleologically endowed enough could invent American Football on the spot.  Within the average shaman’s cosmology there would long have been a development space where proto-votive offering was foregrounded because it had a totally opaque but functional differential advantage.

The second reason is that protocols for rewarding the wolves for their part in the hunt would probably have evolved as functionally effective  rituals.  Most animals and birds are genetically biased towards ritual behaviour, display, mating, taking over at the nest, and such evolved behaviour is the material spatio-temporal locus of all intraspecific interaction.  It can also arise interspecifically, though usually in displays of aggression.  In fact ritual behaviour has evolved from hardwired signing behaviour, the twitching tail, the angle of the ears, that has become part of a usually intraspecific process, what we would call in natural language a social process, such as mating, or greeting.  I read a long time ago an account of an incident where a man who worked with non-wild wolves was talking about wolf etiquette.  One day he’d gone into the enclosure of a big male wolf with which he got on well.  He had gone in to fix a loose fence rail.  He had just bent down to see what needed doing when he was knocked off his feet with tremendous force by a forty five kilogram bolt of fur muscle and bone.  The breath knocked out of his body, doubled up on the ground, he expected the worst, but when he opened his eyes he saw the wolf stalking back to the place where it was doing whatever it had been doing before.  He realised his error.  Carrying tools, absorbed in his task, he had forgotten to make the physical, up close greeting ceremony with this male with a very high sense of self-regard.  He staggered to his feet, went across and did so, and the wolf reciprocated as if nothing disorderly whatsoever had transpired a few moments before.  Protocols, protocols.  Look at our great statesmen on public occasions, and remember baboons.

I’m not suggesting anything teleological.  I’m not suggesting that the shaman might have overridden demotic short-term speciesism because he could foresee that ritual deposition of votive flesh would be long-term functionally better than random chucking of scraps, which produces ambiguity and no dog-trainer would approve of.   I’m merely suggesting that such ritual behaviour would have evolved, with maybe the shaman and his tendency as a significant locus, because its long-term functionality was a differential advantage to both species.

And thus today all well brought up dogs cohabiting with well brought up humans are still ritually fed.  “Sit.  No, no sit. SIT…Oh, alright then.”

This seems to me the most probable way it happened, but there’s no data to work with, so I’m not sure.


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