Moar foxes

As I consider foxes to be Honorary Cat™s, I have no compunction about posting pictures of these fluffy canids. Reader Gregory called my attention to a series of 22 fox pictures on Bored Panda, some of which are great. I reproduce Professor Ceiling Cat’s Favorites below.

By the way, that site also quotes this famous experiment, an experiment about which I’m quite dubious, for I haven’t been able to get the original paper (it’s in Russian, anyway):

The fox is a member of the canidae family, which also includes dogs, wolves and other similar animals. After 50 years of breeding experimentation in the Soviet Union, they’ve also provided us with extraordinary insight into the domestication process. Over several generations of selective breeding (by choosing foxes with less fear of humans), Soviet scientist Dmitry Belyaev was able to breed silver foxes that began to exhibit domestic traits like floppy ears, tail wagging and spotted coats.

I’m not dubious about the efficacy of selecting for tame foxes, as there’s certainly genetic variation for that. What I question is whether the floppy ears and coat color, or even tail wagging, automatically went along with tameness as a byproduct of the “tameness” genes. Science journalists have made a huge deal about selection for tameness being accompanied by these other traits, presumably as ancillary effects of the very genes that made the foxes less wary of humans. (If one gene has multiple effects, those are called “pleiotropic effects,” so color and floppy ears are said to be pleiotropic effects of the genetic variants producing tameness.)

An alternative hypothesis—which might not be checkable even in the original paper—is that the  researchers who selected for tameness also selected simultaneously (and perhaps inadvertently) on ear droopiness and coat color. Those things, too, could have been genetically variable and responded to selection for the traits themselves.

In fact, I’m not even sure why it’s so fascinating to people that genes for domesticity in ancestral canids might have side effects on coat color, even if that were true. Is that supposed to say something about how d*gs became domesticated? Is it supposed to say, for instance, that while humans were selecting the ancestral d*g from the wolf (or the wolf was selecting itself via reproductive advantages accruing to those individuals who could approach humans and their food more closely), the tameness automatically produced some of the traits of modern d*gs? I don’t believe that at all, but I’m open to evidence. And, after all, the closest relatives to modern dogs aren’t foxes but wolves.

Anyway, on to the foxes:

Screen shot 2014-04-17 at 7.30.25 AM

Image credits: Edwin Kats

Screen shot 2014-04-17 at 7.11.37 AM

Image credits: Roeselien Raimond

Screen shot 2014-04-17 at 7.13.27 AM

Image credits:

Screen shot 2014-04-17 at 7.15.15 AM

Image credits: Jim Cumming

Screen shot 2014-04-17 at 7.16.02 AM

Image credits: William Doran

Screen shot 2014-04-17 at 7.32.18 AM

Image credits: Igor Shpilenok

Screen shot 2014-04-17 at 7.34.38 AM

Image credits: Wenda Atkin


  1. gbjames
    Posted April 17, 2014 at 9:09 am | Permalink


  2. Alex T
    Posted April 17, 2014 at 9:13 am | Permalink

    WRT the “tameness” study, I recall reading that they were actually selecting for flight distance alone which would rule out inadvertent selection on coat or these other factors.

    • Posted April 17, 2014 at 9:28 am | Permalink

      Not it they selected inadvertently. It’s hard to tell from the second-hand reports. And it should be repeated. I just don’t trust the results, but that’s a gut feeling.

      • Posted April 17, 2014 at 10:21 am | Permalink

        I’m certainly no expert, but surely flight distance would only be relevant for the earlier generations. As they became more tame and researchers had more contact, I imagine it would become increasingly possible for researchers to unwittingly identify other characteristics as “tameness” when selecting foxes for breeding.

        That said, Dawkins cited this work in The Ancestors Tale (p. 30) – which apparently suggests levels of seratonin and two thyroid hormones as being possible factors.

  3. Posted April 17, 2014 at 9:23 am | Permalink

    Beautiful photos of beautiful animals! Love them.

  4. gravelinspector-Aidan
    Posted April 17, 2014 at 9:27 am | Permalink

    #3 looks like foxy got stuck to a really cold pane of glass.
    Where’s Oksana’s photos of a Siberian fox on the Muravlenko site?

  5. moarscienceplz
    Posted April 17, 2014 at 9:27 am | Permalink

    I would think that tail-wagging is a social communication device, possibly indicating non-aggressive intent and perhaps submission. If that is the case, then it would be rare for humans to observe tail-wagging in wild animals but common to see it in tame(r) ones. No genetics involved, just transfer of the signaling target from other foxes to humans.

    • Posted April 17, 2014 at 12:37 pm | Permalink

      Tail wagging is a trait in at least certain kinds of canids. I know that fox pups and African wild dog pups wag their tails while they greet their parents and beg for food. I do not remember seeing this for wolf pups, but I could easily have missed that. In any case, deliberate or inadvertent selection of tail wagging could be simply selection for persistence of this appeasement behavior.

      • moarscienceplz
        Posted April 17, 2014 at 4:34 pm | Permalink

        I’m not saying there wasn’t a selection for a “wag gene”. I’m just suggesting it’s possible there wasn’t and that wagging increased simply because the tame foxes now think of humans as surrogate parents and wag at the humans just as much as they naturally wag at their parents.

  6. Diana MacPherson
    Posted April 17, 2014 at 9:30 am | Permalink

    What cute foxes. Little arctic foxes are very sweet looking too!

  7. Sastra
    Posted April 17, 2014 at 9:36 am | Permalink

    Given how popular this study was, how relatively simple it would be to do, and how much researchers would presumably enjoy working with cute animals who are supposed to ‘bond’ positively with the researcher, I’m surprised that it hasn’t been replicated. It wouldn’t be hard to try to control for confounding factors.

    That might be a red flag. Or not. It should be replicated either way, though.

    • moarscienceplz
      Posted April 17, 2014 at 9:42 am | Permalink

      Well they would have to work with a lot of wild, aggressive animals, too. It wouldn’t be all cuddles and puppy kisses. Plus, I bet it’s expensive feeding and providing vet care for a large number of fairly big animals.

      • Sastra
        Posted April 17, 2014 at 9:51 am | Permalink

        True, but my suspicion (possibly mistaken) is that there are a lot of animal lovers out there who would throw money at something which they think has something science-y and important to do with d*gs.

        In other words, this is “sexy” research — meaning “publicly appealing,” like black holes. There could be a NOVA special on it, whereas my understanding is that most scientific studies only hope to be fascinating within the field.

        • Posted April 17, 2014 at 12:38 pm | Permalink

          …then there are the larger #s of stock that are selected out. What happens to them? Harder to do in this country, and harder to do now.

  8. frankschmidtmissouri
    Posted April 17, 2014 at 9:58 am | Permalink

    Since the canine genome has been sequenced, we should have an objective measure of the extent to which alleles likely to be associated with tameness (e.g., those for oxytocin responsiveness) would be linked to coat color.

    And no, I’m not going to do it; I have a dissertation exam to go to. But someone should.

    • Diana MacPherson
      Posted April 17, 2014 at 10:02 am | Permalink

      Ok, you’ll do it after the exam then? 😉

  9. William Stewart
    Posted April 17, 2014 at 10:01 am | Permalink

    Dear Professor Coyne and commenters,
    More information on the farm-fox experiment can be found in an article in the American Scientist sometime in 1999.
    William Stewart

  10. Bob J.
    Posted April 17, 2014 at 10:02 am | Permalink

    It would seem that more vaiablity in coat color and ears could come from the fox’s out-sourcing of food gathering, a la Daniel Dennett. That is, due to not needing to hunt, selection pressure on these traits is less. Of course determining juvenile mortality rates of foxes in the wild will be near impossible.

    Great pictures.

  11. Sheila B and Zin
    Posted April 17, 2014 at 10:03 am | Permalink

    Obviously the top photo is good – it’s by Edwin Kats 🙂

  12. Torbjörn Larsson, OM
    Posted April 17, 2014 at 10:34 am | Permalink


    The last pic is a howler:
    – Protect me. bro.
    – Against those scary looking creatures? You are ‘foxing’ kidding me!?

  13. Jacques Hausser
    Posted April 17, 2014 at 10:44 am | Permalink

    Scepticism is a healthy behavior in science, but well, in this case, the study is well known and was subject to numerous publications… yes, I know, few occidental scientists read Russian (I don’t) and until the end of the 90ths Russians did not publish in English. Belyav’s hypothesis to explain color and morphology changes (which didn’t affect all the domesticated foxes) as well as physiological changes was that selection for behavior disrupted general physiology regulation and hence gene expression in several area.

    I had the opportunity to visit Belyaev’s Institute at Akademgorodok, the “Science city” close to Novosibirsk, in february 1992 – I was working on the role of chromosomes in speciation of shrews and I had some colleagues in this Institute. My feeling was that, with rather limited ressources (even if Lyssenko’s era was over, genetics was not in very high esteem in Soviet Union) they did very good and solid science.
    It was a strange time, the Soviet Union just dismanteled, the shops empty and I was the only one, out of several occidental scientists invited to the “First (and obviously last) All Union’s Shrew Conference” daring to attend. Some (pacific) demo0nstrations in Moskow, very interesting voyage in the transsiberian (as my Russian colleagues didn’t have money, I was “smuggled” from Moskow to Novosibirsk with a fraudulous indigenous ticket, less than a tenth of the price for a foreigner), nice colleagues and hot discussions, in both scientific and politic fields !

    For information about modern sequels of Belyav’s researches, see

    and for a general article

    • JoeyM
      Posted April 17, 2014 at 7:42 pm | Permalink

      One other link, a video clip discussing the experiment.

      It’s a nice intro to a discussion on natural selection. I wish I knew what documentary it was from so I could purchase a copy.

  14. D. Taylor
    Posted April 17, 2014 at 10:56 am | Permalink

    This experiment was featured on PBS on Nova’s “Dogs Decoded,” which originally aired Nov 12, 2011 and was repeated on July 3, 2013. Below is an excerpt from the transcript.

    Half a century ago, Soviet scientists set up a breeding program to see if they could domesticate silver foxes. Foxes are closely related to wolves. The project, which has attracted the attention of scientists across the world, is opening a remarkable window on the process of domestication.
    Here on a farm, outside the city of Novosibirsk, the experiment still continues today, overseen by Dr Lyudmila Trut.

    The breeding program began in 1959, when the first foxes were selected from local fur farms.
    DR. LYUDMILA TRUT (Institute of Cytology and Genetics, Russia) [Dubbed]: We approached the animals in the cages and recorded their reaction to us. We could see that some of the foxes showed aggressive behavior, others were frightened; but only one percent of them showed neither signs of fear or aggression.

    NARRATOR: This one percent is selected to become the founding generation of a new population of foxes. At every generation, the selection process is repeated, only the tamest foxes are allowed to breed. Within just three generations, the aggressive behavior begins to disappear.

    LYUDMILA TRUT [Dubbed]: The radical changes came through in the eighth generation, when foxes started to seek contact with humans and show affection to them. The amazing thing was that cubs who had just started to crawl opened their eyes and started showing affection to humans by breathing heavily, wagging their tails and howling.
    This kind of response was a big surprise to us.

    NARRATOR: Half a century and nearly fifty generations later, the foxes are tamer than ever. It’s an accelerated model of how dogs might have been domesticated from wolves.

  15. Heather
    Posted April 17, 2014 at 11:16 am | Permalink

    Couldn’t the co-occurance of tame behavior, mottled fur and floppy ears in both domesticated foxes and dogs be the result of a selective sweep? They’re both in Canidaea, so presumably they have nearly identical chromosomal synteny. Obviously behavior is polygenic, but if genes with the largest phenotypic effect for all three traits, tame behavior, mottled fur and ear cartilage, were near each, or just on the same chromosome, they could be swept together.

    Of course, this assumes there is standing variation for this, in these genes at these loci, and that the association between them is more than a rarity 🙂

    • Posted April 17, 2014 at 2:24 pm | Permalink

      The dog genome project might produce assistance on this. The loci for floppy ears, piebald spotting, curled tails, etc. may be known in dogs, and they should line up to fox chromosomes pretty well by synteny. Perhaps loci for friendly behavior will turn out to be linked.
      There has long been a lot of interest in the fox research, so it seems reasonable that someone will go after this question one day.

  16. Kevin Alexander
    Posted April 17, 2014 at 11:54 am | Permalink

    Fox ears are tools. If a floppy eared fox was born in the wild it would not last long. It might even be a common variation but we don’t see it except in protected animals.

  17. Peter Ozzie Jones
    Posted April 17, 2014 at 5:24 pm | Permalink

    Don’t forget about the dingo. Some originals still survive on Fraser Island.

    Not sure of the baraminology, but they made it back to Australia post flood, possibly chasing the penguins?

  18. lisa parker
    Posted April 17, 2014 at 6:53 pm | Permalink

    Whether for commercial use or just scientific understanding, I find the selective breeding of domesticated animals atrocious. Pretty much all of this type of thing was done well before there was any understanding of genetics or genetically linked traits. This has resulted in animals that look like something the breeders had in mind, but also led to many breeds of domestic animals that are predisposed to various types of physical weaknesses (such as kidney failure, liver disease, heart disease, etc), and often have more than normal susceptibility to disease. I would also like to know if these studies in foxes is purely scientific research to understand genomes, chromosomal behavior, etc, or to produce an animal that is easier to care for and has specific color and coat lengths to help make fur farming more profitable.

  19. sailor1031
    Posted April 18, 2014 at 6:21 am | Permalink

    So why do Samoyeds, whose relation to the wolf is apparent, not have floppy ears and spots? Is the pleiotropic gene only present in the fox?

    • Diana MacPherson
      Posted April 18, 2014 at 6:53 am | Permalink

      Well there is all those years of selected breeding to make different dogs breeds look as they do as well.

      • sailor1031
        Posted April 18, 2014 at 7:05 am | Permalink

        But theses foxes developed floppy ears, spots etc after only 50 years of selective breeding. Samoyeds have been bred for centuries but don’t have such characteristics. And the characteristics of different breeds of dogs have been developed over millenia not 50 years. Granted if you breed a Samoyed with a Dachshund you’ll probably get something very different-looking but what do you get if you remain within the breed? as the Samoyed people did with their Bjelkiers?

        • Diana MacPherson
          Posted April 18, 2014 at 8:16 am | Permalink

          Yes but this was their particular selection that was selectively breeding tamer animals. In creating samoyeds, specific features of appearance may have been selected as well as disposition and the breed would be distinct only when it could “breed true” – in other words, when bred with other samyoeds, the features of samoyeds came out clearly.

          • oxalis
            Posted April 18, 2014 at 6:58 pm | Permalink

            Sammies as we know them are basically a modern western european thing. There were something like 9 founding dogs for the breed, making them one of the most inbred of all. The all white (with some barely tolerated biscuit) is pure european fancy, and to some extent the prick ears, too. The few photo’s I’ve seen of real Nenets dogs from early 1900s have been more colorful (pied dark and white seemed common), and some had floppy ears.

            It’s my favorite d*g breed, probably the most cat like one. The primary flaw is how horrible the coat gets when they’re neutered. Super thick, impossible to comb out, and miserable for the d*g in a warm climate [Sammy people like to pretend that the coat insulates them from the heat, but I asked my d*g, and she went from a tired out arthritic old lady to an ecstatic frisky puppy after I sheared her.]

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