Archive for the ‘Taxonomy’ Category

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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atila and the wolf
Photo by Tanja Askani.

In paleontology, a group of scholars exists largely on the fringe of the discipline. No matter what evidence is provided, they find some way to pump out a paper that says that birds cannot be dinosaurs. An established scholar or two will the publish and beat them down, but there is still an idea in the public mind that there is a debate between dinosaur experts about whether birds are a specific type of theropod dinosaur.

These scholars are known as BAND (“birds are not dinosaurs”), and they do get the attention of the popular press, even if ignored by the mainstream scholarship.

I’ve noticed in that in all my years writing about dogs and their taxonomy that there is a similar group in this sphere as well.  The difference is this group had the backing of one of the leading authorities on dogs in the world, Raymond Coppinger.

Coppinger was certain that dogs had to be classified as Canis familiaris, based upon a very crude ecological species concept. Village dogs that scavenge off human civilization hold a different niche than pack-hunting wolves, ergo, they are different species. Never mind that if we applied that same standard strictly, Arabian wolves, which scavenge a lot and don’t often hunt large prey, would be a different species from arctic wolves or any of the moose, elk, or bison-hunting wolves we have in North America.

If we are to adhere to cladistic classification, though, it is virtually impossible to create arbitrary species for dogs. The reason is best summed up in this paper that compared genomes of many wolves and a few dogs that have origins on different continents. The authors concluded:

 [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.

That pretty much should end this discussion. What these authors found and has been discovered in other papers is that dogs descend from a ghost population of gray wolves, Eurasian gray wolves, to be exact.

Lots of other experts agree with this assessment. Darcy Morey, an archaeologist with a great expertise in the study of Pleistocene wolves and early domestic dogs, has the address for his website as “” He and Rujana Jeger have formulated a conceptual framework of dog domestication that is quite unique. Basing their model upon trophic strategies on behalf of the wolves and shifting perceptions of humans, the authors contend that wolves that became dogs attached themselves to people. These early humans were often already acting as the apex predators in the ecosystem of the Pleistocene, and the wolves that did join up with people were able to take advantage of this niche.  Pleistocene wolves were not operating as apex predators in a faunal guild that included machairodonts, cave lions, cave bears, and Pleistocene spotted hyenas, but when those animals became extinct, the wild wolves became the apex predators of Eurasia.  The wolves that hooked up to people joined humanity in agricultural societies and joined us as apex consumers. When humans began to domesticate other livestock,  wild wolves were seen as competitors and killed off.

The idea that dogs are not wolves does have some currency, especially if you’re quite stuck on Southeast Asian origins for domestic dogs. Vladimir Dinets believes that wild Canis familiaris was some kind tropical Southeast Asian canid that was related to but not descended from Canis lupus.  There is still a massive debate as to where dogs originated, and it should be noted that there are as many good papers that have concluded European or Central Asian origins as have suggested as Southeast Asian origins.

The reason you would go for wild Canis familiaris in Southeast Asia as the ancestor is that Southeast Asia is one of the few places in Eurasia that never has had gray wolves living there. In these schools of thought, much emphasis is placed upon Canis variabilis a possibly being the wild ancestor. Of course, Canis variabilis disappeared from the fossil record 300,000 years ago, and no serious scholar thinks dogs diverged from wolves that early.

The real problem is the genetic closeness between wolves and dogs, and that same genome comparison study mentioned earlier shows a significant gene flow between wolves and domestic dogs. Up to a quarter of all Eurasian wolf genomes likely have some dog ancestry, and in East Asian wolves, the dog component of their genome can be as high as 20 percent. In European and Middle Eastern wolves, the dog component can be as high as 25 percent.

The only thing that keeps dogs from swamping the Eurasian gray wolf population with dog genes is the reproductive and territorial behavior of wolves. Wolves generally allow only one female to raise her pups. Wolves generally kill dogs that wander onto their territories, and they will kill dogs that are in territories they wish to claim.

But dog genes are getting into the wolf population at pretty high rate in Eurasia, a much higher rate than you would think of for two distinct species.

A lot of the people who have a hard time recognizing dogs as wolves are tired of bad dog training advice that is based upon bad wolf science.  They might also be tired of claims from the raw feeding community that say we must feed dogs like wolves.

But just because people misuse the classification does not infer that the classification is wrong.

Cladistically and genetically, dogs represent a now extinct population of Eurasian gray  wolves.  If these terms mean anything, then dogs are Canis lupus familiaris.

These theorists are always going to have a reason to say that dogs are not wolves, just like the BAND theorists.  Indeed, it may be necessary to refer to them as DANW (Dan-double u), for they are they are coming up with reasons to avoid classifying dogs as wolves, no matter how much genetic or archaeological evidence is presented.

In the grand scheme of things, classifying dogs has little effect on our practical understanding of them, but this continuous phylogeny denial makes the dog world seem oddly out of step.

No one would miss a beat if you called a Hereford a domesticated aurochs.  A pekin duck a domesticated mallard? No problem.

But if you say dogs are wolves, which they clearly are, then you’re anti-science.

I’m not, though. You’re the one rejecting cladistics for your special classification model.

I’m adhering to the same model that would be accepted with any domestic species and its wild ancestor.

You’re just rejecting it because you think that’s what the science says. Maybe, but it’s hard to argue with DNA.

But they do it on Maury Povich every day, so why not?

Update: A more recent study that examined the genomes of gray wolves from across their range revealed that 62 percent of all Eurasian wolves have some dog ancestry. That’s much higher than the genome comparison study mentioned above. 

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guardian fail gray fox

I was just perusing the web for photos of a Darwin’s fox (which is a “false fox” from Chile and is probably the most endangered canid species in existence right now), and I cam across this image on The Guardian’s site.

The caption reads:

A Darwin’s fox (Pseudalopex fulvipes) at the Naha-Metzabok reserve, Mexico. Three new Mexican reserves were included in the list of the World Reserve Network of the Unesco Biosphere. The organisation included the Mexican reserves of Naha-Metzabok (State of Chiapas), Marias Islands (State of Nayarit) and Los Volcanes (which has the two highest mountains in the country, the Iztaccihualt and Popocatepetl)
Photograph: Moyses Zuniga/EPA

First of all, there are no Pseudalopex/Lycalopex canids in North America. If your name is Donald Trump, Mexico is in North America.

This animal is obviously the Urocyon, the primitive gray forest and brush dog that ranges from Southern Canada to Colombia and Venezuela. Although it is endemic to the Americas, it is not a Pseudalopex/Lycalopex. It’s it’s own weird little lineage.

Here is actually a good example of parallel evolution:

The Urocyon evolved in the humid forests of what is now South Central United States and the Darwin’s fox evolved in the temperate forests of Chile. They have sort of evolved similar morphologies through living in relatively similar habitats and having relatively similar niches. Dark gray color is also perfect camouflage in a forest habitat.

But the Urocyon is much more adaptable. It’s not even close to being endangered.

But we could very well lose the Darwin’s fox.

Here’s a real Darwin’s fox for comparison:

darwin's fox

I bet if the two species had be discovered at the same time, there would have been a debate as whether they were close relatives or not.

The Urocyon was known by the seventeenth century and fully documented by the end of the eighteenth, while the Darwin’s fox wasn’t even known until Darwin (yes, that Darwin) killed one with a geological hammer. But its exact species status wasn’t fully confirmed until the 1990s. There was a debate as to whether it was forest subspecies of the more common chilla.

So no, there are no Darwin’s foxes in Mexico, but it’s good to know that this Mexican Urocyon has a nice refuge to live out its life.

And seeing as the photo was taken in 2010, it’s probably already moved off this mortal coil.

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bobcat painting

The British zoologist Richard Lydekker writes in The Great And Small Game Of Europe, Western & Northern Asia And America: Their Distribution, Habits, And Structure (1901):

For accurate information regarding this lynx (which was first named by the German naturalist Guldenstadt in the year 1777) and its numerous local races we are entirely dependent upon the writings of modern American naturalists, there being no series of specimens in England sufficiently large to admit of an independent opinion being formed with regard to certain disputed points. By some English writers, notably the late Professor St. George Mivart, the red lynx was regarded as nothing more than a local phase of the common lynx; but this view has been shown by Mr. Outram Bangs to be quite untenable, the skulls of the common and the red lynx being easily distinguishable by certain characters of the hinder part of the palate. To point out the details of this difference in a work of the present nature would obviously be out of place, and the reader must accordingly be content with the fact that such differences do exist. So important, indeed, are these differences considered by the gentle man mentioned, that he refers the common lynx to one subgenus, under the name of Lynx, while he separates the red lynx as a distinct subgeneric group with the title of Cervaria. Mr. Bangs l also considers that the red lynxes of eastern North America are specifically distinct from those of the western side of the continent, regarding the former as the true Lynx rufa (or L. rufus, as, perpetuating an original typographical error, he prefers to spell it), while the latter are assigned to Lynx fasciatus of Rafinesque. He also separates the Florida and the Texas red lynxes as a third species of Cervaria, and the Nova Scotian representative of this type as a fourth. The differences relied upon seem to be chiefly connected with the skull and bodily form. But the possibility of intergradation between these three groups is suggested; and even if this prove not to be the case, they are evidently so closely allied that, in the opinion of the present writer, they seem best regarded as local races, or phases, of a single widely spread and variable specific type. This is indeed the view of Mr. F. W. True, who writes as follows : — “The spotted form of the bay lynx, found in Texas, and the banded form, found in Oregon and Washington, have been described as separate species, under the names Lynx maculatus and Lynx fasciatus. They are now generally regarded as geographical races of the bay lynx.”

According to Mr. Bangs, the red lynx, in addition to the peculiarities of the palatal aspect of the skull already referred to, differs from the common lynx by the smaller relative size of the feet (which is most marked in the Florida race), the larger area of the bare pads on the soles of the feet, the somewhat longer tail, and the shorter pencils of hair surmounting the tips of the ears. The fur, too, is shorter and closer. In the skull the upper jaw-bone, or maxilla, forms a junction of considerable length with the nasal on each side, instead of being nearly or completely cut off from the latter; the auditory bulla on the lower surface of the skull is deeper and longer; and the whole skull is narrower, especially in the region of the muzzle. As regards the teeth, the tusks are said to be stouter and the lower molar smaller than in the common lynx.

As is indicated by its scientific and popular names, this lynx, in the summer coat, is redder than the common species; this red tinge, which in winter is restricted to the flanks, making its appearance in the typical race about February. The backs of the ears are black, with a larger or smaller greyish triangular patch; the upper lip has a more or less conspicuous black mark, and the tip of the tail may be white, with several half-rings of black above, but in other cases is black. The amount of dark spotting and striping on the back varies in the different races.

In the proportionately longer tail, the shorter ear-pencils, and the relations of the maxillae to the nasal bones, the red lynx departs less widely from more typical representatives of the genus Felis, such as the jungle-cat, than does the common lynx. The present species is a more southerly type than the latter, ranging as far south as Mexico.

In habits this lynx is doubtless nearly if not precisely similar to the common species. By American sportsmen it is usually termed the wild cat. In severe weather, according to Mr. Herrick, it is often compelled to prey upon porcupines in order to secure a living, and not unfrequently [sic] pays for its rashness with its life, examples having been killed in which the head and throat were transfixed with porcupine-quills (pg. 408-409).

The terms “red lynx” or “bay lynx” are not commonly used now. Lydekker preferred to use the name “Lynx rufa,” but we’ve since moved to “Lynx rufus.”

I have a field guide that was published in the 90s that calls them Felis rufus, but we now classify the bobcat and the other three species of lynx in their own Lynx genus.

But it was confusing for nineteenth and early twentieth century naturalists. What made it confusing was the American colloquial name for the bobcat.  If you call it a bobcat or a “wildcat,” you’re sort of implying a relationship with the wildcats of the Old World. This is probably because in parts of the South, often aren’t much larger than domestic cat, and if you realize that there are bob-tailed domestics, then you’re already going to think of them as wildcats.

And when most people living in this part of the world came from Britain, which had been free of Eurasian lynx since at least around the year 400, and they had no concept of thinking of a bobcat as a smaller species of lynx. It was easier to think of it as a species of wildcat, when you have no concept of a lynx.

To make matters more confusing, bobcats vary greatly across their range. The largest individuals can weigh over 40 pounds, but the smallest are roughly the size of relatively large domestic cat. Some populations are known for their heavy spots, while others are almost entirely one color (except on the belly and legs).  Some naturalists were of the opinion that these cats represented different species, which we now discount entirely.

Right now, we recognize two species of lynx in North America: the bobcat and the Canada lynx. However, there were always attempts to make the Canada lynx part of the Eurasian species, and I’ve seen them referred to as a type of Lynx lynx rather than as Lynx canadensis. I don’t think that’s entirely accurate. Canada lynx are snowshoe hare specialists, and they actually weigh less than the largest bobcats. Eurasian lynx, however, are quite large cats, much more closely resembling the large species of lynx which is the ancestor of them all. Eurasian lynx are generalist predators, much like giant bobcats.

But these three species are all chemically interfertile. The fourth species, the Iberian lynx, probably is as well, but it is so rare that no one would waste their genetic material with hybridization experiments. But I have seen attempts to put all of these cats into a single species, which almost universally leaves out the bobcat.

Strangely, the only two species of lynx that have been confirmed to interbreed in the wild are Canada lynx and bobcats. Eurasian lynx don’t live in North America, where they could interbreed with bobcats or Canada lynx, and there are no Eurasian lynx near the Iberian lynx’s range.

So to leave the bobcat out of the Lynx genus is pretty silly.

But it was so hard to classify them before we had a broader perspective on the cat family. There is no way you will ever get me to call a bobcat a “wildcat.”  I also think it may have been wiser to hold onto the red lynx name. “Bobcat’ might suggest we have deer-killing Manx in the forest!

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The shell of my canid classification begins with this phylogenetic tree that comes from a complete sequencing of the domestic dog genome.

What I am about to write is a very tentative taxonomic system, and I reserve the right, as would anyone, to revise it.

Here’s how I would classify the dog family in terms of cladistics.

The first thing I would do is that we have to split the dog family into tribes, which will create the clades where I will put the species.

Traditionally, there have been just two tribes for extant canids, but I think a third one is necessary.

The two traditional ones are Canini (Canis, its allies, and the South American wild dogs) and the Vulpini (which inlcudes all the foxes, including the Urocyon gray foxes and island foxes). I would argue, that because the Urocyon foxes are quite divergent from the rest of the dog family, they need their own family. They split from the rest of the dog family 9-10 million years ago.

So my tribes are Canini, Vulpini, and Urocyoni.  In Vulpini, I will put all the Vulpes foxes, the bat-eared fox, and the raccoon dog. The arctic fox and the fennec fox are now both Vulpes foxes. Canini will remain the same.

I do not believe in paraphyletic groupings within my classification, so you will notice some weird things about it very soon.

Let’s start with Canini.

In Canini, we have CanisLycalopex, Cerdocyon, Chrysocyon, Speothos, and Atelocynus. I have done away with Cuon and Lycaon.  The reason I have done so is to keep the genus Canis monophyletic. Traditional classifications have these two species, the dhole and African wild dog, as belonging to distinct genera, but they are likely sister taxa. Further, they are also less removed from the rest of Canis than the black-backed and side-striped jackals are. The rest of the genera are endemic South America canids. Two of these, Cerdocyon and Speothos, occasionally enter Panama, but they mostly a South American species.  North America was the original place of dog evolution and diversification, but today, North America is home to only three genera, the very common Vulpes foxes, Canis dogs, and the Urocyon gray foxes. Endemic North American dogs largely slipped down in South America, which is why South America is home to so many quite diverse species of canid. But all derive from North America ancestors with which they share a common ancestry with Canis.

In Vulpini, I have placed the bat-eared fox and the raccoon dog with all the Vulpes foxes.I have also split the native raccoon dog of Japan, the tanuki, into its own unique species. It has a 2n count of 38, while the mainland raccoon dog is 2n=54.  I have suggested that its scientific name should be Nyctereutes viverrinus.  I also think that evidence is pretty clear that North American red foxes are divergent enough from the Old World red foxes to be given their own species, Vulpes fulva.

In Urocyoni, I would have but a single species, because it turns out that island foxes probably split much more recently that was previously thought.  In this case, I would say that the island fox is a insular dwarf subspecies of the mainland Urocyon, and it will have but a single species in it.

So allow me to list my species in their tribes.

In tribe Canini:

Genus Canis:

  1. Black-backed jackal (Canis mesomelas)
  2. Side striped jackal (Canis adustus)
  3. Dhole (Canis alpinus, instead of Cuon alpinus).
  4. African wild dog (Canis pictus, instead of Lycaon pictus).
  5. Eurasian jackal or golden jackal (Canis aureus).
  6. African golden wolf (Canis anthus, which used to be a golden jackal).
  7. Ethiopian wolf (Canis simensis)
  8. Coyote (Canis latrans)
  9. The wolf/dog/dingo species (Canis lupus).

I do not recognize domestic dogs as distinct species from the Holarctic wolf species, and I also do not recognize the red wolf or Eastern wolf as valid species. Domestic dogs are essentially a specialized wolf that can live with and easily read human beings. Dingoes, despite rather desperate attempts to make it otherwise, are nothing more than an East Asian feral dog that has made its home in Australia. Eastern wolves and red wolves are recent hybrids between wolves and coyotes that occurred after European settlement of the continent. It was only recently revealed that the African golden jackal was quite genetically distinct from the Eurasian golden jackal, and now that species has been divided into Canis anthus and Canis aureus.

So that’s the genus Canis. 9 species. There are potentially two more, because it turns out that two wolf populations, one in India and one in the Himalayas have rather unique mitochondrial DNA sequences. It would be interesting to see how those two populations within a genome-wide analysis.

The rest of Canini are all South American endemics.

Most are in the genus Lycalopex. Here are the species:

  1. The culpeo (Lycalopex culpaeus).
  2. The chilla or “South American gray fox” (Lycalopex griseus)
  3. The Pampas fox or Azara’s fox (Lycalopex gymnocercus)
  4. The Hoary fox (Lycalopex vetulus)
  5. The Sechuran fox (Lycalopex sechurae)
  6. The Darwin’s fox (Lycalopex fulvipes)

The word “Lycalopex” is a combination of the Greek words wolf and fox (“lykos” and “alopex.), and they actually do look like bantamized wolves or coyotes. Indeed, I’ve seen less-informed people use photos of culpeos as coyotes.

They are superficially fox-like. I have seen them called foxes and zorros my entire life, but it is hard to explain to people that these animals are much more closely related to wolves and dogs than they are to foxes.

Also, it has only been since 1996 that the Darwin’s fox has been given full species status. It was previously thought of a subspecies of chilla that lives only in the temperate rainforests of Chile, but when its mitochondrial DNA was compared to several chillas, it was found that they were quite distinct from each other. The Darwin’s fox is most closely related to the Sechuran fox, which also has a very narrow range along the Pacific Coast of Ecuador and Peru. Darwin’s fox is arguably the most endangered of all canids. It lives on Chiloé Island and Nahuelbuta National Park. A small population was also discovered in Valdivian Coastal Range.  There are only about 250 of them on Chiloé Island and about 70 on the mainland. That’s an estimated 320 individuals, which is a bit more than the estimated 350-440 Ethiopian wolves. For some reason, we tend to hear more about Ethiopian wolves than Darwin’s foxes, but they certainly are deserving of our attention.

I should note that the genome-wide analysis found the Darwin’s fox to be a bit more divergent from the Sechuran fox, which was closer to the culpeo. So it does need a bit more work to figure out how those relationships work.

The rest of Canini are what I call the South American weirdos. This is where canid evolution went a little strange. Here we have a dog on stilts, a basset-type dog with the dhole’s trenchant heel dentition, a dog that lives on a lot fish, and one that eats crabs. Also, the only dog species to have gone extinct in historic times is here too.

So here are the species:

  1. The maned wolf (Chrysocyon brachyurus)
  2. The Falkland Islands wolf (Dusicyon australis)
  3. The bush dog (Speothos venaticus)
  4. The crab-eating fox (Cerdocyon thous)
  5. The short-eared dog (Atelocynus microtis)

The Falklands wolf is now extinct. It was an unusually curious animal that was soon killed off to make way for sheep farming. No one knew what it was related to until just a few years ago, when it was discovered that its closest relative was the maned wolf. It has been suggested that this animal was a living fossil that resembled the ancestral South American canid that came down into that continent from North America before branching into the Lycalopex and “weirdo” forms.

The maned wolf itself looks like a large red fox with really, really long legs. Its closest living relative is the bush dog, which is the only native pack-hunting canid in South America. It looks like an unholy hybrid of a basset hound and otter, and it has evolved the dhole and African wild dog’s trenchant heed dentition in parallel. It was even suggested at one time that it was a type of New World dhole, based solely upon the teeth.

The last two are closely related enough that one might be able to put them in a single genus. The crab-eating fox is pretty common in South America, and it is often seen out on beaches and river banks searching for food. It was commonly seen eating crabs, so it got that name. The short-eared dog, its closest relative, is almost never seen and has hardly been studied. It is perhaps the strangest dog still in existence. It has a long, bull terrieresque muzzle and little rounded ears. It has been known to eat quite a bit of fish in its diet, but other than that, very little is known about them. They live in very low densities in the Amazon Basin.

We have finished the Canini tribe, now onto the Vulpini.

The Vulpini go as follows:

First the genus Vulpes:

  1. Old World red fox (Vulpes vulpes)
  2. North American red fox (Vulpes fulva)
  3. Rüppell’s fox (Vulpes rueppellii)
  4. Corsac fox (Vulpes corsac)
  5. Tibetan fox (Vulpes ferrilata)
  6. Swift fox (Vulpes velox)
  7. Kit fox (Vulpes macrotis)
  8. Arctic fox (Vulpes lagopus)
  9. Blanford’s fox (Vulpes cana)
  10. Bengal fox (Vulpes bengalensis)
  11. The Cape fox (Vulpes chama)
  12. The pale fox (Vulpes pallida)
  13. The fennec fox (Vulpes zerda)

The exact relationship between fox species isn’t that clear, but generally, most authorities recognized that kit and swift foxes are distinct species, even though they do have a limited hybrid zone. It is also clear that the arctic fox is a very close relative of those two species, and perhaps the best way to think of the arctic fox is that it is a swift fox that has become specialized to the arctic ecosystems. As I mentioned earlier, a study at UC Davis found that red foxes in North America diverged from the Old World red fox 400,000 years ago.  This divergence is enough to make some consider them a distinct species, and it should be noted that even red foxes that were said to have derived from European imports that were released on the East Coast are also just as divergent. Which means they aren’t derived from these imports, as was initially believed.

The addition of the others in Vulpini are a bit more controversial, but I’ll say that bat-eared foxes are vulpines, as are raccoon dogs. So the rest of Vulpin is:

  1. The bat-eared fox (Otocyon megalotis)
  2. The common raccoon dog (Nyctereutes procyonoides).
  3. The Japanese raccoon dog or tanuki (Nyctereutes viverrinus).

All of these are earlier offshoots of the fox lineage, but they are close enough to foxes to be considered Vulpini. As mentioned earlier, there are good reasons to think that the tanuki is a distinct species, but its exact status is still somewhat controversial. Bat-eared foxes have the most teeth of any canid, and they pretty much eat nothing but harvester termites. They are found in disjointed populations in East Africa and Southern Africa.

And finally, I have made the decision to raise the Urocyon “foxes” to their own tribe, and because of their genetic similarity, I have reduced them from two species to one.

So tribe Urocyoni is:

  1. The North America gray fox or tree fox (Urocyon cinereoargenteus).

So in my classification of Canidae, I have 36 living species, with two more that could be identified in the form of the Indian and Himalayan wolves.

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A side-striped jackal (Canis adustus), the jackal no one talks about.

We’ve always called the smaller wild dogs in the genus Canis jackals.

Historically, there were four species of jackal:  the golden (Canis aureus), the black-backed (Canis mesomelas), the side-striped (Canis adustus), and the Simien (Canis simensis).

Some authorities considered they coyote (Canis latrans) to be a jackal, usually called “the American jackal.”

At one time, they were all placed in the genus Thous.

This, of course, assumed that these animals were all closely related to each other.

However, as we’ve looked at DNA analysis, the relationship between jackals shows that the term “jackal” is actually quite meaningless.

In 1994, an mtDNA study revealed that the Simien jackal had certain mtDNA sequences that were more similar to wolves than other jackals. It was thought to be a relict population of primitive wolves that came into Africa during the Pleistocene.

And from that time forth, the English name of this species was change to “Ethiopian wolf.”  I don’t call it anything else.

However, as more work was performed on jackals, certain facts became evident.

Initial studies of black-backed, side-striped, and golden jackal mtDNA revealed that black-backs in East Africa had huge variances in their mtDNA. Golden jackals had mtDNA that was most similar to coyotes and wolves, while black backs and side-stripes were more similar to each other.

And then the phylogeny of the dog family was drawn from a high-quality sequencing of the dog genome revealed that golden jackals were much more closely related to coyotes and the wolf and domestic dog species than the Ethiopian wolf was. We still call them Ethiopian wolves, even though golden jackals are more closely related to actual wolves than those animals are.

The other issue revealed through this research was that Canis, as it is traditionally classified, is a paraphyletic genus. Modern taxonomy is generally concerned with classifying animals according clades. Clades are, by definition monophyletic. That is, they contain all the animals that descend from a particular lineage.

The dog genome research revealed that two species that are never traditionally classified as being part of Canis, the African wild (Lycaon pictus) and the dhole (Cuon alpinus), actually should be included there.  It turns out that black-backed and side-striped jackals are more distantly related to the rest of Canis than these two species are.

And if we classify Canis with all the jackals, the Ethiopian wolf, and  the wolf and dog species and leave out the dhole and African wild dog, we’ve created a paraphyletic genus that is not useful to modern taxonomy.

Some have suggested giving the two endemic African jackals their own genus.

And this would make Canis monophyletic without including the African wild dog and dhole.

However, the genus that would remain would include several species that are all chemically interfertile with each other (at least in theory). Species complexes exist throughout that part of the Canis, and delineating species is very difficult the species in this lineage.

Although Robert Wayne at UCLA has suggested that black-backed and side-striped jackals might be able to interbreed, no one has confirmed a hybrid between these two species. African wild dogs might be able to hybridize with dholes, but because they live on different continents and because they are both fairly endangered, no one has attempted to cross them. (There are persistent rumors that dholes can cross with domestic dogs. One dog breed, the Bangkaew dog from Thailand, is said to have derived from a dog/dhole cross. However, I don’t believe this claim has ever been tested through DNA analysis.)

All this research has revealed that how we have traditionally thought about the dog family is probably wrong.

The golden jackal is actually a primitive offshoot of the wolf lineage, just as the coyote in the New World is. The Ethiopian wolf is an even more primitive offshoot.

The two endemic African jackals are the two oldest living species in the Canis lineage. They are even more distinct from this lineage than dholes and African wild dogs are.

We do not have a good replacement word for jackal.

I’ve suggested that we call golden jackals “Old World coyotes” almost as a joke.

But I don’t have a good name for either of the two remaining jackal.

Because black-backed jackals are so scrappy, I’ve even suggested that we call them “wild Jack Russells.”

Whatever we call them, the term jackal, if it’s used to reflect close relationships between species, is utterly meaningless.

With the exception of the two found only Africa, it doesn’t refer to any animals that have a close relationship with each other.

It’s just a term we use for smaller wild dogs that are in some way related to wolves.

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African wild dogs (Lycaon pictus or Canis pictus) are more distantly related to dogs than humans are to chimps. African wild dogs cannot crossbreed with dogs, and they are not an ancestor of any domestic dog breed. This species has suffered greatly because of its common name. Many ranchers and pastoralists in Africa have persecuted this species under the suspicious that it is nothing more than a feral domestic dog. It’s not even an ancestor of the domestic dog. It’s a truly unique but endangered species.

Stanley Coren has posted a slide show on Huffington Post entitled Dog Facts: 10 Things You Didn’t Know About Your Pet.

I couldn’t get past the first slide because this is what it said:

How do scientists decide whether dogs started out as wolves? One way is to try to cross breed the species by mating a wolf to a dog. If that mating produces live puppies that are fertile and can have pups, then that means that the wolf and dog are the same species. It turns out that dogs can have puppies, not only with wolves, but also with jackals, coyotes, dingoes, African wild dogs, and even some kinds of foxes. Although genetic research says that the first species that humans domesticated was the wolf, the guess is that dogs are mixture of all of these different wild canines, which probably explains the existence of so many different looking dogs in the world.


One wonders if Coren consulted any of the genetic literature on domestic dogs.

It turns out that their wide variance in phenotype comes from two really basic weird aspects of the dog genome. One of these is that just few genes account for massive differences in phenotype in domestic dogs. Variation on just three genes produces almost all the variance in coat that domestic dogs. That’s somewhat shocking, I know.

Further, it’s been found that all dog species have an unusually high number of tandem repeats in their genes.  Selection selective breeding animals with such high numbers of tandem repeats often results in massive changes in phenotype within just a few generations. That’s why dogs breeds change so rapidly through the years.

Small size in most domestic dogs has been traced to a single gene that likely originated from domesticating the Middle Eastern wolf. Arabian wolves are often as small as 25-30 pounds– much smaller than other subspecies– and have also been implicated in providing a lot of the genetic material to modern domestic dogs.

Now, all the genetic literature points to the wolf as the primary and perhaps sole ancestor of the domestic dog. Coren totally screws the pooch when it comes to classifying the dog and its closest relatives.

Dingoes are not an ancestor of the domestic dog. They are  descended from domestic dogs that came to Australia from Indonesia as domestic animals. In every genetic study I’ve seen, dingoes group with East Asian domestic dog breeds, which also share an affinity with the Chinese wolf.

The best way to classify dogs, dingoes, and wolves is to count all three as belonging to the same species. Conventionally, dogs are Canis lupus familiaris, and dingoes are C. l. dingo. I think that in light of what we’re seeing in the genetic studies, dingoes ought to be classified as Canis lupus familiaris.  The basenji, which is always classified as a breed of domestic dog,  is actually much more genetically distinct than the dingo is.

So Canis lupus encompasses dogs, wolves,  and dingoes (including the New Guinea singing dog).

That species is interfertile with the coyote, and many coyotes have genes from wolves and domestic dogs. Certain wolves in Eastern Canada have genes from coyotes, and the so-called red wolf, which is primarily coyote in ancestry but does have some ancestry from wolves.

Golden jackals can also crossbreed with dogs, and it’s possible that domestic dogs have a bit ancestry from this animal. Evidence for this ancestry has not yet been discovered. Only the Sulimov dogs, which are recent intentionally bred golden jackal/dog hybrids, have been proven to have golden jackal.

Ethiopian wolves can also cross with dogs, and hybridization with domestic dogs in the Bale Mountains has been considered a major threat to their continued survival.

Other than those animals I’ve just listed, there is no proof other than anecdote and lore that these dogs have crossed with any other wild dogs. There is no genetic evidence of a dog and red fox hybrid, though there were always alleged ones. There is no evidence of African wild dogs crossbreeding with any dogs, except from breed lore from Rhodesian ridgebacks and basenjis, and there is no evidence other than unsubstantiated claims that black-backed and side-striped jackals have ever hybridized with dogs.

There is a breed origin story for a breed from Thailand called the Bangkaew dog that claims dhole ancestry. Supposedly, they are derived from a bitch that got impregnated by dhole in the forest. There were no other domestic dogs around, so it had to have been a wild dog.

I think that golden jackals are a much more likely wild ancestor of this dog, if it does indeed have blood from another species. However, as far as I know, no one has looked for the genes of either dholes or golden jackals in this breed.

Except for dogs that were intentionally bred to coyotes or golden jackals, there is no evidence that species other than the wolf have contributed to domestic dogs.

There are always stories about different wild dogs crossing with domestic ones.

However, there has been no evidence of dogs crossing with anything other than the species I’ve listed here. All the rest are nothing more than stories, folk tales, and speculation.

I think one reason why people hold onto the hope that dogs are derived from a wide range of species is they secretly want to vindicate Konrad Lorenz and Charles Darwin, who may have been right about many things.

But they were wrong about the ancestry of the domestic dog.

Dogs, like it or not, are domesticated wolves that have some weird features in their DNA that make them very easy to mold into bizarre shapes through selective breeding.

They did not derive from hybrids of several dozen wild species.

Coren is simply wrong here, and I don’t know why he keeps makingthese claims. He made this same claim in The Intelligence of Dogs, when the genetic literature wasn’t as clear as it is now,and in How to Speak Dog.  Every genetic study shows that dogs are derived from wolves, and modern dog lineages are likely derived primarily from the wolves of the Middle East. No matter how many times Coren or anyone else wants to claim that dogs are derived from many species, the claim will be still be false. And it’s even worse that he makes claims that dogs are derived from species with which they cannot even cross.

There are so many errors in that first slide that I couldn’t get to the rest.

See related post:

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