DIAGRAMS & DESCRIPTIONS OF COAT COLORS/PATTERNS IN THE HARLEQUIN FAMILY

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

Below is a set of descriptions and diagrams of typical coat color variations found in the Harlequin family of dogs (which includes Mantles, merles as well as blacks and whites and, of course, Harlequins). This is written specifically for the Dane fancier wanting to familiarize themselves with some of the seemingly endless array of variation found in our "spotted" Danes, but is also for the individual outside the breed with an interest in "color" dogs in general. The Harlequin variant is in a unique position as this animal carries all three white spotting genes found in the domestic dog: to wit merle (PMEL), "S locus" recessive white spotting (MITF, and other?), and harlequin. For ease of understanding, the old Little terminology is used below. However this will first be augmented and then replaced as more specific alleles & loci are defined and labeled at a molecular level. Note that MITF has been shown to carry the solid and the piebald recessive allele that is associated with deafness, but not our "normal" recessive white spotting gene, called by breeders "irish spotting." The normal recessive spotting allele-the gene NOT associated with defect, may be found on an entirely different gene. And please refer to the above link for more accurate (if also more technical) information on coat color genetics.

HARLS, MERLES, BLACKS, WHITES: FOUR BASIC PHENOTYPES: In Harlequin breeding there are many iterations, endless variation, but essentially four basic phenotypes, which correlate with four basic genotypes & which result in specific and expected variation when they interact. This is what is outlined below. For more information and details on this complex relationship, read THE HARLEQUIN FAMILY OF DOGS: Harls, Merles, Blacks, Whites & more;and there are many other articles on various aspects of Dane coloration gathered up here. What follows below is specifically a demonstration of what occurs when various combination of these three white spotting genes occur. For a discussion of dogs beyond the "black-grey-white standard set," see this article on recessive pigment interaction with harlequin genetics.

"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 different 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.

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EXAMPLES OF GENOTYPES OF TYPICAL COLOR VARIATIONS:

A. HARLEQUIN: A Harlequin is considered a merle-bearing dog with a single Harlequin gene: a double heterozygote (MmHh) with black pigment (E-K-). This is a combination of two "dominant white spotting" genes, and produces a white base coat with black torn patches irregularly distributed over the body, head, neck, legs and tail. Without recessive white spotting genes ("irish" or "piebald"), the Harlequin will usually be referred to as "heavily marked." When the Harlequin has extensive white from the action of the more recessive of the "S locus" alleles, then it will be referred to as "lightly marked." When carrying only "irish" alleles, the Harlequin will be thought close to the ideal under the breed standard. Harlequins are unique: like snowflakes no two are ever just alike. And their markings can be very pleasing or not-so-much when the dogs are identical as to genotype (and so breedability). This isn't a color that will "breed true" in any sense of the word and simply cannot be standardized beyond some basic RANGE of markings. A WIDE range of markings (from heavily marked to lightly marked) is allowed under the breed standard & variations within the standard shouldn't be heavily penalized:: see the GDCA's illustrated standard for details. And note also that ALL Harlequins have some merle markings. The heavier the markings on the Harlequin, the more merle is seen typically, just because the dog has more pigment in general. And merle markings are not always some pale shade of silver or even an obvious gray; merle is referred to as a "mouse" color, as these patches can appear "tarnished," so the merling can be more tan to chocolate in appearance. This does not mean the Harlequin is "showing" fawn or chocolate pigment (a genetic impossibility) and does not mean the animal in question is even carrying for these other recessive pigments. Merling may not be necessarily aesthetically pleasing, but it's a natural part of the pattern, and is simply part of the coat pattern typically seen. So it shouldn't be weighted heavily against the dog, in that it's purely aesthetics and has no real bearing on the dog's breedability as doesn't reflect some change in the basic genotype for Harlequin. And it is worth noting the lighter the dog is, the more likely that hearing and sight will be adversely affected, so it just doesn't make good sense to penalize dogs more heavily for having heavier markings when the dog still fits the standard.

Harlequin #1: A-B-C-D-E-K-MmHh: Harlequin without recessive white spotting genes; more heavily marked (i.e. lots of black markings) with ragged patches of various size, often from very small (ticky) to very large (blanket-like). Take away the harl and merle genes and this is a solid black dog. Typically will have lots of merling. Harlequin #2: A-B-C-D-E-K-MmHh: The category inclusive of "solid-spotting hybrids" is represented here. These animals will be less heavily marked than the SS Harlequin but typically still more densely pigmented than the homozygous Irish dog, and certainly more pigmented than any homozygous piebald-bearing Harlequin. Typically this hybrid will have pleasing markings-take away the harl and merle genes and this dog wll usually be a "mismarked black" or "broken collared Mantle" in breed parlance. Harlequin #3: A-B-C-D-E-K-MmHh: The "Irish" Harlequin: this is the dog that meets the ideal for the breed (standard) most all of the time; a dog with extensive pigment on the head and torso, with white predominating on the legs and neck. Take away the merle and harl genes, and the dog would be a partial-to-full collared Mantle. This gene is stable. Harlequin #4: A-B-C-D-E-K-MmHh: The category inclusive of irish-piebald hybrids is represented here: "spotting hybrids" if you will. These animals will be less pigmented than the typical Irish, but more than the usual piebald (and so may be within acceptable range under the standard). Take away merle and harl genes, and the dogs in this category would be "flashy" Mantles with broken blankets to "plated" dogs with "Pointer" spots. Hybrids don't breed true. Harlequin #5: A-B-C-D-E-K-MmssHh: The "piebald" Harlequin: this is the dog that is predominately white, with what pigment that is left restricted to the head and small areas on the torso. The variation here is an animal from what is referred to as "lightly marked," to one that is sometimes called a "harl-head." Some of these piebald Harlequins can be confused with white Danes, as so much pigment is lost from the combination of harl, merle and piebald genes. And some are reported to have hearing and sight issues; more might have defects than is immediately obvious as CERF and especially BAER testing is rarely done in this breed, and piebaldism alone is associated with deafness, plus merles with extensive white can have both hearing loss and eye defects (e.g. merle ocular dysgenesis). "Harlequins" this white are not "normal" in the sense they are actually either homozygous merles or homozygous piebalds with a harl gene at best; some may not actually even have a harl gene at all.

B. MERLES: A merle under this breed's definition is actually a heterozygote merle (Mm), as the homozygote merle has a special category in the Great Dane and is referred to as a white (White Tiger=weisstiger). Merles are predominately gray dogs (brownish gray "mouse" basecoat) with black markings and some have white markings as well (which are actually largely dependent on the S locus alleles the merle is carrying). Merles are also called "Grey Tigers" (grautiger); Harlequins being Tigerdoggen in the old German parlance. Normally these "normal" (heterozygote) merles are perfectly normal (i.e. do not exhibit deafness or the eye defects that can be associated with the merle gene, and are seen in "double merle" dogs, called white Danes in this breed). Merles are disqualified under the breed standard. Recall merle is MOUSE--a brownish gray, not the gun metal gray of our blues. Merles DO NOT carry Harlequin genes: all are Mmhh. Merle#1: A-B-C-D-E-K-Mmhh: Merle without white spotting genes: "mouse" or solid merle dog (no white trim). Merle#2: A-B-C-D-E-K-Mmhh: Merle "solid-spotting" hybrid with white trim (toes, belly, chest, tail). Merle#3: A-B-C-D-E-K-Mmhh: "Boston" or "mantled" merle (true Irish dog--homozygous for irish spotting). Merle#4: A-B-C-D-E-K-Mmhh: "Spotting hybrid" dogs with extensive white trim: lots of face white, broken blankets, & dogs with big merle "plates" (areas of pigment) where white predominates. Merle#5: A-B-C-D-E-K-Mmsshh: Piebald merle: will appear either as a "merle-head" or a "pointer" marked dog that appears largely white with some round or oval areas of gray, with black spots in the gray areas typically. Sometimes these dogs get called "merlikins" (see whites below for more on merlikins).

WHITE DANES: In the Great Dane, the genotype otherwise called "dominant white," "double merle" or "white merle" is simply referred to as "white" (also called White Tiger=weisstiger) and is a homozygous merle dog. This animal in this breed is given it's own category because it has a seperate set of outcomes to the normal merle when used in a breeding program, as well as has a radically different appearance than the predominately gray dog that a normal, heterozygous merle is, AND because nearly all white Danes are deaf and/or suffer from various eye and other defects, whereas our normal, gray merles do not. White Danes can be referred to as "lightly marked" Harlequins when this predominately white animal has enough black pigment to be confused with a normal Harlequin. White Danes also get called "merlikins" when they have some visually obvious amount of gray pigment predominating. This term came about as the gray patches appear torn & the dog has lost so much pigment he has a white base coat, so looks somewhat like a Harlequin. But merlikins with even extensive pigment do NOT breed as Harlequins, but rather breed as do the more obviously white "white" (MM) Dane. Some genetic merle piebalds get put in this cateory too as they are predominately white. But a piebald merle also only has one merle gene, not two as does a "true" Dane white, and a piebald merle never has any harl genes. There are some distinctions in appearance, but it's not always obvious at a glance which is what genotype. But now there is now a test for merle, any time there is doubt about a dog's genetic status, there is no longer any need to speculate (and no need to test breed), as the dog can be tested directly and it documented if the predominantely white Dane has one, two, or no merle genes. There is no current test (research underway) for the harlequin gene, but harlequin gene homozygosity is typically considered lethal, due to the high percentage of white produced when in conjunction with the merle gene. To read more about white (MM) danes, click here. To see all the various things merle can do in various breeds, click here. There are three broad categories here that represent three major different genotypes, but the one thing they share in common is they have such a massive loss of pigmentation they are at increased risk for defects associated with white dogs, plus they will all be hard (without extensive gene level testing) to identify properly. This is likely why predominately white dogs have always been considered a fault or outright banned under various Great Dane breed standards.

ALL "true" WHITE DANES ARE actually MM "double merles." However there are variations in genotype. Some Whites carry recessive white spotting (S locus) genes, and in fact the "whiter whites" most likely always carry the more recessive alleles. Some Whites carry a harlequin gene, that is they are "harl-factored," but that's not necessarily visually obvious. Some white dogs are actually extreme piebalds without merle or harl genes at all. All predominately white dogs carry an increased risk for defect. Three basic categories of "whites" are presented below. To read more about white (MM) danes, click here. White: "double-merle/dominant white/white merle" = A-B-C-D-E-K-MM(hh or Hh). Dominant-whites which are "harl-factored" could produce harls; dogs without the harlequin gene (merliquins) would be dependent upon their Mantle or Harlequin mate to contribute harl genes for any harl offspring to be produced. Phenotype would appear (nearly?) identical, however the "harl-factored" white is less likely to show color & some suggest less likely to urvive, if what is generally understood about white spotting genes (more white means less viabilty) applies here (as it does all elsewhere to date explored). "Real" merlikins (dogs with merle ragged patches) are these white or "double" merles. White: "hybrid white" or "piebald harlequin" = A-B-C-D-E-K-MmHh. This is essentially an undermarked (overly white) harlequin, just as the piebald white described below is an undermarked Mantle. Such animals will contribute to the production of mismarked (i.e. white bodied, colour-headed dogs) more than more normally marked dogs. " But these "harl-heads" are at least carrying one harl and one merle gene, like a "normal" harlequin, it also recessive white markings that make them poorly marked. The other forms of white here do not do even approach normal Harlequin markings or normal Harlequin genetics. White: "piebald/recessive-white/colored-headed white" = A-B-C-D-E-Kmmsshh. These whites cannot contribute to the production of harl/merle offspring; but will contribute, recessively, to the extension of white and the lack of body markings, thus increasing over time the number of mismarks produced.

For more on excessive white dogs, their variation, and their commonality as to defect, click on the following links:

Color-Associated Health Problems in Dogs (explained by a geneticist and working breeder--with links), Spotting in Dogs, Pattern White Dogs, Double Merle, & DDEAF (Deaf Dog Education Action Fund).

BLACK TO MANTLE TO PIEBALD DANES: Despite a wide range of appearance, this is one category to the Harlequin Dane breeder, as these animals all lack the merle gene, and so are the perfect breeding partner to the Harlequin, as no "double merles" (whites) can be produced, thus is the potential for defective pups (due to color genes) drastically reduced (with the exception of piebald). The variation in appearance of the black, Mantle and piebald is due NOT to dominant white spotting genes like merle and harlequin, but rather to recessive white spotting genes (the S locus; MITF). What is important to also note is while Black/Mantle/Piebald Dane NEVER carry merle, they *can* carry harlequin and thus contribute to a higher percentage of show-marked offspring (i.e. producing more Harlequins) when they do. It's possible that Blacks and Mantles can carry not just one but even two harlequin genes, and so "breed true" (as are homozygous) for this gene (meaning they would have a very low percentage of merle offspring--a desirable outcome in Danes as merles are a breed DQ). Note that for all mismarked blacks and outright piebalds are disqualified under the breed standard, a WIDE range of markings are allowed & variations within the standard shouldn't be heavily penalized: see the GDCA's illustrated standard for details The basic five iterations that correlate with Harlequin are presented below. Note all may NOT come from the same gene even (something at least breeders have to remember).

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PROBLEMS ASSOCIATED WITH HARLEQUIN BREEDING: (problems that do not generally afflict the other colors in the Great Dane breed):
1. Harlequins, being heterozygotes, cannot breed true; i.e. cannot reproduce themselves consistency in either phenotype or genotype.
2. There is a loss of possible progeny due to MM and/or HH reduced viability.
3. There is considerable difficulty in controlling the color patterning & the amount of pigment even when correctly marked individuals are consistently bred to each other; so multiple incorrectly marked puppies therefore are born in most all litters, resulting in a reduced number of show/breed stock to select otherwise correct/superior individuals; high number of pets-by-markings produced..
4. Remaining well-marked animals may lack for breed type, conformation & soundness, so breeding choices and even quality can be starkly reduced when trying to assemble suitable breeding stock.
5. The genotype of the various near-white individuals cannot be determined necessarily on phenotype; test breedings may have to be performed to determine genotype, (if these animals are kept for potential breeding stock, rather than culled): which requires raising & rearing mismarks which may prove useless for breed improvement.
6. All near-white & white individuals should be checked for hearing & eye anomalies if not euthanized at birth as defects are to be expected in the majority of animals. Also animals with such defects may contribute to a rise in defective offspring.
7. Modifiers/other undefined genes inherited independently of recessive spotting (S locus) and dominant spotting (M/H loci) may affect the resulting phenotype and may prove hard to control. Currently it's considered that all these white spotting loci are incompletely dominant and may well interact, leading to unpredictable pattern variations in offspring & mismarks born routinely to even correctly marked stock.
8. All stud contract/co-ownerships must be carefully read & agreement of which color-type individuals will be euthanized at birth/how all mis-marks must be handled (e.g. how culled, euthanized, limited registration, spay/neuter contracts) must be settled between attendant parties given the ethical dilemmas inherent in this situation.
9. It is not uncommon for litters to include only 1 or 2 correctly marked Harlequin offspring. Further, many of the (now acceptable) Mantle offspring may be poorly marked or poorly made for consideration as breeding stock. Successive litters therefor may contain no correctly marked offspring,or the correctly marked offspring present may be the least adequate in all other breed features. The sex desired in also often unavailable in the desired color pattern.The result is few animals of breeding quality being available. So breeding appropriately becomes a certain challenge indeed. Since the serious fancier can easily find the opportunity (such as with fawns producing ~8 pups to choose from in every litter) to more reliably select and breed suitable Great Dane breeding stock, there is little long-term, educated and sustained interest in breeding Harlequins, given the constraints.
10. After careful consideration, most breeders interested in consistently producing quality stock turn to one of the other acceptable colors in Great Danes, where this is more likely to be a reality. This, combined with the low number of animals acceptable as breeding stock means there is a reduced gene-pool in the harlequin family. The color unfortunately attracts many novices who are unable/unwilling(?) to learn enough about the breed & the harl variant to produce correct specimens, or simply abandon the color after some poor breeding results. Given the internal constraints of breeding Harlequins also there is a supply and demand problem naturally met by many rather unsavory or simply opportunity-motivated individuals who exploit the novelty of the color and it's "rarity" to produce entire lines specifically for sale to a ready market. An added burden for the Harlequinn breeder who must judiciously (even zealously) screen to avoid homes who are more interested in owning "the spots" than the breed And limited registrations/spay and neuter options must be rigorously enforced to avoid "spawning" situations where more poor-quality 'spotted' Danes are ultimately bred. Ultimately a few knowledgeable, dedicated individuals with a flair for genetics, a good eye for a dog & a gambler's heart carry the Harlequin variant through history. (Many more who are mating harl-bred routinely and profitably selling these dogs know only enough about the variant to be dangerous to the breed IMO.)
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This message prepared by JP YOUSHA (2007) for educational purposes & permission is given to disseminate this message for that purpose & that purpose only. All copyrights & author's rights are to be respected. For further information contact: jpy@chromadane.com