NOTE THE INCLUSION OF THIS ARTICLE AT INDIVIDUAL BREEDER WEBSITES IS UNAUTHORIZED & THE DISCUSSION OF COLOR & MARKINGS BELOW NO WAY SUGGESTS A SUPPORT OF INDIVIDUAL BREEDER PRACTICES OUTSIDE OF BREED TRADITION.. ALL THOSE WISHING TO PURCHASE A GREAT DANE IN THE USA SHOULD BEGIN BY VISITING THE GREAT DANE CLUB OF AMERICA'S WEBSITE.

**NOTE: The articles in the section on "Great Dane SPECIFIC Coat Color Information" are intended specifically as a practical aid to those breeding & exhibiting Great Danes. They may not always be relevant to other breeds & the information is deliberately not technically minded.

THE HARLEQUIN FAMILY OF DOGS: Harls, Merles, Blacks, Whites & more.

TO See the latest on coat color research in dogs, CLICK HERE.

There always seems to be alot of confusion & "mythos" about harlequins, harlequin breedings & the merles that are born to harls. This little article is intended to answer the basic questions that most newcomers to the breed have, & to hopefully help clear up the confusion that comes from conflicting reports about harls, merles & their family. There is only a little more technical information in the article, but a several links & a list of references is available for those wanting to study the topic. If you don't want to wade thru all this, then remember, you cannot go wrong if you stick to the standard & breed ONLY show-marked & breed-quality Harlequins & Mantles to each other. That's why the standard was written in the first place; to act as a guide to breeders. So you don't need to understand all these details & theories of the genetics of coat color, but you do then need to stick to the standard when breeding. If there is anything you feel that needs to be added, etc. to this article, please email me & I'll do my best to see your question gets addressed. See Problems in Breeding Harlequins for the "short version" of the below.

TO See Diagrams & Descriptions of Various Harlequin Family Dogs, CLICK HERE.

"Let's begin by stating the observable facts that every harlequin breeder has to contend with. For starters, there are four basic colors found in harlequin litters. These are black, harlequin, merle, and white. This outcome is markedly differnt from the other four accepted colors (fawn, brindle, black and blue), which can be selected to breed true (harlequin cannot)." From BMW: A Harlequin Breeders Perspective, 1984.

HARLS, MERLES, BLACKS, WHITES: FOUR BASIC PHENOTYPES. HARLEQUINS all have at least one copy of the "dominant harl gene" (H allele) and at least one copy of the "dominant merle gene" (M allele). No other phenotype has this specify array of alleles. To have the Harlequin *phenotype* you must have a dog who is <HhMm> in genotype. Merles do NOT have the Harl gene (they are hhMm); they _are_ merles as they LACK the Harl gene. Blacks (including Mantles and piebalds) can carry the harl gene sight unseen. They don't have the Harlequin phenotype because they lack the Merle gene. So-called "whites" & "merlikins" in this breed strictly speaking are white merles or double merle: MM dogs. From Leigh Anne Clark's work we now know that genetic whites are MMHh and genetic merlikins are MMhh. Again, here the merle lacks the harl gene. For those wondering "then why do they produce Harlequins?" the answer is as they are bred to Harlequin Black or Mantle dogs that have the H gene, and THAT non-merle parent gives the offspring the ability to "become" a Harlequin. To read more about white (MM) danes, click here. And note below "white" means MM white or double merle, and so lumps genetic whites & merlikins into one category.

Having the harl gene isn't enough to end up a Harlequin; the merle gene (and the genes to end with black pigment) are necesssary for this unique spotting pattern to emerge. So two dogs with the merle gene or two dogs with the harl gene cannot produce Harlequins. One or both parents much contribute BOTH GENES to the puppy in question for Harlequin as a phenotype to emerge, because the Harlequin pattern is unseen when the merle gene is absent. And naturally the same applies with merle--two merles cannot produce a Harlequin as neither parent can contribute the dominant and vital Harl gene to the offspring. Breedings that supposedly have produced "harl from merle" inevitably turn out to have one non-merle parent (who contributes the Harl gene) or a "merle" parent that has been misidentified (usually a nearly white dog, so hard to accurately identify--which is a hint to why these animals are disqualified under the breed standard). The only way to get Harlequin offspring from a breeding with a merle in it is to use as a partner an animal who CAN contribute the Harl gene. And since the Harlequin phenotype required the Merle gene to be present to express (i.e. all Harls have one merle allele), it's perfectly possible for non-harl dogs (others than merles) to carry the Harl gene sight unseen. So anything BUT a merle from a Harlequin litter can, at least in theory, have the harl gene. But a merle is a merle as it lacks the dominant Harl gene.

First off, the basics of what you get when you breed. A note prior to presenting the data & links. Statistically the data are conclusive. This does not mean you will see these exact ratios of pups in litters, as such ratios only appear when large numbers are studied. This has been done. The data are extensive & it's well known what to expect from certain breedings. There is no real mystery to breeding harlequins; it's not "a crap shoot," & it doesn't require the use of mismarked dogs or weird combinations to be successful. There is NO secret recipe involving merles or other mismarked dogs. All this has been done time & again, so PLEASE don't think there is ANY need to try out odd combinations to "see what you get" or prove some elusive point. This is simply not an ethical approach to breeding: you do not care for the breed by "reinventing the wheel" and having litters to learn what has already been discovered, seen & written about. At the end of this article is presented an argument about theories of the harlequin variant. That does NOT mean that the basics are not understood, nor do these arguments have much to do with the practicalities of breeding harlequins. Read before you breed, please, and follow the standard when you do breed. A long list of references is offered at the end of this article & live links are offered thru-out to help you find the relevant articles & information you need. If you want to see the ratios of expected pups from various harl-family breedings, read Jane Chopson's article which gives all the offspring expected from various pairings or read one of the articles I've written, either the basic article or the more technical one. Louise Feddema also has one that outlines what to expect from certain breedings. I am NOT presenting myself here as "the" expert: I am presenting expert information gathered up in one place for the public to see that (hopefully) properly summerizes the available literature. Like I said, email me additions, corrections, etc. The best advice always is: follow the standard for the breed, in color, as in all other things. You cannot go far wrong as a breeder if you use the standard as your guide.

Before discussing color combinations it is critical to not that an important part of Harleqquin breeding involves not color, but PATTERN. Pattern is the stable point in Harlequin breeding, the one place where all good quality breeding stock and correctly marked show dogs have both looks and genetics in common. This simple fact has been very much undervalued and overlooked it would seem. COLOR has been far too much the focus in both judging and breeding, while there has been too little emphasis on PATTERN. Pattern genes are a critical component in getting correct color and arguably the most critical component as pattern, indubitably, is the most certain point of control in producing correctly marked Harlequins & Mantles. Color (Harl, Merle, Mantle, White) just describes how the various necessary genes to produce Harlequins statistically sort in the population/gene pool & there isn't control over this in any real sense (i.e. you cannot find a breeding to rid the gene pool of all but show color). Pattern on the other hand can be managed and maintained: there is one true breeding (the Irish homozygote) show mark and a couple of hybrids (the pseudo-Irish dog that is an irish/piebald heterozygote) that can produce show markings. So the **PATTERN**that results in the show-marked Harlequin & Mantle (and Merle when/if that's accepted) IS THE SAME PATTERN: all show-marked Danes in the Harlequin family have to have, need to have, the IRISH PATTERN to meet the current standard. Therefore it's pattern, not color, that should be the focus in Harlequin breeding. Control the pattern & you create a situation where you can maximize the number of show marks. Consider everything equally breedable as to color/pattern & you lose control.

TO See A Complete Guide to Color Combinations & Resulting Offspring, CLICK HERE.

BREEDING HARLEQUINS: THE BASICS.

First off, regardless of what combos you try, large numbers of mismarked puppies are expected in ALL breedings involving harlequin (& merle) dogs. The best you can ever hope for is about 75% correct color & pattern; 50% is the average if you count both Harlequin & Mantle offspring. You can reduce the expected number of correctly marked pups by at least 1/4 if you want to talk about harlequins specifically. Merles are routinely born to harls, as are various other mismarks, including white merles (MM) also known as (deaf) white (merle) danes. Therefore the problems of euthanasia of defective pups and the careful placement of dogs who fail to meet the standard is a routine part of the responsibility of any ethical harlequin breeder. You can avoid all of this by breeding Danes of any other family but Harlequin. Harls have problems & challenges the other colors of danes do not face. Mismarks in every litter is just one of them.

HARLEQUIN to HARLEQUIN: You can expect, in a litter of 7, to see 2 (likely mismarked) black or Mantle pups, 1-2 merle pups, 2-3 harlequin pups & one deaf white pup (two are statitically conceived, but one of two whites conceived generally dies prior to birth). Naturally people have 7 males in some litters, so some people will have 7 harls..or even 7 deaf whites, but on average, about 1/4 of the four basic colors is what is expected. Of course harls that carry blue or fawn or brindle can produce these "porcelain" dogs, sometimes called bluekin, fawnikin & brindlikin respectively. To read about why fawnikins occur, click here. Flashy harls (with lots of white) could also produce piebalds: white dogs with "mantle heads & harlequin looking bodies" as they have been described. Harl-, merle- or boston-heads; dogs with white bodies & "caps" of color (i.e. color-headed dogs) also can result from the use of piebalds & flashy harls & mantles. These color-headed dogs, also called piebalds, are essentially mismarked Mantles. To read more on piebalds, click here. Any time two dogs with the merle gene (incl. harls) are bred to each other, white merle (i.e. defective, dominant white) offspring are to be expected. The dominant whites that do survive to birth are normally defective & commonly deaf. Ethical breeders have traditionally euthanized most all whites at birth for these reasons. To read more about white (MM) danes, click here.

HARLEQUIN to MANTLE: This is the preferred breeding, all things considered, as it does not produce the Dominant (deaf ) White dane. Expect the litter to be about 1/2 mantle, 1/4 merle & 1/4 harl. So in a litter of 8, expect 2-3 harls, 1-2 merles & 4 mantles. If these black pigmented dogs carry for other pigments, you may get fawns with blazes, or blue instead of a black basecoat. (If that is confusing, read about why fawnikins are here to stay.) Piebald & color-headed dogs can occur, especially when flashy dogs are used.

MANTE TO MANTLE: Produces only Mantle: that is, it cannot produce harl, merle or white danes. Now piebald danes that are mistaken for whites and for harlequins can be born to Mantle parents. This is more likely to happen when breeding flashy Mantles with big collars & broken blankets than with dogs who have more pigment. Mismarked blacks can also be born to this breeding as can piebalds, so this breeding doesn't promise all show marks, it just guarantees having nothing but black and white dogs. It is possible (if unlikely!) that very flashy Mantles bred to each other could produce a puppy so white s/he could be deaf. Possible--not probable--& AFAIK it's never happened in Danes. BUT it does happen in Boxers & Dalmatians.

MANTLE TO (harl-bred)BLACK. Would likely produce mismarked blacks. Mantles & (solid) Blacks also are possible.

BLACK to BLACK: Would likely produce mismarked blacks when harl-bred blacks are used. Mantles & Blacks possible.

BLACK TO HARLEQUIN: Produces as does Mantle to Harlequin, except, since a Black lacks the white collar of the Mantle, expect more mismarks under the AKC standard to occur, both in Harlequin & especially in potential Mantle offspring. Blacks will likely not be solid Black, but mismarked black, sometimes even called "harlequin black" it is so common to black to harl breedings. These "harlequin blacks" are disqualifying under the American standard.

CLICK HERE to see PUNNETT SQUARES with illustrations of colors expected from various breedings.
These above breedings are authorized by the Great Dane Club of America (GDCA) & by various other breed clubs. Note that the Great Dane Club of Germany has banned Harl to Harl breedings in an effort to avoid the production of white danes. To read more about the ban, click here. Some European countries have gone so far as to ban ANY dog who carries the merle gene from being bred to another merle-factored dog & some have banned merles entirely, all because merles produce deaf & defective white puppies & pups who die in embryo. The merle gene is "semi" or "sub" lethal--and kills supposedly about 50% of those pups who have two doses, while is leaves most of the living pups deaf or with other defects. Read more about whites here. The breedings below are NOT authorized by the GDCA. All involve dogs DISQUALIFIED worldwide under the standard for the breed. Disqualification is done specifically to insure that a dog with such a defect will not be bred.

BLACK TO WHITE: (or Mantle to White). Produces only harls and/or merles.* This breeding is done by some people because it sometimes offers the possibility of producing up to 2/3 harls in a litter. Unfortunately a whole litter of merles is more likely than all harls & there are lots of problems with using whites in a breeding program. Whites may all look alike, but they don't all breed alike! And of course most of them are deaf. To read more about whites, click here. White to black breedings that produce black, mantle or piebald pups are not true black to white breedings; the "white" used is actually a mismarked harlequin or a misidentified merlikin.*Note there has been an idea that there are very rare situations where a "back mutation" (reversion to wild type) occurs and a non-merle pup is born to these breedings. But unless this can be proven to be the case, it's best to assume the more normal situation--the breeding actually wasn't MM x mm. And most of the Blacks/Mantles born to Black x White breedings are actually cryptic merles (genetic merles with a black/mantle phenotype). NOTE: Using deaf dogs in a breeding program is reported by some to increase the number of deaf pups thru several generations.

WHITE TO HARL: This would produce a litter usually with 50% deaf whites, the rest split between harl & merle, depending on what kind of white was bred. Again, if this breeding produces black, mantle or piebald pups the "white" used is actually a mismarked harlequin or a misidentified "merlikin."

HARL TO PIEBALD: This would increase the number of mismarked pups, as you'd expect to get harl-heads instead of harlequins & piebalds in the place of mantles, especially if the harl used is flashy or even just has that clean white neck and also mismarks are more likely when there are piebalds used more than once in a pedigree. It also means more of the offspring of this offspring are going to be mismarked as a rule. To read more about the piebald, click here. Piebalds are mismarked Mantles, not a kind of Harlequin.

MERLE TO WHITE: 50% deaf (or dead) white is expected; 50% merle, with reduced litter size due to merle/harl-related deaths.

MERLE (or merlikin) TO BLACK (or to Mantle): This is a breeding often using TWO mismarked dogs (most of the others use at least one parent with correct markings), so will be considered controversial. Since some harl-bred blacks & mantles can carry harl genes (H-factor) unseen & merles obviously carry the Merle gene, recombining the two can occassionally produce harls. But mostly, again, this breeding produces mismarks: black with white markings & merle dogs. And the dogs used in these sort of breedings are all too commonly sub-standard in other ways than color.

MERLE TO HARL: This is like breeding harl to harl, except you reduce your chances of getting harls & increase your chances or getting mismarks and various defects by all reports.

MERLE TO MERLE: This is not deliberately done in Danes, & although it is occassionally done in other breeds, it's always considered controversial. It is universally discouraged & is against the law in some countries. Vague reports of such breedings in Danes make the rounds from time to time. From what is known & extrapolating from other breeds, at least 25% or the litter is expected to be defective, 25% will be a color other than merle, & the rest (50% statistically) will be merle. More on breeding merles below.

WHAT ABOUT THOSE MANTLES--WHAT ARE THEY AND HOW TO BREED THEM?

I published an article in the Great Dane Reporter in 1997 on the action of the S Locus itself (when alone) as it applies to the Mantle (Manteltiger) entitled: "Mantle Dane Genetics: How to Get & Keep Boston Patterned Dogs." There are also somewhat harder to find, but very useful, articles published over the past two decades by such as Paul Hardiman & Laura Kialenaus on the subject of the Mantledane. I tried to summerize their message in the article I wrote in 1997. This article discussed some of the implication of attempting to stabilize a "tux-n-tails" bicolour Great Dane as a breeding partner to the Harlequin. This also is a complex issue, but the basic take home message is:

1) Breed only properly marked Harlequins & Mantles to avoid not only mismarks but an increase (thru loss of pigment) of pups with sensory defects. The breeding of piebald or flashy boston-type dogs (those with extensive white, i.e., more than 1/3 white) will result in a loss of harl offspring (to such as "harl-heads"), & could cause an increase in defective pups due to loss of pigment from these recessive white genes combining with the dominant white gene(s) that make up the merle/harl complex. Therefore, avoid the breeding of mismarks, especially predominantly white mismarks, such as harl/merle & boston heads, whites, mostly white, white headed, piebald & porcelaine (merlikin/fawnikin etc.) dogs whenever possible, as the effects are multi-generational and generally deleterious to both health & color. They simply carry too much recessive white genetically to be safely combined with harldane (merle) dogs. White to black (including Mantle) breedings are not unusual these days in North America. But recall "whites" are often an unknown genetic quality. Also Mantles are now accepted. More reasons to select other choices. Plus, as noted above, using deaf dogs in a breeding program is reported by someto increase the number of deaf pups thru several generations.

2) Attempting to stabilize a full and complete collar and the approximately 1/3 white called for in the ideal of the Mantledane requires one to 'straddle' a gap between two alleles (si=irish & sp=piebald) (or two genes entirely are involved). So many dogs with this same phenotype are going to have VERY different genotypes & most of those genotypes are NOT going to be homozygous (e.g Ssp/se=pseudo irish occurs). The long and the short of that is, that many Mantledanes who look just alike are going to produce very differently, & the more mismarks used in the breeding program (especially predominately (>50%) white dogs), the more mismarks that can be expected, especially "skip generation" recessives. Consider the problem with whites & checks in Boxers, as this is the exact same phenomena-less our problems with the merle gene. What is called "white factoring" occurs in these breeds--i.e. the presense of a piebald recessive white allele--which results in predominately white dogs from predominantly pigmented parent. This phenomenon is present in a variety of breeds from Collies to the Akita. Broken collared dogs & other dogs who are "safely" (at least 2/3) pigmented are a better breeding bet for overall health than rather flashy marked animals; particularly "extreme" bostons and out-right piebalds (Plattenhunde) and most particularly when bred to flashy marked harls.

Checks & whites? If not familiar with this phenomenon, see how "Plain," "Flashy," & "White" phenotypes in the Boxer (roughly) corresponds to Solid Black, Mantle & the Piebald in Danes. Click here for an illustrated tour of this two gene, three phenotype phenomenon typical of incomplete dominants. See also "Spots Before Your Eyes" for an illustrated guide to the complete & complex phenomenon of the recessive spotting (S) locus with its four alleles, incomplete dominance, 10 genotypes, modifiers, & multiple, overlapping phenotypes, which is likely a much closer analogy to the situation in Danes.

3) It would seem inescapable that some & not all Mantledanes carry some sort of allleles, be they "modifiers" of some kind or a more "direct" & seperate H-factor ("harl" locus alleles), that contribute to the production of harls (over merles) in harlequin litters. This is implicit in much of what has been written, but is rarely explicitly stated. It certainly should be addressed at part of the family inheritance, as should the implication that follows directly from this observation; namely that Mantledanes, as individuals, contribute "unevenly" to the production of harls, with some unable to offer any "help" to increase in the percentage of harls in a litter, while others might, at least under certain theories, carry strongly enough for whatever constitutes "harlequin genes" to not only increase the percentage of harls in a litter (over merles) from a harl x mantle breeding, but actually produce harlequins from a merle x mantle breeding.

BREEDING MISMARKS: DANES WITH DISQUALIFYING COLOR DEFECTS:

First off, a Dane with "lesser" markings isn't a mismark really. A mismark is a dog's whose color/markings define him, under the breed standard, as disqualifying--so is a dog of a NON STANDARD COLOR. People often declare: "You should breed the best ones, regardless of color!" That sounds good at first glance: it's practical, seems like then you have your priorities straight, maybe even seems noble. The problem with this is, upon examination, it doesn't look so good. First off, by definition, to breed a mismark means you select as a pup a dog with a disqualifying defect and rear it to adulthood intact. That's already a non-starter for the average reputable breeder with a reasonable litter, as why pick the mismark in the first place? You cannot know it will turn out better, all pups are a gamble, & your litters shouldn't really be so uneven the show marks are awful while this one merle is just wonderful? It's two years or more investment, after all, getting breeding stock is--not many want to start with a dog they cannot show, don't know how will turn out, is disqualified under the standard and so on. Secondly, most who advocate this stance don't just breed that one wonderful mismark; they use this argument to breed MORE of the pups in the litter & sometimes a way to sell them all as breeding stock. So that's not selection at all, nevermind selecting the better dog with a bad pattern over the lesser dog with a good pattern---something anyone, anyway would agree with if it were that clear of a situation. Thirdly, this use of mismarks in a breeding program becomes the most widely abused excuse for not only not showing the dogs, but telling all how wonderful the dog is, when usually noone can confirm or deny that, as noone has access to the animal. So that also means the dog often gets lost to breed history: there are no photos, no recorded data of how she lived or die, and so no. The below goes thru the whys and wherefore of using specific sorts mismarks in a breeding program. However the main take home message has got to be most good breeder don't use them very often. Despite myths, defensive postures, and declarations of their value, most mismarks do nothing for a breeding program but create more mismarks for the breeder to contend with. So if you are trying to breed dogs to the standard, you are trying not to use mismarks.

THE "WHY" OR RATHER THE "WHY NOT" OF BREEDING MERLES:

NOTE that these comments do not necessarily represent my personal views, but provide the new breeder & those curious about this subject a basic reference to the basic aspects of why merles are not recogized under any Great Dane standard anywhere or anytime as acceptable breeding animals.

1) Merles are a disqualification under all standards for the Great Dane--and have always been. That is enough, in many registries, for a dog to be denied "papers," so this is obviously always been thought a very poor breeding choice. (That the AKC will register a dog does not imply approval of it as a breeding choice--the AKC will register *ANY* dog with two registered parents of the same breed. Any dog.) Historically it was thought that if you restricted the gene pool to Harlequins and Blacks(including Mantle), you could rid the breed of the merle gene by denying merles breeding and show status. Breed traditions are often 100s of years old, as is the case with merles being considered a non-standard color. When this breed was codified as a pedigreed breed in the mid 19th century, certain colors were chosen to represent the breed's typical look. Amongst these was the Harlequin Great Dane.

This was just the beginning of the era of understanding genetics and at the time it was thought (by all breeding plants and animals) you could simply select for certain traits & against others by chosing one breeding over another. Merles for centuries have been known to produced defective pups, which was a practical disadvantage for the breeder. This happens in any breed or cross bred dog that carries merle (and it's a gene you cannot always "see" which is another aspect of this topic). Harlequins were widely thought until very recently to be a completely different dog than a merle. Their traditional breeding partner was black (not necessarily solid black), a breeding that produces harl, merle & black(mantle). Discarding the merles as breeding stock was thought to end ultimately in NO merles being born and so raising the standard of puppies in general & creating more "purity" of color. And at this time in the breed's home country blacks were NOT shown, they were simply considered a by-product of Harlequin breeding and used then as you suggest merles be used now--to produce more Harlequins.

This approach in breeding has been used for 1000s of years in all domestic stock and works 99% of the time. Harlequin genetics just happen to not fit the normal mode & it was only when both the merle & harl genes were discovered that it could be finally established that Harlequins always have a merle gene & are actually a merle variant. This is very new information and still has not been fully published. When this data is published and reviewed by such organizations as the GDCA and DDC it may well result in alterations to these traditions. But historically, a variety variations on the merle were used & the results outside of using the Harlequin itself were generally not good which contributed to their being disqualified. In other breeds which allow the merle (very few BTW), merles are not supposed to be bred to each other, to avoid the production of white merles: those defective, predominately white pups. Merles LACK the necessary genetics on their own to produce harls: noone has ever documented a merle to merle breeding that produced harls. So by all reliable reports, merles as a group do not productively contribute to the harlequin-mantle gene pool. That said there are ways to properly use such as the irish-marked merle dog (i.e. mantled merle) effectively & even the "double" merles (whites & merlikin), but the former has only one effective breeding (breed to a Mantle testing positive for the harl gene), and the latter puts the breeder in the murky ethical waters of producing and then using in a breeding program dogs expected to have sensory deficits.

2) It is certainly and always the case that anyone who has not established a *sterling* reputation & can explain to the general dane fancy's satisfaction the reason to break such a general rule of breeding better be prepared to be considered unethical by most if not all others concerned about ethical breeding. This is a case where making an exception to the rule is better NOT done but for the rarest of cases where the person in question has a long track record of good practices, the production of Champions under ethical constraints, AND who has an exceptional & rare enough situation on their hands to warrant a probably once in a lifetime event. People who sell & breed merles (including merlikins) as breeding stock on a regular basis can simply be generally assumed to be commercial (i.e. for-profit) breeders whose goal is cash cropping harl pups, not breed protection. These dogs are also bred in ignorance by people only casually involved with the breed. Needless to say, however well meaning, these breedings do not contribute to breed betterment either.

3) A boston merle, the most likely choice as a breeding partner to a harl, is essentially a "true" boston (i.e.Mantle), with the added problem of producing deaf white & other defective puppies, which does not occur in a Mantle to Harlequin breeding. So a Mantle is always preferable, on this account, to a merle.

4) "Merlikins," a white base-coated dog with black & grey markings & a dog that lacks the harl gene, have been bred on occassion to Mantles to produce harls. But this is a risky breeding for several reasons. Merlikins may be genetic whites (MM), so then are often deaf &/or have eye defects (& without a BAEF/CERF noone can say a dog's eyes or ears "are fine"!), most (Mm) merlikins will NOT produce properly marked harls under any circumstances, & the harls they DO produce are simply the result of their Mantle (boston) mate carrying unseen the factors needed to produce harls. And they are disqualified under the standard. All this means merlikins are established as a poor breeding choice as a rule. So, again, a properly marked animal, in this case a show-marked Harlequin, would be preferable to the merlikin.

5) The long & the short of breeding merles (& other mismarks, for that matter) is, if you have to ask about the whys & wherefores, you probably "can't afford it;" can't afford the potential for dissapointment, even disaster & dishonor likely awaiting you. Unless you are dealing with a publically esteemed breed expert really caught in a corner, you can pretty much assume someone using mismarks (e.g. merles, piebalds, and other mismarks) routinely in their breeding program is less than knowledgeable & less than ethical. This *is* a case of guilty until proven innocent.

(6) Bear in mind this is an internal Dane dialogue & one predicated on tradition. There are several breeds where merles are perfectly acceptable to breed (even if they are not bred typically merle to merle, as all try to avoid the production of "double" merles (white merles), technically known as homozygote merles. And despite Dane tradition, logic and new research suggests that irish-bearing merles ( mantled merles) could arguably have a legitimate place in even responsible breeding circles to help secure the gene pool. By this I mean if a reputable breeder had a show marked merle equivalent to a Mantle and Harlequin (both of whom, to meet the standard, must exhibit the "Irish" pattern essentiall), this might prove useful to the breed.First it would mean the Harl breeder, notoriously restricted as to breeding choices acceptable under the standard, would have a few more puppies to choose from, so could more often choose quality over color. Secondly if mantled merles were accepted under the standard, this would put them on an equal footing with Mantles & Harlequins, which would help demystify this color & that would likely result in a reduction in buyers purchasing pet puppies from poor breeders. Which isn't to say all agree with this argument or that then it's "okay" for all sorts of merles to be randomly bred as cheap alternatives to Harlequins, but that at some time in the future when we know more about the genes involved, we may have to revisit this issue, even consider the Mantled Merle as a useful show/breed dog.

WHITE DANES (actually white merles=homozygous merles) have a page of their own:

WHITES: IS IT WORTH THE RISKS?*OWNING & BREEDING (in two parts-scroll down).

PIEBALD DANES also have a page of their own:

THE PIEBALD DANE: Pintos, Parti-colored, Check: the Colour-Headed or White-factored Dane. (Der Plattenhund)

CLICK HERE FOR UPDATES ON GENETIC RESEARCH AND THE LATEST ON KNOWN FACTS ABOUT THE HARLEQUIN VARIANT

***SOME HARD FACTS TO PONDER BEFORE BREEDING OR BUYING HARLEQUINS***

1) It is never possible to control for a predicably fully white neck, even when two such animals are bred consistently together; the genes (as described above) do not admit of such control. So it is not a trait worth giving an enormous amount of attention to, and the standard for the Mantle doesn't require a fully white neck anyway. But it's aesthetically pleasing & many do make a fetish of it.

2) Even the AKC standards say NOTHING about white "fronts" & chests and forelegs being even desirable, let alone preferred on harls or mantles. Note also the illustrated standard says markings on the neck & legs, etc. of a harldane are NOT to be faulted. So faulting those equally correct dogs is short sighted, even if certain patterns within the accepted range are fashionable or even more aesthetically pleasing. But it happens all the time, in and out of the show ring.

3) Since Harlequin and Mantle dogs of less than ideal pattern are inevitably necessary to a breeding program, it is best to stay within the acceptable range of pattern as documented by the illustrated standard. All sorts of romantic legends about the value of mismarks in breeding programs abound, but it's a simple fact that no truly dedicated breeder would ever prefer a mismark to an animal correct to the standard. Apart from there being absolutely no documentation that mismarks "help" produce harls (in fact the evidence is quite the contrary), it is also obvious far too many mismarks are included in breeding programs simply as a cheap way to make harls. And it's practically an identifying trait of a poorly educated Harlequin breeder to give undo focus to color and to talk of "color pure" dogs and pedigrees.

4) Mismarks make more mismarks. Even if the Harlequin variant, as "obligate double heterozygote" cannot breed "true," they & their Mantle mates cand and will produce more correctly marked animals than mismarks do. And the problem with breeding & getting Harlequins anyway is not license to sell and breed every imaginable mismark that comes out of a harl litter. Many of these myths about the value of breeding mismarks are so obviously self-serving. Many who breed them are simply uneducated about harlequin genetics it would seem. With the possible exception of the mantled merle, the mismark who is of actual breed value is a rare creature from an unusual breeding, so there is no justification for routinely including mismarked animals in a breeding program. Predominately white dogs from harl breedings, if kept, should be BAER & CERF tested, according to DDC (German parent club) recommendations & all should be correctly identified as to genotype, even if that means paying for gene testing. Deaf/defective dogs used in breeding programs are said by some to have an increased incidence of deaf/defective offspring, even several generations later, and many predominately white dogs are not of a useful genotype anyway, but can contribute to problems in the puppies. Something to think about, as losses of normal & useable animals are to be expected when deaf dogs are used.

5) When using dogs either under-marked (lightly marked) or over-marked (heavily marked), it is better judgment (if less the fashion or the common taste) to use dogs with more, rather than less pigment. This was the original intent of the standard of the country of origin & is still so designated in most standards worldwide. The AKC standard for the Great Dane has altered its color description during various revisions for a dog with more white than called for or originally prefered. It is obvious for better health that more white is less prefered.For us to safely breed dogs carry "lethal" & "semi-lethal" genes we need to consider the implications of simultaneously carrying 2-3 white genes well documented to cause serious health defects when "used to excess" to produce predominately white dogs. We should not encourage an increase in defects purely for the sake of a (currently) prefered aesthetic, sale of breeding stock, or ease of breeding choices.

And note: For us to have the CORRECT and ALLOWED range of Harlequin & Mantle coloring/patterning recognized in the show ring, we have to educate our judges by showing them the correct range of acceptable markings; not some individual (or group's) aesthetic choice, or a fashion of the times--but DOGS BRED TO THE STANDARD. This is important for breed preservation.

And as not to forget the art in the science (of breeding), I offer this short poem by Gerald Manley Hopkins, who surely would have included in his beloved "dappled things" the glory that is the Harlequin Great Dane.

Glory be to God for dappled things--For skies of couple-colour as a brindle cow; For rose-moles all in stipple upon trout that swim; Fresh-firecoal chestnut-falls; finches' wings; Landscapes plotted and pieced--fold, fallow, and plough; And all trades, their gear and tackle and trim.

All things counter, original, spare, strange; Whatever is fickle, flecked (who knows how?): With swift, slow; sweet, sour; adazzle, dim; He fathers-forth whose beauty is past change; Praise him.

(Gerald Manley Hopkins)

REFERENCES:
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Carroll-Draper, Nancy. 1981. The Great Dane: Dogdom's Apollo. New York: Howell Book House.
Cattanach, B. (1999). The 'dalmatian dilemma': white coat colour and deafness. J. of Small Animal Practice 40: 193-+.
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Chopson, J. 1992. Inheritance of Great Dane Coat Color. GDCA Color Committee.
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O'Sullivan, Neil. (1988-89) "Harlequin colour in the Great Dane dog." Genetica. 78(3):215-8.
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Robinson, R. 1982. Genetics for dog breeders. Oxford: Pergamon Press.
Schaible, R.H. and Brumbaugh, J.A. 1976. Electron microscopy of pigment cells in variegated and nonvariegated piebald spotted dogs. Pigment Cell. 3: 191-220.
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Willis, Malcolm B. 1989. Genetics of the Dog. New York: Howell Publishing. Yousha, JP. 1996. A Summary of Theories Concerning the Harlequin Variant in the Great Dane. Great Dane Reporter: M/A.

This message written and prepared by JP Yousha for the purposes
of education and can be reprinted to that end.
All copyrights © remain with the author.
CHROMADANE 1998 (revised 2001, 2009)
*multi-titled/certified harlequin family danes*
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