Tag Archives | Klemola

Searching for another founder

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An elderly Finnhorse, descendent of the stallion Eversti

In the previous post about founders, I talked about the search for the origins of the silver dilution. I wanted to present another case study that highlights some of the challenges in tracing color lines, this time with classic splash white (SW1). We tend to talk about this form of splash white as being incompletely dominant, with the heterozygous horses tending to show very little white and the homozygous horses having the more distinctive “classic” form of the pattern. When the pattern was first described in 1931, however, it was called Recessive Pied. This was in contrast with Dominant Pied, which we now call tobiano. Later the author, Valto Klemola, proposed changing Dominant and Recessive Pied to “Piebald” and “Splashed White” respectively.

By visiting the Splashed White Project page, and clicking on the button to display heterozygous SW1 horses, it is easy to see why Klemola would have considered the pattern recessive; the horses in that group do not look like pintos. In fact, there is a growing understanding that in breeds with minimal white markings, horses can carry the SW1 mutation without showing unusual – or sometimes any – white or blue eyes. With that reality in mind, classic splash behaves enough like a recessive that it illustrates how the search for a founder is different when the color is not visible in the carriers.

Klemola noted how the pattern sometimes surprised breeders.

In the native breeds of Northern Europe what at first glance looks like a piebald foal may be produced from quite normally-colored parents. This surprising phenomenon has given rise to many fantastic explanations among the breeders…

That observation comes from the second paper, published in the Journal of Heredity in 1933. The earlier paper, published in Zeitschrift für Züchtung, included pictures of Danish and Swedish horses with the classic splash pattern. But the main focus was on the Finnhorse stallion Eversti, who was known for producing both blue eyes and occasional pinto patterns. Although he was only mentioned in passing in the 1933 paper, the earlier one provided extensive details about both his ancestors and his descendants, all of which point to the likelihood that he was heterozygous for SW1. Eversti was a black horse with white markings, but his paternal great-granddam was a blue-eyed pinto (“glasäugig und bunt”). When dealing with historical records, SW1 horses are rarely identified as pintos unless they are homozygous. Had the color been unique to Finnhorses, she could not have been the founder. One truth about recessive colors is that the founder would not actually be the new color. Remember each specific mutation is a one-time event. To display a recessive trait, an animal must be homozygous – it must have two of the same mutation. So even if his pinto great-grandmother had been the first known splashed white, she could have been ruled out as the actual founder. The real founder appears somewhere on both sides of her pedigree, and he or she was probably quite unremarkable. The original mutation could spread pretty far before two descendants were crossed, and the result was a very obvious pinto.

Klemola knew that splashed white was not unique to the Finnhorse, so that unnamed pinto mare was never considered as the founder. As he noted, the same pattern occurred in other Northern European breeds, so the original mutation happened before those breeding groups separated. In more recent times, the same type of pattern was observed in breeds as diverse as the Pasos of Puerto Rico and the Marwaris of India. When a pattern is found across a broad range of breeds and regions, that usually means that the mutation is old. Like other old mutations – silver, tobiano, leopard complex – it is unlikely that much will ever been known about the horse that carried that first SW1 mutation.

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Oregon Ice On Fire, a descendant of the Morgan mare Royal-Glo, is heterozygous for SW1

The most that may be possible is to identify some of the sources for the pattern within some of the modern breeds. Now that a test is available for the pattern, carriers can be identified even when they have not produced the more obvious classic pattern. With enough testing information, lines that carry SW1 can be identified. This process has already started in the Morgan breed, where Royal-Glo and Lady In Lace, and ultimately their ancestor Rhythm Lovely Lady, have been named as likely sources. It may be possible from there to connect those horses to some of the early American lines that predated the stud books, since some of those were noted for producing blue eyes. This is perhaps as far back as it may be possible to go with the history of the pattern in America. 

In the case of the Finnhorse, Eversti proved to be an influential stallion. His great-great-grandson, Murto, is one of the four male lines in the breed. Both Murto and his son, Eri-Aaroni, were chestnuts with flashy white markings. Intiaani, the first Finnhorse that tested positive for one copy of SW1, carried 21 lines to Murto through the female side of her pedigree alone. Eleven of those were though Eri-Aaroni. And yet that influence is probably the best argument against the color coming from Murto. It is difficult to find a modern Finnhorse without multiple lines to Murto – and by extension, to Eversti. If Murto carried SW1, breeders should have started to see classic splash offspring among his linebred descendants. Yet until the results came back from Intiaani’s test, it was widely assumed that the pattern could no longer be found in Finnhorses. That would suggest that the source for the mutation came through one of the less common lines to Eversti, or even one of the other lines that go back to Eversti’s sire Jalo (grandson of the blue-eyed pinto mare). The flashy white on Murto and Eri-Aaroni may well be unrelated to SW1.

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Eri-Aaroni, a son of Murto and one of the most influential stallions in the Finnhorse breed

There are quite a few horses that have either tested positive for a single copy of SW1, or that have produced the classic pattern, that look like Eri-Aaroni (pictured above). A fair number of the known splash white producers in the Welsh Mountain Pony have similar markings. There are also quite a few breeds where these types of markings are common, like the Arabian, yet SW1 is not believed to be present. That complicates the search for sources in breeds where a variety of white-producing mutations are found – which is actually the case in most breeds. Even the presence of blue eyes in a line may not indicate the presence of SW1 in breeds where it is known to be, because some blue-eyed horses have been testing negative for the (currently) known forms of splash white.

With luck those other blue-eyed horses will prove to have a newer mutation. That is the case with the other four formally identified forms of splashed white. The founder of the most common of those, SW2, is a known individual. While she was not named outright in the original paper, the information given matched that of the 1987 Quarter Horse mare, Katie Gun, dam of the famed reining horse, Gunner. The suspected founders of the other three were all born in the last two decades, which would suggest that mutations of this type are more common than originally thought.

And this is probably a good place to jump over to the subject of my friend’s albino dog, since that involves recessive genes, separate mutations producing similar colors, and the search for founders. I’ll start that topic in the next day or two.

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How I learned to stop worrying and love incorrect color terminology

NoBuckskin

Okay, maybe not love – but certainly stop worrying about it.

Like a lot of people who find horse color fascinating, I once spent no small amount of time online, spreading the Gospel of Proper Color Terminology. Surely if I just presented the facts in a convincing manner, I could save the world from people who were convinced they had palomino Arabians!

Fortunately for my own sanity, my career as a parent – begun just a few years after large numbers of horse people discovered the internet – got in the way of my missionary zeal. Toddlers do not recognize the need for uninterrupted bathroom breaks, never mind enough time to compose an extended explanation about why you really should not call your double-diluted cream an albino. Children also raised my threshold for the type of thing that required immediate action. I could live with someone on the internet being wrong; it was not like they had just painted on my bathroom walls with chocolate pudding.

Looking back, though, I see that my enforced absence from online discussions had unexpected benefits. For a researcher there is a significant downside to spending a lot of energy “correcting” wrong information. If you spend too much time telling people that some common misperception is wrong, you run the risk of having that response become automatic.  It makes it a lot harder to reassess your position, because it is a rare person that can argue a position for a long time without getting their ego involved in being proven right. From there, it is easy to overstate your case. “Your flaxen chestnut Arabian is not a palomino” becomes “there have never been palomino Arabians”, which then becomes “Arabians do not carry any dilution genes.”  The first is – or at least to date has been – true. The next statement is actually open for debate, and the last one is incorrect. (See also, here. Similarly diluted Morgans can be found here.)

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And intriguing painting of the early Twentieth Century Turkish Arabian, Übeyyan. How accurate was this portrait? And what color was he?

The other downside to spending a lot of time correcting errors is that if you automatically dismiss something, it is really easy to overlook important information. Even when people are wrong, they may still hold a clue, a piece of the puzzle you are trying to assemble.

I sometimes get asked why I spend so much time with older documents when so much has changed in our understanding of coat color genetics. Why, for instance, spend time translating Valto Klemola’s 1931 paper on “Recessive Pied” when there are papers written just this year and last on what we now call Splashed White? Surely the new information replaces Klemola’s theory about recessive spotting in horses.

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I am sure my husband also wonders why I need books about horse color published in 1912. After twenty years of losing more and more shelf space to them, he has given up asking.

But the fact is that Klemola – and many of the other earlier authors – were not entirely wrong. They were almost always working from a partial picture, but often the piece that they were seeing was not incorrect. It was simply incomplete.  Read with an understanding of the larger picture, what these older researchers have to say can still provide valuable information. The same is true for owners and breeders who may not have the same grounding in the latest scientific theories. They still have the potential to be valuable observers. It is worth being open to what they have to say, without being excessively concerned about the “correctness” of how it is said.

At the moment terminology – particularly the terms we use when talking about white patterns – are in a state of transition. We are struggling with words that do not completely fit our present understanding. I hope to tackle that in more detail in a future post. It is worth remembering, however, that the real reason for adopting a consistent set of terms is so that we may all communicate more clearly with one another. It may take a little more effort, and perhaps a few more words (and patience) than it once did, but that is ultimately the goal.

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Classic splash

I have gotten a number of messages lately that have made me realize that it might be helpful to clarify the term Classic Splash. I began using that term in place of the commonly used “obvious splash” when I realized that if there were differing views about what was and was not splash, the word ‘obvious’ was probably not particularly instructive. If there is one thing modern testing is teaching those of us who love white patterns, it is that very little is truly obvious! I still needed a way to indicate that I was talking about something very specific, so I opted for the word classic because what I had in mind was very much in line with the pattern as it was described in the original paper by Klemola.

That was not my first exposure to the splashed white pattern, though. Credit for that goes to the pony in the picture at the top of this post. Sometime in the early 1980s, his picture was used to illustrate the entry for the Pinto Horse Association in Western Horsemen’s annual all-breed issue. I was fascinated, because I could not figure out which pattern he had. That particular photo was taken of his other side, and was angled such that it appeared that the dark area of his coat did not start until well after his poll, while much of his neck and body were colored. To someone used to looking at ordinary tobianos and overos, he just looked wrong. Very appealing, but very much like an artist who did not know what they were doing made up his pattern. Needless to say, he went into my artist reference files.

I didn’t know what he was until a few years later, when I acquired a copy of Dr. Sponenberg’s book, Horse Color.  He had a small paragraph about Splashed White, and photos of a Welsh Pony foal with the same kind of pattern. The Klemola paper was included in the bibliography, and that provided still more information and a few more pictures. From that point on, I began to collect images and background information on anything with a similar pattern. Like most artists, I have always collected large quantities of reference images, but my interest in horse color – and patterns in particular – had become a hobby unto itself. All the white patterns interested me, but none so much as the elusive Splashed White.

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My early reference files are filled with advertisements torn or xeroxed from magazines. This gave me a better idea of the range of expression to the pattern, but still the information was limited. I knew that other patterns, like sabino, could occur in such a minimal fashion that the average person did not realize the horse was a pinto until it produced something more extensively marked. If that was the case with the Splashed Whites, it was often a rather big jump from minimal parents to really loud offspring. With sabinos, I could often pinpoint where the color was likely coming from in the pedigree. With splashes, it was often not especially clear. Here is Gambling Man, one of the better-known of the Splashed White Paint Horses from the early 1990s.

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Those are his parents in the inset clipping. So did his color come from his sire, with his blaze and four white feet? Or maybe his dam, with her irregular face marking that covered her nose? My files were full of horses like this, where it was impossible to narrow the source of the color down even to one side of the pedigree. Of course, this was also before the use of the internet was widespread, so there were no online databases or easy access to images, so often the background information was incomplete.

It was actually the format that I used to organize my files that led to the realization that Splashed White was probably incompletely dominant. I always entered horses into my notebooks with as much pedigree information as I could find, because I was usually looking for the color line. That is, I wanted to know where the color came from so that I could more easily rule related horses in (or out) for a given pattern. When color printing became feasible, I began color coding the names to note whether or not the horses in the pedigree were known to have a color or pattern, were suspected of it, or could be ruled out. What made Splashes so maddening was that I couldn’t even rule out one side of the pedigree on any of the entries. It took a while, but eventually I realized that wasn’t the problem; that was the answer. I couldn’t rule either side out because it came from both sides.

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After that, the color began to make a lot more sense. The pattern did not occur on a continuum, like sabino appeared to do. It often did not look like much until the horse inherited it from both parents. That was why I sought out that very specific pattern as “proof” that Splashed White was there. Anything less went into my “maybe” files. I did that because over the years of searching for these horses, I found that some things that looked promising often ended up as dead ends. (Conversely, the horses that actually produced classic splash patterns often looked anything but promising!)

As I mentioned in previous posts, I have classified two sorts of “False Splash” patterns. I should caveat that by saying that it wasn’t that horses with these types of patterns could not have splash. In breeds with multiple forms of white patterning, splash carriers might well look like these horses. But they could also prove to be quite disappointing. That caused me to be rather cautious, because I could not be sure that these horses weren’t carrying something entirely different. Here are some clippings from my files of the two types:

SplashFalse1

These are horses where the bottom part of the pattern – the legs and the underside – look a lot like splash, but the white on the face is more like sabino. That is especially true for the Arabian pictured, Raffon’s Abida. Horses like this don’t usually have blue eyes, nor do they usually produce many blue eyes.

This is the other category of misleading horses:

FalseSplash2

These guys have the right kind of face and the blue eyes, but they don’t have the body white. Their tails are usually dark, too, whereas Classic Splashes tend towards white tail ends.

Horses marked like these are not always disappointing. Sometimes they do produce Classic Splashes. My personal suspicion is that this second type is what Classic Splash (suspected SW1) looks like in its heterozygous state when paired with a white-boosting mutation like sabino. In breeds like the Paint, where white-boosting genes are consistently found, this is what a lot of horses from splash-producing families look like. In breeds where those kinds of patterns are rare or non-existent, heterozygous horses do not seem to look like this.

So some of these horses probably are splashes. That said, I could never be sure that some other combination of white patterning might not also create this kind of look. That was because horses that looked like these pictured above sometimes occur, but they do not produce the classic pattern like the ones seen on the horses at the beginning of this post. It might be that they just haven’t been bred to another carrier, or that the odds haven’t worked out in their favor. But absent a test, I have reserved judgement, just in case it was something else. While it is possible to say something is associated with this or that pattern, there are enough gaps in our understanding that it is hard to know how exclusive those characteristics might be. It is quite likely that there is overlap between the different patterns. My great hope is that the new tests will begin to clear some of that up, even as they raise new questions.

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