Some thoughts on wolf evolution

We spend a lot of time debating about how wolves became dogs. A huge debate exists in the archaeological and paleontological literature about how one can determine whether the remains of a canid represent a wolf, a dog, or a transitional form between wolves and dogs. This debate is why the oldest dog remains are dated to around 14,000 years ago and come from the Bonn-Oberkassel site. Anything older than that, a big debate exists among experts about what can be used to define a wolf, a dog, or a transitional form.

But this debate does not exist solely in relatively recent transition between wolves and dogs. The entire evolution of Canis lupus is a hotly contested and often contradictory, depending upon which source one reads and whether one is looking a source that relies upon paleontological and morphological analysis or one that looks at the molecular evolution of the species.

It is well-accepted in European paleontology that Canis lupus evolved from Canis mosbachensis. Mark Derr paid particular attention to this evolution in his How the Dog Became the Dog. He posits that the extinction of the large hunting dog, Xenocyon lycaonoides, created an ecological niche that could be filled by the Mosbach wolf evolving into the gray wolf.

Yes, the Mosbach wolf was smaller than the modern gray wolf. Individuals from Northwestern Europe were mostly about the size of a modern Indian wolf or a “red wolf.” Indeed, the similarities between some of these mosbachensis wolves and red wolves are the best evidence for a unique red wolf species, for one can argue that red wolves are just a relict form of the Mosbach wolf that held on in Eastern North America. Of course, the genetic data do not agree with this assertion, but it is an interesting idea nonetheless.

My reading is that the Mosbach wolf gave rise to Canis lupus in Eurasia between 300,000 and 500,000 years ago. The coyote, though often posited as a primitive Canis, is actually derived from a divergent form of Canis lupus that got marooned in the American Southwest some 50,000 years ago and evolved to fit a jackal-like niche on a continent already dominated by dire wolves.

The Mosbach wolf disappeared from the fossil record around 300,000 years ago, but there is always a debate as to the possibility that it held on longer. The red wolf and Indian wolf are certainly possibilities for its continued existence today, but as we’ve looked at more wolf genomes both of those don’t come out so distinctive. Every study that I’ve seen that uses Indian wolf genomes finds that they are divergent Canis lupus, and the red wolf is a cross between wolves that are of that coyote type and relict Southeastern gray wolves from a later invasion of the continent. I do think there is pretty good historical data that some smaller wolves that we would define as coyotes lived in the Eastern states at the time of contact, particularly the small brown wolf of Pennsylvania mentioned by Shoemaker and the small “wolues” of Jamestown mentioned by John Smith. My guess is that no one really took stock of what they were killing when they killed off the wolves of Eastern states. It is very possible that coyote-like wolves were killed off in great numbers along with the big ones, and later on, coyotes from the plains came East, crossing with wolves and even relict original Eastern coyotes to form the modern Eastern coyote. The red wolf and the larger Eastern coyote are thus recreations of the Mosbach wolf that have happened in modern times.

In Europe, one potential late surviving Mosbach wolf was thought to have been found in Apulia, Italy, at the Grotta Romanelli site. Wolf remains have been found in the cave that date to between 40,000 and 69,000 years ago and they were often described as belonging to a late surviving Mosbach wolf. A recent morphological analysis revealed that these remains were of a peculiar form of Canis lupus that lived in that part of Southern Italy, and they were not of any kind of Mosbach wolf.

However, the Mosbach wolf is particularly intriguing. Occasionally, it has been posited as a direct ancestor of the domestic dog, but because we don’t have an overlap between the signs of the earliest dog domestication and the existence of Canis mosbachensis in the fossil record, one should be very careful in making such an assertion.

This same caveat should be placed when one sees Canis variabilis posited as dog ancestor. For one thing, there is no such thing as Canis variabilis. Instead, all the specimens listed as this species that come from the Zhoukoudian site in China have now been reassigned to Canis mosbachensis. This reassignment posits them as Canis mosbachensis variabilis, so whenever one encounters that “Canis variabilis” in a paper, just remember that they are discussing a particular East Asian form of the Mosbach wolf.

From my own speculative meta-analysis, it seems that the Mosbach wolf is ancestral to the entire wolf/dog/coyote species complex, which may include the African golden wolf, and the Eurasian golden jackal. A genome comparison study that included dogs, wolves, and one Israeli Eurasian golden jackal found that the divergence between the golden jackal and the dog and wolf species happened just before the anatomically modern Canis lupus replaced Canis mosbachensis in the fossil record. The Eurasian golden jackal could potentially be derived from a diminutive form of Canis mosbachensis that moved toward a more generalist scavenger form.

We also have some evidence of small Mosbach wolves in Europe that could have potentially gone in the direction of the golden jackal. This specimen was found not far from the Grotta Romanelli wolf that were found to be anatomically modern and not Mosbach wolves. It was found at the Contrada Monticelli site in Apulia. It was unusual in that it was quite a bit smaller than the Mosbach wolves found in other parts of Europe, and the authors found that Mosbach wolves were as morphologically variable as modern wolves are.

In North Africa, we also have a recent discovery of a canid that was much like the Mosbach wolf. The authors thought it was a bit different from the Eurasian form, and they decided to call this species Canis othmanii. This African wolf-like canid was found at a site in Tunisia and dates to the Middle Pleistocene, and it could potentially be the basal gray wolf that hybridized with the Ethiopian wolf to make the African golden wolf. More work needs to be done on this specimen, for it very well could wind up like Canis variabilis, a regionally distinct form of the Mosbach wolf.

The really fuzzy part about Canis mosbachensis isn’t that it is the ancestor of the gray wolf. The educated speculations I make about its relationship to the golden jackal and the golden wolf could be debated, and we need lots more data to figure out if I am right or not.

The really fuzzy part is what came before the Mosbach wolf. Most scholars think that Etruscan wolf (Canis etruscus), which makes an appearance in the fossil record around 2 million years ago, is the ancestor of the Mosbach wolf. For years, there was a debate about whether the Mosbach wolf was a chrono-subspecies of the Etruscan wolf or a chrono-subspecies of the gray wolf. All these suggestions would be technically true, simply because we could regard the Etruscan wolf-Mosbach wolf-gray wolf as a species that lasted and evolved over this time period.

However, a bit of a debate now exists as to whether the Etruscan wolf is the ancestor of the Mosbach wolf. An extensive morphological analysis of Etruscan wolf remains and those of another Canis species called Canis arnensis, which compared both to the modern black-backed jackal, the gray wolf, the golden jackal, and the golden wolf, found that our previous delineation between arnensis as being jackal-like and etruscus as being wolf-like were over-simplifications. Some characters of arnensis are much more like modern gray wolves than etruscus is, and it is possible that arnesis gave rise to the Mosbach wolf. Still, the bulk of scholarship thinks that the Etruscan wolf is the ancestor of the Mosbach wolf.

However, because the Mosbach wolf was not included in the analysis, it might be difficult to make such a conclusion. However, maybe the Etruscan wolf or something like it is the ancestor of the Ethiopian wolf. The ancestral Ethiopian wolf must have had an extensive range in Northern Africa for it to have hybridized with Canis mosbachensis, Canis othmanii, or a basal modern gray wolf to form the African golden wolf.

I have focused most of this post on the origins of gray wolves in the Old World, but the first Canis species to evolve were found in North America. Canis lepophagus first appeared in the fossil record 5 million years ago. It was very similar to a coyote or a Canis arnensis of the Old World. This is the part of the story where the molecular data has largely shaken up what we used to believe about coyotes. Lepophagus is thought to have evolved into the larger Edward’s wolf (Canis edwardii), which is sometimes called Canis priscolatrans. These animals might have been the same species or very closely related to the Etruscan wolf. The modern coyote is thought to have derived from edwardii/priscolatrans/estrucus 1 million years ago, but genome-wide comparisons put the existence of most recent common ancestor of gray wolves and coyotes at less than 51,000 years ago.

The dire wolf derived from Armbruster’s wolf (Canis armbrusteri). Armbruster’s wolf derived from Canis edwardii/priscolatrans/etruscus 1.8 million years ago. The dire wolf then evolved from that species 125,000 years ago, which means the dire wolf’s most recent common ancestor with modern wolves and the coyote may have been as far back as 2 or even 3 million years ago.

This analysis is still being worked out. The molecular data is constantly throwing wrenches into the machinery of paleontology, especially the paleontology of canids. The most successful extant canid lineage are full of parallel evolution and phenotypic plasticity, and in this way, it has become quite a challenge to sort out the evolutionary history of these species. At various times, large wolf-like forms have evolved as have smaller coyote or jackal-like forms.

The story of Canis starts with a coyote-like lepophagus, but right now, its likely niche is adopted by the modern coyote, which also very similar to it. But the molecular data suggest that the coyote evolved to adopt this similar niche from a larger Eurasian gray wolf and that it did not directly descend from lepophagus over 5 million years in only North America. Instead, it evolved into wolves that wandered into Eurasia, becoming the Mosbach wolf and then anatomically modern gray wolf. Some of these wolves wandered back into North America and became generalist scavengers in the land of the dire wolf.

Very similar stories likely are lost to us, but we must understand that the history of wolves is not just about getting bigger and developing pack-hunting behavior. That is one part of the story, but another part is about evolving to fit niches, which sometimes means evolving a smaller size and more generalist diet.

Some of my ideas here are very speculative, but I think they are nested in my reading of the available literature. Do not assume that I have the final story of how these creatures evolved, but just understand that the molecular side is so rarely considered in paleontology literature that it is almost like we’re reading evolutionary history of two different lineages.

More work must be done to formulate a synthesis between these two disciplines. Otherwise, there will be continued conflict, and the one using an older methodology and often working with much more incomplete data-set will fall by the wayside. And that is not the one using full genomes.

If we know what problems exist using morphological studies on extant and recently extinct canids, it is very likely that we’re missing important data on many extinct species, one for which there is no DNA to test.

Still, paleontology has much to tell us about the way early wolves lived. It can tell us much about how the ecosystems were and why wolves evolved in the way they did. But its methodologies often miss relationships between extant forms and miss the tendency toward parallel evolution.

I tried for about two years to watch Joe Rogan’s interview with Dan Flores, who wrote a book on coyotes that I think is quite full of misunderstandings about canid taxonomy. When Rogan questioned him about the papers that show a recent origin for the red wolf, Flores pretty much just dismissed those papers because they didn’t look at fossil.

That’s not how it works. Within canids, we know that parallel evolution is a big thing, and it is very possible that coyote-like and red wolf-like canids have evolved more than once on this continent. Indeed, a careful reading of the paleontology and molecular data strongly suggests that this is the case.

In fact, it has always been the case with these wolf-like canids. Big ones evolved from small ones, but sometimes, the big ones become small, because it is a better fit for survival.

Natural History

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New Year’s Resolutions

From the good folks at Yappy. (In case you’re wondering, my resolution is to cook at home more often this year. I make homemade food for the dog, but more often than not, I feed my kids via DoorDash. That’s going to change this year.) Until next time, Good day, and good dog!


Doggies.com Dog Blog

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Is Your Dog Anxious When Traveling? #Giveaway

This post is sponsored by Innovet Pet Products. As always, we only share products that use in our own home with our pets. We all know that travel can be stressful. Whether it’s traffic on your…



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DogTipper

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How to condition a German shepherd

Materials needed:

  1. 1 German shepherd that likes other dogs.
  2. 1 whippet that likes other dogs and is friends with German shepherd.
  3. Ball.

Instructions.

  1. Throw ball for both dogs.
  2. Whippet will beat the German shepherd to the ball.
  3. Whippet will decide to run a victory lap, and the German shepherd will chase him.
  4. No German shepherd alive can catch a whippet, so they will run a whole lot more than if you just threw the ball for the German shepherd.
  5. Repeat once the whippet gets done with the ball and delivers it to hand.

Natural History

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The New Season of MeatEater on Netflix is a Work of Art

As long-time readers of this blog know, I am concerned about the future of hunting, especially because already-established hunters have hitched their wagons to an increasingly narrower and narrower spectrum of America politics. I have implored people involved in mass communications, be it film, television, or the written word, to find ways to reach beyond this narrow base. Indeed, had I not grown up in rural America where hunting and fishing were celebrated ways of life, I probably would be among the most noxious antis, just because my own politics simply don’t align with those of most hunters now.

However, there is at least one hunting communicator who does a great job reaching beyond this narrow base. Steven Rinella is an outdoor writer and science journalist extraordinaire, and what’s more, his art is reaching people who are not conservative in the true wonders that are hunting and fishing and preparation of wild meat. Rinella’s written word, such as can be found in what I consider his classic work, American Buffalo: In Search of a Lost Icon, is a mixture of William Faulkner, Aldo Leopold, and certain Midwestern rusticity that I cannot fully place. and is this writing stye that he applies to narration and story-line of MeatEater.

In 16 new episodes, which you can watch on Netflix, Rinella takes us from Interior Alaska to the Rewa River in Guyana. Lots of things are learned in each episode, from Latvian hunting incantations to the finer art of hand-lining for black piranha, but the most important thing we learn is that hunting is not evil. Not a deer falls in the Nevada desert, when Rinella and Joe Rogan go bow-hunting for mulie bucks, and most of the two-part episode of hunting Sitka black-tails on Prince of Wales Island involves seeing no deer at all. Most of those two episodes involve foraging shrimp, rockfish, and red sea cucumbers from the oceans and simple friendship in magnificent land.

Rinella loves animals. He revels in their biology and habits. To him, hunting is an intellectual exercise in which he matches his skills against their instincts and far keener senses. Animals die, but the carcass is butchered. The meat and organs go not to some banal rough fare, but instead, they wind up being prepared in fine dishes, many of which are cooked right over a campfire.

There are many moving scenes in this new season. When Papa Janis Putelis kills a moose that appeared to have “manifested” itself before his gun as a result of his Latvian “magic,” we learn about his general philosophy of life, which is to enjoy every footstep and savor every breath. We marvel at petroglyphs left by ancient hunter-gatherers in Guyana, ones that even the natives who take Rinella to see the artifacts just call the “ancient ones.” Though these Amerindians who guide Rinella seem almost Stone Age to us, they are inheritors of this world, and their world is ultimately changing as new technologies and foreign capital begin to work their way deeper into their part of the jungle.

However, perhaps the most moving scene in all of this season comes when Doug Duren, a Wisconsin farmer and conservationist, bags a bull caribou out of the Fortymile herd. Duren shoots the bull with pinpoint accuracy, but when he approaches the fallen beast, his emotions are flying. He revels in the excitement of having killed, but there is a look of admiration for the bull on his face that is difficult to describe. He strokes its chocolate and cream-colored fur so tenderly it is as if he’s petting a beloved dog that was just euthanized. There is reverence for this beast as the living being it was. This scene is something I’ve never before seen on a hunting show. Most of these shows have the hunters celebrating and high-fiving as if they have won the Super Bowl.

This reverence for the animals and their habitat mixes so nicely with the realization that ever episode that winds up with an animal’s death ends with that animal being prepared into a fine piece of table fare. Rinella’s narration is mesmerizing and deep, for it is his skill as a writer that really shine. The cinematography is gorgeous, but Rinella’s words pull and prod the images along, striking your aesthetic senses at their very core.

This is the art of MeatEater, and why Rinella is such a fine spokesperson for hunting in this century. He is not trying to feed the prejudices of the already extant hunting show aficionados. Instead, he reaches people at their deepest romantic and aesthetic senses, the part that makes us love animals and nature so much that one could just as easily become an anti-hunter. Indeed, because so few publicly visible hunters try to communicate on this level, these senses remain untapped and untapped at our peril.

I loved this season so much that I was almost in tears when it finally ended. Rinella is truly at master at his craft as a writer and communicator, even more so than he is a master at the art of hunting.

But he is truly a master at both.

Natural History

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Does Your Dog Dig Holes?

From the makers of the iFetch comes the iDig interactive toy for your dog. Two models are available, one to stay at home, and one to fold up and take with you when you travel. The idea is this: you load the dog’s favorite toys or treats into the idig, then close the flaps over […]


Doggies.com Dog Blog

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Siegfried Farnon (Robert Hardy) on the problems of keeping pets

It all makes sense, except for those barking retrievers and that fawn whippet that was Robert Hardy’s own beloved dog.

Natural History

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Our Mexico Trip

Our Mexico Trip

Those of you who follow me on Instagram and Facebook know that we just got back from a glorious (and I mean glorious) trip to Mexico. For the past 12 years, the band my husband Stage Manages for a living has co-headlined a festival called Holidaze that has now been held in Jamaica, Mexico (Tulum and Puerto Morelos), and the Dominican Republic. This year, they returned to Puerto Morelos and it was genuinely blissful for us to get to spend 5 days in the sunshine with so many of our closest friends.

Last year, Essley started throwing up 4 hours before we were to leave for the airport, so the kids and I had to stay home, which was admittedly devastating for all of us. But I think it made us appreciate this year more. (Side note: Essley coughed so hard she vomited the night before we left again this year, and Emmett picked up a cold down there that caused him to cough so hard he vomited on the flight back. I couldn’t make this stuff up.)

Instead of going on and on about the details of the trip (and really, aside from fun in the sun with friends of all ages and lots of music, there aren’t a lot of details), I thought I’d just share some of my favorite photos from the trip, as seen above. These aren’t high quality photos by any means – they were all snapped with my phone. But they certainly do show you how much we fun we had.

Are any of you traveling soon? Who else has been to Puerto Morelos?

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Bubby and Bean ::: Living Creatively

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Happy Holidays + Taking a Break

Happy Holidays from Bubby and Bean!

Happy winter Solstice! Merry almost Christmas! I almost can’t believe I’m typing that. Is Christmas really only four days away? I know it’s super cliche to talk about how quickly time passes but truly, this season has gone by in a blink. It’s been the busiest season of work for me ever (so grateful for this!), we just returned from our trip to Mexico for Robbie’s work Sunday night, we had two school holiday parties yesterday, Essley’s winter dance recital is tonight and her birthday party is Sunday, then family arrives to stay with us, and the following day is Christmas Eve! This is the first year both of my kids are old enough to really get Christmas, and the excitement level is through the roof! The day after Christmas, Robbie leaves town for work, I join him on the 30th in Atlanta, and we fly back together on January 1st (my birthday), which is quickly followed by Emmett’s birthday, then a long (boooo!) winter tour for him with the band.

Honestly, I love being busy, and get sad when things slow down. But I also want to take a breath and enjoy the last few days of the season before it’s gone. So I will be taking my annual break from the blog to spend time with my family and catch up on non-work things, starting today. I’ll likely still be somewhat active on Instagram, but we won’t be back with any new posts here on the blog until after the New Year.

However you celebrate (or don’t celebrate!), I wish you the happiest of holidays and a New Year full of joy, love, and peace.

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Bubby and Bean ::: Living Creatively

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What Do Vegans Feed Their Pets?

Embracing a plant-based lifestyle may start with what’s on your plate, but whether you’re vegan for health, humane, or environmental reasons, for lots of people it’s not just a diet, but a way of life. Being vegan can affect nearly every purchasing decision, so it’s only natural to think about how it affects pets’ diets. 

So, what do vegans and vegetarians feed their cats and dogs?
Due to the distinct differences in their physiology, we have to look at dogs and cats separately. Dogs are omnivores. Thanks in part to living with us for thousands of years (and enjoying food that “falls” off our plates), dogs’ digestive systems have evolved to support a more starch-rich diet. Dogs have a dietary requirement for specific nutrients. Those nutrients can come from meat or they can come from plant-based sources. So, many people who choose not to eat meat choose to feed their dog vegan dog food, like Halo’s Garden of Vegan.

What about vegan cat food?
Unlike dogs, cats are obligate carnivores, meaning there are nutrients cats need that they cannot get from a vegetarian diet. So, from a health standpoint, real, whole meat cat food delivers the nutrients they need. But what about pet parents who are concerned about the treatment of our life-giving animals and the environmental impact of pet food? In the U.S., dogs’ and cats’ diets are responsible for “25-30% of animal production in terms of the use of land, water, fossil fuel, phosphate, and biocides,” according to a 2017 study.

That’s why Halo commits to ethical and sustainable agricultural practices. Our promise of OrigiNative® sourcing means we work with farmers who treat animals with respect and help maintain a regenerative ecosystem by using original animal husbandry and farming practices and rearing animals in their native environments. To ensure the OrigiNative® philosophy is followed, Halo’s meat proteins are GAP (Global Animal Partnership®) certified humane, and our fish is MSC (Marine Stewardship Council®) certified sustainably caught. Plus, all of the fruits and vegetables Halo uses are non-GMO.

Vegans, vegetarians, and other animal lovers and advocates who want pet food they can feel good about can find great choices with Halo. Through our mission to change the way companion animals are fed and farm animals are raised…for the better, we offer canned vegan dog food, dry vegan dog food, and vegan dog treats, as well as dog food and cat food made with third-party certified humane and sustainable proteins.

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