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Posts Tagged ‘gray wolf’

We spend a lot of time debating about how wolves became dogs. A huge debate exists in the archaeological and paleontological literature about how one can determine whether the remains of a canid represent a wolf, a dog, or a transitional form between wolves and dogs. This debate is why the oldest dog remains are dated to around 14,000 years ago and come from the Bonn-Oberkassel site. Anything older than that, a big debate exists among experts about what can be used to define a wolf, a dog, or a transitional form.

But this debate does not exist solely in relatively recent transition between wolves and dogs. The entire evolution of Canis lupus is a hotly contested and often contradictory, depending upon which source one reads and whether one is looking a source that relies upon paleontological and morphological analysis or one that looks at the molecular evolution of the species.

It is well-accepted in European paleontology that Canis lupus evolved from Canis mosbachensis. Mark Derr paid particular attention to this evolution in his How the Dog Became the Dog. He posits that the extinction of the large hunting dog, Xenocyon lycaonoides, created an ecological niche that could be filled by the Mosbach wolf evolving into the gray wolf.

Yes, the Mosbach wolf was smaller than the modern gray wolf. Individuals from Northwestern Europe were mostly about the size of a modern Indian wolf or a “red wolf.” Indeed, the similarities between some of these mosbachensis wolves and red wolves are the best evidence for a unique red wolf species, for one can argue that red wolves are just a relict form of the Mosbach wolf that held on in Eastern North America. Of course, the genetic data do not agree with this assertion, but it is an interesting idea nonetheless.

My reading is that the Mosbach wolf gave rise to Canis lupus in Eurasia between 300,000 and 500,000 years ago. The coyote, though often posited as a primitive Canis, is actually derived from a divergent form of Canis lupus that got marooned in the American Southwest some 50,000 years ago and evolved to fit a jackal-like niche on a continent already dominated by dire wolves.

The Mosbach wolf disappeared from the fossil record around 300,000 years ago, but there is always a debate as to the possibility that it held on longer. The red wolf and Indian wolf are certainly possibilities for its continued existence today, but as we’ve looked at more wolf genomes both of those don’t come out so distinctive. Every study that I’ve seen that uses Indian wolf genomes finds that they are divergent Canis lupus, and the red wolf is a cross between wolves that are of that coyote type and relict Southeastern gray wolves from a later invasion of the continent. I do think there is pretty good historical data that some smaller wolves that we would define as coyotes lived in the Eastern states at the time of contact, particularly the small brown wolf of Pennsylvania mentioned by Shoemaker and the small “wolues” of Jamestown mentioned by John Smith. My guess is that no one really took stock of what they were killing when they killed off the wolves of Eastern states. It is very possible that coyote-like wolves were killed off in great numbers along with the big ones, and later on, coyotes from the plains came East, crossing with wolves and even relict original Eastern coyotes to form the modern Eastern coyote. The red wolf and the larger Eastern coyote are thus recreations of the Mosbach wolf that have happened in modern times.

In Europe, one potential late surviving Mosbach wolf was thought to have been found in Apulia, Italy, at the Grotta Romanelli site. Wolf remains have been found in the cave that date to between 40,000 and 69,000 years ago and they were often described as belonging to a late surviving Mosbach wolf. A recent morphological analysis revealed that these remains were of a peculiar form of Canis lupus that lived in that part of Southern Italy, and they were not of any kind of Mosbach wolf.

However, the Mosbach wolf is particularly intriguing. Occasionally, it has been posited as a direct ancestor of the domestic dog, but because we don’t have an overlap between the signs of the earliest dog domestication and the existence of Canis mosbachensis in the fossil record, one should be very careful in making such an assertion.

This same caveat should be placed when one sees Canis variabilis posited as dog ancestor. For one thing, there is no such thing as Canis variabilis. Instead, all the specimens listed as this species that come from the Zhoukoudian site in China have now been reassigned to Canis mosbachensis. This reassignment posits them as Canis mosbachensis variabilis, so whenever one encounters that “Canis variabilis” in a paper, just remember that they are discussing a particular East Asian form of the Mosbach wolf.

From my own speculative meta-analysis, it seems that the Mosbach wolf is ancestral to the entire wolf/dog/coyote species complex, which may include the African golden wolf, and the Eurasian golden jackal. A genome comparison study that included dogs, wolves, and one Israeli Eurasian golden jackal found that the divergence between the golden jackal and the dog and wolf species happened just before the anatomically modern Canis lupus replaced Canis mosbachensis in the fossil record. The Eurasian golden jackal could potentially be derived from a diminutive form of Canis mosbachensis that moved toward a more generalist scavenger form.

We also have some evidence of small Mosbach wolves in Europe that could have potentially gone in the direction of the golden jackal. This specimen was found not far from the Grotta Romanelli wolf that were found to be anatomically modern and not Mosbach wolves. It was found at the Contrada Monticelli site in Apulia. It was unusual in that it was quite a bit smaller than the Mosbach wolves found in other parts of Europe, and the authors found that Mosbach wolves were as morphologically variable as modern wolves are.

In North Africa, we also have a recent discovery of a canid that was much like the Mosbach wolf. The authors thought it was a bit different from the Eurasian form, and they decided to call this species Canis othmanii. This African wolf-like canid was found at a site in Tunisia and dates to the Middle Pleistocene, and it could potentially be the basal gray wolf that hybridized with the Ethiopian wolf to make the African golden wolf. More work needs to be done on this specimen, for it very well could wind up like Canis variabilis, a regionally distinct form of the Mosbach wolf.

The really fuzzy part about Canis mosbachensis isn’t that it is the ancestor of the gray wolf. The educated speculations I make about its relationship to the golden jackal and the golden wolf could be debated, and we need lots more data to figure out if I am right or not.

The really fuzzy part is what came before the Mosbach wolf. Most scholars think that Etruscan wolf (Canis etruscus), which makes an appearance in the fossil record around 2 million years ago, is the ancestor of the Mosbach wolf. For years, there was a debate about whether the Mosbach wolf was a chrono-subspecies of the Etruscan wolf or a chrono-subspecies of the gray wolf. All these suggestions would be technically true, simply because we could regard the Etruscan wolf-Mosbach wolf-gray wolf as a species that lasted and evolved over this time period.

However, a bit of a debate now exists as to whether the Etruscan wolf is the ancestor of the Mosbach wolf. An extensive morphological analysis of Etruscan wolf remains and those of another Canis species called Canis arnensis, which compared both to the modern black-backed jackal, the gray wolf, the golden jackal, and the golden wolf, found that our previous delineation between arnensis as being jackal-like and etruscus as being wolf-like were over-simplifications. Some characters of arnensis are much more like modern gray wolves than etruscus is, and it is possible that arnesis gave rise to the Mosbach wolf. Still, the bulk of scholarship thinks that the Etruscan wolf is the ancestor of the Mosbach wolf.

However, because the Mosbach wolf was not included in the analysis, it might be difficult to make such a conclusion. However, maybe the Etruscan wolf or something like it is the ancestor of the Ethiopian wolf. The ancestral Ethiopian wolf must have had an extensive range in Northern Africa for it to have hybridized with Canis mosbachensis, Canis othmanii, or a basal modern gray wolf to form the African golden wolf.

I have focused most of this post on the origins of gray wolves in the Old World, but the first Canis species to evolve were found in North America. Canis lepophagus first appeared in the fossil record 5 million years ago. It was very similar to a coyote or a Canis arnensis of the Old World. This is the part of the story where the molecular data has largely shaken up what we used to believe about coyotes. Lepophagus is thought to have evolved into the larger Edward’s wolf (Canis edwardii), which is sometimes called Canis priscolatrans. These animals might have been the same species or very closely related to the Etruscan wolf. The modern coyote is thought to have derived from edwardii/priscolatrans/estrucus 1 million years ago, but genome-wide comparisons put the existence of most recent common ancestor of gray wolves and coyotes at less than 51,000 years ago.

The dire wolf derived from Armbruster’s wolf (Canis armbrusteri). Armbruster’s wolf derived from Canis edwardii/priscolatrans/etruscus 1.8 million years ago. The dire wolf then evolved from that species 125,000 years ago, which means the dire wolf’s most recent common ancestor with modern wolves and the coyote may have been as far back as 2 or even 3 million years ago.

This analysis is still being worked out. The molecular data is constantly throwing wrenches into the machinery of paleontology, especially the paleontology of canids. The most successful extant canid lineage are full of parallel evolution and phenotypic plasticity, and in this way, it has become quite a challenge to sort out the evolutionary history of these species. At various times, large wolf-like forms have evolved as have smaller coyote or jackal-like forms.

The story of Canis starts with a coyote-like lepophagus, but right now, its likely niche is adopted by the modern coyote, which also very similar to it. But the molecular data suggest that the coyote evolved to adopt this similar niche from a larger Eurasian gray wolf and that it did not directly descend from lepophagus over 5 million years in only North America. Instead, it evolved into wolves that wandered into Eurasia, becoming the Mosbach wolf and then anatomically modern gray wolf. Some of these wolves wandered back into North America and became generalist scavengers in the land of the dire wolf.

Very similar stories likely are lost to us, but we must understand that the history of wolves is not just about getting bigger and developing pack-hunting behavior. That is one part of the story, but another part is about evolving to fit niches, which sometimes means evolving a smaller size and more generalist diet.

Some of my ideas here are very speculative, but I think they are nested in my reading of the available literature. Do not assume that I have the final story of how these creatures evolved, but just understand that the molecular side is so rarely considered in paleontology literature that it is almost like we’re reading evolutionary history of two different lineages.

More work must be done to formulate a synthesis between these two disciplines. Otherwise, there will be continued conflict, and the one using an older methodology and often working with much more incomplete data-set will fall by the wayside. And that is not the one using full genomes.

If we know what problems exist using morphological studies on extant and recently extinct canids, it is very likely that we’re missing important data on many extinct species, one for which there is no DNA to test.

Still, paleontology has much to tell us about the way early wolves lived. It can tell us much about how the ecosystems were and why wolves evolved in the way they did. But its methodologies often miss relationships between extant forms and miss the tendency toward parallel evolution.

I tried for about two years to watch Joe Rogan’s interview with Dan Flores, who wrote a book on coyotes that I think is quite full of misunderstandings about canid taxonomy. When Rogan questioned him about the papers that show a recent origin for the red wolf, Flores pretty much just dismissed those papers because they didn’t look at fossil.

That’s not how it works. Within canids, we know that parallel evolution is a big thing, and it is very possible that coyote-like and red wolf-like canids have evolved more than once on this continent. Indeed, a careful reading of the paleontology and molecular data strongly suggests that this is the case.

In fact, it has always been the case with these wolf-like canids. Big ones evolved from small ones, but sometimes, the big ones become small, because it is a better fit for survival.

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Gordon Buchanan hanging out with fully wild wolf named Scruffy on Ellesmere Island. These Ellesmere wolves and those of Greenland were recently found to be a very genetically distinct population of North American wolf.

Long-time readers of this blog know that I am particularly fascinated by studies on wolf DNA, and I was surprised that I missed this little gem that came out in PLOS Genetics last month.

The authors used 40 genome sequences of gray wolves, Great Lakes wolves, proposed Eastern and red wolves, and coyotes. The authors found further evidence to show that red, Eastern, and Great Lakes wolves are various mixtures of coyotes and gray wolves. The paper also found that all gray wolves derived in North America do derive from a single ancestral population and thus represent a single monophyletic clade within Canis lupus. 

The most interesting part of this paper though dealt with the genomes of wolves from the Queen Elizabeth Island, the famous arctic wolves, which are known for their white coats and curious nature around people.

The authors found that there were three distinct populations, which the authors define as East Arctic, West Arctic, and Polar.   The first two had some evidence of admixture with mainland gray wolves, but the ones defined as “Polar” did not. 

The wolves whose genomes came back that distinct were from Ellesmere and Greenland, which are the most northerly distributed of all North American wolves. The authors found that these wolves are relatively isolated from other wolf populations, and they do not have much genetic diversity.

These wolves have long fascinated me. They are curious and even socially open with people, and I think could give us a clue about how wolves could have hooked up with people in those Pleistocene days. 

But the discovery that they really are a genetically distinct population is also of great interest. Even more, we have full genomes from these wolves now, and maybe we can do a comparison study of these curious wolves that have never been intensely persecuted by man, normal gray wolves, and domestic dogs.  Maybe we can see what sorts of genes dogs and these polar wolves share that do not exist in other wolves, and maybe we could find out that my hypothesis is correct. 

This hypothesis is the one that states that the original wolves of Eurasia behaved more like these polar wolves than the timid and fearful wolves of lower latitudes. If these polar wolves share genes associated with tameness that are also associated with domestic dogs and their general behavior, then we might see evidence some evidence that the original wolf of Eurasia would have had the temperament that could have led to domestication.

But that will have to wait for another paper, which I am waiting for. 

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brush wolf

It cannot be overstated how much the discovery that coyotes are not as distantly related to wolves as we believed ultimately questions our entire understanding of the evolution of the Canis species.

The traditional understanding Canis species evolved from some form of Eucyon dog some six million years ago. Wang and Tedford, who wrote the most important book on the paleontology of the dog family, believe this was Eucyon davisi, which was the first of its genus to enter Eurasia. The genus Eucyon is where the common ancestor of the Canis dogs (including Lycaon and Cuon) and the South American wild dogs would be located. Eucyon dogs were small. Imagine them as being something like a black-backed jackal or a Hoary fox rather than a coyote.

Then, 5 million years later in the Southwestern US and northern Mexico, a coyote-like Canis evolved, which was called Canis lepophagus. This animal is sometimes considered the common ancestor of wolves and coyotes. It may be, but considering how close we now know wolves and coyotes are now, it’s not the most recent common ancestor. Canis lepophagus did migrate into Eurasia, where it either founded or is identical to Canis arnensis.

In Eurasia, several smaller jackal-to-coyote forms evolved. One of these was Canis estruscus,  which then evolved into Canis mosbachensis (which is called Canis variabilis in China).

Ron Nowak believed the red wolf was an offshoot of this wolf that wound up colonizing North America and then becoming isolated from the rest of Canis mobachensis when the ice sheets expanded. There was also a competing view that the red wolf was actually a remnant version of Canis edwardii or Canis priscolatrans (which were probably the same animal). This animal was roughly the size of a red wolf, but Nowak rejected it as a red wolf ancestor because it lived too early for what he thought were red wolf fossils.

The Eurasian wolf species evolved mosbachensis-variabilis, but the two forms of wolf shared habitat and likely exchanged genes, making it very difficult

The coyote’s evolution was never clear. It was thought to have evolved out of Canis lepophagus. It was thought that lepophagus evolved into edwardii, and then it began to become more gracile and smaller, eventually becoming the now coyote.  It’s now pretty clear that it evolved out of the Eurasian Canis lupus and not these endemic North American “wolves.”

It either evolved from the modern wolf, which evolved into roughly its current form 800,000 years ago, or it came from a late surviving mosbachensis-type wolves that were regularly crossing with modern wolves before they came into this continent. Maybe the remains that Nowak had been considering “red wolves,” were actually these ancestral wolves that were evolving into the modern coyote.

Maybe when this wave of wolves came back across from Eurasia, perhaps 50,000-100,000 years ago, it came into a world already dominated by a dire wolves, which already occupied the niche for large, pack hunting canids and this wave of Canis lupus evolved as the American jackal.  After all, the bobcat is just a diminutive Eurasian lynx that found itself in a very similar position when it came into this continent, and it evolved to be a smaller animal that generally hunts smaller quarry than its larger ancestor. Of course, the modern bobcat didn’t reach its current form until about 20,000 years ago, but it still was forced to adapt to a slightly different niche than its Eurasian ancestor.

In literature on the paleontology of Canis, there is a heated debate as to how these animals all fit. The conventional view is that the wolf evolved from Canis mosbachensis/variabilis through Canis etruscus, which may be the same thing as Canis edwardii/ Canis priscolatrans. Wang and Tedford contend that the coyote and wolf split from Eucyon.  The modern wolf evolved from Canis chihliensis, which was a large wolf-like canid. It spread into North America to found Canis armbrusteri, which then evolved into the dire wolf (Canis dirus) in North America and Canis gezi and Canis nehringi in South America.  In the Old World, another offshoot of chihliensis gave rise to Canis falconeri, which the supposedly gave rise to the Xencyon, which is supposed ancestor of the dhole and African wild dog. Another view holds that the Armbruster’s wolf (C armbrusteri) is descended from edwardii/priscolatrans (which may be the same as etruscus). This lineage then gave rise to the dire wolf and the two sister species in South America, thus descending solely from North America wolves.

All of these ideas come from paleontology, and they pretty much are done without looking very deeply into the studies that are examining the DNA of these species. It is pretty obvious from that literature that the notion that coyotes and wolves split at the time of the Eucyon ancestor is quite wrong. For that hypothesis to work, African of  wild dogs and dholes would have to be genetically closer to wolves than coyotes and golden jackals are. They aren’t.

But if the genome-wide analysis shows that coyotes are so much more closely related to wolves is true, then all these fossil and subfossil canids that are said to be the most recent common ancestor of wolves and coyotes simply aren’t.  Instead, all of these species that are classified in Canis are likely a mix of evolutionary dead ends, like the dire and Armbruster’s wolf, or could be hidden ancestors of extant canids that aren’t wolves or coyotes.

For example, black-backed and side-striped jackals diverged from the rest of Canis and its allies at about the same time that Eucyon was diverging from Canis. It is possible that there are many relatives of these particular dogs that are hidden in this vast sea of Canis fossils.

The new discovery about the coyote’s split from the wolf also means that any remains of North American canid that are listed as coyote that date to 1 million years before present are not coyotes. What they actually were is a very good question.

We’ve spent a lot of time assuming that coyotes and wolves were quite divergent. We know now that they really aren’t, but when we look into the past at all the “wolves” and “coyotes” that came before, we see how this genus became so successful. It can easily evolve into big game-hunting forms, but the real success is in its ability to assume the size and shape of the generalist predator. Phenotypic plasticity is a wonderful thing for a lineage to possess.

But the real message of the new discovery about wolves and coyotes should be is a cautionary tale about paleontology. Paleontology is a wonderful science, and it makes amazing discoveries every day, but when its faced with a lineage of animals where phenotypic plasticity and tendencies toward parallel and convergent evolution are commonplace, it is bound to make errors. Paleontologists aren’t examining flesh and blood that can have its molecules tested for relationships. They are measuring anatomical characters and determining phylogenetic relationships based upon the similarities of these characters.

Which works well.

Until you get something like wolves and coyotes, where there are many ancient fossil and subfossil remains that look like they could be ancestors of either.

But the DNA says they aren’t.

And paleontology would have problem catching the inverse. There are many species that we’ve discovered only through DNA testing. African butterfly fish in the Congo and Niger basins look identical to each other, but they have been isolated from each other for 57 million years. I have yet to see this species split into two, but if they were mammals, you could bet they would be placed in distinct species in heartbeat.

Paleontology is missing some really important things we’ve since found out through molecular analyses.

And paleontologists know this.

They are working with the data they have, and by definition, it’s going to be more incomplete than genetic studies.

Science is provisional. Different disciplines and methodologies are going to come up with different answers. It’s pretty amazing that one genome-wide assay study can wipe out so much literature in paleontology.

These debates have been raging for years.

And it turns out that everyone was actually wrong.

Update 21 August 2016:  It turns out that I missed a paper that actually did some limited DNA analysis and found that Canis nehringi was pretty much a South American dire wolf, as in it was likely the same species as the North American dire wolf. Canis gezi, however, was  more closely related to the modern maned wolf and had been incorrectly identified as a wolf. So let this stand as a correction to the error above.

 

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west virginia coyote

I’m currently reading John Lane’s excellent book, Coyote Settles the SouthIt is an excellent book, and I will be reviewing it here very soon. The whole time I’ve been reading it I thinking about my encounter with the male Eastern coyote I called in back in March.

He’s not exactly the same coyote that Lane is writing about. He’s a coyote of the gray woods, not the subtropical pine forests and river bottoms.

But in some ways, he is the same. He is the same creature that has adjusted to all that Western man can throw at him and thrived.

And he’s thrived at the expense of the wolves that once roamed over the Northeastern US and the South. He’s just the right size to live on a diet of rodents and rabbits but also has the ability to pack up and hunt deer. He can be an omnivore, enjoying wild apples and pears that fall to the ground, almost as much as he would if he came across a winter-killed deer.

The coyote is a survivor. I’ve written on this space several times that the reason he has thrived is because he has been here far longer than the wolves that once harried his kind. Until last week, it was assumed that the coyote split from the wolf some 1 million years ago. This million year split has been used for virtually every study that has examined the relationships between different populations or species in the genus Canis. It is used to set the molecular clock so that we can figure out when wolves and dogs split and perhaps give us some idea as to when dogs may have been domesticated.

This assumption has been directly challenged in a new study that was released in Science Advances last week. The paper examined full genome sequences of several different canids, and it can be argued that it pretty much ended the debate as to whether the red wolf and Eastern wolf are species. They aren’t. Instead, they are the result of hybridization between wolves and coyotes. Most of the media attention has paid attention to this discovery in the study.

It’s the most important practical implications, because the US Fish and Wildlife Service delisted the gray or Holarctic wolf in most of the Eastern, Southern, and Midwestern states in favor of protecting the Eastern and red wolves. Red wolves are called Canis rufus, and  Eastern wolf is Canis lycaon. With them being recognized as hybrids, this greatly complicates the issue of how to conserve them under the Endangered Species Act, which, as its name suggests, is meant to conserve actual species and not hybrids between species.

The authors of the study feel that these hybrid populations are still worth conserving, largely because the red wolf contains the last reservoir of genes belonging to the now extinct wolves of the Southeast.

But in order to make this work, we’re probably going to have to rewrite the Endangered Species Act, and that is not going to happen any time soon.

However, the finding in the study that is worth discussing more is that not only showed that red and Eastern wolves were not some relict ancient species of wolf. It is the finding that coyotes and wolves split only 50,000 years ago.

Using a simple isolation model and a summary likelihood approach, we estimated a Eurasian gray wolf–coyote divergence time of T = 0.38 N generations (95% confidence interval, 0.376 to 0.386 N), where N is the effective population size. If we assume a generation time of 3 years, and an effective population size of 45,000 (24, 25), then this corresponds to a divergence time of 50.8 to 52.1 thousand years ago (ka), roughly the same as previous estimates of the divergence time of extant gray wolves.

This finding means that the studies that use that 1 million year divergence time to set the molecular clock for all those dog domestication studies need to be reworked. This is going to have some effect on how we think about dog domestication, and although the domestication dates have been moved back in recent years, the actual split between dogs and wolves is likely to be much later than when we see the first signs of domestication in subfossil canids.

That’s one important finding that comes from this discovery that wolves and coyotes are much more closely related.

The other is that yes, it did pretty much end Canis rufus and Canis lycaon as actual species, but it probably also ends the validity of Canis latrans as a valid species. Coyotes could be classified as a subspecies of wolf. Indeed, they are much more closely related to wolves than Old World red foxes are to New World red foxes, which split 4oo,ooo years ago. And there is still some debate as to whether these two foxes are distinct species, because we’ve traditionally classified them as a single species. Plus, if we start splitting them into two species, we’re likely to find the same thing exists with least weasels living in the Old and New World. And the same thing with stoats.

And then it’s not long we’re fighting over the house mouse species complex.

But if we’re going to lump red foxes, it’s pretty hard not to lump coyotes and wolves. It is true that wolves normally kill coyotes in their territory, but it also found that wolves in Alsaska and Yellowstone, wolves that were thought to be entirely free of any New World ancestry, also had some coyote genes.

So the coyote, like the extinct Honshu wolf and the current Arabian wolf, could be correctly thought of a small subspecies of wolf. We know from paleontology that in both North America and Eurasia there were various forms of canid that varied from jackal-like to wolf-like, and although we know the jackal-like form is the earliest form, these two types have ebbed and flowed across Eurasia and North America. We’ve assumed that the jackal-like forms gave became the coyote and the larger wolf-like forms have become the gray, red, and dire wolves.

But what we’re looking at now is the coyote isn’t the ancient species we thought it was. It’s very likely that some ancestral wolf population came into North America, and instead assuming the pack-hunting behavior of Eurasian wolves, it tended toward the behavior of a golden jackal. When this ancient wolf walked into North America, it would have found that the pack-hunting niche was already occupied by dire wolves. There were many other large predators around as well, and evolving to the jackal-like niche would have made a lot more sense in evolutionary terms.

This is what the coyote is.

The pack-hunting modern wolf came into the continent and took it by storm, and the coyote exchanged genes with it. They lived together as sort of species-like populations in the West, but when wolves became rare from persecution following European settlement, the coyote and wolf began to exchange genes much more.

So with one study using complete genomes, the entire taxonomy of North American Canis is truly blown asunder.

And the implications for dog domestication studies and for the practical application of the Endangered Species Act could not be any more consequential.

Very rarely do you get studies like this one.

It changes so much, and the question about what a coyote is has become unusually unsettling but also oddly amazing.

I will never think of a coyote the same way.

The mystery is even more mysterious.

 

 

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Eastern North American wolf subspecies (Canis lupus lycaon). This is the wolf that likely lived in West Virginia.

From The Annals of Webster County, West Virginia (1941). The story is by Daniel Stoffer Hamrick, who was 17 years old when he killed this, the last West Virginia wolf.

During the last 40 years there has been a lot of controversy about the wolves still roaming the woods and hills of West Virginia. The following article may set at ease some minds which still believe they hear the cries of the wolf at night.

On January 8, 1897,the last gray or timber wolf was killed in West Virginia, according to my belief. Some eight or ten years before that date, wolves were playing havoc with sheep in Webster, Pocahontas, Nicholas, Braxton, Greenbrier, and Randolph Counties. About 1891 or 1892, John Gregory killed the mother wolf and one or two of her young.

From that time on for five years the male wolf was the lone representative of his tribe in the West Virginia mountains. He ravaged sheep far and wide. One night he would kill only one and the next night a great many.

On the night of Jan. 20, 1895, the wolf killed 27 lambs owned by my brother, Jacob Hamrick, on Point Mountain, Randolph County. The wolf was often hunted and traps were set for him to no avail. The county courts of Randolph and Webster Counties offered separate bounties of $100 each to the person who would bring the scalp of this wolf to them. The farmers of the mountain region were discouraged. They said they could not afford to raise sheep as the wolf killed the most of them.

On New Year’s morning, 1897, Uncle John Hamrick, who was living on a farm near Whittaker’s Falls of the Elk River, came down to my father’s house and said that on the previous night the wolf had killed five of the best sheep that he had, a loss of at least $50.

There was two feet of snow on the ground and it was very cold. Nevertheless, we started immediately to organize a wolf hunt. Every man we could coax into the hunt was sought. I still remember clearly my father saying: “Sonny, you put the saddle on my horse and go as quickly as you canto Joe Sharp’s place and tell him I have sent you for him and his dogs.”

This I did and I found Mr. Sharp in a willing mood. When we returned with the dogs we learned that the wolf had been tracked into a thicket on Point Mountain, just above where Currence Chapman then lived.

I do not remember all who started the first day but the second morning when we met halfway up Mill Run there were 15 men and boys in the party, namely: John, Robert and George Rose; Calvin Hamrick, Spencer Hamrick, W.S. Hamrick, David W. Hamrick, Lilly Hamrick, Francis Cowger, John Dodrill, Garfield Dodrill, Lee A. Hamrick, Adam Hamrick, Joe Sharp and myself. We followed that wolf track for eight days, in subzero weather and over four counties, but most of the time in Webster and Randolph. One evening just at dark we found ourselves at the mount of Flint Run on the Back Fork of the Elk, where Lilly Hamrick said to me: “I can’t travel any more.”

We built a campfire but we didn’t have anything to eat. Brother Jacob said that if someone would go after it, he could get a horse at his place on which Lilly could ride. Brother Lee said he would go and I went with him.

We walked three miles up Point Mountain to the farm. Shortly after we left the campfire someone produced a bottle of hot drops and told Lilly that if he would take some he would be all right. Adam, who had been the leader of the party from the start, remarked so Lilly could hear him, that there was nothing wrong with him except that he had given up. Either the hot drops or Adam’s insult did the work for when we started down the mountain with a pair of horses we met the whole part coming with Lilly in the lead.

One morning we were on stands on the Upper Elk. The wolf came within 50 yards of John Dodrill and stopped. He had a new gun, either a Winchester or a Marlin. John took dead aim and pulled the trigger but the gun didn’t go off. He tried again, and again the gun refused to fire. The wolf had enough of the fooling around and disappeared.

When we came up, John said: “Men, that wolf is not to be killed.” We hunted in the snow and found the two cartridges which had failed to discharge. We put them back in the gun and they both went off this time.

Then John said: “The hand of the Almighty is against us.” We tried to convince him that the reason the gun didn¹t discharge was because ice was in the gun. John finally said: “I will still go with the hunt but don’t place me where you think the wolf will come.”

On the night of Jan. 7, we were cared for by the good Dutch[German] people on Turkey Bone Mountain in Randolph County. The next morning while I was loading my gun, Mr. Wooftner said to me: “One cartridge is enough.” Ir eplied that I could carry cartridges better in the magazine of the gun than in my pocket. (Wooftner knew Hamrick was a good marksman.)

We went that morning to the spot where we quit the night before, the head of Back Fork of  the Elk. Adam told Jacob Hamrick and Milton Hull to hold the dogs for one hour and 20 minutes until he could have time to place the men on stands. We all went down to the fork of the stream and took stands. Lee Hamrick stood close to the creek. Alva Sharp stood above him, and I was next. Laben Hull was next above me and John Rose above him. The rest were strung out in like manner on up the mountain. I could see Mr. Sharp below me and Mr. Hull above me.

Mr. Sharp got so cold he was building a fire when the wolf came straight to him. The wolf must have winded him for he changed his course and came between Mr. Hull and me. I was standing on a log that was lying against a large maple tree when I sighted the wolf and fired the shot that brought his depredations to an end. I fired past Lee Hamrick and I know he jumped at least three feet when the bullet zipped past him. (Apparently Lee Hamrick pursued the wolf when it appeared before Sharp.)

The others came up and measured the distance of my shot at 187 steps. It was then one o’clock. There was much rejoicing that our enemy was dead and that our cold and painful tramping was ended. We walked from there to Laben Hull’s and danced all right. The next day everyone within a radius of several miles came to see the wolf. We weighed the wolf at Mr. Hull’s and discovered it weighed 87 pounds. It had eaten nothing in the eight days except one grouse. We didn’t give it time to kill any sheep in that period.

I took the scalp to the county court at Webster Springs but the court refused to pay the bounty. After finding a man for my guardian, and I was only 17, ‹I employed the late Senator E.H. Morton, a very able attorney. He brought suit against the county court and secured judgment in justice court. The court appealed the case to the circuit court. But when we came up for trial the court compromised and paid the bounty.

If there are any of those who would scoff at the above story, I refer them to the above men now living and to the court record of Webster County (pg. 255-258).

This story reads almost like a piece of Russian literature. Heavy snow. Deep forests. Men with guns.  And a fell beast that was destroying flocks of sheep, which was a great source of income, wool, and meat for small farmers in those days. All it needs is a sled

This region in West Virginia is deep within the heart of the Allegheny Mountains. It doesn’t surprise me that this was the last redoubt of the wolf in the state. Not very far from here, at Valley Head in southern Randolph County, the last of the Eastern wood bison, which ranged as far north and east as Western New York, was killed in 1825.

This wolf was part of a breeding pair, but his mate and pups were killed. Forced to make a living on his own, he began to regularly haunt the sheep pastures.

Natural prey must have been quite scare in those days, for the only contents of the wolf’s stomach was a paltry grouse.

A grouse is pretty hard for a dog or wolf to catch. After all, they do fly when something tries to pound upon them.

So this wolf would have been pushed to his limit to be forced to waste energy hunting ruffed grouse just to survive.

In West Virginia, people still claim to see wolves. In reality, they are almost certainly seeing the larger Eastern coyotes, which do have some wolf ancestry.

But a wolf was recently killed in Missouri. It was later confirmed to have come down from the Great Lakes wolf population— most likely Minnesota, if my geographic sense is correct. A wolf of a very similar size to the one Hamrick killed was shot for killing sheep in Massachusetts.  This wolf was believed to have wandered down from Quebec. It is also very likely that wolves occasionally show up in the remote haunts of northern Vermont, New Hampshire, Maine, and the Adirondacks of New York, which is a proposed site for a wolf reintroduction.

So it’s possible that a wolf will one day wander into West Virginia, but by the time they get here, they will have already established themselves in other parts of the East.

I don’t for a minute think the (contrived) red wolves of North Carolina will come up here. That population is already tightly managed and is largely restricted to a peninsula on the Carolina coast.

I don’t think the wolf that Hamrick killed was a red wolf or an Eastern wolf. The best science on the genetics of these animals shows that they are recent or relatively recent wolf and coyote hybrids.

This wolf was likely part of or closely related to the Eastern North American wolves (Canis lupus lycaon) that are now most common in Quebec and Ontario. The Massachusetts wolf that was killed in 2008 was of this subspecies, and it would be the subspecies that would be reintroduced into Northern New England and New York State.  Many members of this subspecies have coyote mtDNA, which was once used to declare them a new and unique species.

The truth is that many of them derive from a female coyote ancestor. That’s why they have coyote mtDNA. Their nuclear DNA shows that they are wolves with quite a bit of coyote in them.  The red wolf, by contrast, is a coyote with some wolf in it.

I’m sure that many states have some record of their last wolves, and I’m sure that most of them involve stories about wolves tripping traps and slaughtering whole herds of cattle.

But I don’t know of many states that have one about a 17-year-old crack shot bringing the last wolf down.

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In all the different accounts of people using wolves to do the tasks that are normally reserved for dogs, I had not come across anyone using a wolf as a gun dog. Audubon met a hunter in Kentucky who used a wolf to trail deer, and a French hunting hound pack included a wolf as a member.

There is also the story of Big Jim, one of the wolves raised to be released upon Isle Royale that happened to bond very strongly with a retrieving water spaniel. He learned to retrieve ducks from her, but I don’t know if he was ever of any use as a hunting dog.

But there actually was a wolf that proved to be an excellent retriever and gun dog.  He was actually a multipurpose gun dog, for in addition to being a great retriever of grouse and ducks, he also would bring down deer that had been wounded by the hunter’s bullets. He would sometimes kill  the deer, and then he would allow the hunter to collect his venison.

The story of this wolf comes from Blackfeet and Buffalo: Memories of Life Among the Indians by James Willard Schultz, also known as  Apikuni. Schultz grew up in the Adirondacks, where he learned woodcraft and hunting methods from the local mountain men, but as a young man in the  late 1870’s and early 1880’s, he moved into the Montana Territory, where he worked as a trader with various Indian tribes– most notably the Pikuni or Piegan. The Pikuni are nation within the Blackfoot Nation, and they are the only members of the Blackfeet to live in the United States. Schultz would chronicle much of their culture and daily life, selling accounts of his experiences to Forest and Stream. He would also become associated with the region that eventually became Glacier National Park, where he became well-known as an outfitter.

The story of his hunting wolf is rather simple. It begins with Schultz taking a wolf pup from a den, and he then trains it just like anyone would train a gun dog.

This wolf clearly was not a Native American dog, for the author makes it very clear that most Native American dogs, which probably were never encouraged to do much playing, didn’t want to play with him. The author makes a clear distinction between the two.

Further, it suggests that the wolves that became dogs could have participated in hunts with hunter-gatherers. If one reads the part where Schultz’s wolf was of great utility in bringing down wounded carefully, one could see how such a wolf could have been used even by people using more primitive weapons.

There is a general tendency in the popular conception of dog domestication generally tends to deny this possibility.

It’s stories of wolves like this one and the one that Audubon saw accompanying the Kentucky deer hunter that lead me to consider this possibility.

It’s also of note that this particular wolf fits almost exactly what  Raymond Coppinger derisively calls the “Pinocchio Hypothesis.”  Coppinger contends that no one ever tamed a wolf pup that has reached the weaning stage, so it must be impossible that hunter-gatherer man ever collected wolf pups and tamed them in this fashion. Hunter-gatherers just didn’t have puppy milk replacer, and they simply wouldn’t have been able to raise tamed wolves at all. Thus, dogs had to have come from wolves that self-domesticated through evolving to fit the niche as scavengers in human settlements.

Schultz was able to take an already weaned wolf pup– and not only was he able to tame it, he was able to use it as a hunting dog. This pup was eating bison meat, not suckling from bottles filled with Esbilac.

One should note that nowhere in Coppinger’s book is their any consideration for good historical records of wolves that were very useful as working dogs. These records are ignored or are not considered at all. It is almost as if they don’t exist.

Not only was this wolf working very nicely as a hunting dog. It was retrieving. Retrieving in the Coppinger model is an inherited predatory motor pattern. This inherited motor pattern is a truncation of normal predatory behavior, and dogs that inherit it are unable to engage the full predatory sequence. A dog that retrieves can’t  kill, dissect, or consume. It can only grab something and carry it back– and it exists within specialized breeds. One certainly wouldn’t find a wolf that did it, and if one did, that wolf would be unable to kill anything. Of course, this wolf not only retrieved shot birds, he actually killed wounded deer.

Modern wolves may no longer possess this aptitude. Decades of persecution have certainly changed wolf behavior. They are now much more emotionally reactive than the ones that lived on the Great Plains during the nineteenth century.  Most Native Americans were not wolf persecutors in the same way that Europeans were, and accounts of wolves in that region generally discussed how docile and curious they were.  Meriwether Lewis described the wolves he encountered during the Corps of Discovery Expedition (1804-1806) as being “extreemly gentle” and that Captain Clark had managed to walk up to a wolf and kill it with his espontoon.

Decades of persecution removed the curious and docile wolves from the population, leaving behind only those that are too emotionally reactive to handle to produce the next generation. Making comparisons with dogs and trying to generalize how domestication could have happened using these wolves is a major methodological error.

It is likely that the wolves of the Old World were much like these wolves. They had to have been very easy for Paleolithic hunter-gatherers to interact with. A wolf like “Big Mouth” would have been almost impossible not to domesticate.

The other factor that drives much of the conversation on wolf versus dog behavior is that there has always been an assumption that wolves must be handled roughly to get them to behave.

Did you read in Schultz’s account any place where he used lots of domination and confrontation to train his wolf?

He didn’t.

But modern wolf experts will write out all of these domination exercises that people must use to keep them under control.  A few months ago, I saw an Animal Planet program in which a wolf expert claimed that the reason why a woman’s captive wolf and wolf hybrid pack killed her is because she didn’t bite them. With animals that are already really emotionally reactive, such methods might exacerbate some of the aggressive behaviors that wolves might be exhibiting in already stressful captive situations.

Making comparisons with modern wolves in these high stress captive situations and domestic dogs and then trying to promulgate a domestication theory from these comparisons is really methodologically stupid.

All we really have are these historical accounts, and from them, we might be able to glean some idea of how domestication might have happened.

We know from modern examples that scavenging is not domestication.

Wolves in the Middle East and Italy have been scavenging off people for a very long time, but no one has seen them develop floppy ears or a curled tail or start barking or herding or pointing.

The idea that scavenging alone was the main force behind dog domestication is really quite dubious. Lots of animals have scavenged off us– everything from spotted hyenas to raccoons to marabou storks.  Black-backed and side-striped jackals have been scavenging off people ever since people learned how to hunt successfully. And they haven’t become dogs either. They also have contributed nothing to the domestic dog gene pool,  simply because they are not chemically interfertile with dogs, wolves, coyotes, golden jackals, or Ethiopian wolves. Scavenging alone will not make an animal evolve into domestication.

I think in the end that the real reason why people have issues with these notions is quite simple. Deep down, people are uncomfortable with knowing that dogs are wolves. They are wolves that experienced different selection pressures, but the two animals have not speciated. There are dog-like wolves and wolf-like dogs, and the two populations have exchanged genes and have continued to exchange genes. Wolf people tend to think of dogs as debased wolves, while dog people like dogs to be different so we don’t have to have a discussion about dominance hierarchies. Never mind that the dominance model that has been used to understand both wolves and dogs has largely been falsified through new scholarship. When one says dogs are wolves, one is not also saying Cesar Millan is a genius. Of course I’m not. But I’m not going to deny what dogs are in terms of their phylogeny, just because of the failed dominance model.

I think that much of what we think about wolves and dogs has unfortunately become too reductionist. I’m not saying that the typical family should be keeping a pet wolf, and I do recognize that there are tendencies in which wolves– in general– do differ from domestic dogs. The unfortunate aspect of this reductionist line of thinking is that has created a dichotomy in which dogs are dogs and wolves are wolves– and it has always been so. The truth is that dogs are derived from wolves that were very easy to domesticate. The nervous and emotionally reactive wolves we have today are not that easy to domesticate at all, but assume that they have always been this way I think is very faulty.

This model makes sense only if we ignore many examples of wolves that succeeded as working dogs in the past. Such a model does create contours, which easily fill out into a meme.  But meme like this one can be as blinding as it might be helpful, and this one is no longer of any use. It makes excellent fodder for documentaries, but the simple reality is that dog domestication is much too complex a subject to be reduced to such broad contours and generalizations.

Schultz was able to do something that many experts today would say is impossible.

He took a weaned wolf from a den, and he trained it to be an excellent hunting dog.

None of those things can happen, if we are to believe the popular literature on dogs and wolves.

But they did.

Simple as that.

***

Many years later, Schultz would train a coyote to be a retriever and turkey flusher, but because dogs and wolves are the same species, I felt that it was more appropriate to discuss the dog and wolf dynamic in this post. I will have a separate post for Smokey, Schultz’s duck retrieving coyote.

***

Many of these issues are discussed in Mark Derr’s How the Dog Became the Dog, which will be out this month.

See related posts:

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I was previewing John Bradshaw’s Dog Sense on Google, and I came across this profound passage. Please read it and then spend a few minutes reflecting upon it.

It is so poignantly true. The wolf that hitched its wagon to our star has thrived– thrived beyond what one would expect from any large species in the order Carnivora. Most large carnivores exist in very small numbers and at low densities.  Not so with Canis lupus familiaris.  It is everywhere.

And generally thriving.

No other large predatory mammal exists in these numbers.

Becoming the dog was truly an evolutionary breakthrough.

***

My only complaint about the quote is that last line about wolves “staying” wolves. That implies that wolves themselves have no experienced selection pressures but only dogs have.

Much is made about the selection pressures that produced the domestic dog, but what about the selection pressures that produced the modern wolf?

Many of our assumptions about wolf behavior come from an assumption that the ancient wolves that were ancestral to dogs and modern wolves were exactly the same as modern wolves in terms of their behavior.

This assumption causes some problems. Modern wolves are actually quite difficult to domesticate, but the ancestors of domestic dogs had to have been very easy to domesticate. Modern wolves must be habituated to people by 19 days of age to imprint upon people. After that age, they are next to impossible to domesticate.  This means that it would have been hard for ancient hunting people to domesticate any wolves, because a wolf isn’t weaned at 19 days. It is possible to tame adult wolves that have had no prior experience with people, as Woolpy and Ginsburg were able to demonstrate. However, this tame process would have also been hard for hunter-gatherers to use, for their taming process involved what might be called “taming by Stockholm Syndrome.”  These researchers isolated adult wolves, and the only other living beings they saw were people. The wolves became quite tame to people, and they were able to generalize their affections to all humans, even retaining this tame aspect 18-22 months after human contact had been discontinued.

It seems likely that the original wolf population was quite like other wild dogs in non-persecuted populations– opportunistic and quite curious.  They were probably very interested in people, and people became interested in them. One can see possible parallels in this sort of relationship with Timothy Treadwell and his “pet” foxes. If Treadwell had focused his attention on his camp foxes instead of brown bears, he might have produced something really interesting, for within his fox family, I think we can see the contours of what the first tame wolves were like. For the foxes, Treadwell was company and a food source. For Treadwell, the foxes were a curiosity, and they sometimes barked warnings about approaching bears or wolves. I found his footage of his camp foxes far more compelling than the “grizzly” stuff, because I think that through his relationship with the foxes, he told us how dog domestication could have happened.

These foxes were curious about people because they had been fed (illegally) by park tourists– and because they were park foxes, they were never persecuted by man.

Persecution could cause selection pressures on wild populations. I seriously doubt that I could go out and tame wild red foxes in West Virginia, where they are heavily hunted by hound and have been extensively trapped for their fur.

The wolves that did not become dogs experienced some similar to these red foxes. The effects of man’s nearly crazed persecution of wolves on the evolution of modern wolves have not been fully considered. But one effect of this relentless trapping, poisoning, and shooting is that the only wolves that have been able to survive have been those most emotionally reactive and “paranoid.”

One of the overlooked aspects of the Belyaev experiments that domesticated silver foxes is that the critical period for socialization in these  domesticated foxes increased simply through the selection for tameness. These experiments selected foxes for tameness and approachability, and the researchers bred from those foxes. Over generations of selecting for tameness and approachability, the foxes became very dog-like in both behavior and appearance. Because these experiments are used as analogies  of the domestication process, they are almost always mentioned in dog domestication literature– but never the part about the critical period.

It seems to me that wolves have experienced a Belyaev experiment in reverse. Persecution was a selection pressure against tameness and approachability, and the animals that survived the cull were largely those that were most nervous and emotionally reactive.   It is so severe that many wolves won’t cross roads. Roads and virtually anything else that appears novel are too much for them. This is one reason why it was found that many captive  wolves won’t eat beef. They were raised eating deer and elk, and beef  is just too novel and too scary. Perhaps one of the reasons why these wolves are so nervous is that nervousness and paranoia are the result of a selection pressure that chooses wolves with shorter critical periods for socialization. Just as Belyaev selected for tameness and got longer critical periods, man could have selected for only those wolves that were paranoid and emotionally reactive– and this may be in some way correlated or associated with a shorter critical period.  Maybe it goes like this:  If you have a short critical period for socialization, one has only a limited opportunity to learn which things are safe, so almost everything else in the world is scary. Conversely, if you have a longer critical period, one can learn that more things aren’t scary and one’s brain develops very differently. Both of those courses of development would have profound influence on how one’s brain would develop, and perhaps, it can explain many of the differences between dogs and wolves.

Such animals would have had a very hard time colonizing almost all of Eurasia and North America, so it seems likely that the ancient wolves that existed during the time of dog domestication were not nearly as paranoid. They were likely curious and opportunistic and maybe even a touch more socially tolerant.  These animals could be more easily tamed, simply because they were not overwhelmed with fear or fear-based aggression. They also would have been more willing to exploit the new opportunities in becoming a camp wolves. These wolves would have regular access to large prey, which they may have helped their human counterparts bring down, and they would have humans around to help protect and raise their offspring, which means that the pair bond system that virtually all wild dogs seem to use would no longer be as efficient a way of passing on genetic material. The Casanova wolf strategy would be the best way for these camp wolves to breed, and thus more wolves in the population would be breeding. Early domestication would have had the advantages of good food and freedom from a pack structure in which only a few individuals breed.

One part that is always missing in the wolf and dog comparison is the tendency to ignore the simple reality that wolves have continued to evolve after dogs were domesticated. The dog didn’t descend from the wolf running wild today, but both descend from a common ancestor.  Both may be the same species, but that same species exists in two distinct populations. One has been selected by both nature and man to be very close to humans. The other has been selected to fear humans at all costs.

The error is assuming that the latter has always been this way.

Doing so creates too much confusion in trying to understand how dogs could have been domesticated from wolves.

It wasn’t confusing. It was likely so easy that a caveman could do it.

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