Genetic bottlenecks, environmental pressure, and the development of dog breeds

I read MANY blogs. Somewhere in the range of a couple hundred. News, photography, scrapbooking (to make me feel guilty), and of course animal behavior, sheltering, all the Cardi blogs, legislation blogs, Gizmodo and Freakonomics and Lifehacker and Dooce, etc.

Some I read to keep me honest–Nathan Winograd is good at that.

Some I read because deep down some part of me wishes I lived the life of that blogger–the Daily Coyote comes to mind.

And there’s one I read because after I read his stuff I have to turn off the computer and go stare at the sky. Stephen Bodio is a hawker, a dog owner and breeder, a cattle rancher, a man who has dust in his boots and an incredible amount of deep and true thought in his head.

He wrote recently and touched on a few of the current genetic arguments surrounding dog breeds, which I thought dovetailed neatly into what I’ve been talking about in terms of genetic diversity in breeds. So I’m not going to quote him but I am going to use a few of the example breeds that he brought up.

Breeds as landraces are very, very old. A landrace is just as useful as a breed but it tends to be defined solely by the job it can do and by its environment–if you think about the Border Collie before it was ever registered, you get an idea. Their appearance can vary–if you mean appearance like “show quality” you could say the appearance varies a lot. Some Border Collies are substantially larger, some smaller, some have prick ears, some drop, color and coat type and leg length can vary. But in many other ways they are very similar–most people can still reliably identify them as Border Collies, and they all work in basically the same way.

The difference between a landrace and a breed is that a breed generally has a closed population, where individuals are bred only to other members of that breed. When you’re working with a landrace utility is king–if a dog shows up that kills rabbits really well, you’ll want to keep him around to breed to your other rabbit-killing dogs even if he doesn’t look exactly like them. But he’s unlikely to look totally different from them either, because there’s a certain body type and size and speed that naturally lends itself to successful rabbit killing. So among landraces there is a certain amount of sharing of genetic material–if the rabbit killer also happens to be a decent fox chaser, he may be used by people who like chasing foxes too.

Landraces probably developed along the same lines that dogs became domesticated hundreds or thousands of years prior–the dog wanted it and the human wanted it, and dogs have a tremendously plastic body and brain and from that symbiosis came dogs that were more useful at doing certain dog-human things than their ancestors had been.

So very, very long ago we can see what we think of as the major types of dogs. The Far East pioneered the idea of the dog purely as a companion, and they like the very stylized appearance of the flat-faced small dog, and so the flat-faced companion dog begins to spread along the trade routes coming west. This was happening thousands of years ago–the flat-faced small companion dog is very ancient.

Another group, probably in Germany, liked a type of dog that had very short legs, because short legs enabled the dog to do some things that long-legged dogs could not do. They probably began using them as hunting dogs but when the trade and conquest routes brought the short-legged dogs to other areas where hunting was not the predominant activity that dogs did, the owners of those dogs realized that short legs made some of their activites easier and made those dogs very useful. So, and again this is thousands of years ago, we already have a corgi landrace herding in Wales, a Vallhund landrace herding and hunting in Scandinavia, a Dachshund landrace hunting in Germany, etc.

The earliest true breeds, where pedigrees were maintained and it was understood that you only breed members of that breed to each other, are also extremely old. They grew out of an increased specialization of the landraces, a desire to have greater predictability in terms of exactly how big, small, tall, short, or useful that dog was.

There are only a very few of those true early breeds still around. Most of the breed-origin myths that are droned over the picture at Westminster are romantic fiction. But some of them DO still exist. One is the Sloughi (not Saluki), one is the very primitive Basenji, another is the Chow, and I think the Pekingese makes the list as well.

So the vast majority of the breeds we know today became closed, where individuals were not bred outside the breed, only in the last couple hundred years. It’s important not to therefore conclude that all dogs looked the same before that point, or anything close to that. If you traveled through Europe in 1600, you’d see an incredible number of distinct-looking dogs and probably hundreds of what we’d call breeds. But there was still communication between the breeds, sometimes slight and sometimes heavy.

When identification of breeds and registration of individual dogs, and eventual closing of studbooks, became fashionable in the 1800s, two things happened: Existing breeds were identified and codified and registered, and older breeds that were seen to be in danger of extinction or in come cases had already become extinct were re-created. So the Ibizan Hound and the Pharoah Hound were re-created, as were the Irish Wolfhound, the Elkhound, and many others. It seems unlikely that any of these were created from absolutely nothing; it’s a lot more likely (and certain in a few of them that I know the history of) that people passionate about a disappearing breed or landrace gathered the last remaining members of the population and then bred them to dogs that were as similar as possible to them in appearance and function.

At any rate, we have between 1850 and 1900 a whole ton of breeds being gathered, registered, and from then on bred only to each other.

By the 1930s there are a whole bunch of breeds that have their stud books closed (in other words, no more dogs are ever going to be let in).

Here’s where it gets sticky. In the 1920s and 30s distemper was killing dogs off in huge numbers. And then the advent of World War II and the extreme levels of starvation and hardship in Europe dealt a horrible blow to a large number of dog breeds.

Sussex Spaniels, for example, almost completely died off or were killed. The population dropped to just eight individuals. Shar Peis dropped to incredibly small numbers even more recently, as of the late 1970s. Lundehunds were reduced to six total. Cavalier King Charles Spaniels almost disappeared and were only saved by intense inbreeding in the 40s. Flatcoat Retrievers almost died out as well. The list of breeds that was reduced to only a bare handful of individuals is pretty long.

This is the definition of a genetic bottleneck. If, after the event or environmental pressure that kills off the vast majority of the population is over, no more founding individuals are added, then the entire breed (or, in the case of certain animals like the Asiatic lion, the entire species) becomes an expression of only those six or eight or whatever individuals.

A founding population that is that small is extremely risky. The likelihood that you can develop a genetically diverse and healthy ongoing population from six or eight or twelve already related members is small.

And, in fact, we do see that risk played out in these breeds. Sussex with their low fecundity and heart issues, Cavaliers with heart and brain problems, Lundehunds with mysterious digestive issues and early deaths, Flatcoats with cancers that effectively cut the breed’s lifespan in half.

So we have moved, in most breeds, from a situation of quite a bit of genetic information coming into and leaving the landraces to gene pools that are entirely based on a startlingly small number of dogs and where zero genetic material comes into or leaves the population.

We have effectively made each breed into an endangered species.

I can tell you, from far too many late nights up studing population ecology, that one of the things that is considered an immediate emergency in an endangered species with no available genetic material from the outside is keeping the heterozygosity of the population absolutely as high as it can be. It is considered so critical that the at-risk populations are constantly monitored for COI and also a direct measure of heterozygosity called MLH, and increasing heterozygosity is a key sign that the population has hope and may succeed and survive.

There are very few, if any, endangered species experts monitoring dog populations. That’s because the “dog” is not in any way endangered. Taken across the whole world it’s in great shape. But don’t fool yourself into thinking that individual breeds are not at critical danger of collapse and extinction. Both in absolute numbers and in size of gene pool, there are a strartling number of breeds that should be considered critically endangered. Even in the more numerous breeds, the representation of founding members is extremely poor and the average mating is equivalent to full-sibling.

If we don’t pay attention to this, we WILL pay the price, or the next generation of breeders will pay the price. I don’t know how much more plainly to put this–we’ve been breeding in a way that is very unlikely to be sustainable. And since we’re the ones making the decisions, it’s up to us to do our best to fix it.

High coefficient of inbreeding and disease

One of the things I’ve read recently by a breeder of a different breed (not Cardis) is that high COIs are absolutely essential to keep disease from spreading through a population.

Here’s how that line of reasoning works.

If I have a population of dogs that does not have Spotted Snarkle disease in their pedigrees, and I know that, I should keep breeding those dogs only within their own lines because breeding outside their lines will spread Spotted Snarkle disease.

The logical outcome of this is that the entire breed becomes very segregated, with some lines “known” for Spotted Snarkle disease, others for Awl Border disease, others for Maniac Mouth, and so on.

Increasing the genetic crossover, the exchange of genes between these lines, is therefore “dangerous” because then ALL the lines have all three diseases! Oh noes!

Here’s why that’s wrong.

Let’s create two populations.

Population A has a thousand dogs. 900 of them are free from Spotted Snarkle and 100 of them have pedigrees that are “known for” it. Of the 100, fifty are carriers of the disease and 25 actually have it. Every dog who actually has Spotted Snarkle is culled. The populations are rigidly separated.

Total population year 1: 1000

Incidence of Spotted Snarkle: 25

All affected dogs are culled, but the carrier population remains segregated. As a result,

Total population year 2 (assuming identical numbers of births and deaths): 1000

Incidence of Spotted Snarkle: 25

Population B has a thousand dogs. 900 are free from Spotted Snarkle and 100 are in pedigrees that are known for it. Same situation; fifty carriers and 25 affected dogs. The populations are allowed to freely interbreed. Again, all affecteds are culled.

The 50 carriers move out into a population of 925 other dogs. Therefore, the chance of a carrier meeting a carrier moves from 50% to around 5%.

Total population year 2: 1000 dogs.

Number with Spotted Snarkle: 2.5

So has Spotted Snarkle moved out into the whole population? Yes. Is the population still better off? YES. There are many fewer affected dogs; the problem has radically improved.

That’s how maximum genetic diversity works. Even though the incidence of deleterious genes is wider spread, the chance of identical deleterious genes meeting is MUCH lower than in an inbred population.

Proponents of high COIs and rigid segregation of lines forget some very key population truths.

One problem is that they’re assuming “success” based on only a tiny, tiny, TINY fraction of the number of diseases, genetic disorders, and weaknesses that actually exist. You can’t claim success if you’ve avoided Maniac Mouth when you have no idea what other 500 genes you’ve concentrated in your little insular population. High COI is not just associated with creating or avoiding the “sexy” diseases, the big flashy ones like Addison’s disease. It’s associated with an increased risk for low-grade, old-age, late-onset diseases. Mastitis. Kidney disease. Smaller litter sizes. Lower growth rates. Senility. Worms and other parasites. Vitamin deficiencies. Lower resistance to infections. The list is VERY long.

The second, and most sad, of the things that advocates of high COI ignore is the fact that breeders lie. They lie a LOT. I am thankful to have mentors in both Danes and Cardis who have been honest with me, but by the end of my time in Danes I could have written a book about breeders lying about pedigrees and about health. I hope things are different in Cardigans, but when when I was breeding my last Dane litter and was looking at pedigrees of stud dogs between a third and a half of them I was rejecting because I knew that at least one dog, or one health result, was false or at least very suspect. The tragedy of this, beyond the moral downfall of that breeder and the sadness and pain she causes to everyone who breeds based on a lie, is that you really can’t count on your “safe” pedigree. Not to any great extent.

If the goal of an entire breed is to keep COI low, the overall incidence of disease will be lower even if you were deceived in a pedigree or by a breeder (or that breeder’s breeder, ad nauseam). You will keep disease expression lower across the board, protecting future generations from the hundreds of diseases we can’t test for and that don’t make the headlines. And you will keep the population’s ability to respond to new, unknown threats at its highest level.

Another, more sentimental, benefit from my point of view is that spreading genes around keeps you concerned about the whole breed, and links you to and bonds you with many more breeders. It keeps the world small. You won’t be as willing to cut people down if you know they’re part of your pedigrees and you’ll be concerned with the eradication of disorders across the whole breed because your dogs participate across the whole breed.

Followup on A Football Field of Dogs

I wanted to address comments specifically in a post, because I know that not everyone reads comments. I am not including identities because I want to be VERY CLEAR that I am responding to issues, not people. If I use “you,” it’s the collective you and includes myself. I am making no statements about individuals or breeders with names.

I don’t know. Your posts along these lines always give me a lot to think about, and I certainly agree on some levels. For instance, the fact that every dog that is sound, mentally fit, & healthy has a place in a breeding program if so desired. And I think it takes a trained eye to be able to tell what faults will eventually cause unsoundness. But I am nonjudgmental when someone breeds a dog that isn’t “typey” or has some glaring faults, as long as the dog is sound. I could care less about a dog having the CH in front of their name. Some of the best Cardigans in history have come out of less than stellar parents, and I think that is a lesson worth studying.

Yes, I think it really is. I suspect, however, that those “best” Cardigans were not really anomalies when you look at the whole pedigree–at least in terms of soundness. You can have a bitch with a wonky topline and still breed her wisely if you know that the bad topline is not throughout her pedigree, or if you know that the stud dog you’re considering for her reliably corrects toplines. It ends up coming down to our two tests: Does it affect her life? Yes. Will it affect and hurt future generations? If you’re fairly sure the answer is no, it’s a good and ethical decision to breed her. 

To a certain extent I think this mindset follows down to “fault judging” versus finding the virtues of a dog. Many people fault judge and will decide that so-and-so shouldn’t be bred because of xx fault. I also think that what you should consider for the whelping box is vastly different than what you should consider for the show ring.

Yes, yes, and yes. If you have a dog who is likely to produce well (and by “well” I mean my oft-repeated phrasing about a happy, healthy, long, pain-free life), it is a solid contributor to the next generation even if it is not shown, or shown heavily. Breeders SHOULD show, and I think it can’t be a low priority. As political and unfair as it is, showing makes you put your money where your mouth is. It’s a peer review activity, where you “show” the products of your breeding program. But it’s not invalid to say that the products of your breeding program don’t have to be every single dog you’ve ever kept. Many good breeders keep back animals, especially bitches, that they feel will produce well but would not necessarily be the best choices to represent them in the ring. Now you have to be very careful–there’s a fine line between keeping a dog back because it is a solid producer and keeping an unsound or nasty dog out of the show ring but thinking up ways to justify breeding it. Keeping dogs out of the ring can’t be an invitation to kennel blindness. But as a strategy, yes, it’s valid.

And, of course, the issue with trying to do only the “valid” and relevant tests is that it’s not always clear which are which. I’ve heard many arguments that hips are not a relevant test in Cardis, but having lived with an OFA “mild” who DID show symptoms (while his OFA “moderate” dam did not), it’s hard for me to accept those arguments. Like above, I don’t get judgmental of those who choose to breed borderline hips when the dog seems very sound, as long as they do so thoughtfully. But I also can’t accept that idea that we should give up on trying to improve hips just because the tools are flawed. 

For me, the question on hips again comes down to whether it hurts the individual dog and whether it hurts the next generation. In some breeds that answer is completely obvious. In ours I really think it’s not. You saw it yourself in dogs that did not follow the “rule” of severity of dysplasia equalling pain level. And the answer to the second one, hurting the next generation, seems to be VERY poorly understood. I think we can say in Cardigans that an OFA-type view has moderate–not super, but moderate–value when it comes to analyzing the health of that dog as he or she stands there. It does not seem to have a lot of value when it comes to predicting how that dog will produce the next generation. The question is not whether the tools are imperfect–the question is whether they work AT ALL.

And I also feel that because of the AR activists, we need to be very careful about differentiating ourselves in as many ways as possible from mass-market breeders. That’s a tricky line to walk.

I want to address this one more fully below, because I think it is a VERY VERY BAD IDEA to be thinking along these lines. But since it’s repeated below, let me write about it once and not twice.

Comment 2:

“EVERY DOG WHO IS REMOVED FROM THE POPULATION HURTS THAT POPULATION.” – No, I won’t go there. And then you don’t go there either as you go on to talk about eliminating dogs with unsound structure or temperament from the gene pool. However, structurally sound dogs with cosmetic problems (such as mismarked or coat length) need not be eliminated in my opinion as well. And color? Well, most know how I feel on that issue.

Removing any animal from a population hurts that population. That’s how it works. It’s not my personal agenda; it’s ecological fact. No change is neutral, no removal without cost. The question is whether the removal benefits the population more than it hurts it. 

Nature performs this task with incredible efficiency and also with incredible conservation. She never unnecessarily removes an animal; she leaves the maximum number who can survive to reproduce. Barring a bottlenecking event like a flood or a volcanic eruption or something that kills a ton of animals in an unnatural way (i.e., in a way that doesn’t prove whether or not they would have survived in their environment), populations will stay at their maximum possible, breeding as widely as possible, maintaining the richest gene pool possible. The extent to which we screw with that process is the danger we put populations in. 

I am sure you know the term “no sacred cows.” We need to make sure we’re not falling into groupthink or conventional wisdom; we have to tell the truth even if nobody else is. For some reason, breeding has become something we view almost as a necessary evil, and it’s really better to not breed. That’s how the majority of “breeders” feel, or at least how they behave. I’ve heard people brag that they’ve been in a breed for 30 years and only bred four litters in that time, and they really do think that makes them a better breeder than someone who has been producing three or five litters a year over that span. 

That is, in the words of Trollope, a damnable lie. It is utterly contrary to the way you behave if you want to produce and maintain the healthiest possible population. We need to stop thinking that the best way to be good breeders is to not breed! We need to be breeding the largest possible number of dogs to the largest possible number of dogs or our gene pool will disappear. It should be “I neuter wisely,” even more than “I breed wisely.”

I strongly agree with [the above] statement about needing to be careful in today’s political AR climate. We need to be the guardians of our breeds and do our best to raise the bar, not lower it.

Now, see, here’s where I get the major heebies.

 

It is utterly vital to realize that the HSUS and the more generalized animal rights agenda has absolutely nothing to do with discovering who has the healthiest puppies. If you are laboring under the delusion that we have ANY kind of defense against their agendas because we do four health tests instead of one you are VERY VERY wrong.

And, if we’re honest, no matter how careful we are we can’t guarantee health. We can’t even guarantee that the puppy we’re selling is going to live a longer or better life, or have a better temperament, than the most raddled Malti-Poo from Petland.

If you’ve been reading my blog for any length of time you know that the dog we are all foolishly and totally in love with is Ginny, a genetic nightmare of a designer dog with a mouth that can’t even close properly. If I had a houseful of Ginnys I’d be in heaven. And the worst experience I’ve ever had with a dog I owned was a purebred with a pedigree as perfect as you could ever imagine.

That’s why I think it’s a really terrible idea to even pretend we can “promise” a product, or to say that our dogs are “better” than the worst reject from a puppy mill. Owners love their dogs, and what makes dogs “better” from their standpoint has nothing to do with the way we tend to define it. We can say they are sounder, we can try to educate them about conformation, we can talk about the ability to do a job. Ninety-nine percent of that will go in one ear and out the other. And then we’ll sell them a puppy, they’ll make a hundred dumb mistakes, they’ll create a fear-biting dog, and they will be convinced that we’ve ripped them off. Promising “better” is a dead end.

What we can do is WARRANTY health, stand behind our dogs; fix problems and replace puppies. But we should be doing that just because it’s the right thing to do, not because it will decrease litigation or liability. I’m afraid that ship has already sailed, and we’re going to be in court whether we like it or not and it will have nothing to do with whether we have healthy dogs.

The HSUS and its ilk make no differentiation between responsible and irresponsible breeding; their only goal is to end breeding altogether. The HSUS is asking for lemon reports to prove that unhealthy puppies come from breeders, that breeders produce unfit animals (and they do-I don’t care how many tests breeders do, if you have more than a couple of litters you will produce puppies that die young and even horrifically, sometimes due to genetics but usually due to the fact that they’re living things and some living things die young), that breeders create animals with bad temperaments or bad behavior, and that breeders treat their animals cruelly, and therefore you should never buy from a breeder.

If we breed with the HSUS’s threat as a motivator, or with some mythical definition of perfect health as the qualification for responsible breeding, we WILL fail. Don’t forget that we’re breeding dogs with a deformity, and even though we know that their quality of life is not hurt we’re automatically viewed as sickos who like deformed dogs. In other words, if we cater to that approach we will be neatly forced into not breeding at all.

Think about this carefully: If you were taken to court and asked to prove that the puppies you’re selling are “better” than a group of ten Aussie-doodles, could you do it? Because that’s what you’re saying you can do. You’re saying that because you health-test and somehow breed only “elite” dogs, raising the bar, you’ve differentiated yourself as “better.”

The prosecutor leans over and says, “So you’re saying that none of your dogs have ever shown any kind of reactivity or aggression toward other dogs? How about kids-is every single one of the dogs you have in your house completely trustworthy with children? Will they happily approach the elderly and disabled? Has any dog you’ve ever sold bitten any other animal or human? Has any dog you’ve ever bred been diagnosed with any genetic health problem? OK, well, plainly you’re in trouble there, so let’s go on to our expert witness. Dr. Wilson, can you show us a study that establishes that the defendant’s dogs are healthier than these mixed-breed dogs? OK, well, are the defendant’s dogs able to run normally? Oh, they have a deformity, yes. Why would anyone choose to breed dogs with a deformity? Well do they have any hip dysplasia? Oh, these deformed little dogs have hips that are twice as loose as the mixed breeds’ are? So… in other words, there is a documented history of bad temperaments, bad behaviors, bad health, and they’re congenitally deformed and damaged.”

You could NOT defend yourself. You would not have a single leg to stand on. “Raising the bar” is what we tell ourselves to make ourselves feel better, to tell ourselves that when someone comes knocking we’ll be safe. WRONG. This is one thing I CAN guarantee: If breeding Cardigans were put on trial according to the animal rights agenda, our breed would be shut down without hesitation.

So forget the animal rights organizations-they do not respect you, they do not make ANY differentiation between you and the guy who has a thousand dogs in rabbit cages full of filth. They will work just as hard to destroy you as they work to destroy him.

You breed for the BREED. For the DOGS. Not because somebody has a carrot or a stick. You find out the truth-about genetics, heterozygosity, soundness, movement, health, testing, all of it-and you breed to hand off the best and best-prepared population to the next generation of breeders.

Being a guardian of the breed needs to be something you take very seriously, and that means understanding and owning your decisions and working to understand the situation and the actions that will benefit the entire breed.
We’ve GOT to stop defining it as “doing more health testing than my neighbor does.” Even if that were a positive, it’s about five percent of what makes a healthy population. What about disease resistance, heterozygosity, population dynamics and geographical diversity, founding members, 200-year projections, growth rates, fecundity, fertility, lifespan, survival rates, biomechanical fitness, and the hundreds of other topics that we know about PINE TREES, for crying out loud, that every property manager has to know about his CLUB MOSS but we conveniently ignore in dogs because ooo, we’re such great breeders because we x-ray hips?

The long-term health of this precious, precious population, this endangered species, this cup of wine so close to spilling, think of it however you want. Bringing it from |here| to |there| demands every single bit of us; it demands tearing down the sacred cows and looking at the truth. It demands actions that are defensible scientifically and morally. It demands seeing the whole picture. That is the ONLY motivation; nothing else will stand the test of time and nothing else is fair to the dogs.

 

A football field of dogs (health testing…yeah, you know I’m going to stir this pot)

OK, sports fans, imagine standing on the 50-yard line and looking at an entire football field full of Cardigan Corgis. Tens of thousands of dogs, representing the entire national population of the breed. 

It is your job to get from this football field to the next field, the field ten years from now. You can use any of the thousands of dogs in this field, and success will be measured by whether you have a result pool (the ten-years-from-now field) that is at least as long-lived, healthy, athletic, sound, happy, and sane as your current pool. Bonus points will be awarded if you can improve on at least one of those axes without hurting any of the others. 

So how do you do it?  Stop for a minute and really plan it out before you read on.

My instinct is that most people thought to themselves “I should be as picky as possible, health-test everybody, prove that each dog is healthy, make sure that only the ones who are incredibly high-quality in terms of conformation and show success are allowed to breed. I should build the next ten thousand dogs from the most elite pool of this one.” That’s the conventional wisdom, the way “good breeders” do everything, right?

I want to suggest to you that a strategy like that will bring most breeds swiftly and inevitably to the grave.

Because here’s what is NOT being taught as conventional wisdom, and the entire breeding community needs to be smacked upside the head with it.

EVERY DOG WHO IS REMOVED FROM THE POPULATION HURTS THAT POPULATION.

That is FACT. It’s population ecology 101. A huge proponderance of all animal behavior is designed to create a population that is the most unrelated it can be–where the genes are as much UNlike each other as possible. That’s why bachelor males are kicked out of packs and herds; it’s a huge reason that animals try to get away from each other and form territories; it’s why we evolved different genders and all the millions of behaviors that govern breeding.

Maximum genetic variation is essential to a population that can withstand stress. If you lose genetic variation, you end up with substantially lower resistance to disease and you stand a good chance of concentrating deleterious genes. Loss of genetic variation is why we have such huge problems with cancers in Flatcoats, or epilepsy in Poodles, or Fanconi in Basenjis. 

One of the ways that breeders continually shoot themselves in the foot is by eliminating founding lines–if there were ten founding stud dogs of the breed, back in 1930, and they together produced ten thousand dogs that are living in 2008, are they evenly represented? Or are eight thousand of the puppies the descendants of just two of the founding dogs, two other dogs have disappeared entirely and their genetic material is now gone forever, and the other six have just a few hundred puppies in the entire world that are now living? 

Because of what is winning and what is in fashion in terms of hot kennels and top stud dogs, the entire world will rush to just a few dogs, like people running to the side of a sailing ship. This overweights the gene pool and it decreases the ability of the population to respond to threats and diseases. 

If, for example, we end up with a ton of heart disease, and Cadno and his descendants represent a pool of dogs with no heart disease, even if Cadno Cardigans have, for example, longer legs than we’d like we’ll find them extremely valuable. If that line was abandoned in 1970 because the Golden Arrow (or whatever) descendants were tearing up the green carpet and had such glorious short legs, we’re going to be stuck.

Why is the conventional wisdom so different from this? Well, one HUGE problem is that we have a collective guilty conscience, and we’ve bought a certain amount of conventional wisdom that comes from other breeds, and we’re under the thumb of a lot of groupthink that is actually coming from animal rights, so we have made it a virtue to remove every single dog from the gene pool that we possibly can. 

That’s where you get the “I know it isn’t perfect, but it’s SOMETHING” line that is used to justify neutering dogs based on everything from the DM test to the fluff test. In the back of that is a thought, however subconscious, that it’s good to neuter and iffy to breed, so the more stringent, even nonsensical, we make the requirements the more moral we are as a group of breeders.

We in Cardigans have a wonderful, healthy breed with very few issues. One of the best ways to KEEP it that way is to breed toward maximum genetic variation–in real-world terms, that means breeding as many individuals as possible to as many individuals as possible, spreading the genetic material as far as we can. Sharing the wealth. It’s not good to neuter but iffy to breed; it’s BAD to neuter and GOOD to breed. 

I know this is already making people itchy, but I challenge you to prove me wrong. It’s supported in every population study I have ever read–loss of breeding animals is a bad thing.

So the question is NOT how to choose the best from this football field. The question is how to REMOVE the weakest.

After all, that’s what happens in nature; it’s how all living things evolved. The term “survival of the fittest” is a little misleading; evolutionary pressure doesn’t choose which animals survive. It’s “death of the weakest.” Nature kills those that are not strong, leaving behind every single individual that WAS strong enough to make it. Those are primed to breed as widely as is practical for the population, keeping the population at its maximum level of genetic richness.

So how does this apply to our field of dogs? It’s our job to wisely remove the weakest. It’s not plucking the very “best” out and elevating them–if you do, in just a few generations your population will lose the great majority of its genetic material. It’s deciding who “dies” in the population, who does not get to reproduce. Thankfully we can do it by sterilizing or separating and we don’t have to actually kill them, but the effect on the population is the same.

Remember, every loss to the population is a negative. It is NOT a neutral decision, ever. That means that the benefit to the population of removing that dog must outweigh the negative effects of removing him or her. If it does not, you are hurting the population and sending your breed to the pit.

So, for a moment, forget anything to do with health testing and let’s just try to choose which dogs to “kill.” It’s honestly better to think of it as killing than neutering, because it correctly communicates the gravity of the decision. It is a great and terrible responsibility to remove dogs from a breeding population and you SHOULD do it with no little fear and trembling.

If you’re going to kill a dog, you need to make sure that you’re doing so based on two criteria: The “fault” needs to hurt that individual dog AND that fault needs to be reliably communicated to the next generation. If the faulty dog won’t pass on that fault, he or she should not be removed. 

I would say that the first dogs we remove are those that have broad issues of unsoundness. We are absolutely sure that major issues with body shape a) hurt that individual dog, and b) are reliably communicated to the next generation and therefore hurt that generation.

So if a dog cannot run freely, walk without pain, eat its food, and live to a normal old age, its genes should be killed off. Very unsound bites, fronts that end up painful and arthritic, swaybacks, terribly unsound movement, etc. I would also add congenital shyness to this list; a dog who is born so shy that it cannot be happy in normal society would never survive if it had to live in a community of dogs or run down game. Ditto with ingrained reactivity and willingness to ignore the bite-humans taboo. Environment is ALWAYS king of behavior, but you know what I’m talking about here. That bitch who bites judges and you know that three of her puppies also bite people? Don’t be blaming the owners; look in the mirror for that one. Perpetuating what is in effect a mental illness is bad for the population.

If we’ve killed off the unsound dogs–please note that I did NOT say the “untypey” dogs–we should be left with a group of dogs that is basically able to make a next generation that will succeed. They all have strong, sound bodies and would be considered at least average to good in conformation and movement. We now start applying the kill criteria that are much, much more slippery. These are health testing and selecting for “type.”

I want to talk about type first because I am terribly worried about the fact that so many dogs are “killed” for totally superficial reasons. Jon Kimes got here first, but maybe he’ll allow me to expand on this.

The proper Cardigan head is called proper because it is a SOUND head, a healthy, long-living head that allows the dog to do its job. So if a dog has a tremendously clunky head, a Lab head on a Cardi body, that’s not a superficial fault. It goes to soundness and, while it is nowhere near as unsound as a very forward front or a straight shoulder or cowhocks, there are good valid reasons to try to remove it from our healthy population.

Similarly, the extremely foreshortened radius and ulna in the front legs are sound. Thick, short dwarfed bones are actually healthier than longer, thinner dwarfed bones. So, as with heads, a dog up on too much leg is quite possibly still sound enough to  breed, but it’s a genuine fault. 

Markings, on the other hand, or coatedness. Let’s examine them according to our two criteria. Does having white around one eye hurt the individual dog? It MAY, to a very marginal extent, if the dog does not have good pigment otherwise. But clearly white in and of itself doesn’t hurt working dogs; every livestock guard dog, the big hounds, the big sporting dogs–they all have a ton of white around the eyes. So it certainly isn’t a slam dunk. Now #2: Does having white around one eye hurt the next generation? Since white-headed dogs can clearly produce lots of color when bred wisely, the answer to that is no, unless you breed stupidly. 

The “off” colors are even more superficial. They have no detriment to the individual dog and are not passed along unless, again, you don’t understand color genetics or how to breed the standard colors.

Coat is similar. A long coat is NOT disadvantageous automatically–the long coat we call “poor” is no longer, softer, or more open than many breeds with distinguished working records have. And coated dogs can clearly produce dogs with standard coats.

Both of those–markings and coats–are part of the “game” of showing dogs. If you want to show dogs, you follow some rules just because they’re part of the game. So Labs can’t have any white, but Tollers need it. Rotties can’t, but Berners must. All of them are just part of the stuff we accept when we decide to show. They are NOT reasons, and I’d challenge you to prove me wrong, to kill off dogs from the population. If a dog is sound, healthy, built to live a long and good life, has working conformation and a sane temperament, it’s really not very defensible to kill it when it can clearly produce worthy contributors to the next generations. 

So back we go to our football field. We’ve taken out some ugly heads and the really bad tails (we’ve probably left in the tails with hooks, but we’ve taken out the tails that go way up over the back because they’re a sign that the dog can’t move the way a working dog should). We’ve taken out long, weak legs, and a few other un-typey dogs. 

It is only NOW that we’re at the place where many breeders begin–at health testing. That is VERY deliberate. The dogs who were going to hurt the population because they were so unsound were “killed” long before we’d even consider winnowing them via health testing. 

And here’s where I make my big pot-stirring statement:

I think that many people get the whole motivation behind health testing completely backwards. They feel that they’re doing it to “prove” that their  bitch or their dog “is healthy.”

If you go that route, then there is no end of testing that you can and should do. In fact, there is no end to the testing you MUST do. After all, just because I know that his hips are healthy doesn’t mean his heart is healthy, and doesn’t mean that he doesn’t have one or more factors for von Willebrands, and doesn’t mean that she doesn’t have autoimmune thyroiditis, doesn’t mean she doesn’t have a rare storage disease, doesn’t mean he has healthy patellas, doesn’t mean that she doesn’t have elbow dysplasia, and on and on and on it goes. If proving that a dog is healthy is what breeders are supposed to do, then you need to line up a hundred or more tests and you can’t be making any excuses.

Others will say that they are doing it to prove that their dog or bitch is healthy, but only until and up to a certain dollar amount. Again, bad idea. That encourages you to do a lot of cheap tests rather than better, more expensive tests; it also sets up a false expectation of, among other things, puppy prices (at least to a certain extent, health testing is passed along to the puppy buyer; that’s just reality, so if we have to get expensive testing done we should just do it and charge more for puppies rather than whining that we can’t do it because we don’t make it back in the puppy sales).

The only reason that makes ANY sense is that health testing is to remove the right individuals from the breeding population. It MUST be a removal process is because that keeps the focus on what actually WORKS. If a health test protects the integrity of the population by reliably removing dogs who will hurt that population, it is worth it and not only worth it but mandatory. No matter how much it costs or what kind of trouble it is or what we have to pass on to puppy buyers.

So, for example, because Danes have cardiomyopathy, a disease that typically begins to show between the ages of two and four, and because cardiomyopathy is reliably genetic but there’s no DNA test yet, I did serial echocardiograms on my stud dog, to the tune of $400 each time, and his prospective mates got echoes too. Because I was working with a line that had clear GENETIC hypothyroidism (where you can trace it down the pedigree very reliably), everybody got full thyroid panels; because hips are at least somewhat genetic in Danes everybody got PennHIP tested. I routinely had to put over a thousand dollars into each breeding partner before the breeding. It hurt a huge amount, financially, but tough for me. If I wanted an “in” to that breed, that was what I was buying. If I wanted to get into Cavalier King Charles Spaniels, I would feel that I had to do an MRI on each breeding dog to check for syringomyelia. No matter what it cost, it would be absolutely necessary.

Now here’s the part that people get mad about: the opposite also applies. If a test does not improve the population more than it hurts it, then we shouldn’t be doing it, or if we do it we shouldn’t be removing dogs based on its results, because we’re skewing the population–running to the side of the sailboat–for no reason; we are hurting our future generations.

Because Cardigans are so healthy and don’t have the huge issues that other breeds have, we’re in that situation with the majority of our health tests. The stuff we see as health problems are only somewhat genetic and the tests are very unreliable, but the tests exist and they will multiply. I’d put money on the fact that in 15 years we’ll be able to test for twenty or thirty disorders that are only weakly connected to the gene test. 

I believe in testing, believe strongly in it, but for those disorders where there is no clear link between the result and the life of the dog (DM is a great example of this) I think we need to be ignoring the results of the test for a long time, breeding as usual, until we can say “Yes, we now know that dogs with positive DM tests live, on average, four years less than dogs with a negative DM test.” That’s a real reason to start killing those dogs off. If, as I suspect, the line is more like “Dogs with positive DM tests live, on average, three months less than dogs with a negative test,” then we are REALLY foolish to be chucking thousands of dogs from the gene pool–if we “kill” carriers and at-risk dogs, probably more than half the population of the breed–for that. 

So if you are looking at the available battery of tests for your dog, please first be serious about whether the dog is sound and can live a normal life, is safe and sane and happy. If the answer is yes, then by all means do the testing, gather information. But don’t be a slave to the tests if they do not clearly pass the two criteria: They must affect your individual dog and they must reliably change the next generation for the better. Do whatever research you have to do to make sure you really know, genuinely UNDERSTAND, the answers to those criteria. Because we really do, as a community of breeders, have to fill that ten-years-from-now field. And every single breeding decision changes that result. We should be doing so very, very thoughtfully.

Being a good breeder

When I started out in Great Danes, I was thoroughly convinced that I had the answers. I don’t know why on earth I thought I had the right to be that arrogant, and thankfully I didn’t talk about it too loudly and make myself look really idiotic, but I was SURE that somehow I could, by following all the recipes I had read about and researched, actually get a better result than anyone else. I really did think that the problems in the breed were because others had become complacent. I “knew” that if I never bred without an OFA number, never failed to test thyroids, did a yearly echocardiogram on all my dogs, that I was a good breeder. Not just a good breeder, a BETTER breeder than the other good breeders.

I bred my first litter with the feeling that nothing could possibly go wrong; my ducks were all in a row; my co-owner had signed off on the stud dog; it was going to be a litter for the ages.

I

Was

An

Idiot.

Looking back on it I just cringe. Nothing terrible happened, although seven hours into the delivery, covered in blood and placenta and trying unsuccessfully to bring back a dead puppy, I had a BADLY needed moment of “OH LORD WHAT HAVE I DONE?” Those puppies did well, I placed most of them well and some of them spectacularly and a couple of them horribly and got them back and, much wiser, rehomed them better.

But I can barely stand to look back on it just because I can remember what it felt like, that sense of total justification and assurance that I was DA BOMB because I was so terribly serious about following the rules. I want to go back there and smack myself upside the head and knock some of that hubris out.

I was a fault judger, of breeders.

Fault judging is that thing that everyone says separates the new breeder from the old, and when you can stop fault judging you are finally on the road to somewhere. So when you look at a dog you don’t say “short neck,” you say “gorgeous topline, fluid mover” while also realizing that yes, the dog is lacking in layback and could use a longer neck. When you fault judge that one fault is the ONLY thing you see, and it disqualifies the dog for you, often completely nonsensically.

And oh, heck yes I did that. I have a catalog from a Nationals, my first Nationals, where beside the name of a dog who was far and away the strongest and soundest and most short-coupled stallion of a dog in his class, I have three words: “not enough stop.” I really thought that I was all that and a bag of chips because I could see that fault, and at that moment you could not have paid me to breed to that dog, because look at that ugly ugly slide down from the ears to the nose.

Like I said, idiot.

We make the same mistake when we fault-judge breeders. Ignore the fifty champions and the couch full of 14-year-old dogs at their house, because they don’t OFA. Dismiss their multiple ROMs and ranked stud dogs because so and so got a dog with bad elbows from them, and I’ll never make bad elbows, no sir I won’t. Because I follow the RULES, man.

I am not far enough removed from that moment when I was up to my elbows in goo and realized exactly how little I knew, and I never will be, to say that now I am some kind of guru. If anything, the longer I stare at dogs and the more I know about them the more I realize that they are perpetually slightly astonished at how stupid we are. I’ve read that “Please, God, make me the person my dog thinks I am” bumper sticker and it makes me giggle, because my dog thinks I’m a clumsy ox with a speech impediment, a useless nose, and a magic pocket that grows hot dogs. She hopes that if she plays with me enough and speaks loudly and slowly enough I’ll improve, but she hasn’t seen a lot of hope for me.

So I am hardly “there” yet. But I maybe have moved far enough from that moment to be able to be slightly more accurate, at least for myself, about what makes a good breeder.

The rule: Good breeders use health testing organizations (like OFA).

The truth: (and if you’ve been reading my blog over the last week you can skip this part, because I’ve devoted thousands of words to it already):

It’s absolutely correct that good breeders have extremely high standards and do every breeding carefully.

However, I would object to the idea that there’s an epidemic of Cardigan breeders out there ignoring the facts and breeding stupidly.

We DO NOT HAVE a method for making perfection if by “perfection” you mean dogs with hips that look like sighthound hips (which is not really my version of “perfection,” but I’ll grant the point). The tools, honestly, suck. Relying on OFA ratings produces a result that is not statistically greater than zero. In other words, it works no better to produce healthy puppies than not using it at all. So why on earth does it make you a bad breeder to realize that? PennHIP is a pretty good method for the breeds upon which it is based (Lab, Golden, Rottie, Shepherd) but a careful look at the statistics provides no such assurance for Cardigans.

Insisting that good breeders use OFA or PennHIP to improve quality of life is like giving someone a fork and telling them to build a house–the tool may make them feel like they are doing something, but the end result is not going to satisfy the requirements.

The response often given is “Well, it’s SOMETHING. So we should use it anyway.” I’d disagree. You don’t make decisions that have the potential to dramatically shift the gene pool without good reason. As I said earlier, the numbers should serve us; we don’t serve the numbers. Our goal is supposed to be to produce healthy, happy, pain-free dogs, not a carat on a number line. If the carat doesn’t reliably equal those healthy pain-free dogs, it’s foolish to elevate it above any other piece of information we have about the dogs in question.

The rule: Good breeders never do questionable breedings.

This generally means, or is meant to imply, a breeding that doesn’t follow rule 1 above. I’ve rarely heard anyone say that about, say, breeding to a stud dog who is cowhocked. It’s code for “she bred without an OFA number” or “He knowingly bred a dog with one dysplastic hip.”

That was one of my huge hangups; I developed big statements about certain breeders in that breed or in others, statements like “Well, I always thought he was a great breeder, but now I know THIS juicy fact and my opinion of him is ruined!” Again, I was an idiot. And I was also nasty and mean-spirited and I am ashamed I ever built those air castles of meanness.

The truth: Listen–every single breeding we ever contemplate, every single breeding we do, no matter what letters or numbers are before after, or under their names, is “questionable.” We’re producing living things who don’t tend to follow neat little patterns. And that’s not rationalization; that’s fact. Breeders who break the “rules” generally know about fifteen thousand percent more about the breed than I do, and they know far better than I do how much you have to give up and let go and sometimes breaking that rule is the best possible thing you can do for your breeding program.

So let’s put aside health certifying agencies for a moment. Let’s look at breeding choices apart from that.

The rule: Good breeders only have a small number of litters, maybe one or two a year at MOST. (This often becomes “I’ve been in the breed for 28 years and only bred six litters in that time!” or similar)

The truth:

This is a really bad one, a really insidious one, calling people bad breeders if they don’t apologize for every litter (which is what I sometimes think is being implied–that good breeders only breed if you have no other choice, or only breed with extreme reluctance).

The breeders who have done the greatest good for this breed, from the latter half of the nineteenth century to now, didn’t fall into the belief that fewer breedings are somehow automatically better. That’s an extremely effective lie that has permeated even the good breeder community. The fact is, good breeders DO test breedings, do “experimental” breedings, see what happens when Joe is bred to Mary because maybe Joe would be good for Marsha as well. Many of them would say that you don’t even know what your stud dog does or doesn’t do until you’ve seen five or six litters grow up, so it’s only after those litters that they start really using him with wisdom and intent. Fewer breedings is not better; those that breed a litter only once every five years are not the ones that change the breed. They may go along with the breed changes, make a few really nice dogs, but they’re not a force for improvement.

The rule: good breeders do not compromise.

Unlike “questionable,” which the secret decoder ring translates to “She didn’t submit those x-rays to OFA,” “compromise” means a breeding that the speaker personally thought was a bad idea. Generally one or more of the pair was ugly, or had some screaming fault, or in some other way the breeder did something that the speaker is super-super-sure she’d never do. “Oh yes… I think it’s so sad the way she had to compromise to breed that bitch.”

The truth:

Good breeders know that there is NO SUCH THING as a breeding that does not force a compromise. You will NEVER find a dog for your bitch that is absolutely perfect; you will never ever feel that there is not something you’re giving up, or something you’re really hoping doesn’t show up in the puppies. There will be some breedings that you are absolutely thrilled about and some that you aren’t (for example, breeding to a male that you think is ugly because he’s got a beautiful sister), and good breeders know that sometimes the very best puppies come from the latter and sometimes those oh-so-fabulous breedings are a genetic disaster.

If you DON’T go into a breeding agonizing about what might show up if the thing that makes the male less than perfect and the thing that makes the bitch less than perfect happen to meet up in the puppies, you have no business breeding. Breeding IS agonizing. If you don’t lie awake at night worrying about puppy owners standing in your driveway with pitchforks and torches, you shouldn’t be breeding. Your conscience should be screaming at each and every breeding, because that’s what keeps you from getting careless and foolish.

The rule: Good breeders make happy puppy buyers.

This usually gets whispered as “I heard that Bigshot So and So got a puppy back because it was a fear biter and it had to be put down!” or “I heard that Top Winning So and So sold a puppy and it was CRAP!”

The truth: The number of times I have seen the downfall of a puppy and can absolutely say that it was the breeder’s fault is very, very, VERY small.

Puppy owners are like any other buyers of any other products, except that most people who buy a fridge understand how to use a fridge, whereas the majority of people who own dogs are well-meaning but completely clueless.

No matter how well you screen, there will be a few complete dipwads who buy your dog and then blame you for everything they do wrong, and no matter what you do they will try to bad-mouth you every chance they get.

And, if you breed long enough, there will be at least a couple of evil, abusive dipwads you’ve got to get dogs back from, and you’ll have to threaten lawsuits and police involvement, and those owners don’t generally take out an ad in Variety talking about how great you are.

What makes a good breeder? What’s the REAL rule?

What makes you a good breeder is that you clean up your own messes. You own your decisions and the buck stops with you. You DO x-ray hips and you DO look at eyes and you DO keep track of the longevity of your dogs so you can make truth statements about your breeding program.

If you produce a puppy that is a disaster of one kind or another, you replace it. If one of your puppies needs a home, you take it. If an owner is dissatisfied, you make it right. If an owner is a danger to the dog, you move mountains to get your dog back.

You answer every phone call and you stay up until five a.m. talking an owner through the death of a young, healthy dog, a death that you will then blame yourself for and cry over for weeks. You listen patiently as an owner says that their vet says that it’s your fault that Buffy is now crippled. If it’s true, you don’t argue. You stand ready to accept that blame. Because I don’t care if you do a full-body MRI screening and only breed to dogs with genitals of solid gold, you WILL produce a crap puppy every now and then, and probably far more often than that.

One more thing: the Cardigan CAN AND WILL die out as a breed if we’re not careful. It is teetering on the brink of complete collapse in every country but this one. Three hundred puppies a year is what the UK has determined is a self-sustaining number, and I would add that those three hundred should be largely unrelated to one another for true health of the population. That means without the US population, there are not enough unrelated well-bred individuals in all the other countries put together to avoid the extinction of the breed or (virtually as bad) such a severe genetic bottleneck that the health and soundness of the breed would be destroyed. Because the US population exists we’re in acceptable shape, but not good shape. We’re definitely not in a position where we can make the gene pool even smaller. We should not be looking for excuses not to breed; we should be looking to use the largest number of dogs that are as unrelated as possible as wisely as possible.

What IS true is that the Cardigan is not a breed in a health crisis. “Despite” our very best breeders not OFAing–go to the ofa site and look at the names that are NOT there–we have a breed that lives a ridiculously long, happy, healthy life. Before we start saying that people shouldn’t be breeding, we need to have a good reason to stop them from doing so–we need to be able to say “Your breeding practices are proven to make unhealthy puppies that don’t live a long time.” Until and unless we can do that, we have no basis on which to criticize.

Charlie of Finnshavn Cardigans, who is an amazing owner, breeder, and repository of information as well as a true gentleman, calls this tack of criticism “the faceless theys.” As in “If we don’t make this rule, THEY will destroy the breed.” Evidently these trollish THEYS spring up and take over if we don’t beat them back with a sword made of PennHIP scores over the fiftieth percentile.

All those horrible THEYS makes it really quite incredible that you ended up with the super-nice dog that’s at your feet right now. After all, that dog is the result of decades and centuries of breeders who didn’t follow the rules as well as you’re going to, you there with the sword.

I was that person, standing on someone else’s shoulders, holding a puppy they and the fifty people whose shoulders they were standing on had handed to me, beating them around the head and criticizing them for not doing as good a job as I was sure I was going to do. You can understand why I now cringe and hate to think about it.

Like I said, I am nowhere near where I want or need to be to even BEGIN to be a good Cardigan breeder. But the one huge thing that has changed is that I am trying to know, and tell, the truth. Not the rules, the truth. Even though the truth is a lot harder and a lot more dangerous and allows in a lot more mistakes and even a few breeders who aren’t in it for the good of the breed. Because I think that the only way I can do this with a clear conscience is to make sure that I am not acting out of fear, or guilt, or envy, or the desire to make myself better by labeling others as worse. The dogs, and my friends, and my puppy buyers, deserve the very best of me. The part of me that acts, to quote a very wise book, “reverently, soberly, discreetly,” and above all with love.