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Posts Tagged ‘gray wolf’

I love reading old breed books. Getting into German shepherds now means that I have a whole new selection of books to read, and in older German shepherd books that are written in English, there is a strong desire to distance this breed from wolves.  At one time, the breed was banned in Australia because of its supposed wolf ancestry, and Australian sheep interests were quite concerned that the breed infuse wolf phenotype and behavior into dingoes if they got loose and crossbred. So there is a tendency to downplay any relationship between this breed and wolves, and this tendency sometimes gets quite ridiculous.

In the first few pages of Jane Bennett’s book on the breed, which had its last printed in 1982, I noticed this image of a wolf.

Jane Bennet Wolf German shepherd

If you cannot read the full caption, check it out here:

Tomarctus wolf Jane Bennett

So I don’t expect to see accurate zoology or paleontology in dog books, especially from old ones. And to be honest, I am skeptical that German shepherds are especially wolf-like dogs with close wolf-like ancestry.   It is possible that some of the Thuringian sheepdogs in the breed’s ancestry had some wolf crossed in, but I don’t think they are wolfdogs in the same way that a Czechoslovakian vlcak is.

But the idea that the most recent common ancestor between a wolf and German shepherd was Tomarctus is not at all accurate. In some of the old dog books I have, Tomarctus is sometimes mentioned as an ancestor of modern dog species.

However, current paleontology places Tomarctus in the Borophagine subfamily of Canidae. Not a single living descendant of the Borophagine dogs exists. These dogs lived only in North America and all were extinct by the end of Pliocene. Tomarctus went extinct about 16 million years ago, which would be in the Miocene.

So it was not even a late surviving Borophagine dog, and it certainly was not the most recent common ancestor of wolves and German shepherds.  If it were the most recent common ancestor, then Czechoslovakian vlcaks, Saarloos wolfhonden, and Volksoby would have been impossible to create. 15-16 million years is more than enough time for two mammalian lineages to lose chemical interfertility, and dogs and wolves simply are chemical interfertile right now.

The most recent common ancestor between a German shepherd and a wolf could have been a wolf kept at the Frankfurt zoo that some think is behind the Thuringian sheepdog Hektor Linksrhein/Horand von Grafrath, which is the foundation dog for the modern German shepherd breed. Or it could have been a wild wolf that mated with a sheepdog somewhere in Germany, and that sheepdog line got mixed into the breed. Further, dogs in Eurasia, some of which may have German shepherd in them, are interbreeding with wild wolves at a much higher frequency than we might have imagined. 

I honestly don’t know, if the GSD breed has close wolf ancestry, and reasonable people can disagree on this issue.  I have not seen definitely proof either way, so I do remain agnostic on this issue. The temperament of the breed, though, is of very trainable herding dog.

But whatever the truth is, I don’t think anyone thinks the most recent common ancestor of the German shepherd and the wolf was a species that outside the lineage of both.

This claim isn’t as bad as the claim that chow chows are derived from extinct digitigrade  bears or from an extinct predatory species of red panda.

Jane Bennett’s book includes lots of good information in pedigrees and care of a German shepherd, but that page of the book indicates a strong desire to distance the dog from the wolf in a way that those of us living in the era of molecular biology and modern cladistics would find a bit bizarre.

The current thinking from full-genome comparisons is that all domestic dogs are derived from a now defunct lineage of Eurasian gray wolf. To keep Canis lupus monophyletic, we must keep the dog as part of that species.

So I have noticed a theme in many of these older books to keep German shepherds as distant from wolves as possible, even if it means making a claim that could easily disproved with a simple look at the Czechoslovakian vlcak or the Saarloos wolfhond, which both existed when this book was last printed.

Jane Bennett bool

 

 

 

 

 

 

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black coyote

For most of my life, scientists believed that the present era was still the Holocene.  Glaciers retreated with a global warming trend around 11, 650. Man went from being the apex predator over much of the world and became the apex consumer. Agriculture allowed our populations to expand, and we started to give up our wandering ways and became “civilized.”

It was generally believed that the past few centuries are but a continuation of this age, but now a growing number of scientists believe we have left the Holocene and entered into the Anthropocene. Several scholars have issues with this new distinction, but I think it is quite useful. In this era, human activity is the main factor affecting climate and ecology, which is why the age is named for the Greek word for human (anthropos).

The best argument I’ve seen for when this era began is 1610.  In this scenario, the era is dated to when European disease and conquest killed off enough Native Americans and enslaved and enough African had been enslaved to allow forests to grow back in former agrarian fields.  This process started in 1492, but by 1610, enough of those trees had grown to remove enough carbon dioxide from the atmosphere to push the planet into the Little Ice Age.

Darcy Morey and Rujana Jeger have a great model for understanding dog domestication as a change in trophic strategies.  In the Pleistocene ecosystems, most wolves were mesopredators, playing second fiddle to an ecosystem full of cave lions, machairodonts, large bears, and cave hyenas. When these wolves hooked up with people, though, they hitched their wagon to the species that often behaved as the apex predators in the ecosystems.  When humans switched to agrarian lifestyles during the Holocene, humans became apex consumers, and dogs joined us as beneficiaries of being allied to that apex consumer species.  During the Holocene, many wolves became apex predators, as the cave lions and other large predators became extinct.

I’ve always liked the framework that Morey and Jeger derived in this paper, but now that we’re entering into a new geological age, maybe we need to look at the change in trophic strategies of wolves in this new age.

Morey and Jeger don’t have a good framework for what happened to wolves in the Anthropocene, but across Eurasia and North America, wolves were gone from many human-dominated landscapes by the first decades of the twentieth century.  They existed only in isolated areas in Western Europe, and in the  lower 48 states of the US, they lived only in Minnesota and in an isolated region in East Texas and Louisiana, where the taxonomically controversially red wolves were located.

Large pack-hunting wolves were really in quite a bit of trouble.  In the United States, the coyote population began to expand out of its Western core range into the Great Lakes States. They eventually made to New England and the Maritimes of Canada, and they hybridized with relict wolves and the expanding population of domestic dogs.  Coyotes eventually colonized all the Eastern states, and as they did so, they largely became the apex predators in many parts of their range.

But in the 1960s, attitudes about wolves began to change. Many nations protected wolves, and there were often introduction plans in the works.  By the early decades of the twenty-first century, wolves were making significant comebacks in Germany and Italy. The wolves in Italy were often living very much like stray domestic dogs, living large at garbage dumps. Wolves live near large cities in Germany, and how these wolves are going to adapt to living in such human dominated environments is going to be a major question for researchers.

And in throughout Eurasia, we began to see that domestic dogs were mating with wolves.  Indeed, it is now estimated that a majority of wolves in Eurasia have relatively recent dog ancestry. 

Similarly, as coyotes expanded in North America, their genes began to work their way into the wolf population.  Yes, coyotes in a large part of the US have wolf ancestry, but we also have discovered that wolves across North America have coyote ancestry. Indeed, one interesting thing about these genome comparisons is that coyotes and wolves are much more closely related than we initially gleaned form mitochondrial DNA analysis. The calculation is that the gray wolf and the coyote last shared a common ancestor around 50,000 years ago.  This recent common ancestry has a taxonomic implication, which is that coyotes are themselves a divergent form of gray wolf in much the same way domestic dogs are.

In the Anthropocene, the wolves that have done the best have been the domestic dog and the coyote. The domestic dog’s ability to ingratiate itself into human society or live very nicely as an opportunistic scavenger/hunter on the periphery of humanity is a great gift.  The coyote can live as an opportunistic scavenger/hunter as well, and it also can live very nicely as a mousing fox or pack up and hunt deer.

Gene flow among wolves, coyotes, and dogs has made these entities much more fuzzy than we once thought they were. Dog genes are working their way into both the coyote and wolf population.  Strange pelt colors are popping up in the wild animals. The black coloration in domestic dogs was conferred onto the North American wolf population during the Holocene, but this same mutation for melanism has entered the coyote and Italian wolf population in very recent years. Dogs have introduced dewclaws on the hindlegs to some wolf populations, and I have seen photos of Eastern coyotes that have those hind dewclaws as well, which likely were introduced through breeding with domestic dogs.

Coyotes in the East are evolving larger size to become better predators of deer, but becoming larger and more effective ungulate hunters will have a trade-off. As carnivorous mammals grow larger, they become more and more dependent upon large prey to survive.  Very large wolf-like coyotes will lose their ability to live well on small prey and garbage.

So in the Anthropocene, dogs remain allied to the apex consumers. Some coyotes operate as apex predators, and some wolves live as opportunistic scavengers.

And as these creatures adjust their trophic strategies in a much more predator tolerant world, the pseudo species barriers that exist among wolves, coyotes, and dogs can break down. Hybridization among these creatures is likely to be a major feature of their continued evolution, a definite feature and not a bug.

These canids  thus make the leap with us into this human dominated age, an age that is experiencing a mass extinction of amphibians and great retrenchment of large sharks and big cats.

Yet they are still there. Evolving as the winds change. Winds that we ourselves are changing and are only now starting to understand.

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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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