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The myth of neoteny

September 20, 2011 by retrieverman

St. Bernards developed their domed heads as a result of selective breeding, not some magical neotenization process.

I am about to explain something that is very hard for people to grasp. It is almost as difficult for dog people disabuse themselves of this common meme in the dog world as it is the dominance model.

This one actually popped up in response to the dominance model, and it’s just as bizarre. It’s one I used to accept without thinking about it, mainly because the only book on dog domestication that I’d ever read was Raymond and Lorna Coppinger’s book, which is where this notion got started.

This theory has largely been falsified for me, for a very simple reason:  the dates don’t add up.

Basically, the theory goes this way:

About 12,000 years ago, man started to do small scale agriculture and was able to set up the first permanent settlements.  This coincided with a period of scarcity of large prey, which brought wolves into the camps to scavenge. Those wolves that had less fear of people were better adapted to scavenge, and these began to evolve into village dogs. In the village, they adapted to the less high quality food. They evolved smaller brains and jaws, and because they didn’t hunt, they lost their ability to do much hunting to survive.  They also became smaller animals, developed floppy ears and spots, and started coming into heat twice a year.  And because dogs evolved from this village dog population, we should consider dogs a different species– and we shouldn’t use dominance theory to dominate our dogs.

In this model, the dog became  a perpetually juvenilized species compared to the ancestral wolves, and thus, it has become very common  for people to call dogs neotenous wolves. Neoteny is the retention of juvenile morphology into adulthood, and it’s normally used in relation to various species of salamander, like the axolotl, that retain the larval gills into adulthood.

Different species can have different neotenous features. The dodo retained much of the morphology of a juvenile pigeon, even though it weighed as much as 50 pounds. Humans have several neotenous features in comparison to other apes, including our broad skulls and hairlessness. The epicanthic eye fold that is present in some human populations, such as in Asia, is also sometimes considered a neotenous trait. However, it’s more like a fetal trait, because it is present in other humans only at the fetal stage.

I’m not denying that some traits dogs have are neotenous. Floppy ears and flattened faces are definite neotenous traits in dogs.

What I think is a bit screwy is suggesting that this is the process by which they were domesticated.

My biggest problem with Coppinger’s theory is the dates simply do not line up. He has no answer for dog remains that have been found in Germany, Russia, and Switzerland that all date to at least 14,000 years before present.  That’s 2,000 years earlier than Coppinger thinks they would be found.

Furthermore, 14,000 years ago, there were no villages, even of the type that existed among the earliest agriculturists. These 14,000-year-old were all the product of hunter-gatherer camps, not semi-permanent villages. Strike one.

Then, we have another problem.  The genetic data points to an even older origin for the domestic dog than the 14,000-year-old specimens, and the Russian specimens have had a least one dating that the put them at 17,000 years before present.  The study that revealed the dog genome calibrated the dog’s split from the wolf to 27,000 years before present. Some of the early mtDNA studies have pointed to an even older origin than this– over 100,000 years before present. However, I’m a bit skeptical of such an old date for dogs, because we don’t have any record of them from this age. We do have records of what might be dogs or early domesticated wolves that are 31,000 years old in Belgium and 33,000 years in the Altai Mountains.  These discoveries are a bit controversial, for the simple reason that we can’t assume that domestication always yields major morphological changes between wild and domestic populations. Assuming that domestication always changes morphology is a major hindrance to finding out exactly when dogs began to diverge from wolves. It is possible that many “wolves” found in caves near humans were actually wolves that were domesticated.

All of those dates are older than the 12,000 years that Coppinger needs for his theory to have a basis in the actual history of the earth.

Now, that would be enough for me to really question this theory, but there is more.

What does he do with the odd wolf that acts like a dog?

These exist. I’m not making this up.

There was one in Alaska recently named Romeo, who did all he could to become a dog.

And then there was Wags, the wolf that Adolph Murie took from a den as part of his study of the wolves at Denali.

Then there are the dogs that are very much like wolves, and not all of these are primitive dogs with a wolfish phenotype. Jess at Desert Windhounds has an Azawakh that dislikes being touched and has bonded to only her husband. These are traits we would normally see in wolves that have been raised in captivity, not in a domestic dog breed that was developed where there are no wolves (as far as I know).

The truth is that although there are differences between dogs and wolves, these differences are not hard and fast. There will always be dog-like wolves and wolf-like dogs.

In Coppinger’s theory, physical neoteny, which is there at some level, is accompanied by behavioral neoteny. Wags looked like a fully adult wolf, not like a cuddly little puppy, but she acted more or less like a Labrador retriever. Jess’s Azawakh has drop ears and a hound-dog appearance, both of which are manifestations of physical neoteny.

You cannot predict among dog breeds which ones are friendly or not by their appearance. Kuvaszok look like golden retrievers, but they will guard sheep and property just as wild wolves protect their territories and pups from intruders from other packs. German shepherds and Malinois look like wolves, but they are very easy to train.

And then there are the wolves that live by scavenging today.

If Coppinger suggests that all domestication required was for wolves to scavenge off of human settlements, then why aren’t the wolves of Israel and Italy becoming dogs? I’ve never heard of a floppy-eared wolf in Israel or in Italy. Nor have I heard of one with spots on it.

Then, there is the big one.

Last year, Mark Derr sent me an e-mail with the title “neoteny death nell.”   In it was an attachment of a study on St. Bernard skulls by Abby Drake and Christian Peter Klingenberg. The authors discovered that over the past 120 years, the St. Bernard’s skull changed to a more rounded and more dome-like shape as the result of selective breeding, not any magical process that was the result of breeding for greater docility or trainability. The results of this study and a full critique of Coppinger’s neotenous dog model can be found in Mark Derr’s forthcoming book, How the Dog Became the Dog, which will be out in October (but you can pre-order from Amazon today!)

It seems to me that the apparent neotenous features we see in domestic dogs are the result of selective breeding, not this magical neoteny process.

One of the big flaws in Coppinger’s analysis is the reliance upon the Belyaev fox farm experiment to generate most of this theory. I agree that this is an interesting study, but it can’t tell us everything there is to know about dog domestication. It’s not a good 1:1 comparison. In the end, I think the most important thing to come from the experiment is the change in the critical period time between non-selection and selection for tameness foxes.  That change in critical periods can have really major effects upon the development of any animal, because a longer period gives an animal much longer time to become social to more creatures. Further, the floppy ears and spots have occurred in farmed foxes for many years, and there is no evidence that these foxes are tame. There are whole strains of non-tame farmed foxes that are spotted. And then, there’s the case of a line of Mexican wolves that were kept at Carlsbad Caverns. After generations of keeping them in captivity, they started to develop dog-like features, even though they had never been selected for tameness.  Because of these features, the wolves were thought to be hybrids with dogs and were euthanized. However, the DNA analysis of surviving wolves from this lineage revealed that they were pure Mexican wolves.

Even being raised in captivity can change the morphology of Carnivoran skulls, for there are plenty of examples of zoo lions and wolves having less powerful jaws and smaller heads than wild ones. The larger head appearance is the result of the jaw muscles developing on top of the head, and the jaw power is the result of those muscles being used to kill and dissect large prey.

And then there is the whole theory about motor patterns that Coppinger posits. In Coppinger’s model, domestic dogs that use predatory behavior for “work” have been selected to show those motor patterns only in a truncated manner. Herding dogs can chase and stalk, but they can’t kill. Pointers can only stalk.  Retrievers can catch but they can’t kill. Because dogs are derived from non-predatory village dogs, they were selectively bred to show these truncated predatory motor patterns. And that’s all they can show. They can’t kill or dissect prey.

Now, I read this as I watched a golden retriever with very strong retrieving motor patterns– that would retrieve anything at anytime with a very soft mouth– go out, run down and kill a rabbit, dissect it, eat it, and then crap it out in little furry turds in the yard. My own eyes falsified that part of Coppinger’s model.

The Coppinger model is beguiling because, like the dominance model, it is very easily explained on television. It’s easy for the layperson to follow it.

And it’s also a model that many people who reject the dominance model use, because if dogs are derived from village dogs and not wolves, then we don’t need the wolf dominance paradigm at all.

However, when you take on the Coppinger model to refute the dominance model, you wind up shooting yourself in the foot. There are very real problems with the neotenous dog model, as many as there are with the dominance model.

We need to recognize that dogs are wolves,  but they are domesticated. However, the process by which they were domesticated was much more complex than the model provided in Coppinger’s book.

To turn neoteny into this bizarre, almost magical process that creates domestic animals is an oversimplification of what was a very dynamic process that happened over tens of thousands of years.

 

 

 

Posted in dog behavior, dog domestication | Tagged dog domestication, How the Dog Became the Dog, Mark Derr, neoteny, Raymond Coppinger | 26 Comments

26 Responses

  1. on September 20, 2011 at 8:36 pm Jess

    I never pre-order books. I wait until they show up on Half.com for cheap.

    I’ll happily take review copies, though.


  2. on September 21, 2011 at 2:19 pm M.R.S.

    I don’t mean to argue, but as I read Coppinger, he is saying that certain behavioral patterns are strengthened, or are minimized, in certain breeds/strains. Not that they are necessarily completely absent. He does say that certain parts of the pattern are considered faulty in certain dogs, such as the grab-bite or bite-kill behaviors in herding dogs.

    I think it’s also true that different parts of the pattern may be expressed in certain circumstances and not in others, and also that they may be modified in individual dogs by experience and by training. And certainly also, modified in populations over generations by selection in breeding.


    • on September 21, 2011 at 3:29 pm Dave

      If “grab-bite” is considered as a faulty behaviour, then I feel sorry for the cattle-dogs; or the hog-dogs, which by the way, can learn the different behaviours required of them with all the different livestock.


      • on September 22, 2011 at 11:38 am M.R.S.

        Dave– I should have been more explicit. You are correct, that at times a good bite is definitely needed (although preferably one that does not unduly damage the stock).


    • on September 21, 2011 at 4:13 pm retrieverman

      He actually does say this. If you read the part on livestock guardian dogs, he claims that they cannot dissect dead lambs and eat them. They have no predatory behavior whatsoever.

      Mark Derr did a great critique of this in his book, showing how even cheetahs have to learn how to dissect dead prey.


      • on September 21, 2011 at 5:40 pm Dave

        Those Tobets from Kazakhstan pretty much debunked the no drive thing.


        • on September 21, 2011 at 6:11 pm retrieverman

          Thank you, Dave.

          That post is basically Coppinger Cryptonite, as is Lindsay Tompkins’s herding Caucasian Ovtcharka.


          • on September 21, 2011 at 6:56 pm Dave

            In the western world, it would be considered as a fault. So, Masakado Shiba wouldn’t be the silver bullet. However the footage of the aboriginal dogs are.

            Unless the Ovtcharka crowd wants to pull the Saluki card and claims the non-Westernized dogs are impure.


            • on September 21, 2011 at 6:59 pm Dave

              Footages*.


            • on September 21, 2011 at 7:30 pm retrieverman

              There are other herding Caucasian ovtcharkas. There’s another one on youtube.


              • on September 21, 2011 at 7:34 pm Dave

                Yeah, but you get the Paddy Burns of the world who say collies can’t be gundogs; and if they are used, the Labrador Retriever is a more refined, better breed for the job.


                • on September 21, 2011 at 7:35 pm Dave

                  Or should that be, Paddy Burnses? Not sure how people with the same surnames are handled in the English language.


  3. on September 21, 2011 at 4:00 pm Peggy Richter

    grab-bite is something that you need with difficult livestock. I’ve had dogs that were run over by stock when they wouldn’t grip when necessary. I’ve seen plenty of sheep that WOULD NOT MOVE if the dog didn’t give a sharp nip or so, and a few that would charge a dog that didn’t have the confidence that comes from knowing it was allowed to bite. Plenty of older herding books give advice on what to do with a dog that is too inclined to use it’s teeth when unnecessary– few of these recommnedations are to get rid of the dog. I never bought off on Coppinger’s theory. It’s rather like saying every city has beggars and therefore we are all derived from beggars. It may have sounded plausable, but as a theory, from the beginning there were serious holes in it.
    I think if someone seriously tried a “wolf domestication project” and used as their goal an animal that might be able to spook or drive prey towards human hunters, maybe follow a scent trail to find game and maybe sound an alert for threats near the home(cave, etc), you would find that it took only a very small number of generations to achieve, even with wolves that have been selected for generations to fear man. But if you expect that a wolf will suddenly be able to run in a modern herding or field trial, you would be disappointed. The assertions that wolves can’t be “tamed” or domesticated are all pretty much based on what happens with the first pups taken from wild parents, and have mostly been pretty unrealistic in their expectations.
    Peggy Richter.


  4. on December 1, 2011 at 6:57 pm The truth about cats and dogs « Retrieverman's Weblog

    […] delayed” wolves. This notion can be traced to Raymond Coppinger’s work on neoteny. Dogs are not neotenous wolves. All of their bizarre shapes come from selective breeding by humans, not some magical juvenilzation […]


  5. on January 3, 2012 at 2:14 pm “The booted lynx” « The Retriever, Dog, & Wildlife Blog

    […] of these cats evolved to scavenge and hunt rodents near human settlements– which means that Coppinger’s flawed theory on dog domestication might actually work for […]


  6. on May 28, 2012 at 2:11 pm Ann D.

    A large number, up to 70%! of the ‘tame’ foxes in Belyaev’s experiment had NO phenotypical change from the wild type. This result demonstrates that neoteny is NOT a prerequisite to domestication. Like Romeo and these silver foxes, domestic acting canines can still have the wild phenotype.

    Check facts with:
    Lyudmila Trut
    trut@bionet.nsc.ru


  7. on November 14, 2012 at 2:55 pm Lena A.

    Many interesting points here, but I wonder why we have so many (appearance wise) rather distinct groups of dogs bred for different purposes. Spaniels, retrievers, waterdogs (even setters) look very similar. It’s hard to imagine that it is so only because hunters who have bred these breeds over last centuries in different parts of Europe just have happened to prefer that type of dog over say a spitz or terrier type. I think it’s more plausible that if you select for certain hunting traits that appear in wild canine’s development at a certain age, you some what arrest the dogs development on the whole to that level. Not entirely but for many parts. So you are not likely to produce a dog that behaves like a LGD or a spaniel but looks like a husky, although you can have exceptions to any direction (and thus the need for selective breeding).

    “You cannot predict among dog breeds which ones are friendly or not by their appearance. Kuvaszok look like golden retrievers, but they will guard sheep and property just as wild wolves protect their territories and pups from intruders from other packs.”

    Kuvasz might also be acting like a small puppy who keeps very close to his den and littermates, and who reacts with aggression or fear to all things unfamiliar to him. Golden retriever might be “a wolf puppy” just one developmental step ahead of Kuvasz, one that has grown out of the fear stage and is showing the first steps of hunting behaviour. Also Kuvasz should be very gentle with his family and rather sensitive to instructions and praise from his owner, so I don’t think it’s entirely true to say they are not friendly. They should be, just not towards things they haven’t been socialized to.

    Also what we see in dogs is that some breeds on average develop faster than others. Newborn puppies look the same, but a Husky puppy reaches developmental stages faster than a Spaniel puppy. LGD’s are probably the slowest ones to mature. This seems to somewhat follow the pattern of how the breed behaves or looks as an adult.

    “The authors discovered that over the past 120 years, the St. Bernard’s skull changed to a more rounded and more dome-like shape as the result of selective breeding, not any magical process that was the result of breeding for greater docility or trainability. ”

    I’m sure one can greatly modify almost any features by selective breeding. But I wonder if this study can show that the docility or trainability of St. Bernard have NOT increased during the last 120 years as well?


    • on November 14, 2012 at 6:03 pm retrieverman

      You can take a West Siberian laika and teach it to do everything that a retriever or spaniel can do, and it will do it just as well.

      And West Siberian laika looks like a Siberian husky. They often have some wolf in them.

      So I am not buying that what a dog looks like has anything to do with its performance.


      • on November 15, 2012 at 11:47 am M.R.S.

        Hmm…will a Laika find and carefully retrieve live baby wild ducks for banding (as retrievers have done)? Or do the sort of work required in today’s field trials (either UK or North American)? I don’t see guide dog schools using Laikas (or huskies) for their purposes, but they do use a large number of retrievers.

        Will a St Bernard compete equally with a Border Collie in Agility or Herding? “Looks” in this instance include physical properties that do affect performance.


        • on November 15, 2012 at 5:45 pm retrieverman

          Because they don’t know about them yet.

          A West Siberian laika is a golden retriever in wolf’s clothes.

          Many Siberian hunters train their West Siberian laikas as bird dogs. They also retrieve ducks.


        • on November 15, 2012 at 7:52 pm Dave

          Russian trial rules:

          https://sites.google.com/site/laikarticles/documents/pravilapovodoplavausheiptice


        • on November 15, 2012 at 9:04 pm Dave

          Training manuals and books on Laika and Waterfowls:

          https://sites.google.com/site/laikarticles/nataskaohota/dich/nataskapovode

          https://sites.google.com/site/laikarticles/nataskaohota/dich/nautok

          https://sites.google.com/site/laikarticles/nataskaohota/dich/lajkautina

          https://sites.google.com/site/laikarticles/nataskaohota/dich/ohotanautok

          https://sites.google.com/site/laikarticles/nataskaohota/dich/nautokskarelkoj

          https://sites.google.com/site/laikarticles/nataskaohota/dich/trenirovkautka


  8. on November 15, 2012 at 1:36 pm Lena A.

    I’m familiar with laikas and some other northern spitz type hunting breeds. They are often more likely to aggressively guard the kill from people (especially from anyone other than their owner) than eagerly retrieve it.

    Also I think that if it was true that you can take any hunting breed and train it to behave like any other type of hunting breed, we should see way less correlation between behaviour and appearance within any breed type. We should have at least some LGD’s with pointed ears, because in many areas LGD’s have existed and worked with herding dogs. We should see breeds that hunt like terriers but look like spaniels, because these breeds have co-existed in the same areas, such as the UK. And so on.

    In general we should have way more “allrounder”, versatile dog breeds, because why would people of the past have created and sustained several breeds or types of dog, if one breed could have done any of the jobs required. It’s actually quite rare for a dog breed to have history of being used both as a hunting and a herding dog. There are some, but usually they are no match for more specialized breeds.


    • on November 15, 2012 at 5:46 pm retrieverman

      Actually, you aren’t familiar with West Siberian laikas. They very commonly used to flush and retrieve birds. They are not particularly dog aggressive, unlike the Karelian bear dog or the Russo-European laika.


    • on November 15, 2012 at 5:53 pm retrieverman

      I want to disabuse people of the notion that only specialized breeds can do their jobs and that only those specialized breeds can do those jobs well.

      I also want people to understand that even a dog as theoretically specialized as a golden retriever can be trained to hunt many things, including treeing raccoons and tracking foxes.

      The idea that dogs should be hyper-specialzied is very new, and it’s also uniquely Anglo-Saxon. Most Continental European “bird dogs” (Brittanys and Short-hairs) are actually used in a wide variety of hunting situations. It was this way in America, too. It used to be that the main farm dogs were a “shepherd” (a collie-type) that was used to hunt just about everything and herd and tend the stock. In other areas, that dog was the cur– blackmouth, Catahoula, etc.

      There are livestock guardian dogs with pointed ears. Have you seen the dogs the Masai and Navajo use? Mark Derr in Dog’s Best Friend points out a particular prick-eared type of dog that the Navajo use to guard their sheep. The Navajo prefer little mongrel dogs, not great big European mastiff-types. I can’t recommend that book enough. It will get you thinking much more skeptically about some of the claims people make about dogs.

      Mark Derr on the prick-eared livestock guardian dog: http://www.nytimes.com/books/first/d/derr-dog.html


  9. on November 15, 2012 at 6:08 pm Lena A.

    You’re right about the Laikas, all three of them can be used as flushing/retrieving dogs.
    Thanks, and I’ll try to look into Dog’s Best Friend!



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