The wolf is iconic and charismatic. We see him on t-shirts, on posters, and in fantasy novels. Conservationists do battle with ranchers to preserve populations of wolves. The coyote, on the other hand, is neither iconic nor loved. A newcomer to suburbia, he is feared as a suspected predator of cats, small dogs, and even small children. He is rarely seen on t-shirts; his name is not used to designate a rank of Boy Scout.
But now that we have the genetic tools to look at these animals’ genomes, it turns out that many of the populations of coyotes in North America are actually coyote-wolf hybrids, as are many of the populations of wolves. Unable to draw clear lines between these species, biologists have dubbed the populations of hybrids “Canis soup.”
What’s a Canis?
The term “canid soup” has also been used for this mess of wolf, coyote, and even dog genes that we find in some populations of canids. So what does Canis mean, and what is a canid?
These are terms related to the scientific classification of the species in question. Going through the hierarchy, we have Kingdom Animalia, Phylum Chordata, Class Mammalia, Order Carnivora, Family Canidae (canids), and Genus Canis. Wolves, dogs, jackals, and foxes belong to the family Canidae, but only wolves, dogs, and jackals (not foxes) belong to the genus Canis. We call the wolf-like canids “canines” and the fox-like canids “vulpines.”
As foxes do not interbreed with wolves, dogs, or jackals, what we’re talking about here is correctly Canis soup, or perhaps canine soup, but not canid soup.
Is it Canis or is it soup?
The more you dig into wild canines in North America, the more unclear it is where any species lines should be drawn. So who makes up our cast of characters?
The first ingredient in Canis soup is the charismatic North American gray wolf or timber wolf, Canis lupus, sometimes known as Canis lupus lupus to differentiate it from the dog and the dingo, who belong to subspecies. The gray wolf is the largest wild canine, at a 79 pound (36 kg) average weight. (Domestic dogs of some breeds, of course, weigh more than that.) Its coat coloring can vary from white through blond, brown, grey, and black. It is found in the western parts of North America.
Next is the Western coyote, Canis latrans. This animal is also known as the American jackal or prairie wolf, suggesting that there has been some confusion about how to distinguish canine species for some time. The Western coyote is a significantly smaller animal than the gray wolf, weighing in closer to 20 pounds (7-14 kg). Its coat color is less varied than the gray wolf’s, almost always a grey-brown as you see in the image here.
The range of the Eastern wolf or Algonquin wolf, Canis lycaon, is Ontario, Canada. This wolf is smaller than the gray wolf, and has a distinctive grey-red coat with black hairs along its back. We believe that this wolf was the original North American canine, and that Canis lupus and Canis latrans immigrated over the land bridge from Europe. There’s a lot of debate about the species status of C. lycaon, as many Eastern wolves appear to have significant C. latrans heritage. Some people suggest that the Eastern wolf is in fact a C. lupus/C. latrans hybrid, or, alternately, a subspecies of the gray wolf, C. lupus lycaon.
The Eastern coyote, spreading along the east coast of the United States, is significantly larger than his Western counterpart. It turns out to be a coyote/wolf hybrid, and it has been argued that it should more accurately be called a coywolf. His wolf ancestors seem to be Canis lycaon — but then again, there is debate about whether C. lycaon is really different from C. lupus at all.
The red wolf or Southeastern wolf is subject to truly intense debate about species status. Is it his own species, Canis rufus? A subset of the gray wolf, Canis lupus rufus? Or a population of Eastern wolf, Canis lycaon? It has a beautiful red coat, and is smaller in size than the gray wolf. Its range was historically the southeastern U.S., but it went extinct in the wild by 1980. A founder population of 19 animals survived in captivity, and a reintroduction project in North Carolina was begun in 1987. Here the red wolf is today enthusiastically interbreeding with coyotes, leaving conservationists to wonder what they are conserving.
The three species of wild canines in North America today, then, are Canis lupus, Canis latrans, and Canis lycaon. But we really have just two soup ingredients, wolf and coyote. There are pure wolves (Canis lupus) and there are pure coyotes (Canis latrans), and there are populations that are mixtures of more or less wolf and more or less coyote (Eastern wolves, Eastern coyotes, and red wolves). There appears to be some dog mixed in there, too. You can think of gray wolf and Western coyote as ingredients, and everything else as soup.
Coyote flavor versus wolf flavor
The 2011 paper “A genome-wide perspective on the evolutionary history of enigmatic wolf-like canids” analyzed the various soup flavors out there and presented their findings in some easy-to-understand charts (below). Here, the different colors represent different amounts of each ingredient. The first chart describes the Eastern wolf, here referred to as the Algonquin wolf, which is mostly gray wolf (green) and joint wolf/coyote (yellow), but also has significant coyote (red). The second chart describes the red wolf; at a glance, it is obvious that the red wolf has a much larger percentage of coyote genes (again, red in this chart). These charts both use τ to denote the number of generations since the most recent admixture with another species.
The two coyote recipes pictured below describe two subpopulations of what I have described as the Eastern coyote; this particular paper considers them split into Northeastern and Southeastern coyotes. At a glance, these populations are mainly pure coyote (red), with big dashes of mixed coyote/wolf (yellow), and small but notable amounts of our friend the dog (dark blue, light blue, and pink).
Wild canine populations challenge us to let go of our obsessive need to categorize. Instead of slotting a canine population into a single species category, we might instead think of it as existing on a spectrum from “wolf-like” to “coyote-like.” A strongly wolf-like canid would be larger, sixty to ninety pounds. It would require a larger range, and would be a deerivore, subsisting off of larger game. It is likely to be a shyer animal, found only in more rural or wild areas. Conversely, a strongly coyote-like canid would be much smaller, fifteen to thirty pounds, with a smaller range. It might eat deer as well as rabbits and et cetera (probably a lot of et cetera, as coyotes are more willing to scrounge than wolves are). It would be more likely to be found in suburban areas, with a greater tolerance for human proximity. A given population of canines might fall anywhere on the spectrum between the two. The fact that a spectrum actually exists is beautifully demonstrated by the Eastern coyote, who has mixed coyote/wolf ancestry, is mid-sized between coyote and wolf, and has a mid-sized range.
What’s your preferred flavor?
Does the intermixture of various ingredients in the formation of soupy populations matter as more than a gee-whiz story? To some people, the answer is very much yes. The conservationists who are committing significant resources to the preservation of the red wolf don’t want to see the wolves that they reintroduce interbreed with coyotes. If the reintroduced wolf population blends into a coyote population, then are these resources actually being spent just to support a bunch of coyotes (who have been doing fine on their own)? At the same time, evidence shows that the founder population of 19 red wolves was already significantly coyotified, and we’re not sure how long it’s been since there have been any pure Canis rufus specimens in North America.
It is, of course, possible to think about the problem without asking for genetics to provide the complete answer for us. The red wolf is a red wolf, a beautiful, iconic animal that has lived in the southeastern United States throughout living memory. We know what the red wolf looks like (and that hasn’t been changing much, no matter what is happening to his genes). We also know that it is important in a particular environmental niche, and that hasn’t been changing much either.
Practically, the mixture of coyote genes into fragile wolf populations may be a good thing. Because coyotes are better at living on smaller ranges and in closer proximity to humans than wolves are, they are better adapted to the realities of North America today. As their genes mix into wolf populations, these populations become demonstrably more robust, more able to tolerate human presence, and able to survive on smaller ranges. It is possible, in fact, that coyote genes are exactly what are eventually going to allow a red wolf population to flourish without human assistance.
Conclusions, if we can make any
Does it matter that some of what we think of as wolves have coyote genes? I think the answer comes down to a cultural perception of the wolf as a romantic and charismatic creature, and of the coyote as a pest. Perhaps any mixture of the two is perceived as diminishing the wolf. A friend of mine once made this analogy: if you have an entire bottle of fine wine, and you pour just a teaspoon of sewage into it, now you have a bottle of sewage. Does any amount of coyote, no matter how miniscule, make the wolf impure, and less worth conserving than it was?
As a culture, I hope we can come to appreciate the strengths that the coyote brings to Canis soup, in its ability to coexist with humans in the modern world. It may be what saves populations of charismatic wolves from permanent loss. As we look at populations of canines in North America, we should learn to say that one is more coyote-like and another more wolf-like, on a spectrum from one flavor of soup to another, and appreciate the benefits of both.
Canis soup has been used before as an example of the blurriness of some species lines and the inadequacy of many existing definitions of a species, but it also provides some interesting insights into the fluidity of canid morphology and behavioral characteristics. How did something as large and wild as a wolf become something as variably-sized and tame as a dog? Moreover, how did this change happen (presumably) without a carefully planned breeding program? Why is it so easy to breed types of dogs with such different behavioral and physical characteristics, especially compared to the much more limited variety of breeds of cat, horse, or cow? The canine genome clearly has the capacity for expression across a startlingly wide array of phenotypes. The evidence of this variety has always been right before our eyes, but we are just beginning to understand its implications.
References:
· Adams J. R., Leonard J. A., Waits L. P. Widespread occurrence of a domestic dog mitochondrial DNA haplotype in southeastern US coyotes. Molecular Ecology. 2003;12:541-546.
· Adams J. R., Kelly B. T., Waits L. P. Using faecal DNA sampling and GIS to monitor hybridization between red wolves (Canis rufus) and coyotes (Canis latrans). Molecular Ecology. 2003;12:2175-2186.
· Hailer Frank, Leonard Jennifer A. Hybridization among three native North American Canis species in a region of natural sympatry. PLoS ONE. 2008;3:e3333+.
· vonHoldt Bridgett M., Pollinger John P., Earl Dent A., et al. A genome-wide perspective on the evolutionary history of enigmatic wolf-like canids. Genome research. 2011;21:1294-1305.
· Way Jonathan G., Rutledge Linda, Wheeldon Tyler, White Bradley N. Genetic Characterization of Eastern ”Coyotes” in Eastern Massachusetts. Northeastern Naturalist. 2010;17:189-204.
· Wilson Paul J., Grewal Sonya K., Mallory Frank F., White Bradley N. Genetic Characterization of Hybrid Wolves across Ontario. Journal of Heredity. 2009;100:S80-S89.
· Zimmer Carl. What Is a Species? Sci Am. 2008;298:72-79.
Images: Gray Wolf (Image courtesy of vargklo at Wikipedia and Flickr); Western Coyote (Image courtesy of Rebecca Richardson at Wikipedia and Flickr); Eastern wolf (Image courtesy Christian Jansky at Wikipedia); Eastern coyote/coywolf (Image from Eastern Coyote Research); Red wolf (image from True Wild Life); Two recipes for wolf flavored Canis soup (vonHoldt, 2011); Two recipes for coyote flavored Canis soup (vonHoldt, 2011)
But now that we have the genetic tools to look at these animals’ genomes, it turns out that many of the populations of coyotes in North America are actually coyote-wolf hybrids, as are many of the populations of wolves. Unable to draw clear lines between these species, biologists have dubbed the populations of hybrids “Canis soup.”
What’s a Canis?
The term “canid soup” has also been used for this mess of wolf, coyote, and even dog genes that we find in some populations of canids. So what does Canis mean, and what is a canid?
These are terms related to the scientific classification of the species in question. Going through the hierarchy, we have Kingdom Animalia, Phylum Chordata, Class Mammalia, Order Carnivora, Family Canidae (canids), and Genus Canis. Wolves, dogs, jackals, and foxes belong to the family Canidae, but only wolves, dogs, and jackals (not foxes) belong to the genus Canis. We call the wolf-like canids “canines” and the fox-like canids “vulpines.”
As foxes do not interbreed with wolves, dogs, or jackals, what we’re talking about here is correctly Canis soup, or perhaps canine soup, but not canid soup.
Is it Canis or is it soup?
The more you dig into wild canines in North America, the more unclear it is where any species lines should be drawn. So who makes up our cast of characters?
The first ingredient in Canis soup is the charismatic North American gray wolf or timber wolf, Canis lupus, sometimes known as Canis lupus lupus to differentiate it from the dog and the dingo, who belong to subspecies. The gray wolf is the largest wild canine, at a 79 pound (36 kg) average weight. (Domestic dogs of some breeds, of course, weigh more than that.) Its coat coloring can vary from white through blond, brown, grey, and black. It is found in the western parts of North America.
Next is the Western coyote, Canis latrans. This animal is also known as the American jackal or prairie wolf, suggesting that there has been some confusion about how to distinguish canine species for some time. The Western coyote is a significantly smaller animal than the gray wolf, weighing in closer to 20 pounds (7-14 kg). Its coat color is less varied than the gray wolf’s, almost always a grey-brown as you see in the image here.
The range of the Eastern wolf or Algonquin wolf, Canis lycaon, is Ontario, Canada. This wolf is smaller than the gray wolf, and has a distinctive grey-red coat with black hairs along its back. We believe that this wolf was the original North American canine, and that Canis lupus and Canis latrans immigrated over the land bridge from Europe. There’s a lot of debate about the species status of C. lycaon, as many Eastern wolves appear to have significant C. latrans heritage. Some people suggest that the Eastern wolf is in fact a C. lupus/C. latrans hybrid, or, alternately, a subspecies of the gray wolf, C. lupus lycaon.
The Eastern coyote, spreading along the east coast of the United States, is significantly larger than his Western counterpart. It turns out to be a coyote/wolf hybrid, and it has been argued that it should more accurately be called a coywolf. His wolf ancestors seem to be Canis lycaon — but then again, there is debate about whether C. lycaon is really different from C. lupus at all.
The red wolf or Southeastern wolf is subject to truly intense debate about species status. Is it his own species, Canis rufus? A subset of the gray wolf, Canis lupus rufus? Or a population of Eastern wolf, Canis lycaon? It has a beautiful red coat, and is smaller in size than the gray wolf. Its range was historically the southeastern U.S., but it went extinct in the wild by 1980. A founder population of 19 animals survived in captivity, and a reintroduction project in North Carolina was begun in 1987. Here the red wolf is today enthusiastically interbreeding with coyotes, leaving conservationists to wonder what they are conserving.The three species of wild canines in North America today, then, are Canis lupus, Canis latrans, and Canis lycaon. But we really have just two soup ingredients, wolf and coyote. There are pure wolves (Canis lupus) and there are pure coyotes (Canis latrans), and there are populations that are mixtures of more or less wolf and more or less coyote (Eastern wolves, Eastern coyotes, and red wolves). There appears to be some dog mixed in there, too. You can think of gray wolf and Western coyote as ingredients, and everything else as soup.
Coyote flavor versus wolf flavor
The 2011 paper “A genome-wide perspective on the evolutionary history of enigmatic wolf-like canids” analyzed the various soup flavors out there and presented their findings in some easy-to-understand charts (below). Here, the different colors represent different amounts of each ingredient. The first chart describes the Eastern wolf, here referred to as the Algonquin wolf, which is mostly gray wolf (green) and joint wolf/coyote (yellow), but also has significant coyote (red). The second chart describes the red wolf; at a glance, it is obvious that the red wolf has a much larger percentage of coyote genes (again, red in this chart). These charts both use τ to denote the number of generations since the most recent admixture with another species.
The two coyote recipes pictured below describe two subpopulations of what I have described as the Eastern coyote; this particular paper considers them split into Northeastern and Southeastern coyotes. At a glance, these populations are mainly pure coyote (red), with big dashes of mixed coyote/wolf (yellow), and small but notable amounts of our friend the dog (dark blue, light blue, and pink).
Wild canine populations challenge us to let go of our obsessive need to categorize. Instead of slotting a canine population into a single species category, we might instead think of it as existing on a spectrum from “wolf-like” to “coyote-like.” A strongly wolf-like canid would be larger, sixty to ninety pounds. It would require a larger range, and would be a deerivore, subsisting off of larger game. It is likely to be a shyer animal, found only in more rural or wild areas. Conversely, a strongly coyote-like canid would be much smaller, fifteen to thirty pounds, with a smaller range. It might eat deer as well as rabbits and et cetera (probably a lot of et cetera, as coyotes are more willing to scrounge than wolves are). It would be more likely to be found in suburban areas, with a greater tolerance for human proximity. A given population of canines might fall anywhere on the spectrum between the two. The fact that a spectrum actually exists is beautifully demonstrated by the Eastern coyote, who has mixed coyote/wolf ancestry, is mid-sized between coyote and wolf, and has a mid-sized range.
What’s your preferred flavor?
Does the intermixture of various ingredients in the formation of soupy populations matter as more than a gee-whiz story? To some people, the answer is very much yes. The conservationists who are committing significant resources to the preservation of the red wolf don’t want to see the wolves that they reintroduce interbreed with coyotes. If the reintroduced wolf population blends into a coyote population, then are these resources actually being spent just to support a bunch of coyotes (who have been doing fine on their own)? At the same time, evidence shows that the founder population of 19 red wolves was already significantly coyotified, and we’re not sure how long it’s been since there have been any pure Canis rufus specimens in North America.
It is, of course, possible to think about the problem without asking for genetics to provide the complete answer for us. The red wolf is a red wolf, a beautiful, iconic animal that has lived in the southeastern United States throughout living memory. We know what the red wolf looks like (and that hasn’t been changing much, no matter what is happening to his genes). We also know that it is important in a particular environmental niche, and that hasn’t been changing much either.
Practically, the mixture of coyote genes into fragile wolf populations may be a good thing. Because coyotes are better at living on smaller ranges and in closer proximity to humans than wolves are, they are better adapted to the realities of North America today. As their genes mix into wolf populations, these populations become demonstrably more robust, more able to tolerate human presence, and able to survive on smaller ranges. It is possible, in fact, that coyote genes are exactly what are eventually going to allow a red wolf population to flourish without human assistance.
Conclusions, if we can make any
Does it matter that some of what we think of as wolves have coyote genes? I think the answer comes down to a cultural perception of the wolf as a romantic and charismatic creature, and of the coyote as a pest. Perhaps any mixture of the two is perceived as diminishing the wolf. A friend of mine once made this analogy: if you have an entire bottle of fine wine, and you pour just a teaspoon of sewage into it, now you have a bottle of sewage. Does any amount of coyote, no matter how miniscule, make the wolf impure, and less worth conserving than it was?
As a culture, I hope we can come to appreciate the strengths that the coyote brings to Canis soup, in its ability to coexist with humans in the modern world. It may be what saves populations of charismatic wolves from permanent loss. As we look at populations of canines in North America, we should learn to say that one is more coyote-like and another more wolf-like, on a spectrum from one flavor of soup to another, and appreciate the benefits of both.
Canis soup has been used before as an example of the blurriness of some species lines and the inadequacy of many existing definitions of a species, but it also provides some interesting insights into the fluidity of canid morphology and behavioral characteristics. How did something as large and wild as a wolf become something as variably-sized and tame as a dog? Moreover, how did this change happen (presumably) without a carefully planned breeding program? Why is it so easy to breed types of dogs with such different behavioral and physical characteristics, especially compared to the much more limited variety of breeds of cat, horse, or cow? The canine genome clearly has the capacity for expression across a startlingly wide array of phenotypes. The evidence of this variety has always been right before our eyes, but we are just beginning to understand its implications.
References:
· Adams J. R., Leonard J. A., Waits L. P. Widespread occurrence of a domestic dog mitochondrial DNA haplotype in southeastern US coyotes. Molecular Ecology. 2003;12:541-546.
· Adams J. R., Kelly B. T., Waits L. P. Using faecal DNA sampling and GIS to monitor hybridization between red wolves (Canis rufus) and coyotes (Canis latrans). Molecular Ecology. 2003;12:2175-2186.
· Hailer Frank, Leonard Jennifer A. Hybridization among three native North American Canis species in a region of natural sympatry. PLoS ONE. 2008;3:e3333+.
· vonHoldt Bridgett M., Pollinger John P., Earl Dent A., et al. A genome-wide perspective on the evolutionary history of enigmatic wolf-like canids. Genome research. 2011;21:1294-1305.
· Way Jonathan G., Rutledge Linda, Wheeldon Tyler, White Bradley N. Genetic Characterization of Eastern ”Coyotes” in Eastern Massachusetts. Northeastern Naturalist. 2010;17:189-204.
· Wilson Paul J., Grewal Sonya K., Mallory Frank F., White Bradley N. Genetic Characterization of Hybrid Wolves across Ontario. Journal of Heredity. 2009;100:S80-S89.
· Zimmer Carl. What Is a Species? Sci Am. 2008;298:72-79.
Images: Gray Wolf (Image courtesy of vargklo at Wikipedia and Flickr); Western Coyote (Image courtesy of Rebecca Richardson at Wikipedia and Flickr); Eastern wolf (Image courtesy Christian Jansky at Wikipedia); Eastern coyote/coywolf (Image from Eastern Coyote Research); Red wolf (image from True Wild Life); Two recipes for wolf flavored Canis soup (vonHoldt, 2011); Two recipes for coyote flavored Canis soup (vonHoldt, 2011)
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