Saturday, December 31, 2011

The hearty ingredients of Canis soup

[Reposted from my post on Scientific American’s guest blog.]

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?

Gray wolf
(Image courtesy of Wikipedia)
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 him 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.) His coat coloring can vary from white through blond, brown, grey, and black. He is found in the western parts of North America.

Western coyote
(Image courtesy of Wikipedia)

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). His coat color is less varied than the gray wolf’s, almost always a grey-brown as you see in the image here.

Eastern wolf
(Image courtesy of Wikipedia)
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 his 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.

Eastern coyote/coywolf
(Image linked from
Eastern Coyote Research)
The Eastern coyote, spreading along the east coast of the United States, is significantly larger than his Western counterpart. He turns out to be a coyote/wolf hybrid, and it has been argued that he 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.

Red wolf
(image linked from True Wild Life)

The red wolf or Southeastern wolf is subject to truly intense debate about species status. Is he his own species, Canis rufus? A subset of the gray wolf, Canis lupus rufus? Or a population of Eastern wolf, Canis lycaon? He has a beautiful red coat, and is smaller in size than the gray wolf. His range was historically the southeastern U.S., but he 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.

Two recipes for wolf flavored Canis soup
(vonHoldt, 2011)

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).

Two recipes for coyote flavored Canis soup
(vonHoldt, 2011)

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. He would require a larger range, and would be a deerivore, subsisting off of larger game. He 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. He 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). He 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 he 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 his ability to coexist with humans in the modern world. He 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.


Adams, J., Leonard, J., & Waits, L. (2003). Widespread occurrence of a domestic dog mitochondrial DNA haplotype in southeastern US coyotes Molecular Ecology, 12 (2), 541-546 DOI: 10.1046/j.1365-294X.2003.01708.x

Adams, J., Kelly, B., & Waits, L. (2003). Using faecal DNA sampling and GIS to monitor hybridization between red wolves (Canis rufus) and coyotes (Canis latrans) Molecular Ecology, 12 (8), 2175-2186 DOI: 10.1046/j.1365-294X.2003.01895.x

Hailer, F., & Leonard, J. (2008). Hybridization among Three Native North American Canis Species in a Region of Natural Sympatry PLoS ONE, 3 (10) DOI: 10.1371/journal.pone.0003333

vonHoldt, B., Pollinger, J., Earl, D., Knowles, J., Boyko, A., Parker, H., Geffen, E., Pilot, M., Jedrzejewski, W., Jedrzejewska, B., Sidorovich, V., Greco, C., Randi, E., Musiani, M., Kays, R., Bustamante, C., Ostrander, E., Novembre, J., & Wayne, R. (2011). A genome-wide perspective on the evolutionary history of enigmatic wolf-like canids Genome Research, 21 (8), 1294-1305 DOI: 10.1101/gr.116301.110

Way, J., Rutledge, L., Wheeldon, T., & White, B. (2010). Genetic Characterization of Eastern “Coyotes” in Eastern Massachusetts Northeastern Naturalist, 17 (2), 189-204 DOI: 10.1656/045.017.0202

Wilson, P., Grewal, S., Mallory, F., & White, B. (2009). Genetic Characterization of Hybrid Wolves across Ontario Journal of Heredity, 100 (Supplement 1) DOI: 10.1093/jhered/esp034

Zimmer C (2008). What is a species? Scientific American, 298 (6), 72-9 PMID: 18642545

Wednesday, December 21, 2011

Book review: Control unleashed

I recently finished Control Unleashed: Creating a focused and confident dog, by Leslie McDevitt. This book is designed for people training dogs in agility who are having issues with their dogs’ ability to focus, so you might imagine that the book isn’t useful to people with fearful dogs like mine. But it turns out, unsurprisingly, that if your dog is having issues focusing, he may well actually be nervous about something in his environment, such as strange people or other dogs. This book is full of exercises for helping your dog be more comfortable and relaxed, whatever his reason for being distractable. It was incredibly helpful to me in thinking through exercises for my shy dog Jenny, to help her learn to trust the world a little more. Even if you don’t do agility with your dog, this book may be helpful in making you a better trainer. It includes many examples of specific training exercises to try, and stories about situations in which they were particularly helpful. It was also an enjoyable enough read that I got through it while working long hours on some difficult rotations — I looked forward to finding ten minutes to read it before bed. I recommend this book to everyone who has has basic understanding of positive training and learning theory, and wants to learn more.

Sunday, December 4, 2011

I have become a chef. For owls

Some days it seems like none of your patients will eat. I asked the intern at the wildlife clinic where I was doing a two week rotation what his advice was for convincing the tiny screech owl (214 grams of cute) to actually ingest food on her own. We had tube fed her; this worked, but you don’t really want to have to do it every day, and she really doesn’t want you to do it every day. We had force-fed her mouse bits with tongs, which was even worse. The intern suggested peeling the skin off of baby mice “so she can see the meat.“


In vet school, you get a little inured to gross things. I cut up mice for raptors every day on this rotation without getting squeamish. But I found peeling the skin off of two day old mice (thankfully already dead) to be a bit much. The intern was a little bewildered by my reaction. Anyways, it didn’t work.

The next day I went to a different vet for advice. “Sometimes,” she said, “they like to have the mice split open and the liver displayed... You can leave the skin on.” I did this. I was artistic about it. HERE ARE THE LIVERS. PLEASE EAT THEM.

And that night, she ate.

My next problem patients were two barred owls who had recently been transported a fairly long distance, which we were examining before they were placed into permanent captivity, as they were not releasable. These guys really didn't want to eat. I laid the mice out enticingly next to a branch on the ground. I split them open and DISPLAYED THE LIVERS. No.

The intern said, “They don’t want to come off their perch. They’re scared. Put the mice on the perch.” Easier said than done (he said it, I did it). One of the owls inevitably reacted to my entry into the pen with OH HOLY CRAP I AM GOING TO DIE, flying back and forth at high speeds. It wasn’t a very big pen. I covered my head with my arms and crept at a snail’s pace towards the perch. Veeeeery slowly I balanced the mice on the perch. And crept away again. That night, the owls ate.

The next day, as I was artistically displaying peeled mouse haunch for an Eastern box turtle (the mealworms weren’t working as they would just escape into his cage and set up shop there) I thought to myself: how did this become my life?