Posts Tagged ‘black wolf’


Photo by Jiangou Xie

Yep. It’s a real photo!



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One of the pleasures of my Alaska trip was meeting Nick Jans. Nick Jans is the author of A Wolf Called Romeo, which is the story of the black wolf that came out to play with free-running dogs at Juneau’s Mendenhall Glacier.

I may have written a few things on Romeo on this blog before. He was just that fascinating an animal. Most wolves want to off the dogs they encounter. This one decided to become friends with them and even tolerate the humans who came with them.

The book is a wonderful discussion of wolves and dogs and people and what they truly mean to us and what we mean to them. It also tells the story of an odd wolf, who lived out six incredible years running and playing with the local domestic dog contingent.

The story does not have a happy ending, but the story of a wolf coming to trust people and dogs is something so amazing that you would have to look into the fiction of Jack London to find something even remotely similar.

But this is a true story.

If you would like to know more about Romeo, Jans gave a talk on the ship about the book that was an abbreviated version of this one:

My friend Bronwen Dickey wrote a review of the book in the New York Times I just happened to have been the one who mentioned the book to her over two years ago, and I guess I played a tiny role in getting this book the wonderful review it received.

I received a copy of The Giant’s Hand, which is Jans’s new collection of short stories about life in the Inupiaq Village of Ambler  and his experiences in Alaska’s far north.



The prose in each of these stories is so beautiful. He really can capture the essence of a place with words in a way that very few modern writers are able to match.

I particularly love the stories that include the exploits of Clarence Wood, an Inupiaq hunter and wolf trapper. He is a man of particular genius about the land and its wild inhabitants, but his way of phrasing things is just so perfect if a bit eccentric.

My favorite is: “Too much think about bullshit. That’s what makes you nervous.”

I think I may have to put this on a rock somewhere.

My favorite story in the book thus far is “Crossing Paths.” It is a kind of future warning about Romeo. In the story, Jans meets a red fox near his home, and wanting to get to know it better, he starts leaving out bits of food for it. Things go well until a neighbor shoots it for fear it might be rabid.

Jans has a philosophical discussion in the story about how much wild even Alaskans are willing to tolerate. The truth is that everyone has some limit.

Romeo was not fed to bring him near to humans. He merely came by to socialize with dogs and a few select people.

But Romeo wound up like that poor red fox in the arctic. He wasn’t taken because there was a fear he might be rabid. He was killed by two poachers who just wanted to cause trouble.

As a species, we have a very odd relationship with the wild. We admire it. We want to be part of it. But we also want it to be on our terms.

Like it or not, we’ve long since left the garden. We can only be visitors here, but some of us can truly be at home for a while.

And that’s the best we can do. Unfortunately.

It goes without saying that this book is the best souvenir I’ve ever brought home. I mean I do have a t-shirt from the Chilkat Bald Eagle Preserve, but nothing can compare to this book.

This was the trip of a lifetime.





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Not all wolf and dog interactions are adversarial.

Take the story of Romeo.

Romeo was a black wolf who liked to hang out at the Mendenhall Lake near Juneau, Alaska.

Lots of people walked their dogs there, and Romeo joined up with them.

According to most stories, Romeo lost his mate, and then he tried to bond with a female Labrador retriever that was brought to the lake on a regular basis.

He eventually extended his friendship toward virtually all dogs that were walked there.

In 2010, Romeo went missing.

Although his exact fate has never been confirmed, it has been claimed that a hunter killed Romeo.

But while he was alive, Romeo’s interaction with many different domestic dogs became something of an internet sensation.

A simple Google Image search for “Romeo the wolf” will reveal dozens of photos of a striking black wolf standing next to an assortment of common domestic dog breeds.

The juxtaposition of a Northern-type wolf with mostly Western dog breeds is quite striking.

One can see how much more robust this sort of wolf is.

One can tell that his kid of Canis lupus evolved to hunt the big game. Their massive teeth and jaws help them drop moose and bison.  Their massive heads anchor enormous jaw muscles that allow them to deliver punishing bites and break thick bones.

If the latest genomic data is correct, the modern strains of domestic dog are derived from an entirely different sort of wolf, the more primitive and slightly-built “Southern” wolves of South Asia and the Middle East.

Romeo is the sort of wolf who would have contributed his genes to domestic dogs.

In the thousands of years that existed before humans regularly spayed and neutered and confined dogs, wild genes had a way of working their way into the population.

Unattached wolves like Romeo would sometimes mate with the domestic bitches they encountered.

Dog and wolf initially started out as a cultural distinction.

Humans regarded the wolf as wild and the dog as tame.

They are still the same species, but now each has adapted to its niche.

Just as the Romeo’s subspecies has adapted to hunting large game in frigid environments,  the wolves who became dogs adapted to living in the human world.

Here are two types of wolf that have specialized and adapted to very different environments.

One of them has hitched its evolutionary wagon to the Homo sapiens star.

And it has thrived beyond any reasonable expectations for a relatively large carnivoran.

The other subspecies is doing okay.

But it will never have the success or the range of the human-adapted one.


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All black wolves that have been examined in modern times have been found to be dominant blacks. The dominant black mutation first originated in domestic dogs and was transmitted through crossbreeding between wolves and dogs. However, there is at least one record of a wolf that was carrying recessive black.

Recessive black is most commonly found in German and Belgian shepherds. It can also be found in pulik, Samoyeds, schipperkes. Shetland sheepdogs, and the so-called “American Eskimo dog,” which is actually an American variant of the German spitz.

It’s one of two ways that a dog can be solid black, but it’s far less common than dominant black.

The mutation that causes dominant black originated either in dogs or the wolf population that became dogs, because the mutation is older in domestic dog populations than in wolves. This black coloration wasthen transmitted to Italian and New World wolves through cross-breeding with domestic dogs.  All wolves that have been examined in North America thus far have turned out to be dominant blacks, as have those in Italy.

However, there was at least one case of a wolf carrying recessive black in the literature.

The Soviet zoologist and dog expert N.A. Iljin carried out several experiments crossing various dogs with wolves. In 1941, he reported on the progeny of a male wolf that was bred to a female mongrel sheepdog.  In the first litter, there were black and “zonar gray” (wild wolf gray puppies). If the dog in question were a dominant black, then the entire litter would have been black, but getting gray puppies suggested a very different conclusion.

After breeding from the offspring for several generations, Iljin discovered that the black was being inherited as a recessive allele, which means the dog in question was a recessive black– and the wolf was a carrier!

Now, results of Iljin’s study have been used to show that wolves carried recessive black from the beginning.

However, since the time of Iljin’s work, no one has found a recessive black wolf.  The team of geneticists at UCLA have found only dominant black in wolves.

So it’s possible that this wolf was not actually “pure,” and at some point, one of its ancestors was a recessive black dog. I would not be surprised if someone had crossed a recessive black German shepherd into captive Russian wolves at some point. Iljin himself was very much into breeding German shepherds to wolves, and his studies on wolf and German shepherd morphology are pretty much classic literature for those interested in wolves and dogs.

So maybe recessive black did exist in certain Old World wolves from the beginning, but it’s just not been confirmed in the genetic literature in the same way that dominant black has.

I don’t know of another species besides Canis lupus that has two separate genetic variants for melanism. Coyotes have inherited dominant black from breeding with either dogs or wolves, and golden jackals and Ethiopian wolves could also inherit both types of melanism through similar hybridization.

So it’s very interesting that we have this one case of a wolf carrying recessive black, but we need more information to see where this color came from.

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Stories of easily tamed wolves are not unique to North America. In 1866, a British colonel and big game hunter named Alexander Angus Airlie Kinloch–who was then  serving in the King’s Royal Rifle Corps in Ladakh in northern India– managed to procure two black wolf pups that grew up to become very tame adults. He recounts the story about the tame wolves in his treatise on the big game animals called Large game shooting in Thibet, the Himalayas, and northern India (1885):

On the 5th June, 1866, I was encamped at the foot of the Lanak Pass, between the Tsomoriri Lake and Hanle, when one of my servants brought in a young Black Wolf apparently about three weeks old. He had procured it from some wandering Tartars, and informed me that they had another one. I at once recognized the value of my prize, and sent off a man to secure the other cub, which arrived next morning. I had only heard of one other Black Wolf having been met with by Englishmen, and that had been shot the previous year in the neighbourhood of the Mansarovara Lake. I was, therefore, particularly anxious to keep the young Wolves alive, and in this I was fortunately successful. Emptying a ‘kilta’ I converted it into a kennel for the cubs, which I fastened to opposite ends of the only dog chain I possessed. I made the middle of this fast to an iron tent peg, which was driven into the ground, and thus the little beasts were secured. They fed ravenously on raw meat, and before long became pretty tame. When I marched they were bundled chain and all into the ‘kilta,’ the lid of which was then tied on, and thus they journeyed to the next halting place, the ‘kilta’ being slung horizontally either to the pack saddle of a Yak, or behind a coolie’s shoulders. On camp being pitched they were taken out and pegged down. One night they managed to draw their peg, but they were fortunately discovered next morning, the chain having become entangled in a bush, about a mile from my tent. They accompanied me for more than two months, and before that time had become a good deal too large for their abode: they gnawed holes in it, and used to travel with their heads sticking out at opposite ends.

As I was quartered at Meerut, whither I had to return by the 15th of August, I was afraid that the heat of the plains would be too much for them ; so I left them in charge of a friend at the hill station of Kussowlie, near Simla, till the end of October, when I had them sent down to me. By this time they had immensely increased in size, but although they had not seen me for so long, they recognized me, and also my greyhound, of which they had previously been very fond. They soon became much attached to me, and would fawn on me like dogs, licking my face and hands; they were always, however, ready to growl and snap at a stranger. I took them down to Agra at the time of the great Durbar there, and used to let them loose in camp with my dogs, so tame had they become.

I presented them to the Zoological Society, and they reached the Regent’s Park gardens in safety: they lived there for eight or nine years, and produced several litters of cubs.

All the cubs were black, a fact which, I think goes far to prove that the Black Wolf is a separate species, or at any rate a permanent variety, and not a mere instance of melanism, as some naturalists have supposed (pg. 40).

Kinloch was simply wrong that the black wolves represented a unique species. It is simple melanism, but it is not clear if these wolves got their melanism through crossbreeding with domestic dogs, as is the case with modern black wolves in North America and Italy, or are black as the result of another mutation. In dogs, we have two forms of black. The most common– and the one that was transferred to modern wolves and coyotes– is called dominant black. It is very easily transmitted through a population–just because it is a simple dominant trait. The other is called recessive black, and it is pretty rare in domestic dogs. It has not been observed in any wild canids, and because it’s recessive, it is pretty difficult to get transmitted intergenerationally. These black wolves could have had an entirely different genetic basis for their color, but if it were dominant, it could be transmitted in a single population, which would give it the appearance that we were looking at a different species.

I hate to burst everyone’s bubble, but despite all of this speculation that has gone on for centuries, the extensive nuclear DNA studies are suggesting that there is only one species of wolf– unless one counts the Ethiopian wolf as a wolf. However, it is more distantly related to the wolf than the golden jackal . Domestic dogs, dingoes, and New Guinea singing dogs are also part of this species. The s0-called red wolf is a recent hybrid between the coyote and the wolf, and the proposed Eastern wolf species is a wolf with some coyote ancestry. Although some Indian and Himalayan wolves have unique mtDNA sequences, genome-wide assays have not found any unique characteristics.In at least one nuclear DNA study,  Indian wolves were found to be closely related to domestic dogs, which probably means that their supposed uniqueness may exist only in their mtDNA, which reflects nothing more than an ancient matriline.

The newly discovered African wolf may be truly unique, but only its mtDNA has been examined. Nuclear DNA analysis might reveal it to be something unique, but it could just as easily turn out that it is part of this species.

There are many accounts of people taming wolves. The black coloration is usually mentioned as a prerequisite for taming, but it could be that black wolf pups were those most likely to be stolen and kept as pets.


NB: The Indian wolf I’m referring to in the post is Canis lupus pallipes, which is a smaller, primitive wolf subspecies. The wolves that Kinloch procured in northern India were likely Canis lupus chanco.  The pallipes wolf ranges from India to Israel and Turkey, and it may be an ancestor of the domestic dog. Some pallipes wolves in India have a unique mtDNA sequence, which has led to some speculation that that they are a distinct species.

As we have seen with East African black-backed jackals, possessing quite divergent mtDNA sequences is not necessarily indicative of unique species status.

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FromChapters on Animals (1877) by Philip Gilbert Hamerton:

It happens from time to time that an attempt is made to bring up a wolf like a dog. These attempts succeed up to a certain point. One of the most remarkable instances occurred in the neighbourhood of Bordeaux, where a grand veneur brought up a black wolf-cub, a bitch, along with his young dogs, in perfect liberty. She went out hunting with the dogs, and enjoyed the chase extremely, except when the purpose of the expedition was a wolf-hunt, to which she had honourable objections. She behaved charmingly in the kennel, and her only fault was sheep-killing, a crime she committed whenever the opportunity offered (pg. 167-168).

So it was not only frontiersmen in Pennsylvania and Kentucky who had wolves that were fine hunting dogs.

At least one wolf in France proved to be a decent hunting dog.

But like those in the New World, this French wolf was also black.

The black coloration is likely indicative of some dog blood in every one of these cases.

But I don’t know if any of these animals had a significant amount of dog ancestry. There is simply no mention of it.

Modern black wolves aren’t necessarily tractable, and Wags, the very friendly tame wolf owned by Adolph Murie, was a normal gray phase wolf.

The Beast of Gevaudan and possibly the killer wolves of Paris were likely wolf dogs or wolves with a certain percentage of dog ancestry. I don’t think they would be quite as nice as these wolves that proved excellent hunting and working dogs.

So it’s a complex question to which there may be no simple answer.

See earlier posts

But again, we do have more evidence of people in modern times using wolves as hunting dogs.






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The Kentucky frontier wasn’t the only place that wolves were used as hunting dogs. Wolves were also commonly used as hunting dogs on the Pennsylvania frontier. They were also crossbred with “improved” Western dogs to make superior working animals, but the practice was largely discontinued when it was decided that every farm ought to have a “purebred” collie or shepherd.

Henry Wharton Shoemaker was a polymath of sorts.  He was a Columbia graduate, who worked on Wall Street and for the US foreign service before moving to Pennylvania to work as a newspaper publisher. He became well-versed in the folklore and local history of the mountainous regions of Central and Western Pennsylvania, and it is from his writings on this subjects that we can find out what the original settlers thought of different animals. He was an ardent conservationist, and he worked as writer for Gifford Pinchot’s campaigns for the US Senate and for governor of Pennsylvania.

His interests as an historian, conservationist, and folklorist brought him to write two volumes on the history of the extinct animals in Pennsylvania. One volume would cover wolves and “panthers” (the creatures also known as cougars/pumas/mountain lions/catamounts), and another would cover the other extirpated species.  These two volumes are often compiled into a single volume called Extinct Pennsylvania Animals, which is traditionally the name for the second volume. The second section of the first volume is called Wolf Days in Pennsylvania, and it was originally published in 1914.

Through his research and interviews with those who had first hand accounts with these animals, he found that the settlers believed that there were three kinds of wolf in Pennsylvania.  There was a large gray wolf and a large black wolf, but these appeared to breed true, even though it wasn’t unusual for a black wolf to whelp a gray pup. And there was a small brown wolf, which sounds suspiciously like an account of the existence of an eastern population of coyotes. I think the corpus of the evidence– particularly the genetic evidence–suggests that coyotes did exist in the East, but they were extirpated with the wolves.  As the larger wolves were removed from other parts of their range, the smaller coyotes were able to file back into the East again. (The so-called red wolf is largely a fictional animal.)

The settlers in Central and Western Pennsylvania considered the black wolf to be a separate species from the gray.  The gray and “small brown wolves” were easily killed, but the black ones were much more cunning and wary.  In what might sound like a contradiction, it was very common for these black wolves to be socialized to people and then used as hunting or working dogs.

Shoemaker writes:

As far as intelligence went, the black wolf was far the superior of the others. It was susceptible of domestication, and would have made the ideal hunting dog of Pennsylvania….Dr. W. J. McKnight, in his “Pioneer Outline history of Northwestern Pennsylvania,” states “the pioneer hunter would sometimes raise a wolf pup. This pup would be a dog in every sense of the word until about two years old, and then would be a wolf in all his acts.” Audubon in his “Quadrupeds of North America” says: “Once when we were traveling on foot not far from the Southern boundary of Kentucky, we fell in with a black wolf, following a man with his rifle on his shoulder. On speaking with him about this animal, he assured us that it was as tame and gentle as a dog, anr) that he had never met a dog that could trail a deer better. We were so much struck with this account and the noble appearance of the wolf, that we offered him one hundred dollars for it, but the owner said he would not part with it for any price.” What was the case in the West, was equally true in the Seven Mountains and in Clearfield and Jefferson Counties. One or two of the earliest hunters trained black wolves to act as hunting dogs and companions. These and wild black wolves bred with dogs owned by pioneers, producing a really worthy progeny. St. George Mivart has said “hybrids between the dog and the wolf have proved to be fertile, though for no long period.” The writer remembers that in his early boyhood about twenty years ago he saw several of these wolf-dogs. They were intelligent and kindly, and highly prized by their owners, farmers in some of the valleys adjacent, to the Seven Mountains. The craze for handsome sheep dogs or collies which struck the valleys about this time resulted in ending the breeding of the wolfish clogs, which to those not in sympathy with them, were technically mongrels, and they eventually disappeared. There are probably few of them now in existence. Their owners declared that they never showed the slightest tendency to revert to a wild state. In September, 1898, the writer visited a farmer, who tilled some back lots at the foot of the mountains on the South side of Brush Valley not far from Minnick’s Gap. This old fellow, Abe Royer by name, kept some turkeys, half wild, which were the result of his tame turkey hens crossing with wild gobblers which lived on the mountain back of his cabin. He had preserved several wild pigeons until 1895, to be used as “stool pigeons” in the event of the great flocks “returning.”

He also kept several wolf-dogs. These animals had dun and grey coloring not unlike collies, but had the shorter hair and longer legs of wolves. There was no trace of black in their coloring, although their owner stated that their grand-sire had been a black wolf which coupled with a shepherd bitch some ten years before when he was lumbering for Ario Pardee in High Valley. He said that neither turkeys nor dogs had the least inclination to revert to the savage proclivities of their ancestors. If the grey wolves and the brown wolves had any of the admirable characteristics of their black relatives, the old hunters sayeth not. “Crafty and mean” is the general verdict expressed about the grey wolves, “nasty like little cur dogs,” is the general run of remarks relative to the brown wolves. Doubtless these uncomplimentary characterizations are unjust to the animals, but they were certainly not up to the standard of the black wolves. If all are of one variety these attempts at specialization are hardly worth the time to read. At the same time it may show that color in animals has much to do with habitation, character and disposition. It may help to reveal the secret of why some men are blonde and others dark (pg. 24-25).

The black coloration in North American wolves is thought to have originated through crossbreeding with Native American dogs, but the dog ancestry in these wolves probably wouldn’t have been a high enough percentage to have affected their temperament. Modern black wolves are, for all intents and purposes, very much wolves.

The black coloration is a simple dominant trait, so it is possible that certain populations of wolf in the Eastern United States were consisted of almost of nothing but melanistic individuals. This dominant black trait in dogs, wolves, and coyotes comes from a mutation that controls the protein beta-defensin 3. This protein does regulate the amount of melanin that appears on the dog, wolf, or coyote’s coat, but it also is associated with immune response.  Having this particular mutation might provide the wolves with some advantages  in fighting off viral and bacterial infections, which may have been more common in the temperate and subtropical forests of North America.

I don’t think these black wolves were a different species at all, but for whatever reason, this particular type of wolf was easily domesticated. It may have been that this was a sort of intermediary animal that included the genetics of both wolves and domestic dogs, and there was some continual hybridization between the two forms. And that might be why these wolves were so easily domesticated.

It is possible that the black coloration could have been associated with a tendency toward domestication. A very similar finding was found with black deer mice, which were found to be more docile than the more common agouti deer mice.

But black deer mice have a different genetic basis than black wolves or dominant black dogs, so one should be careful about making generalizations from that study and trying to apply them to the tame wolves of the Pennsylvania.

These domesticated wolves appear to have been very much a part of life in America in the early days of settlement.  These settlers of the Pennsylvania mountains didn’t have access to the best lines of Western hunting dogs, so they improvised. They found that the black wolves were a good outcross to their curs and shepherds.

So here we have another account of modern people keeping wolves as working and hunting dogs.

See earlier posts

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The following account of a tame black wolf that was used as a hunting dog can be found in John James Audubon’s The viviparous quadrupeds of North America, Volume 2 (1851):

Once, when we were travelling on foot not far from the southern boundary of Kentucky, we fell in with a Black Wolf, following a man with a rifle on his shoulders. On speaking with him about this animal, he assured us that it was as tame and as gentle as any dog, and that he had never met with a dog that could trail a deer better. We were so much struck with this account and the noble appearance of the wolf, that we offered him one hundred dollars for it; but the owner would not part with it for any price (pg. 130).

One might claim that this was a Native American dog, but I doubt it.

Audubon and the people living in Kentucky at this time could tell dogs from wolves, and the deer hunter didn’t claim this animal as a dog. The people living in Kentucky during the early nineteenth century lived very close to Native Americans and to the natural world as a whole. They were fully aware of what animals existed there.

I think this animal is what Audubon and this hunter said it was.

It was a deer hunting wolf.

So yes, people in modern times have used wolves as hunting dogs.

See earlier posts

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A wolf-like animal depicted in a Roman mosaic in Syria in the fifth century. It is said to be a hunting dog, but it looks a lot like a robust golden wolf.

This description of the five species of wolf native to the Roman Empire comes from Cynegetica, a poem on hunting that is attributed to Oppian of Apamea.

Of wolves there are five species—the first of a yellow hue,—swift, audacious, and by shepherds named the archer,—the next of superior magnitude and swiftness, known by the two names of the hawk and the plunderer; he seeks his prey with the dawn, and dwells in the lofty mountains—but when snow covers the ground, he assumes greater boldness, and in quest of prey approaches even the city walls. The third species inhabits the mountains of Taurus and Cilicia—an animal superior to the race of wolves, named the Golden, of prodigious strength, and able to resist the unspent brass and the pointed iron. He dreads the rising of the dog-star, and during the prevalence of its heat, lies concealed in his shady cavern. Of the two remaining species, the one from his white colour is named the Hoary Kite. The other is of smaller size,—black,— hirsute,—preys on hares (Cynegetica, Book Third).

This poem was written in the third century, and it includes an analysis and description of just about every game animal within the Roman Empire and the areas adjacent to its borders. The notion of there being more than one species of wolf within Eurasia is something we don’t  generally accept today, the supposed full species status of the Indian and Himalayan wolves not withstanding.

There is also a lot of exotic information about wildlife in this piece. Within lines adjacent to the discussion about wolves, we learn that hyenas change their sex every year and that male hyenas “become fruitful dams.” This is obviously a reference to the bizarre genitalia of female spotted hyenas, which would have been known from African specimens.  Spotted hyenas lived in the Middle East as recently as 8,000 years ago, but that would have been thousands of years before the poem was written. However, female striped hyenas sometimes have similar genitalia morphologies, which disappear as they mature, and striped hyenas are the species Oppian clearly describes in the text.

The author also points out that wolves mate with panthers, producing a creatures called “Thoes.”  This word is very similar to “thos,” the Classical Greek word for jackal. I don’t think that anyone thinks a jackal is a hybrid between a black leopard and a wolf.

Despite its inaccuracies, this piece of zoological literature is still useful.

For one thing, this section on the wolves tells us that wolves were once quite diverse in behavior and phenotype– even more so than they are now.

This small black wolf was around for a while.  As recently as the nineteenth century, there were small black wolves in the mountains of Syria that was a “Derboun.”  It may have been a pariah dog, or a type of naturally occurring wolfdog hybrid. Or it may have been a naturally occurring melanistic Arabian or Iranian wolf that may or may not have any relationship to modern melanistic wolves.

The big golden wolf could have been a striped hyena, but it also could have referred to an unusually powerful wolf. However, Oppian clearly describes the striped hyena as an animal with stripes. If the golden wolf had been a striped hyena, it would have been unlikely that he would have clearly defined the striped hyena without pointing out this similarity.

However, one should keep in mind that it is unlikely that Oppian had first hand knowledge of every animal in the text, but he might have encountered them via trade routes that crossed through the Levant. The Romans were obsessed with keeping exotic animals in menageries and for their circuses. Oppian may have been a noted sportsman of the region– perhaps someone who was responsible for taking notables out on hunting forays in the region–for the book is full of advice on how to hunt various animals.  It is unlikely that Oppian hunted these animals himself, but he could have known from talking to those who hunted them and were transporting them across the empire.

But I do think this piece suggests something we already knew. Wolves are a highly variable species, and in ancient times, they were so varied in appearance that they were thought of as different species.

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I just saw remarkable documentary on National Geographic Wild about a particular wolf living in Yellowstone. It was called The Rise of Black Wolf.

Some of the best nature documentaries are biopics about a particular wild animal, and this particular wolf was quite an interesting character. He lived for about ten years in Yellowstone National Park, which is actually quite long for a wild wolf.

He also sired a lot of puppies.

But how he was able to achieve both was truly remarkable.

Most of us know about wolf packs that are based upon a breeding pair. They are two, usually unrelated, wolves that form a pair bond. They are fairly monogamous.  The male in the pair does he can to keep his sons and interlopers from breeding with his mate, and the female also does everything to keep other females away from her mate, including social suppression of estrus. Yellowstone packs are unique in that they often have multiple breeding females.

However, these females don’t tend to breed with wolves in their own pack. Very often, they are the daughters of the breeding pair, and because wolves have very strong inbreeding avoidance behavior, they would not mate with their father. And they certainly weren’t going to mate with their brothers, who very often leave the pack by the time they are two or three years old.

To breed, they often consort with lone male wolves that come near the pack territories during the mating season. These wolves are often called “Casanova wolves.” They are young males that have left their natal packs in search of new territories and mates, but very often, neither can be procured. So while thy are without mates or territories, they often take to mating with the daughters of an established breeding pair. In normal wolf packs, bitches other than the breeding female typically don’t raise their pups to maturity. They often don’t have access to dens or food for their offspring.

Yellowstone is different. Because the wolves live in an area with relatively abundant prey, packs often have enough resources to allow these females to raise their litters.

Because it is not normal for more than one female wolf in a pack to raise a litter to maturity, such breeding are not a sure strategy.

The wolf in this particular documentary did something unusual:  He based his entire reproductive strategy upon being a Casanova.

He found that he could hang out at the margins of the territory of the Druid pack and mate with the female daughters of the breeding pair. He pair-bonded with none of them. He was a love ’em and leave ’em kind of guy.

But over time, he wound up siring more puppies that the original breeding male in the pack and his successor, who were both forced to mate with only their mates. During the tenure of the first breeding male, the Casanova had a hard time breeding with his daughters. The breeding would attack the Casanova if he caught him, but the Casanova figured out that the breeding male had a taboo about going near a road that ran through the edge of the pack’s territory. The Casanova used the road as his base from which he would launch his romantic forays. When another breeding male took over after the death of the first breeding male, the Casanova was tolerated, even allowed to become fully part of the pack. He was still breeding with all the females in the pack, except for the breeding male’s mate.

And such a strategy is very successful, but it is entirely reliant upon a wolf pack allowing more than one female to raise her offspring.

And that typically happens only in areas of material abundance.

As you may be thinking, dogs typically employ a Casanova strategy. Dogs can pair bond, but it is not the norm.

Dogs are able to utilize this strategy for the same reason that the Casanova wolf did:  Most female dogs living even within the margins of human society are able to raise at least some of their puppies to maturity.

Dogs that are able to utilize this strategy are better able to spread their genes than those that employ are pair bonded within a pack system. From a gene-centered view, the only reason why wild dogs pair bond or form packs based upon the offspring of a bonded pair is so that more resources can be devoted to raising litter. (See kin selection).

In a situation where a pack can raise more than one litter– as is the case in Yellowstone and with domestic dog population– it makes much more sense not to pair bond.

That Casanova wolf in the documentary wound up siring more puppies than any wolf in Yellowstone. That is not a trivial influence upon the population genetics of any population. If animals can produce more offspring without forming an almost entirely exclusive pair bond, then that animal will spread its genes into more descendants.

That reason alone is probably why virtually all domestic dogs have given up the pair bond behavior. A male dog can spread his genes to more descendants if he’s not tied to one bitch.

Of course, when domestic dogs become wild again, as has happened with the dingo, the pair bond returns.

It is really remarkable that Canis lupus has developed these two very different strategies for reproduction. Neither is useful every situation. If a wolf pack allowed all its females to raise a litter in areas where prey was often scarce, the pack would likely never raise any litter to maturity.

Casanova wolves probably tell us how and why dogs gave up the pair bond. It’s just not an efficient way of spreading genetic material in ecological niche in which many females can breed and raise their offspring to maturity.

In those situations, it is more efficient to sleep around.

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