Archive for the ‘dog domestication’ Category

hooded vs carrion crow

In Europe, there are crows. The most famous of which is the carrion crow, which looks and behaves quite like the American crow.  It is usually black, fairly omnivorous, and it often regarded as an agricultural pest.

At one point, it was believed that carrion crows existed in two distinct phases, the common all black form, which is common in Western Europe, and the phase that is marked with gray on the neck and body.  This form exists in the northern parts of the British Isles, where it has been called the grey grow, the hoodie crow, or the hooded crow. This form is found in Northern Europe, Central Europe, and the Middle East.

However, ornithologists began to notice that where their ranges overlapped, it was quite unusual to see hooded crows paired off with black carrion crows, but the traditional taxonomy still thought of the hooded and black forms as being distinct phases of the same species.

Because the two forms were rarely seen mating or paired off, it was decided to call the carrion crow and hooded crow distinct species, and this is the current understanding. The hooded crow is Corvus cornix and the carrion crow is Corvus corone. But the two birds are otherwise quite similar in terms of ecology, vocalizations, and general phenotype. All that really separates them is the coloration.

Scientists that surely there was a rather deep genetic divergence between the two species, which is why there is a species barrier between the two forms of similar crow.

However, when the genomes of carrion and hooded crows were sequenced, it was revealed that they were almost identical.

Less than .28 percent of the genome varied.  That variance was related to the fact that carrion crows have gray plumage on their neck and torso.

But that little variance is enough to create a species barrier between the carrion and hooded crows. Birds are highly visual, and when young crows imprint upon their parents, they imprint heavily upon what they see. If a young crow is raised by parents that are gray hooded, they will look for mates that are gray hooded. IF their parents are all black, they will look for mates that are all black.

However, it has also been suggested that these crows look for mates that appear not to have aberrant mutations, and this keeps the crows looking for mates that generally look like those belonging to their general family group and social circle.

Whatever the case, we have two very closely related species that do not hybridize. They probably became distinct during the heavily glaciation cycles of the late Pleistocene. One form evolved a gray hood and one evolved an all-black form. Maybe founder effect is the only real reason for this difference in plumage, for this difference in plumage is awfully random.

But that difference in plumage color is enough to create a species barrier, which, if it holds, will lead to greater and greater speciation between hooded and carrion crows.

This discovery about crows is quite interesting for what it tells us about dog taxonomy. Domestic dogs and wolves live together over a broad swathe of Eurasia, and for many centuries, dogs and wolves were regarded as distinct species. However, we have recently found that there is an extensive gene flow between Eurasian gray wolves and domestic dogs across Eurasia, and this gene flow is so significant that the majority of Eurasian gray wolves are estimated to have some relatively close dog ancestry.

Carrion and hooded crows have a clear species barrier that is likely only going to intensify as the two lineages continue to diverge with very limited gene flow. Dogs and gray wolves are not experiencing such a species barrier. Indeed, it looks like the gene flow between dogs and wolves is only going to increase as wolves move into human-dominated lands in Western Europe, and the Eurasian dog population continues to increase along with the human population.

So here we have two crows that are diverging, and the wild and domestic forms of Canis lupus that are continuing their gene flow.  Closing down gene flow is a major part of speciation, and the crows are clearly on their way.


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feral dog

Most feral domestic animals revert to a form that is roughly similar to their wild ancestor. You can see this quite dramatically in feral pigs. They generally evolve into a form that is about the same size and even coat type of the Eurasian wild boar. City pigeons look very much like the rock dove or “rock pigeon” that is their wild ancestor after just a few generations of breeding without human care.

Because village and pariah dogs tend to be mid-sized, it has been assumed that the wild ancestor of these dogs surely would have been on the smaller side as well. Therefore, the gray wolf simply could not have been an ancestor.

What actually drives the size of freely breeding and feral domestic dogs isn’t that they have some ancient alleles that force them into returning to an ancestral form.  The truth of the matter is that ecological niche and caloric restraints have a lot more to do with this phenomenon.

Dogs are unique among domestic animals in that they are the only domesticated form of large carnivoran. We have never domesticated any other species of large predatory mammal except for those Pleistocene Eurasian wolves that are at the base of domestic dogs.

Most domestic dogs are poorly adapted to living as predators, and they really don’t have to be. When dogs go feral in societies with extensive agriculture, they readily scavenge and hunt small prey. They dabble in various levels of omnivory.  Some dogs might be good at hunting deer, but deer are a lot harder to catch than garbage and groundhogs.

There is an extensive literature on mammal predator size and prey choice. The best known researcher looking at these issues is Chris Carbone, and in a 2007 paper called “The Costs of Carnivory,” which was published in POLS Biology, Carbone and colleagues looked at body mass of mammalian predators and their prey choices. If a predator weighed more than 20 kg, it hunted large vertebrates. If it weighed less than that weight, it hunted invertebrates or small vertebrates.

Larger predators get a much higher net energy gain by targeting large prey, and this large prey allows them to maintain their larger body size.

Feral and freely-breeding domestic dogs are not hunting large vertebrates. It is much easier for them to scavenge as mid-sized creatures. Natural selection would favor a moderate size, because any dogs that retained the large dog or large wolf alleles in the population would have a harder time feeding itself efficiently on these resources alone.

There are, of course, exceptions to this rule. In Uruguay, there was a population of feral mastiff-type dogs, which are called Cimarrón Uruguayo. These dogs were introduced by Europeans as working dogs, but some of them went feral. They were able to maintain their large size because they hunted livestock and game, and they were such a problem that the government placed bounties on them.  These dogs were living in a feral existence for at least 250 years, but they were able to retain a large mastiff phenotype. The feral mastiff is now being transformed into a standard breed.

However, the general rule is that village and pariah dogs tend to be significantly smaller than wolves, but this smaller size cannot be used to deny that dogs are derived from gray wolves. This smaller size is just more efficient for the ecological niche of feral and village dogs.

And it is poor reasoning to assume that dogs cannot be wolf derivatives simply because they do not evolve back into a wolfish form once they go feral.  Dogs have been domesticated for a long time, and their domestication is quite unique.  As the only large predator we have domesticated, ecological pressures create a different sort of animal than the original wild ancestor.

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4,000 year old dog

The skull of a 4,000 year-old dog has been used to reconstruct what its head may have looked like.

The dog was quite wolf-like and described by the BBC as being about “the size of a large collie.” In the UK, “collie” almost always refers to border collies, so this dog probably would have been on the large side of a medium-sized dog.

The dog’s skull was found at Cuween Hill Chambered Cairn on on Mainland, Orkney, off the northern coast of Scotland. 3D images of the skull were use to make the reconstruction, which was created in clay. The artist then made the fur look like that of a European wolf, which is not entirely unreasonable given the morphology of the skull itself.

The dog may have been used to tend sheep or guard settlements. It was clearly a respected creature in the society that interred in the tomb. Maybe it was a valued working animal or simply a totem of its people.

Whatever it was, it was clearly more wolf-like that one might have expected from a dog from this late a date. The Ancient Egyptians, who were contemporaries of these Orkney cairn tomb builders, were already breeding dogs that were quite distinct from wolves.

But the truth of the matter is that this dog was significantly smaller than most modern and contemporary European wolves, and the mainland of Scotland was full of wolves that were probably still interbreeding with domestic dogs on occasion at this time.

So the Orkney Islanders from 4,000 years ago clearly had dogs, but I imagine this dog as being something like a Norwegian elkhound, a laika, or one of the old German herding dogs, like the Thuringian sheepdog.

I would love to see more reconstructions from ancient dogs skulls.  I would love to see the Goyet Cave canid and the Razboinichya Cave “dog” undergo a similar reconstruction.

Yes, this is art, but it is art that is informed through science.


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Humans and the various canids belonging to gray wolf species complex possess the most complex relationship of any two beings currently living on this earth.  At one point, they are our cherished companions, often closer to us than we ever could be with other people, and on another point, they are the reviled predators that might take a child in the night.

We have clearly defined relationships with other predators. Leopards and cougars, well, we might hunt them for sport or photograph them in the wild. But we never become closely aligned with them, except for those eccentrics who dare to keep such dangerous predators as pets.

People living in the Eurasian Pleistocene brought some wolves into their societies.  Wolves and humans should have been competitors. We should have had the same relationship with each other as spotted hyenas and lions do in Africa now.  But at some point, humans allowed wolves in.

Raymond Pierotti and Brandy Fogg demonstrate that many humans throughout the world have had some kind of relationship with wolves. In some cases, it is or was a hunting symbiosis. In others, they were totemic animals.

In their work, Pierotti and Fogg contend that the relationship between humans and wolves broke down with the rise of Christianity in the West. I don’t think that’s when it broke down. It started to become complex when humans began to herd sheep and goats.

In Kazakhstan, wolves are hunted and revered at the same time. The Kazakh people herd  livestock, so they must always worry about wolf predation. Stephen Bodio documents this complex understanding of wolves in his The Hounds of Heaven.

“They hunt them, kill them, chase them with hounds and even eagles, take puppies and rear them live, identify with them, make war on them, and claim descent from them,” writes Bodio. This description sort of fits modern humanity’s entire relationship with this gray wolf complex. We pretty much have done and continue do almost all of these things.

Wolves, coyotes, and dingoes have killed people. So have domestic dogs. In the French countryside, wolf hunts were considered a necessity to protect human life, largely because has the longest and best documented history of wolves hunting people. The dispossession of rural peasants and the depletion of game in the forests created conditions where wolves would consider humans easy prey.  Lots of European countries have similar stories. And when Europeans came to North America, they knew about the dangerous nature of wolves, even if they had never even seen one themselves.

Humans have declared war on wolves in Eurasia and in North America. The wolf is extirpated from much of its former range in Europe. They live only over a limited range in the lower 48 of the United States.

Man fought the coyote with the same venom and lead he threw at the wolf. The coyote’s flexible biology and social behavior meant that all that effort would come for naught.  The coyotes got slaughtered, but they rebounded. And then some. And the excess coyote pups found new habitat opened up with big ol’ wolves gone, and they have conquered a continent, while we continue our flinging of lead and setting of traps.

In Victorian times, Western man elevated the domestic dog to levels not seen for a domestic animals. They became sentient servants, beloved friends, animals that deserve humanity’s best treatment.

And in the modern era, where fewer and fewer Westerners are having children, the dog has come to replace the child in the household. Billions of dollars are spent on dog accessories and food in the West.  Large sectors of our agriculture are ultimately being used to feed our sacred creatures.

A vast cultural divide has come to the fore as humans realize that wolves and coyotes are the dog’s wild kin. Wolves have become avatars for wilderness and conservation, and coyotes have become the wolves you might see out your front window.

Millions of Americans want to see the wolf and the coyote protected in some way. Dogs of nature, that’s the way they see them.

The rancher and the big game hunter see both as robbers taking away a bit of their livelihood. Humans are lions. The canids are the spotted hyenas. And their only natural state is at enmity.

Mankind’s relationship to these beings is so strangely complex. It greatly mirrors our relationship towards each other. We can be loving and generous with members of our own species. We can also be racist and bigoted and hateful. We can make death camps as easily as we can make functioning welfare states.

And these animals relationships with each other are just as complex. Wolves usually kill dogs and coyotes they find roaming their territories. But sometimes, they don’t. Sometimes, they become friends, even mates.  Hounds can be trained to run down a coyote, but sometimes, the coyote and the dog become lovers in the forest.

Social, opportunistic predators that exist at this level of success are going to be a series of contradictions. Dogs, wolves, and coyotes certainly are. And so are we.

It is what we both do. And always will.

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One of the great exercises on the internet among those who wish to be taken seriously as “dog people” is to say that dogs are not wolves. In one sense, they are quite right. Dogs are not wild canids, and they are certainly not the mostly fearful and reactive wolves of the middle latitudes of Eurasia and North America.

But in another broader sense, they are dead wrong. I’ve been following this debate for some time. At one time, there was a great emphasis on the so-called Canis variabilis that were contemporaries with Homo erectus at the Zhoukoudian cave system in China. The remains date to 500,000 years ago, and it’s quite a leap to say that Homo erectus began dog domestication.

It should be noted now that Canis variabilis is no longer an accepted scientific name for these early wolves. They have since been reclassified as a subspecies of the Mosbach wolf (Canis mosbachensis). Their new name is Canis mosbachensis variabilis, and although the Mosbach wolf is ancestral to the modern gray wolf, the Chinese subspecies is now not regarded as leading to the modern one.

So this idea that these Chinese specimens are ancestral to the domestic dog is quite faulty. Even if we were to say that Canis mosbachensis were the ancestor of dogs, we would have a real problem on our hands. The Mosbach wolf disappears from the Eurasian fossil record no later than 300,000 years ago, when it was replaced by modern gray wolves. The earliest domestic dog that has been proposed dates to 33,000 years ago in the Altai Mountains.

Somehow, you have to get a species that went extinct hundreds of thousands of years before the formation of the earliest domestic dog to become its ancestor. The chronology makes no sense.

Now, we do have some ancient mitochondrial DNA of a Siberian Canis cf. variabilis that appeared to show a connection with the origins of the domestic dog. This specimen is probably a ate surviving Siberian variant of the Mosbach wolf, and it is possible that the reason for this mitochondrial DNA similarity is that domestic dogs have a mitochondrial DNA lineage that very close to this extinct wolf. The real problem with this study is it is a mitochondrial DNA study, and if we could somehow get a full genome comparison from these remain, which would not be easy, then we could get a better picture of how the Mosbach wolf relates to wolves, domestic dogs, and coyotes. Yes, the discovery that gray wolves and coyotes shared a common ancestor only around 50,000 years ago means that coyotes descend from the Mosbach wolf as well.

So when you see someone claiming that Canis variabilis is wild Canis familiaris, just understand that this person hasn’t looked at the most recent literature on these Middle Pleistocene wolves. But I’ve seen this repeated enough that I do think I need a place on this blog where I can easily link to the problems with this assertion

The real problem with all of this is that in dogs, at least in the English speaking world, there is a real problem with phylogeny denial. So many people are caught up in this “dogs are not wolves” idea that they invest lots of mental gymnastics in trying to create another wild ancestor for the domestic dog.

So many people got worked up with the discovery that no extant population of gray wolf is ancestral to the domestic dog that they had to make it about how dogs were not derived from wolves.

Again, the gray wolf species is at least 300,000 years old, and no one has found a relationship between dogs and wolves that posits their divergence as being greater than 33,000 years. There is an old mitochondrial DNA estimate that is largely not accepted that puts their split between dogs and wolves at something like 135,000 years ago, but that’s still after the gray wolf existed as a species.

So let’s talk about why saying dogs are not wolves is an exercise in phylogeny denial:

One of the implications of our modern Darwinian synthesis is monophyletic descent. All organisms derive from ancestors, and it is impossible to evolve outside one’s ancestry. If we were to go back in time to see when the most recent common ancestor of dogs and gray wolves, you would have a hard time describing that ancestor as anything other than a form of Canis lupus.

Dogs have evolved through their Canis lupus ancestor, just as modern wolves have evolved through theirs. It is accurate to say that domestic dogs are not derived from extant wolves, but it is not accurate to say that dogs did not derive from wolves. It is also not accurate to say that dogs are a different species from Canis lupus, because dogs are still part of a Canis lupus lineage.

Further, we have lots of data about the extensive gene flow between dogs and wolves in Eurasia. We know that livestock guardian dogs in the Republic of Georgia have exchanged genes fairly extensively with wolves. But we now have data that shows an extensive gene flow between domestic dogs and wolves across Eurasia.

So dogs and wolves are continuing to exchange genes. They are not becoming reproductively isolated from each other in a way that would lead to speciation, even now.

I’ve never understood why this line of thinking has ever been popular, except that wolves people have indeed abused dogs under the assumption that their social systems are much like those of captive wolves. Further, it is quackery of the worst order to assume that dogs should be fed only full raw carcasses of meat because that is what wolves eat.

But those problems are not adequately addressed by promoting another scientifically dubious prospect. Dogs do behave somewhat differently from wolves, but that is because dogs are domesticated. Wolves behave differently because they are a wild form, and as a wild form, they have undergone a selection for extreme timidity and wariness as we have tried to wipe wolves off the face of the earth.

The argument that dogs are part of Canis lupus is well-supported by science. Indeed, an analysis of gray wolf, domestic dog, and dingo genomes revealed that creating a separate species for the dog, the dingo, or for both would make the entire species polyphyletic and thus not in keeping with Neo-Darwinian principles.

So it is scientifically correct to say that dogs are wolves, but one should say that dogs are domesticated wolves. And just leave it at that.

And drop the phylogeny denial.

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Mark Derr has been writing critiques of the Coppinger model of dog domestication for decades. His best known work on the subject matter is How the Dog Became the Dog, but his ideas can be read in a more succinct place at this article at The Bark (but do buy the book!).

Derr recently posted about Jung and Pörtl’s “Active Social Domestication Model” on his blog at Psychology Today.

His analysis is worth your time to read. He largely agrees with this model, but he contends that it needs to be placed in a larger framework of Derr’s own work and that on Wolfgang Schleidt and Michael Shalter, which is linked on Jung’s website.

Derr contends that Jung and Pörtl’s ideas need to be placed in this line of scholarship to which I’d also add the work of Darcy Morey and Pat Shipman. They don’t agree with each other on some important particulars, and they may have quibbles with this new model. But both scholars have been publishing outside of the Coppinger model for quite some time.

What I find most interesting is how this scholarship is pretty well-known outside of the English-speaking countries, but in the US and the UK, the generally accepted model by virtually everyone is the Coppinger model. Part of it may be that North America is home to wolves that are not particularly admixed with dogs. Indeed, they are probably the most “pure” wild wolves anywhere in the range of Canis lupus. And Great Britain and Ireland have no wolves, and there is no chance of any wolves coming back to those islands, at least by their own volition.

However, in Germany and Russia, wolves are admixed with dogs, and in the case of Germany, it is not at all impossible to see wolves living near large urban centers.

So they have a much more practical understanding of what it is like to live near wolves that have quite a bit of gene flow from domestic dogs, and they are less likely to buy into models that see dog and wolf as fundamentally distinct entities.

North Americans are much more accepting of a dichotomy model, and we have a hard time with gene flow between Canis populations. Our laws want hard and fast species, but the thing about Canis is that none of them are hard and fast species.

So it is easy for North Americans to posit that wolves are unable to be domesticated, because modern North American wolves (for the most part) are reactive and timid predators that do kill both dogs and coyotes they find on the trail. They do, but they also do the same with other wolves. And sometimes they mate with those wolves, just as they will mate with dogs and coyotes.

The Coppinger model requires an assumption that all wolves living in history and in the present are these shy timid ones, but that’s not what the historical record shows. And it is certainly not what is seen on Ellesmere or Baffin Island, where the wolves have never been persecuted by man.

The Coppinger model requires us to create the gray wolf as a Neanderthal dog in which it is big in size, big-brained, and meant to hunt only large prey, and posit the dog as the modern human with a smaller brain and more flexible diet.

We need a model that can place the origin of dogs before the Mesolithic, and this Active Social Model goes along way in that direction.

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I finally got around to watching Alpha, a film that depicts a fictionalized account of dog domestication. It is set 20,000 years ago in Europe, which means that it posits a European origin for the domestic dog, and because of this early date, it sets domestication in the time of Paleolithic hunter-gatherers.

The events in the film are backed in at least some of the literature on dog domestication. The oldest dog subfossil remains in existence that are not hotly debated as to belonging to a dog and not a wolf are the Bonn-Oberkassel dog, which dates to around 14,000 years ago. Yes, that’s about 6,000 years later than the events in Alpha, but we do have that paper positing European origins for the domestic dog that suggests an even older domestication event on that continent.

I should note right now that I remain agnostic whether dogs originated in Europe or Central Asia. Both are likely candidates based upon the best genetic data. I am open to there being two domestication “events,” but other than that one well-publicized paper, this hypothesis has not been well-received.

So it is potentially possible for the events in Alpha to be in keeping with some of the literature on dog domestication.

The film’s protagonist is Keda, who speaks a language that is called Solutrean, which is the name archaeologists have given to an industry culture in some parts of Paleolithic Europe. Because of the similarity between spearpoints from this Solutrean culture and some aboriginal North American peoples from roughly the same time period, some experts posited that European hunter-gatherers came across the North Atlantic following marine mammals along the ice flows, suggesting a European origin for some North American native people. Of course, this hypothesis has been dumped in light molecular data that show a Siberian origin for the indigenous people of the Americas.

Keda is the son of a Solutrean chief who wants his son to be a powerful and skilled hunter of big game, especially steppe bison, which are his people’s predominant prey species. Keda, though, is a bit soft with quarry. He refuses to finish off a wild boar that has been speared, and his father takes great umbrage with his temerity when it comes to the kill. But this is Keda’s first big hunting expedition, and his father really wants to teach his son the ways of his craft.

This world is not meant for those who cannot be crafty and tough. While sitting at a campfire, a cave lion absconds with one of the men. A memorial cairn is placed where the lion took the man, and they move on in search of meat in the hoof in a way that shows they accept their mortal fates in such a hostile environment. So Keda’s softness and excessive empathy might not be well-matched for such a lifestyle.

The hunting expedition reaches its climax when tribesmen and their close allies come across a herd of steppe bison. They stalk them in close, and they use their spear throws to drive them towards a cliff. Many fall off the cliff and die, and the Solutrean tribes can have their meat and hides and bones for another year. However, as they drive the bison off the cliff, one unusually recalcitrant one charges Keda and lifts him up with its horns and throws him off the cliff. He falls to a rock outcropping, where the tribe is forced to leave him for dead. A cairn is left in his memory on the cliff face, and his father and the hunters leave him to the ages.

It is at this point that film begins to play around with artistic license. Keda awakes when a griffon-type vulture tries to scavenge his carcass, and when Keda realizes that he is just resting on a ledge, he tries to climb down the steep cliff side with his badly injured leg. At that point, it starts to rain and the ravine below the cliff fills with water. He then jumps into the torrent below. Miraculously, he survives the fall and the rushing water, which I found a bit implausible. He then put his wounded leg back in order and begins looking around for a bit of food and shelter.

While he limps around, a pack of wolves shows up, but he manages to fight them off with the spearheads he’s carrying on his clothing and escape in a tree. He badly injures one of the wolves, which lies wounded beneath the tree.

I should note here that I am quite skeptical that Pleistocene wolves were particularly dangerous to people. As I’ve noted before, the wolves of Ellesmere and Baffin Island, which have never been intensely persecuted, are unusually curious and socially open with people. It seems to me that Eurasian wolves living 20,000 years ago would have had a similar curiosity about people, and thus, they would have been relatively easy for people to habituate them to our presence. I don’t think that an animal that was a much a threat to people as cave lions would have been trusted at all, and there would have been little of the empathy between people and wolves that could have led to any kind of partnership.

The empathy that Keda showed the boar, though, begins to cast upon the injured wolf. He muzzles the wolf and carries it to a cave, where he puts maggots on its wound to eat out any infection. He feeds the wolf a bit of the rabbit he manages to kill, and although the wolf is growling and surly through their initial interactions, the wolf eventually comes to trust Keda and within just a few scenes becomes as tame as any dog. Again, this is artistic license, but I think even those socially open wolves from the Pleistocene would still have had clearly-defined boundaries. They might have been friendly with people, but it is unlikely that they would have become that trusting at the first point of contact.

However, this part of the film does fit nicely with the recently posited Active Social Domestication Model in which human social interactions with wolves are the main catalyst behind creating the domestic dog. This model is relatively new, but it has a lot of explanatory power, especially when compared to the Coppinger Model, which just posits that scavenging wolves in at the middens Neolithic camps begat neotenic village dogs that were later selected for their working and hunting abilities and became the breeds of dog we have today.

Keda eventually realizes that the wolf is not going to leave his side, so he begins to show it even more empathy. He gets an idea that he could train this wolf to help him the hunt, and there are several scenes where he teaches the wolf to do commands. I doubt that this happened very much early on the domestication of the wolf, especially if we are to assume that humans and wolves developed a hunting symbiosis.

Indeed, the way Keda hunts with the wolf Is awfully unlikely. He uses the wolf to herd and flush wild boar, which he kills with a spear. Wolves can hunt wild boar that way, but it seems to me that the best way to use a wolf is as a “bay dog,” in which the wolf circles and distracts the quarry, holding it in one place so that a spear can be thrown.

Keda names the wolf “Alpha,” though he uses the Solutrean word for the term, and the two begin their journey to Keda’s home grounds. On their way, the man and wolf develop a tight bond. They play at the lake together, just the same way we would with our own dogs.

One night, a pack of wolves shows up at Keda’s campsite, a black wolf lures Alpha away from Keda. The black coloration is interesting, because it is quite rare in modern European wolves. It has only been introduced to wild wolves in modern times through crossing with dogs, including in North America where it was introduced to wolves from a dog living in either the Yukon or Northwest territories several thousand years ago. What we do know, though, is that this black coloration, which is conferred in wolves and is most dog breeds through a dominant allele, originated first in the population that led to domestic dogs.

As Keda makes his way back, the Pleistocene winter sets in. He now must travel without much food through the driving, blinding snow. Through a series of misadventures, he finds himself falling through the ice on a lake, and Alpha hears his distress and runs to rescue. The whole scene where the wolf comes to rescue Keda from the ice requires lots of artistic license, but it is visually spectacular.

Alpha and Keda are reunited and begin their long journey back to the home grounds. On their way, a pack of cave hyenas (a type of Pleistocene spotted hyena) chases them into a cave. Alpha and Keda await the hyenas in the cave. The snow piled up on the entrance of the cave quakes with their footsteps, but they do not enter. We soon learn the reason for their reluctance, a cave lion has been lurking deep with in the recesses of the cavern. It charges Keda and Alpha. Alpha fights the lion bravely, and Keda takes a deep breath and throws the spear, hoping it hits only the lion. It does, and Alpha survives, though pretty badly wounded. The lion provides some sustenance, and Keda and Alpha continue to make their way to the home grounds.

When Keda arrives home, he is quite ill, and he shows up with a very sick wolf on his mother and father’s doorstep. I can only imagine what it would have been like if such a scene had occurred in real life. If these people had no concept of partnering with a wolf, I bet their first compulsion would have been to club the wolf in the head for meat and fur. But Keda tells them they must care for Alpha too.

In the final scene, Keda is healed from his illness, and he sees a neonatal wolf puppy being held up by a priestess in an induction ceremony to the tribe. We then see Alpha nursing a litter of puppies. So, it turns out that after 90 minutes of film, we learn that Alpha is a female wolf. I wish there had been some mention the wolf’s sex earlier in the film, because I was honestly not prepared for the puppies.

The father of the puppies was apparently the black wolf, because one of the puppies is black, and thus, this detail would fit with the black coloration originating in the population that led to domestic dogs. The puppies grow up in the Solutrean tribe, and the final image of the film is the Solutrean hunting party going out on a hunting expedition with their wolves walking among them.

I will give this film props for doing quite a bit of research on some of the literature that puts dog domestication in the Paleolithic hunter-gatherer societies of Eurasia. It posits a possible origin for the domestic dog that came from the coming of age story of one of these Pleistocene hunters.
However, the actual domestication of the dog from wolves in these societies had to have been a bit more complex than the film states. In one flashback of the film, Keda’s father discusses wolves howling around their camp and how men must behave like wolves if they are to be good hunters. Of course, the father uses the toxic memes of alpha wolves in both the canid and human society. I honestly could have done without either.

But this flashback does hint that these hunter-gatherer people had some empathy with wolves, and it is this empathy that humans had with wolves that allowed us to form this partnership. I am reminded very much of Schleidt and Shalter’s hypothesis that wolves showed man how to hunt ungulates in Eurasia more effectively.

Humans and wolves have sort of convergently evolved as cooperative hunters, and it is very likely that humans would have seen much of themselves in wolves. Plus, if Pleistocene wolves behaved like the wolves of Ellesmere, humans would not have feared them in the way they would have feared cave lions or hyenas. This is a large social carnivoran that hunts big game but does not typically target us, and those features provided just enough space curiosity, empathy, and even reverence to develop.

I think that the hunting symbiosis hypothesis for dog domestication is essentially correct, but I don’t think it came about because someone managed to tame a wolf and hunt with it. I think it simply happened because humans, as opportunistic hunters and scavengers, figured out that following wolves was a great way to get good fatty meat. Wolves constantly test ungulates by harrying them. Those that are healthy stand and fight. Those that are weak run. The wolves usually kill the weak ones. The healthy ones that stood to fight would have been easier targets for the spear, and the healthy ones are full of fat that our big brains need.

So we would have figured out that if we followed wolves we could get the good meat we needed to survive, and the wolves likely would have figured out that we were the ticket to getting an easier meal. We probably drove the wolves off the carcasses at first, but we probably left enough meat for the wolves get a lot of reward for their effort.

Further, reliance upon human societies would have allowed one wolf reproduction strategy to operate quite well. In wolf packs, a single mated pair does all the approved breeding. If another female gets pregnant, the main breeding female (sometimes called the alpha female) kills the puppies of the other female or steals them to add to her litter. Usually, this main breeding female comes in heat first, and her older puppies easily outcompete the other female’s pups.

But these females do get pregnant. That’s because on the outside of the pack territory, there are unpaired males roaming about. These females are usually the daughters of the main breeding pair, and because wolves have some inbreeding avoidance behavior and because their mother will beat them down if they try to mate with her mate, they will often try to mate with these unpaired males that roam outside the pack

One notable male wolf in Yellowstone, the so-called “Casanova,” wound up living most his life as an unpaired male that mated with these unpaired females. In the early days of the Yellowstone reintroduction, the main breeding females of several packs allowed these unpaired females to raise their litters. Prey was abundant and naïve, so there was no need to kill the pups of these females.

Keeping a single litter is a lot of work for a wolf pack, so there is a very strong need for the litters tying them down each year to be reduced to one or none. When you have more prey, these pressures are released.

It is very possible that humans provided a space for that Casanova strategy to work more often. Some of the first wolves that may have hooked up with humans on a more intimate basis could have been females that wanted to have their litters away from their murderous mothers, and humans could have felt empathy towards these female wolves, tossing them food and protecting them from predators while they raised their litters. Humans could have provided a space for wolves that bred this way to reproduce efficiently, and if you’re just mating with a male and not engaging in all the social suppression of estrus and litter culling and purloining behavior, genes can spread much more rapidly. Perhaps the wolves that had lessoned genetic tendency toward pair bonding behavior became the basis for the domestic dog, and these genes wound up swamping the entire population of wolves that became domestic dogs, which is why pair-bonding behavior is uncommon in most domestic dogs

So basing the domestication story upon a female wolf is pretty wise.

My other quibbles with the film have more to do with the depth of characterization. I never really got to know Alpha as a wolf or a dog or anything. She was just a straight-up heroic figure, but I didn’t find the whole process of her transforming from a predator that would hunt humans to an extremely dog-like wolf particularly believable. I also wanted better CGI of the cave lions and hyenas. I am a bit of a Pleistocene mammal nerd, and I really wanted more of them. But they are like phantasmal entities that lurk in as agents of death and nothing more.
The writers did do quite a bit of homework on dog domestication, but I think they could have done more and pushed for an even more compelling narrative.

Finally, the “wolf” in the film is a Czechoslovakian vlcak, a breed of dog derived from Czechoslovakian working German shepherds and European wolves from the Carpathian Mountains. They were originally bred as an “improvement” to the working GSD of communist Czechoslovakia, but they never really got off the ground in that way. However, because they are mostly German shepherd and were selected hard for temperament, they are the most successful wolfdog breed ever produced. I do wonder, though, if people watching the film will realize that this breed is what was used to portray a wolf in the film. We are already going through a bit of a boomlet with Siberian huskies in North America, which is somewhat attributed to their wolfish features. If people do realize there is a breed of wolfdog that is recognized by many kennel clubs, then I can see this breed being mass-produced and sold to gullible people without much regard for temperament. This film gives the public this bit of information, then we could see lots of these dogs in shelters and pounds, which I’m sure no responsible breeders of vlcaks wants.

No, the performance of vlcaks in Alpha was not as compelling as Jed the wolfdog, but the public is now being exposed to this breed’s existence. It is something we need to think about very carefully.

I do give Alpha major props for trying to posit the origins of the dog within European hunter-gatherer societies and to give some credence to the Active Social Model for dog domestication, but in story-telling, I just couldn’t allow myself to follow that much artistic license with Keda’s miraculous escapes from danger.

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