Archive for the ‘wolves’ Category

Great Lakes gray wolf on Wisconsin’s Stockton Island.

An extensive camera trap survey of the Apostle Islands in Lake Superior has revealed a great diversity of carnivorans, including gray wolves and American martens. These islands, located just off Wisconsin’s Bayfield Peninsula, are among the few places left in the Eastern and Midwestern US that still retain an intact predator guild, and because all but one of the islands is protected as a national lakeshore, these islands will be protected from most exploitative development.

I love living in a country that still has room for wolves and wild places. Our natural heritage is every bit as important to us as a nation as our constitution and rule of law, and it should be protected with the same ferocity that we use to protect our republicanism.

Also of importance is the Great Lakes wolf in the trail camera capture that was part of the survey. This wolf looks to be one of wolves with some amount of coyote introgression. Great Lakes wolves can have quite a high amount of coyote ancestry, and the earliest estimated introgression of coyote genes into gray wolves that has been documented is from these Great Lakes wolves. This introgression happened as early as 963 years ago.

Coyotes and gray wolves are found on the islands now, and one wonders if they still occasionally interbreed. The Great Lakes are also the region that experienced the beginnings of the hybrid swarm we call the Eastern coyotes.

So these islands could be a new Isle Royale from which to study the new and evolving wolf and coyote of North America.

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Why coyotes are not basal Canis

The traditional understanding of coyote evolution is that coyotes are basal wolf-like canids. This understanding comes from the hypothesis that coyotes directly evolved from Canis lepophagus in North America alone. Coyotes look and behave a lot like jackals of the Old World, and because we know that the larger wolf-like canids evolved from jackal-like ones, we just assumed that the coyote was a primitive form.

One problem with this positioning has always bothered me. Jackals tend to have proportionally smaller brains than wolves, but coyotes have proportionally larger brains than wolves. Domestic dogs have evolved smaller brains from wolves, although wolf and dog brain size comparisons aren’t as cut and dry as people think. 

No one thinks of dogs as basal forms of Canis, so it is possible for animals in this lineage to lose brain size, just as it is possible for a primitive lineage of canids known as coyotes to evolve a larger brain.

Please note that my discussion on brain size here isn’t really a discussion about intelligence, because the literature on which form is most intelligent is quite all over the map. Domestic dogs kept in Western countries in the modern way do appear to have social cognitive abilities that virtually all other species lack, while wolves are much better at working with each other to complete tasks.

But coyotes have proportionally larger brains than either wolves or dog do, and in this lineage, larger brains are generally a derived characteristic.

However, the really important data about coyote evolution is the discovery that they shared a common ancestor with gray wolves much more recently than commonly suggested. A genome-comparison study of various North American canids found that the common ancestor of both gray wolves and coyotes lived around 50,000 years ago. Because anatomically modern gray wolves replace the Mosbach wolf in the fossil record between 300,000 and 500,000 years ago, the ancestor of both had to have been a form of gray wolf from Eurasia.

The coyote is thus a jackal that has evolved in parallel out of the gray wolf lineage, which means it is not a primitive canid at all. It likely evolved this jackal -like morphology and behavior because the form of gray wolf that it derives from was unable to compete with the dire wolf, the American lion, the short-faced bear and the machairodonts as a top-level predator. It was forced to evolve a smaller body that could be fed on carrion and small prey.

We know now that there is a big difference in what prey predators target once they exceed 20 kg. Predators that weigh more than that mass target large vertebrates, while those that are smaller than that weight target smaller prey. Although coyotes do cooperatively hunt deer, they primarily feed on rabbits and mice. So by becoming smaller, coyotes were not directly in conflict with dire wolves or the other large predators of Pleistocene North America.

Only through analyzing full genomes of coyotes and gray wolves did we realize that our assumptions about their evolution were wrong. Earlier studies that looked at mitochondrial DNA alone found that coyotes fit within a basal position of the wolf-like canid lineage. However, recent full genome comparison of various wolf-like canids that looked at the role hybridization played in their evolution found something interesting. The lineage that leads to wolves, dogs, and coyotes experienced some introgression from a ghost species that was closely related to the dhole. The authors think that the reason why coyotes turn up so basal in these mitochondrial DNA studies but appear so wolf-like when their full genomes are compared is coyotes have retained a mitochondrial line that comes from that ghost species.

So the generalist coyote is a re-invention out of the gray wolf lineage. It is not basal to the wolf-like canids. It just merely resembles the basal forms in some of their ecology, in some of their behavior, and in their odd mitochondrial inheritance.

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Long-time readers of this blog know that I am quite critical of Dan Flores’s Coyote America, a book that has been hailed as a sort of definitive source for the natural and cultural history of the animal. The good parts are where the author talks about native peoples and their relationship and understanding of the animal. The bad parts are where he misrepresents the molecular research on coyote evolution, most notably where he contends the genetic difference between a gray wolf and a coyote is equivalent to that of humans and orangutans (page 27, if you’re looking for it). I’ll give Flores a pass in that he didn’t do his research for the book until after the coyote, gray wolf, Eastern wolf, and red wolf genome comparisons came out, and found that all these animals were as closely related to each other as humans from different continental origins.

But I don’t know of anyone who thought that coyotes are to wolves what humans are to orangutans. At best, we thought coyotes were to wolves what our species of human was to Neanderthals.

So that was my beginning of great distrust in Flores’s account of how coyotes evolved in North America.

I do remember one part that I thought might have been true, simply because it mirrors the way coyotes moved into the eastern parts of North America. Flores contends that coyotes did not make it into Southern Mexico and Central America until after European colonization took place. The clearing of the tropical forests and the introduction of sheep and goats made all of this possible.

This made sense to me, but then I thought, well, I should look it up.

It turns out that Flores was dead wrong about Southern Mexican and Central American coyote populations. A 2004 paper that looked at the paleontology and sixteenth century accounts of coyotes in the region found that coyotes were in the region before European conquest.

So coyotes have lived in Guatemala and El Salvador long before Europeans felled the forests and turned out sheep and goats. Their recent range expansion into southern Panama may eventually lead to their arrival in Colombia, and they will have the Southern Continent to colonize.

This book gets so many facts wrong about the evolution and natural history of coyotes that I do worry a lot about its impact. It is written as a popular natural history, so it needs to be understood in that vein. However, the author seemed to choose which scientific facts he wanted to present without looking deeper into the fullness of the literature that exists on them.

And as a natural history writer, I find such errors to be problematic, but I always find some way to make sure you know that I am not the final authority on any subject. Because I blog, I can show you my evolution in thought more easily. Books are far more permanent inscriptions. That’s why you will see me hedge about certain subjects where I know more research is being done, such as what the African golden wolf actually is or where dog domestication happened.

The challenge is to make natural history subjects interest and to make your interpretations fit the literature, both of science and of prose.

And yes, it took me a month to read Flores’s coyote book. I had that many problems with it.

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One of my favorite cryptozoology rabbit holes is the story of the “onza,” a supposed late-surviving coursing cougar or “American cheetah.” Onza is Spanish for cheetah, and there have been skinny, long-legged cougars from Mexico that have been claimed as onzas.

One of the earliest indications that such an animal existed in historic times supposedly come from Bernal Diaz del Castillo, one of the conquistadors who took over Mexico with Hernan Cortes. While living in relative luxury on his own personal fiefdom in what is now Guatemala, Bernal Diaz wrote an extensive account of his exploits in the taking of”New Spain” from the Aztecs and other Meso-American peoples.

His account of the court of the Aztec Emperor Moctezuma II (known as Montezuma in the US) also includes an account of the emperor’s menagerie. Advocates for the onza as a late surviving American cheetah point a line the the text where Bernal Diaz discusses the “tygers [jaguars] and lions [cougars], one of which resembles a wolf.”

I don’t know exactly what Bernal Diaz meant by cougar that looked like a wolf, but the really interesting thing is that this is not the full line.

The full line is “tygers [jaguars] and lions [cougars], one of which resembles a wolf, called here Adive. “

“Adive” is a word that entered Spanish through North African Arabic, and it refers to the golden jackal. Well, the African golden wolf as we know it now.

Bernal Diaz is talking about wolf-like creatures called “adives” in New Spain, and although no one can make sense of his lion that looked like a wolf description. In an aside, he actually made mention of the coyote, and this must be the earliest mention of coyotes in a Western text.

My guess is Bernal Diaz was talking about a jaguarundi, which does look like an holy hybrid of a wolf and a cougar. This animal is sometimes called in Onza in part of Latin America, and it is also part and parcel of the late surviving American cheetah legend.

He could have also seen a Mexican wolf, and for whatever reason, he didn’t think of it as a wolf like existed in Spain. Instead, he may have classified it with the cougar, for it seemed so exotic to be a true wolf.

Bernal Diaz would have known these adives from his life in Guatemala, and he would have seen them in the Yucatan and during his adventures in Central Mexico.

Bernal Diaz wrote his account in 1576, when he was in his 80s. This account, even though it is just an aside, is the first mention of a coyote, and it predates the most famous early account of the “Indian fox” in Francisco Hernandez de Toledo’s Plantas y Animales de la Nueva Espana . Hernandez de Toledo was on his scientific expedition for the King of Spain at roughly the same time that Bernal Diaz mentioned the adive, but his account of the animal was not published until 1651, when he had been dead for decades.

So I would argue that Bernal Diaz was the first person to mention a coyote in Western literature, even if almost everyone missed his account as a mere weird word in an account about cougars and jaguars.

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

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

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

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

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

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

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

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

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

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

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

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

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Charles Darwin was an early proponent of humans belonging to a single species. The concept was not popular in Darwin’s imperialist nation, where many of the best and brightest were dead certain that “lesser peoples” of the empire were entirely different species. Monogenists versus the polygenists this was the debate by even those who accepted Darwin’s controversial thesis that species evolved through natural selection and that humans evolved from African great apes.

We know now that the monogenists won out. Only the basest racist frauds believe that humans represent multiple species deriving from distinct evolutionary lineages. The only exceptions are that we do have evidence that humans have trace amounts of other extinct humans in their DNA, such as Neanderthals and the Denisovan hominin.

The monophyly of a species is a concept that should be axiomatic. Much of the “rewriting” of taxonomy and systematics comes from us discovering that a clade is either paraphyletic or polyphyletic, but what does sometimes happen is we do run into situations where we have thought species were really distinct but now we think of them as being subspecies. Black-tailed and mule deer come to mind, as do Cape and African forest buffalo. We have to combine them in a single species to retain its monophyly.

I think something now must be done with gray wolves and their closest kin. In recent years, it has become difficult to retain that domestic dogs and dingoes remain distinct species from the Holarctic gray wolf, especially if we wish to keep the species monophyletic. But I think now a good case can be made that coyotes belong to the same species as Canis lupus. They diverged from a common ancestor only recently, around 50,000 years ago, and the much debated Eastern and red wolf “species,” which appear to be hybrids between coyotes and gray wolves can also similarly be folded into Canis lupus. Whether there was a coyote sister population in the Eastern forest that gave rise to red and Eastern wolves is a moot point. That “forest coyote” population is no more distinct from gray wolves than those of the West are.

Further, we do have evidence of continued gene flow between gray wolves and coyotes. Eastern coyotes have wolf ancestry, and many also have wolf ancestry that comes from domestic dogs.

This new way of describing Canis lupus forms has some benefits. One is that we can come up with a strong legal foundation for the protection of red and Eastern wolves without them being distinct species. Their hybrid status is no longer problematic. We no longer must strain credibility talking about them being ancient North American wolves. Instead, we just go the Endangered Species Act’s statute language which defines an “endangered species” as being a “subspecies” that requires conservation action. We currently protect the Florida panther as an endangered species using this definition, and what is more, the Florida panther was given genetic rescue when Texas cougars were released into its range. The Florida panther exists as a hybrid that holds onto alleles of a population that no longer exists in its pure form, and in the case of red and Eastern wolves, one can very easily make this case.

The problem with going this route is that coyotes themselves do not experience many protections in the United States or Canada. Most states have a very open hunting season on them year-round, and some places have bounties on them. Eastern coyotes are hard to tell apart from what are called Eastern wolves and are even harder to tell apart from red wolves. There is an overlap in phenotype that comes from both forms having similar hybrid origins.

Further, because red wolves living in the North Carolina recovery range are all derived from 19 individuals, and wild wolf-like canids have inbreeding avoidance behavior, it is quite common for red wolves to pair up with Eastern coyotes. Much energy has been devoted to killing off and removing coyotes in red wolf recovery zones, including the euthanasia of crossbred puppies, but it seems that this won’t work long-term.

Recently, researchers at Princeton discovered that the coyotes of Galveston Island are quite closely related to the red wolves in the recovery program. The authors think that many southeastern coyotes might hold the alleles of these red wolves, but I think a better explanation is that we simply under-estimated what the coyote’s range was in the United States at the time of contact. John Smith described the “wolues” of Jamestown as not being much larger than English foxes, and I initially thought he was talking about gray foxes. But then I read original text, and the gray fox is described in complete detail, including the fact that it lacks the red fox’s odor. Henry Wharton Shoemaker, the naturalist and folklorist who chronicled much of the lore of Pennsylvania’s wildlife, wrote about a “small brown wolf” that lived along the Susquehanna River, which made a yipping howl that resembled that of the prairie wolf or coyote. Shoemaker postulated that these small brown wolves were the same species as the coyote, but very few scholars have taken his speculations seriously.

Perhaps, there were coyotes living in the forest alongside Eastern wolves and a endemic Southeastern form of wolf that was very often melanistic. All three exchanged genes, and all three were killed off for their furs and for bounties without any regard to what they were. It was only when settlers in Indiana noticed that some of the wolves in their area were quite small that they began to wonder if they were a distinct species. Later explorers into the West began to call them prairie wolves, which was their name until Americans adopted the Nahuatl-derived name for them.

The most important bit of text I’ve read in any of these North American wolf taxonomy papers is one in a response paper to another one that still argues for multiple species of wolf in North America. The authors write that “[the] genetic differentiation between red wolves and other North American [wolf-like] canids is comparable to the amount of genetic differentiation found between different continental human groups, which of course are not considered to be distinct evolutionary lineages.”

And at that moment, we are back to Mr. Darwin’s debate with the polygenists. He was right to oppose their claims back then, and maybe when it comes to North American and Eurasians wolves, dogs, dingoes, and coyotes, one should be a monogenists.

That’s where I am planting my flag. I think that is where it will all end up once we’ve sorted it all out.

I find the idea of gray wolves representing a phenotypically and behaviorally diverse species awfully intriguing. This diversity is greatly exaggerated in the domestic form, but this diversity is also reflected in the wild, where we have 15-pound coyotes and wolves that weigh 130 pounds.

It may even be that the African golden wolf, which is derived from the ancestor of the gray wolf and coyote mating with the ancestor of the Ethiopian wolf, may be close enough to extant gray wolves that we might also regard it as part of this very diverse species. It would be a coyote-evolved in parallel out of the gray wolf lineage, for the coyote likely evolved in its smaller and more generalist form from the ancestral Eurasian gray wolf because it could not compete as a pack hunter in a North America dominated by dire wolves. African wild dogs once ranged over almost the whole continent of Africa, and they could have provided competition for that niche that would force these ancestral gray wolves to evolve in something like an African coyote.

The recalibration of the evolution of Canis needs to be done with light of the knowledge that gray wolves and coyotes aren’t so genetically distinct from each other. We need new papers that look at he full genomes of golden and Himalayan wolves to get an idea of when they may have split from the rest of the gray wolf swarm. It may be very well be that they all belong to this wide-ranging phenotypically diverse species that over parts of four continents

Yes, this is controversial, but I think it is parsimonious. So many scientists now have no problem thinking that pugs and Yorkshire terriers are Canis lupus familiaris, so why is it so difficult to think of coyotes as being Canis lupus latrans?

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We don’t really think of antlers as being practical weapons. No species of deer that has them has them year round, and in only one species, caribou/reindeer, do both sexes have antlers.

We tend to think of antlers as being used to attract the opposite sex and for ritualized combat between conspecifics, usually just competing males during the rut in all species but caribou/reindeer. In that species, the females retain antlers well into the winter, which they use fight for preferential feeding areas, a great asset in feeding the calves they carry through the long northern winters.

But we don’t normally think of them as weapons to be used against predators. After all, predators are a problem that plagues deer all year, not just during the rut, and if they were widely used in fighting off predators, one would think they would evolve to hold onto them permanently or at least for longer periods of the year.

Well, a recent paper in the journal Nature examined how these factors work with regard to elk living in Yellowstone, where wolf predation is a significant factor in elk survival.

The authors found that bull elk that lose their antlers relatively early tend to be in better physical condition than those that retain them, and those that lose their antlers early tend to grow larger antlers than those that retain them, simply because they have more time to grow their new antlers in the coming year.

The authors found that there is a massive trade-off for how long elk hold onto their antlers. Those bull elk that lose their antlers are preferentially targeted by wolves. Yes, even though they are in better physical condition than those that retain them, the wolves go for elk that lack antlers as weapons.

Predation from wolves could be driving elk in Yellowstone to hold onto their antlers longer, and it could explain why elk in general hold onto their antlers long after their breeding season.

This study has some interesting implications, because wolves could indirectly be selecting for smaller antler size in elk, simply because the elk that lose their antlers sooner tend to have bigger racks in the following year. Further, because the elk hold onto their antlers longer when they are in poorer physical condition, the wolves could be selecting for weaker elk that are much poorer foragers than they might otherwise be.

These questions were not addressed in the paper, but I’m sure the questions did arise as the researchers looked at their data. More work is going to have to be done, but it is clear that wolf predation is a lot more complex in how it selects for fitness in the elk population than we might have assumed.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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It’s not unusual for people who are trying to deny evolution or promote creationism or both, to come up with a common question:

“If evolution is true, then why don’t dogs have something that isn’t a dog every once in a while?”

This question would not be so much of a problem if we, who think we know better, would stop trying to create a species called Canis familiaris.

Canis familiaris made sense when we didn’t know what dogs were derived from, and it might have made sense if we thought there were hard and fast reproductive barriers between dogs and wolves.

But it turns out that they really aren’t such distinct animals. We’ve learned this when we’ve performed more complete assays of domestic dog and wolf genomes. Since then, we’ve found that the majority of Eurasian wolves have some domestic dog ancestry, and black wolves in North America got their black coloration as the result of a single cross with a black dog that mated with a wolf thousands of years ago in the Yukon or Northwest territories.

A recent genome comparison study of wolves and dogs that attempted to put together a phylogeny of the species clearly states:

[W]ithin the Old World clade, wolf and dog represent sister taxa. Therefore, suggestions that the dog or dingo are a separate species (Canis familiaris) (e.g., Crowther et al. 2014) would cause gray wolves to be a polyphetic taxon; and consequently, our results support dogs as a divergent subspecies of the wolf. This result has societal significance as legislation in some countries and regional governments consider wolves and dogs as distinct species restricting the possession, interbreeding, or the use of vaccines and medications in wolves or dog–wolf hybrids if they have only been approved for use in dogs. In this sense, analysis of evolutionary history informs law and veterinary practice, as dog lineages are nearly as distinct from one another as wolves are from dogs, and the justification for treating dogs and wolves differently is questionable.

The monophyly of the species is one thing that I think everyone should agree is worth preserving in any taxonomic system, but the genomes clearly show that if we create a special species for the dog or the dingo, we wreck the monophyly of Canis lupus.

I would also contend, perhaps a bit more controversially, that in light of a similar study of North American wolf-like canids’ genomes, that the coyote is also part of Canis lupus. This study found that gray wolves and coyotes have exchanged genes across North America and that gray wolves and coyotes last shared a common ancestor only around 50,000 years ago. That ancestor was probably an ancient Eurasian gray wolf that came into North America and evolved for a more generalist, jackal-like niche in the mid-latitudes of North America.

When someone claims that dogs are not wolves, they can only mean it in the same way that pugs are not Siberian huskies or that Great Danes are not dingoes. They are not wild Canis lupus, but they clearly are within that species, if we wish to keep the species monophyletic.

The reason why people want to claim a special species for the dog is because of Raymond Coppinger’s ideas still hold a lot of sway with people who wish to be learned about dogs. It’s not that everything that Coppinger said was wrong. It is what he was wrong about seems to be all that people know.

Coppinger argued that domestic dogs were obligate scavengers and thus must be placed as their own ecological species. An ecological species is the best argument for Canis familiaris. But it has limits for our understanding of evolution, and it can be turned into an absurd concept. For example, there are two sharp-tailed grouse subspecies that live in slightly different but adjacent habitat but do not readily interbreed. If we were to adhere to the same sort of species concept, then these two subspecies would have to be distinct species, even if it busted up the entire monophyly of the sharp-tailed grouse species.

Coppinger is ultimately quite wrong about the obligate scavenger status for domestic dogs. In India, for example, predation by feral and free-roaming domestic dogs is a major conservation issue. And Italian wolves are big time dump denizens. So both dogs and wolves can be predators or scavengers based upon available prey and refuse resources.

Because the ecological species concept is muddled when comparing wolves to dogs and keeping an arbitrary Canis familiaris species destroys the monophyly of Canis lupus, it would make more sense to drop Canis familiaris entirely.

One could raise dogs to Canis lupus familiaris, but Raymond Pierotti and Brandy Fogg have argued in their book, called The First Domestication: How Wolves and Humans Coevolved, that there is no set of behavioral, physical, or physiological traits that define all dogs as a taxonomic entity. They instead argue that we should just call them “domestic Canis lupus,” in which they also group the dingo, which is “feral domestic Canis lupus.

I remain agnostic about what we should call dogs, but Pierotti and Fogg’s quibbles are difficult to ignore. Perhaps we could have the subspecies for the dog, but there must be some acknowledgement that all we are doing is defining a domestic and feral population of a species.

If this blog post looks familiar, I wrote almost this exact same post in March, but I sometimes feel that I have to explain the very real scientific reasons why we don’t say that dogs are a unique species. It is not anti-science to do so, despite what Facebook dog experts tell you. If we want a monophyletic Canis lupus, then dogs have to be part of it.

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