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The term “African wolf” often has been used to refer to the Ethiopian wolf of the Ethiopian Highlands and Xenocyon lycaonoides, the extinct ancestor the African wild dog or “Painted wolf.”

But now, it looks that name is going to refer to some animals that were previously classified as golden jackals. Now, the exact taxonomic status of the wolf-like “jackals” of Egypt has long been contentious.  When I was first reading about wolves as a child, the maps of wolf subspecies always included a subspecies that was native to Egypt and Libya. This animal was always called Canis lupus lupaster. However, around the year 2000, I noted that these wolves had been reclassified as golden jackals and were referred to as Canis aureus lupaster.

I always had some issues with this classification.  These North African golden jackals were larger than the golden jackals of Europe, which would have been the second largest subspecies if the Egyptian jackal would have been considered a golden jackal. This classification meant that golden jackals break Bergmann’s rule, something that other wolf-like canids follow rather closely. Bergmann’s rule says that warm-blooded animals tend to be larger in colder environments than those found in warmer ones. Wolves follow this rule fairly closely, although the wolves of the forested regions of Alaska tend to be larger than the arctic subspecies. The Balkans are a much colder place than Egypt, and one would have expected the European golden jackal to be the largest subspecies.

My other problem with this classification was that these Egyptian and Libyan canids looked very similar to the Arabian wolves, which had long been recognized as the subspecies that lives in the Sinai. If an Arabian wolf could live in the Sinai, why couldn’t it live in the rest of Egypt or North Africa?

The original Mitochondrial DNA studies on these canids found that they were more similar to the golden jackals of Israel than Arabian wolves, which are part of the Holarctic wolf species. It was decided to move these animals into the golden jackal species and that has been where they have been classified for several years now.

I came up with the possibility that the wolf-like appearance of these jackals came from fact that there were once Arabian wolves in Egypt and Libya, and these wolves became heavily persecuted. Just as the wolves of Eastern and Southern US bred with female coyotes when their populations dropped, I surmised that these wolves did the same with golden jackals. And that is why these wolves have golden jackal MtDNA.

However, I was wrong.

A study was released today that found that the lupaster jackals are actually wolves. This study, released in the journal PLoS One, used a boostrapping methodology to compare the MtDNA sequences of golden jackals, Holarctic wolves (most modern wolves are of this ancestry), the Indian wolf, and the Himalayan wolf. The last two are considered older lineages than the main Holarctic wolf lineage, leading some taxonomists to conclude that they are unique species (not by this armchair taxonomist, however). The study found that the lupaster jackals were within the wolf species– but, like the Himalayan and Indian wolves, represent an older lineage.

The study also found that supposed golden jackals in the Ethiopia were actually the same species as the lupaster jackals.

That means that Canis lupus has an African range that exends from Egypt and Libya to Ethiopia.

Contrary to what some media reports suggest, the findings of these study did not discover a new African species of wolf– unless one thinks that the Himalayan wolf and Indian wolf are separate species. All the researchers found was that there are Canis lupus wolves in Africa.

The authors suggest that we combine the lupaster population with those wolfish jackals in Ethiopia into a single subspecies of wolf– and call it Canis lupus lupaster. The authors conclude that more golden jackals in other countries need to be tested to see if they are actually wolves.  The wolf’s range in Africa may be more extensive than we originally thought.

So the diversity of Canis lupus is even greater than we thought. The wolves in the Ethiopian population of the African wolf are actually smaller in size than the Ethiopian wolves with which they share a habitat.  The animals are actually closer to the golden jackal in size, which means that this is the smallest extant subspecies of wolf. These wolves are well within the Southern clade of wolves, which are the most primitive of subspecies.

And very much in keeping with Bergmann’s rule.

This finding is very much in keeping with our understanding that the diversity in domestic Canis lupus is reflected in the wild population. The little African wolves of Ethiopia are the same species as the big wolves of Alaska, which are the same species as the chihuahua and the extinct bone-crushing wolf of the Pleistocene.

The diversity of this species never ceases to amaze me. And I am so glad that this remarkable species has been able to form such an intimate relationship with us.

***

Thanks very much to Jess for sending me this study and making my day!

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The species from yesterday’s query was an Egyptian jackal. It comes from The living animals of the world; a popular natural history with one thousand illustrations; Volume 1: Mammals (1902) by C.J. Cornish, Frederick Courteney Selous, Sir Harry Hamilton Johson, and Sir Herbert Maxwell.

The animal is listed as a “North African jackal,” but the subspecies is better known as the Egyptian jackal.

However, its exact taxonomy is still being debated. Many sources, including Peter Steinhart’s The Company of Wolves, list this animal as a subspecies of Canis lupus. If listed as a wolf, it is called Canis lupus lupaster. Morphological studies found that it was most similar tot he Arabian wolf subspecies (arabs), which ranges into the Egyptian Sinai. Its skull is said to be almost exactly like that of the Indian wolf (pallipes). It is also larger than the golden jackal subspecies that are found near it. It is even larger than the European golden jackal, which is the largest accepted subspecies of Canis aureus.

The most recent genetic evidence suggests that this animal is a golden jackal. The most recent analysis of MtDNA that compared Egyptian jackals with other members of the genus Canis. The study found that the MtDNA was most similar to the Israel golden jackal, and because of this finding, the Egytpian jackal is no longer considered a wolf.

However, the variance was quite high between Israeli golden jackals and Egyptian jackals. They differed by over 4 %, which is much higher than the variance between dogs and wolves. Of course, golden jackals are a much older species than wolves are, and one could expect to find a bit higher variances. Black-backed jackals have intraspecific MtDNA variances that are as high as 8 percent– even within contiguous populations. The black-backed jackal is the oldest subspecies within the genus Canis.

However, because the study that found that Egyptian jackals were golden used only MtDNA, one needs to be a bit skeptical. We know that wolves can interbred with golden jackals, as can domestic dogs. It seems to me that it is at least plausible that there was once a North African wolf that was very similar to the Arabian, Iranian, and Indian wolf subspecies. That animal became threatened through persecution, so it interbred with golden jackal bitches. For a wolf to have golden jackal MtDNA, it would only require a single female golden jackal ancestor.

In fact, this cross-breeding could have happened at any time. We know that wolves evolved in southern Asia, and when the first wolves entered Africa, the only mates they could find were golden jackal bitches.

No one has examined the nuclear DNA of an Egyptian jackal. They are not included in the .large genome-wide studies. They need to be.

Like the red wolf, their exact taxonomy is not clear until their full nuclear DNA is examined.

It is possible that the Egyptian jackal is a distinct species. A very similar large golden jackal-like animal with wolf characteristics was found in the Danakil. It is called the wucharia by the locals, who do not consider it to be the same thing as the golden jackal. Perhaps both of these wolfish jackals are new species or they are the descendants of relict populations of wolves that lived in Northern and East Africa.
We don’t know. Just as the coyotes of the Eastern US and Canada are quite wolf-like, there are also golden jackals that have wolfish characteristics.

 

 

 

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The endangered African wild or "painted wolf" is a descendant of the larger Xenocyon lycanoides-- the so-called "African wolf."

Xenocyon lycaonoides.

The way the Greek rolls of the tongue suggests something like a Spartan general who led his phalanxes against the Persians.

It was actually a large  canid that lived between 1. 8 million and 126,000 thousand years ago. Although its name suggests a totally African range, it was also found in Eurasia.

It was larger than the typical modern wolf, and it was probably a major predator of  all sorts of wildlife. It is even suggested that this hulking wolfish creature preyed upon early man.

It was not a true wolf as we would know it today. It came from a different line of large wild dogs.

Its most likely descendant is the African wild dog or “painted wolf” (Lycaon pictus). However, Xenocyon was a much more robust animal than these multicolored carnivores.

Xenocyon was a very successful species from which several different forms descended. The so-called Sardinian dhole, which lived on the island that became Sardinia and Corsica, was probably descended from Xenocyon. It is often suggested that the Sardinian dhole (Cynotherium sardous) was nothing more than a dwarf insular form of Xenocyon.  Two extinct Javanese dogs may have also been descendant of Xenocyon.

Current research of the modern dhole’s molecular evolution suggests that it did not descend from Xenocyon. It is an early offshoot of the line that gave us the genus Canis. It’s more closely related to wolves, jackals, and coyotes than the African wild dog, which it superficially resembles.

There is a move in some academic circles to move Xenocyon and its variants into the genus Lycaon.

Of course, we could put all of them in the genus Xenocyon, but it would make more sense to put them in the same genus as the living species.

After all, it is now accepted that the African wild dog is the only living descendant of Xenocyon, and it would make sense that we would move all of these related dogs into the same genus.

I am trying to imagine what Xenocyon may have looked like.  A piece of me sees it as mottled in different colors with rounded ears, as is the case of the African wild dog.

But another part of me sees it as a more robust animal.

Maybe dire wolf crossed with African wild dog is a better way of imagining it.

The African wild dog evolved as a specialized form of Xenocyon that could prey almost exclusively on antelope and other ungulates native to the savannas of Africa.

The robust Xenocyon went extinct, as did all of those possible insular forms, leaving only the specialized Lycaon pictus as a relict of what was once a more diverse line.

The African wild dog is truly a unique species, and to think of its story within this context should increase the urgency to conserve this species.

It’s truly a remarkable animal.

***

One should note that there are other species that are called African wolves. All of these are members of the genus Canis and are in no way related to African wild dogs or Xenocyon.

The most famous of these is the Ethiopian wolf (Canis simensis), which is no longer classified as a jackal. The original molecular genetic studies of this wolf found that it was a descendant of Canis lupus.   A more recent study that they were not actually derived from the wolf, but they were very closely related to both wolves and coyotes.

The golden jackal is more closely related to wolves and coyotes than to the other two species of jackal,  so it could also be called an “African wolf.”

And there are two populations of golden jackal that are quite wolf-like.

One of these is the Egyptian jackal. It may be its own species (Canis lupaster), a subspecies of wolf (Canis lupus lupaster), or part of the golden jackal species (Canis aureus lupaster). It is found in Egypt and Libya, and it looks more like an Arabian or Indian wolf than a golden jackal. Because of this similarity, it was always classified as a type of wolf. Recent MtDNA studies show that it is some form of jackal, but the variance in the MtDNA sequence is pretty high from the closest golden jackal population.

Of course, MtDNA studies can be limited in their scope.

MtDNA is inherited via the mother, and we know that the only hybridization between dogs and wolves and golden jackals is between male dogs and wolves and female jackals. Male jackals just are unable to subdue female dogs or wolves to mate with them.

It could be that there was once a true Egyptian wolf.  Through intense persecution, it was reduced to very small numbers.  Male wolves were unable to find bitch wolves with which to breed, so they mated with female golden jackals.

And that could explain why these wolf-like dogs come out with golden jackal MtDNA.

Or maybe the hybridization happened very long ago with some canid that was closely related to the golden jackal, which would explain why the MtDNA of Egyptian jackals varies so much from Israeli golden jackals.

To make matters more complicated, a similar wolfish jackal has been found in the Danakil in Eritrea. It is called the wucharia, and the people who live there recognize it as something unique from the more common golden jackals that also inhabit the desert.

It may actually be a population of Arabian wolf, or it may be the same species or subspecies as the Egyptian jackal.

Both the Egyptian jackal/wolf and the wucharia could be modern day versions of African wolves. Maybe we should look to them to see where the African wolf population went.

We do know that Canis lupus did invade northern Africa at some point, but no one is sure how far south they got.

Or what happened to them.

When the original molecular genetic study came out on the Ethiopian wolf, it was though that the Ethiopian wolf was last surviving population of African Canis lupus.

I think it is very likely that there were African wolves, but these became extinct. Perhaps due to persecution. Perhaps due to competition with other carnivores.

A few individuals survived in North Africa and maybe in parts of the Horn of Africa, where they interbred with golden jackals or a close relative of that species as their numbers dwindled and these wolf populations became isolated.

Whatever these wolves were, they were likely very closely related to the Middle Eastern subspecies. They were probably the same subspecies that we call the Arabian wolf or maybe the Iranian wolf.

If this is true, then these would have been the first wolves that man encountered.

We now know dogs were first domesticated from Middle Eastern wolves.

However, those Middle Eastern wolves could have been living in North Africa when they first encountered humans.

It is possible.

And worth considering.

But the archaeological evidence shows that the domestication most likely happened in the Middle East, not North Africa.

It’s  still an idea to keep in mind.

***

Arabian wolves are well-documented in Egypt.

They are found in the Sinai, which is the land bridge that connects Africa and Asia.

If any wolves could live in Africa, they would be something similar to these animals.

They aren’t big.

They really aren’t bad.

They are nothing like Xenocyon lycaonoides— the big, bad African wolf.

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strange wild dog

The wild dog from yesterday has a somewhat controversial identity. Historically, it was called the Egyptian wolf and was recognized as Canis lupus lupaster. For those of you have old wolf books, this animal is the supposed wolf that lived in Egypt (but not the Sinai, where the Arabian wolf lives) and Libya. I need to repeat that this is not the subspecies that lives in the Sinai. That animal is definitely the Arabian wolf (Canis lupus arabs).

However, recent analysis of its MtDNA has changed the Egyptian classification. It turns out that its MtDNA is very similar to Golden jackals, and thus, most taxonomists refer to it as the Egyptian jackal (Canis aureus lupaster).

But that is not the end of the story.

It turns out that this animal does have some unique characteristics. It is a very wolf-like jackal, with longer legs and slightly larger size.

Some morphological studies suggest that this animal represents a unique population of Canis that was isolated from other jackals. It is believed to have existed in two separate populations: one in the Sahara and one in the Danakil, where it is called “the wucharia.”

However, both of these animals could represent wolf populations that once lived in Africa, which then became stressed through persecution. As the wolves became more stressed, they started breeding with golden.

We know that the same thing has happened with the “red wolf” and “Eastern Canadian” wolf, which some authorities consider separate species from C. lupus. The fact that we get golden jackal MtDNA is exactly what we would expect. MtDNA comes from the matriline, and in “red wolves” and “Eastern Canadian wolves,”  we have evidence of coyote MtDNA. In the New World wolves, we know that female coyotes breed with male wolves. The reverse has never been seen in the wild. It would make sense that a similar situation would occur here.

Just as I consider C. rufus to be C. lupus rufus, even though it has coyote MtDNA, I think it might be correct  refer to the wucharia and Egyptian wolves as subspecies of C. lupus.

Of course, it may be that both represent a unique species, which we would call C. lupaster.

It is another one of those canids like the red wolf and Ethiopian wolf that no one can fully classify. I am very hesitant to create species, especially when a species appears to be an intermediate between two known species.

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strange wild dog

The wild dog from yesterday has a somewhat controversial identity. Historically, it was called the Egyptian wolf and was recognized as Canis lupus lupaster. For those of you have old wolf books, this animal is the supposed wolf that lived in Egypt (but not the Sinai, where the Arabian wolf lives) and Libya. I need to repeat that this is not the subspecies that lives in the Sinai. That animal is definitely the Arabian wolf (Canis lupus arabs).

However, recent analysis of its MtDNA has changed the Egyptian classification. It turns out that its MtDNA is very similar to Golden jackals, and thus, most taxonomists refer to it as the Egyptian jackal (Canis aureus lupaster).

But that is not the end of the story.

It turns out that this animal does have some unique characteristics. It is a very wolf-like jackal, with longer legs and slightly larger size.

Some morphological studies suggest that this animal represents a unique population of Canis that was isolated from other jackals. It is believed to have existed in two separate populations: one in the Sahara and one in the Danakil, where it is called “the wucharia.”

However, both of these animals could represent wolf populations that once lived in Africa, which then became stressed through persecution. As the wolves became more stressed, they started breeding with golden.

We know that the same thing has happened with the “red wolf” and “Eastern Canadian” wolf, which some authorities consider separate species from C. lupus. The fact that we get golden jackal MtDNA is exactly what we would expect. MtDNA comes from the matriline, and in “red wolves” and “Eastern Canadian wolves,”  we have evidence of coyote MtDNA. In the New World wolves, we know that female coyotes breed with male wolves. The reverse has never been seen in the wild. It would make sense that a similar situation would occur here.

Just as I consider C. rufus to be C. lupus rufus, even though it has coyote MtDNA, I think it might be correct  refer to the wucharia and Egyptian wolves as subspecies of C. lupus.

Of course, it may be that both represent a unique species, which we would call C. lupaster.

It is another one of those canids like the red wolf and Ethiopian wolf that no one can fully classify. I am very hesitant to create species, especially when a species appears to be an intermediate between two known species.

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