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by Scottie Westfall

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« Dogs moving north, dogs moving south
The enigma of Hagenbeck’s wolf »

Canis oriens is not a good species

August 25, 2016 by SWestfall3

eastern coyote

Within Canis, there are lots of bogus species proposed. A few years back,  a team of researchers in Australia made some skull measurements of dingoes and decided that we should declare the dingo a new species distinct from the domestic dog and wolf. Never mind that every single genetic study on dingoes clearly puts them within the East Asian dog clade. A dingo is a feral dog very closely related to things like chow chows and akita inu.

Comparative morphology once declared the Japanese chin a distinct species, complete with their own genus, Dysodes. Never that such a thing wouldn’t pass the smell test now, it is still being tried on dingoes.

Within the genus Canis, there has always been a desire for some to split up species. Morphological variation is really great in the more wide-ranging species, but thus far, every proposed new species has come out lacking. Molecular techniques have discovered one species in this genus, the golden wolf of Africa, and there might be a distinct species of wolf in the Himalayas.

But all the rest have come up short. A recent study of wolf and dog genomes revealed that if you make dogs and dingoes a species, the entire species of Canis lupus becomes paraphyletic. So unless we want to abandon cladistics as our classification model, we pretty much have to keep dogs and dingoes within the wolf species. The red and Eastern wolves have also come up short in these studies,

And, I would argue, so has the coyote. Because coyotes split from wolves only very recently, I think a case can be made to classify them as Canis lupus latrans.

But that’s not where some people want to go. In fact, as of March this year, there was a paper that came out calling for classifying the Eastern coyote as Canis oriens.

I think this is quite unwise. For one thing, this ecomorph of coyote, which does have both wolf and domestic dog ancestry, is pretty new. Further, there is no evidence that this population is fully reproductively isolated from dogs, wolves, or the original Western coyote population. There might not be a lot of crossbreeding with domestic dogs.

But Western coyotes that are free of dog or wolf blood can still come into the East.There are no massive barriers that stop these coyotes mating with coyotes that might have wolf or dog in them.

Roland Kays, who was part of one of the original genome-wide studies of North American wolves, recently wrote a piece arguing against calling the Eastern coyote “a coywolf” or a distinct species. In the piece, Kays argues that this population of coyotes is not reproductively isolated, so it really is premature to call them a species now.

I would argue that the Eastern coyote is actually an ecomorph that is evolving to live in the human-dominated world that was once woodlands of Eastern North America, and an ecomorph is not a species. It could become one, but it takes quite a bit of time and isolation in order to do so.

In fact,  I think the big take away from all the most recent study is that coyotes are a small type of wolf.

And that means that all this splitting we’ve done in Canis isn’t really all that helpful in understanding their exact biology and natural history.

That doesn’t mean there aren’t species of wild dog to be discovered. The case to split red foxes into two species is quite strong, and there is also a strong possibility that the gray foxes of the  West and of the East are distinct species. The new species of wild dog will be found in creatures in like these, not in the larger dog species.

But Canis is where the charismatic dogs are. Wolves and their kin capture our imaginations. They are the closest relatives of the domestic dogs, and the domestic dog is descended from the Old World wolf. Within domestic dogs, we’ve been splitting them up into different varieties for thousands of years, and in the last two hundred, we’ve been doing so almost insanely. This has had to have had some effect, perhaps subconsciously, on how we view their closest relatives.

At one time, people used to go nuts naming things. Clinton Hart Merriam named dozens of species of bear in North America, which we now all recognize as belonging to one species. There is a herpetologist in Australia who does much the same thing with snakes and lizards in that country.

We live in a time when most of the larger fauna have become known to science. Pretty much the only way new species can be discovered is through trying to race molecular evolution. We don’t live in those times of gentlemen naturalists taking ships up the Congo River in search of new species of leopard.

We’ve just cataloged so much nature since that time. We don’t know it all, but we know a lot more than we did in 1880 or 1920.

And while I’d argue that the term “species” has to have a subjective element to it, it can’t be so subjective that it become squishy and useless.

And that’s unfortunately what we’re getting with things like Canis dingo and Canis oriens. 

They really aren’t that much better than Dysodes pravus.

 

 

 

 

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Posted in wild dogs | Tagged Canis oriens, coyote, Eastern coyote | 30 Comments

30 Responses

  1. on August 25, 2016 at 8:05 pm Margaret - from Oz

    The dingo was sometimes called the ‘white footed wolf’ by old timers here !


  2. on August 26, 2016 at 12:51 pm massugu

    Agreed and the golden jackal should be probably be called C. lupus aureus.

    This is another good one Scottie. Keep ’em coming, they’re such a pleasant change from all the political dreck we all post, see and share.


  3. on August 26, 2016 at 9:36 pm nebbie916

    Just like you would call the Eastern coyote an ecomorph of the coyote (Canis latrans), wouldn’t you call the Scottish wildcat an ecomorph of the European wildcat (Felis silvestris silvestris) and call both the domestic cat and the Gordon’s wildcat ecomorphs of the Near Eastern wildcat (Felis silvestris lybica)?

    Alternate names for wildcat subspecies

    1) European wildcat: Forest cat
    Ecomorph a) Scottish wildcat: Highland tiger
    2) Near Eastern Wildcat: African wildcat, North African wildcat; Desert cat
    Ecomorph a) Domestic cat: Housecat, Moggie
    Ecomorph b) Gordon’s wildcat: Arabian wildcat
    3) Southern African Wildcat (Felis silvestris cafra): African wildcat
    4) Central Asian Wildcat (Felis silvestris ornata): Steppe cat, Asian steppe cat, Asiatic wildcat, Indian Steppe cat, Indian wildcat
    5) Chinese Mountain cat (Felis silvestris bieti): Chinese desert cat

    You’re great at posting about the subject of taxonomic subjectivity.


    • on August 26, 2016 at 10:08 pm retrieverman

      I really need to do a cat piece. I just confess that I have a lot of ignorance about cats.


      • on August 27, 2016 at 8:53 pm kittenz

        Feline taxonomy is always in a state of flux. Just when (nearly) everyone reaches agreement on exactly how many species of cats there are and what to call them. along come those pesky new DNA analyses to shake things up again.
        :D


  4. on August 27, 2016 at 2:44 pm dobermann

    What needs to be understood is the difficulty involved in pigeonholing populations of organisms into neat categories called species. Sometimes the delineation between species are clear and other times not. This has always been a problem and while DNA has shed light on the relationships between the populations of some species, it has muddied the waters in other cases.

    The 50,000 year old split between wolf and the coyote may be more recent than previously thought but it isn’t exactly yesterday. 50,000 years is plenty of time for a new species to evolve. Reclassifying the coyote as a subspecies of Canis lupus is crazy talk. We would end up with new world gray wolves forming one subclade of closely related subspecies, old world subspecies forming a second subclade with perhaps the Tibetan wolf forming a subclade of it’s own. Where is the coyote? The coyote is in a subclade of it’s own dangling off to the side. Taxonomists would be debating whether the coyote ought to be classified as a species in it’s own right rather than a subspecies of C. lupus and we would be no wiser.

    To give this some sense of scale, when humans and the two chimpanzee species are compared by DNA alone the results indicate that we ought to be classified in the same genus. So either humans ought to be renamed ‘Pan sapiens’ or the chimps ought to be included in the genus ‘Homo’. By extension that would include all the Australopithecines such as ‘Lucy’ being included in the genus Pan or Homo as well. However the human ego won’t let that happen. We are above all those other apelike creatures after all or so we like to tell ourselves. “DNA doesn’t lie”, where have I heard that before?

    Canis oriens might be a good name for the eastern coyote, let’s see what their DNA looks like in a couple of thousand years time. I suppose the authors wanted to propose a species name before anybody else did.

    The Roland Kays article refers to the mating of the eastern coyote with the wolf 100 years ago and the dog 50 years ago as “events”. In other words these matings were something of note which can be measured and not everyday occurrences. Everybody has heard that those of us whose ancestors lived outside of Africa carry a little Neanderthal DNA. What is not so widely known is that all the Neanderthal DNA in people of European ancestry can be accounted for by only two Neanderthals. In East Asians there is evidence for the ancestry of three Neanderthals and for Melanesians it’s only one Neanderthal. Two ‘events’, three ‘events’, one ‘event’ maybe as few as three ‘events’ in total if they are shared.

    To illustrate the difficulty in classifying animals into nice neat groups called ‘species’, google ‘Ring species’ and read the Wikipedia article for a summary. Then see if you can get a hold of a book titled
    “Mountain Sheep: A Study in Behavior and Evolution” (1971) by Valerius Geist, a bit dated but still worth a read.


    • on August 27, 2016 at 6:12 pm retrieverman

      The reason I think that Canis latrans should be Canis latrans lupus isn’t “crazy talk” is because the variance between wolf and coyote is roughly the equivalent of the difference WITHIN wolf and coyote. Read the paper, please. There are small subspecies of wolf, like the Arabian wolf, which is coyote sized, and the extinct Honshu wolf which wasn’t much bigger. The only reason we regard wolf and coyote as different is because it’s culturally accepted and we were led astray by all that fossil evidence.

      Also coyotes and wolves have exchanged genes to a greater extent on this continent than wolves and domestic dogs have in the Old World, but you can’t honestly still want to use Canis familiaris anymore.

      Good species don’t have gene flows between them like exist between wolves and coyotes, and for an animal that has a 2-5 year generation time, 50,000 years isn’t that long a time period. This 50,000 year divergence is based upon 2-3 year generation time, but within wolves, 5 years is probably more accurate.

      I have no problem considering two species of red fox. There is no gene flow between the two, and the divergence time is great. I have a big problem with the island fox being considered a species, when it is probably part of a cryptic species, just like the North American red fox.

      I’ve not read Geist’s take on it, but I’m of the impression now that there is only one species of mountain sheep in North America, too. Stone sheep have lots of evidence of hybridization.


      • on August 27, 2016 at 7:43 pm cyborgsuzy

        Also, saying “Google it, also read a book from the ’70’s” is not the best material to use if one is arguing against a well-referenced article about a 2016 scientific paper.


        • on August 27, 2016 at 7:48 pm retrieverman

          It’s Geist too. Geist has some good ideas and some not so good ideas. I think his analysis about when wolves become dangerous to people is good, but he thinks wolves are an invasive species in North America (never mind that we’ve had wolves of some sort for at least a 2 million years. Canis edwardii was a 75 pound “wolf” that lived very nicely in North America. He also led me astray about mule deer, and I’ve got to do a revision to the post I wrote about it. The truth is I don’t have a good answer on how to classify mule vs. blacktail deer. There could be some hybridization, but I don’t think the mule deer actually is a hybrid species anymore.


      • on August 27, 2016 at 9:10 pm nebbie916

        Similarly, the reason I think that Felis silvestris catus (or Felis catus depending on the source) be left as Felis silvestris lybica isn’t “crazy talk” is because the variance between the wild Near Eastern wildcat and the domestic cat is virtually equivalent to the difference between two wild Near Eastern wildcats or between two domestic cats.

        Agreed, whereas subspecies have gene flows, good species don’t have gene flows.

        An example of good species is the human and its closest living relatives, the chimp and the bonobo. Humans have no gene flow with chimpanzees and bonobos. Also, humans diverged from chimps and bonobos 7 million or 13 million years ago depending not on the source.


    • on August 27, 2016 at 6:14 pm retrieverman

      If you’re going to make coyotes a species, then you’re going to have start splitting up wolves. Let’s try to keep species as monophyletic as possible.


    • on August 27, 2016 at 6:17 pm retrieverman

      I do know what ring species are. If you use the search function, I think that dogs will evolve into a ring species, not based upon distribution geographically, but because of size. There is isolation between the smallest dogs and the largest ones, but the in between sizes will still have some gene flow.


    • on August 27, 2016 at 6:35 pm retrieverman

      Also calling analysis based upon scientific studies “crazy talk” is the kind of ad hominem nonsense for which I have very little tolerance.


    • on August 27, 2016 at 6:52 pm massugu

      Is the coyote able to reliably reproduce with dogs and wolves? Yes. Are dogs, coyotes, Algonquin Wolves and N. American gray wolves reproductively isolated? Obviously not as witness the gradations of both dog and wolf genes in “Eastern Coyotes”-(30% eastern wolf and 10% domestic dog), as well as coyote genes in “Red wolves” (as much as 76% coyote), Great Algonquin wolves (42% coyote), and even Mexican wolves, not to mention the melanistic form of N.Western wolves being derived from an ancient cross with dogs. If isolated long enough, coyotes would almost certainly become a distinct species (they already don’t readily breed w/ N. Western wolves.) But they haven’t been, they’re not and given the ubiquity and propensities of domestic dogs (which are most definitely a subspecies of Old World wolf), not likely to be. Wherever the 3 types of canids coexist, there is genetic exchange, the amount varying with the circumstances. Based on the rule of genetic isolation alone, species status for the coyote is “crazy.”


      • on August 27, 2016 at 9:09 pm kittenz

        I wonder whether any specimens of what used to be called Canis niger currently exist from which DNA could be obtained for analysis? The “black color phase of the red wolf”, which was endemic from the lower Mississippi Valley eastward to southern Florida. I suspect that wolf was either a species itself, or a distinct subspecies of Canis lupus, depending on how you define species. The few photos that exist of them sure look more wolf- than coyote-like. It would be interesting to compare the DNA of that Mississippi Valley wolf, most individuals of which were, by contemporary accounts, black, with other wolves, particularly with the black wolves from western North America.


        • on August 27, 2016 at 9:13 pm kittenz

          I should have made it clear that I mean I wonder whether any PRESERVED specimens of what used to be called Canis niger currently exist from which DNA could be obtained for analysis, because that animal is almost certainly extinct.


          • on August 27, 2016 at 10:12 pm retrieverman

            Canis niger really was something special. It was probably Canis lupus niger, but imagine a subtropical black wolf! I’ve called it the Florida black wolf.


            • on August 28, 2016 at 3:06 am kittenz

              I agree. The photos that Tappan Gregory shot of those wolves in the 1930s definitely show something special. I’m drooling over the book – really just a 35-page museum pamphlet – that he wrote about that expedition. But I’ve only found one copy for sale, from an antiquarian bookseller in Austria, and it’s beyond my reach at the moment. I think that Gregory said, in one of his other books, that he got 5 photos altogether of the black wolves. I have never seen any photos of them other than Gregory’s; by the time that a few people had begun to realize these wolves might be special, most of them were gone.


              • on August 28, 2016 at 8:16 am retrieverman

                Bartram knew all about them in Florida. In Florida, he never say anything but black wolves, though some would have some white on the chest, which is to be expected.


  5. on August 27, 2016 at 10:51 pm dobermann

    So I guess you all favor placing humans and chimps in the same genus as well huh?

    My point was that trying to pigeonhole species into categories is always problematic. It is partly subjective and all parties will never be satisfied. It has been problematic in the pre-DNA years and it is true now. In fact in the past some scientists have considered doing away with the concept of species altogether and just look at populations. The trouble is it is not very useful from a practical point of view.
    If you think the species concept is still useful then how long does it take a species to evolve and at what point can we say a population has crossed the line and become a fully fledged species and when is it still a rather distinct population (subspecies) of the parent species?

    @massugu
    You wrote:
    “If isolated long enough, coyotes would almost certainly become a distinct species (they already don’t readily breed w/ N. Western wolves.)”
    Exactly the point, “(they already don’t readily breed w/ N. Western wolves.)”
    You also entirely missed the point of Roland Kays mention of the crossbreeding of coyote with wolves and dogs being historical “events” and not business as usual.

    You can take a common canary and mate it to the red siskin of Northern South America and get fertile male offspring, the females are sterile. In fact that is where the gene in red factor canaries came from. Despite having considerable genetic and geographical distance between them, by your definition massugu you would not consider canaries and red siskins “good species” because they can still hybridize and produce fertile young.

    Don’t take this as a personal attack on your ideas Scottie, rather see them as points of discussion. It is the Prussian in me on my father’s side that makes me speak my mind.


    • on August 27, 2016 at 11:03 pm retrieverman

      The only good reason humans and chimps aren’t in the same genus is it messes up all those transitional fossils. It’s the only good reason not to include Cuon and Lycaon in Canis. It also messes up the name of the fossil record.


    • on August 27, 2016 at 11:04 pm retrieverman

      Bird species are very different from mammals. There will always be a lot of fertility between very distinct species. Often, it’s just the song that keeps them separate species as it is with Carolina and black-capped chickadees (I live in a hybrid zone).


    • on August 27, 2016 at 11:12 pm retrieverman

      How long does it take? Well, that’s a good question. It can never be a hard answer, but when the variation between two supposed species is close to the variation within those two species and they hybridize a lot, then you’ve got a real problem.

      I have not seen the proposed dates on when Canada lynx and bobcats split, but I am willing to admit they are different species, but only because they have a very narrow and limited hybrid zone.

      The coyote and wolf hybrid zone goes from Alaska to Texas to the Maritimes. That’s not even a hybrid zone. That’s where-ever you have the two together, you get gene flow, and in the case of Great Lakes wolf, there was a gene flow between the two that happened about a 1,000 years ago.

      I think there are some good questions about why some wolves more readily mate with coyotes than others, as they still do in parts of Eastern Canada. This has traditionally been seen as these wolves being a unique “coyote sister taxa” species. We know that’s not true now.


    • on August 28, 2016 at 12:39 pm massugu

      No, I didn’t miss your point at all–I’m saying that they aren’t there yet, and given the nature of what’s been happening in the way of cross-breeding, they may never get there. Seasonality and bonding seem to be the major deterrents to successful naturally occurring dog x coyote and dog x wolf crosses; size and territoriality for naturally occurring wolf x coyote crossings. Neither set of deterrents is a totally effective reproductive barrier.

      As for that crossbreeding being limited to historical events, I would refer you to the plethora of contemporary photos of “coyotes” regularly trapped or shot and displayed on-line that show obvious dog breed coloration (classic collie markings being one) as well as the presence of rear dew claws (characteristic of many dog breeds but not of either coyotes or N. American wolves.) It has not only happened in the past, it continues to happen, despite the protestations of many experts. Googling with the search terms “images coyotes with dog markings” or “images odd-looking coyotes” will net you a number of examples along with linked descriptions.

      In re your canary x siskin example, I note that the females of the cross are sterile–not a good example of ‘reliable’ reproduction since the hybrid can’t stand on its own but must cross back to one of the parent species to reproduce.

      That is not the case with a coyote x wolf, coyote x dog or wolf x dog cross.

      The one is a hybrid, as in mules, ligons, leapons, wolphins, etc. The other is a cross like the grolar bear (yes it could be argued that the Polar Bear is actually a highly specialized subspecies of the Brown or Grizzly Bear. They’re estimated to have split about 150kya, 3 times the newest estimate for the split between coyotes and wolves.)

      Crosses (both sexes) are reliably interfertile with each other, hybrids are not—but by breeding back to one or both of their parent species, hybrids do ensure introgression of genetic traits between/among species, as apparently happened with H. sap vis-a-vis a number of other hominins.

      Time elapsed since the split of two related populations seems to be the major determinant of whether they can cross, hybridize or are no longer genetically compatible.

      This wolf/coyote/dog issue is a prime example of the huge amount of time, in relative terms, that it takes for a true species to emerge.


      • on August 28, 2016 at 2:08 pm retrieverman

        The deal is that only a tiny percentage of the siskin-canaries cocks were fertile. All hens are sterile.

        So they are like mules or savannah cats.


      • on August 28, 2016 at 2:13 pm retrieverman

        A species CAN evolve really rapidly, but you have to take in lots of different factors. Like the cichlids of Africa’s Great Lakes probably do still possess a lot of chemical interfertility and they did evolve fairly rapidly. However, the issue is there isn’t much gene flow between species, not like what exists between wolves, dogs, and coyotes.


    • on August 28, 2016 at 2:52 pm retrieverman

      Also ring species don’t exist in nature as people commonly think: https://whyevolutionistrue.wordpress.com/2014/07/16/there-are-no-ring-species/


  6. on August 28, 2016 at 1:55 am dobermann

    OK, so let’s say the consensus is that the coyote is a subspecies of Canis lupus. What becomes of the 19 recognized subspecies of Canis latrans? Do some or all of them become individual subspecies of Canis lupus as well or are they deemed not distinct enough to warrant individual subspecific status within Canis lupus?

    If the subspecies of Canis latrans are not worthy of subspecies status then what of the named subspecies of wolf found in North America? Do some of them become null and void because they are not as genetically distinct as Canis lupus latrans? The yardstick by which Canis lupus is measured has been changed. By extension this could affect the classification of old world wolves as well. The African golden wolf Canis anthus could be welcomed as a subspecies of Canis lupus and some subpecies of old world wolf could be rendered null and void as lacking distinctiveness.

    I see an open can on the table and a lot of worms crawling out of it. For the time being I think it is best to think of the coyote as a species allied to the wolf. Allied in terms of blood and genes, not because they are good friends. :)

    Going back to birds, the genus to which the red siskin belongs had traditionally been ‘Spinus’, the same genus the North American goldfinch belongs to. DNA studies caused taxonomists to reclassify the genus ‘Spinus’ as a subgenus of ‘Carduelis’. Further studies caused them to reverse their decision and undo that and set ‘Spinus’ out as a separate genus again. There is a lesson in that FOR US.

    Either way, the canary belongs to the genus ‘Serinus’. The canary can produced fertile hybrids with members of the genus ‘Spinus’. Some breeders had experimented with crossing canaries with the North American goldfinch because they thought it produced a sweeter song. In contrast, crosses of Canary x the European Goldfinch (Carduelis carduelis) have only ever produced sterile hybrids called “mules” in the bird game.


    • on August 28, 2016 at 8:05 am retrieverman

      Those subspecies become either ecomorphs or they become like breeds of dog. We don’t have subspecies created out of dachshunds or Afghan hounds. They are all Canis lupus familiars.

      You cannot have the same rules for song bird taxonomy as you do higher mammals. Because they rely so much on song to determine species recognition, species can evolve in birds rapidly. Same goes with the cichlids of the African Great Lakes. None of those animals have 2-5 year generation times, and there is also isn’t a precedent of having smaller subspecies in the Old World. If the coyote is a species, then maybe we should just say the Arabian wolf is a species. It is smaller, and if you’re just going to ignore that coyote is very close to the wolf, then why not ignore that the Arabian wolf is and just make it a species?

      The reason I’m a stickler for this we don’t classify animals without paying very close attention to evolutionary history. Cladistics has come to the fore since the 70s, and it’s important.

      My guess is that the red siskin actually diverged a lot longer than coyotes and wolves did. If you look at the divergence times for avian species they are often astronomically early, yet chemical interfertility isn’t lost. There has to be something unique about avian evolution and mutation rates, because you can get hybrids out of birds that are more genetically divergent thatn you ever could get from mammals.


    • on August 28, 2016 at 8:11 am retrieverman

      Also the fact that you get fertile hybrids between canaries and this siskin would raise real questions if we’re classifying canaries and this siskin properly. If we think that canaries are closer to goldfinches, then we’re wrong.

      Bird taxonomy is generally FUBAR, because lots of lineages diverged a long time ago but still can interbreed.

      There is very little to learn from them until we get some high resolution genomic studies on birds, which really don’t exist, These studies on carnivorans and primates are really new.

      King of red siskins or red herrings. I can’t decide.



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