BEAUTIFUL BICOLOURS - TUXEDO AND MAGPIE CATS
Copyright 2002 - 2011, Sarah Hartwell

Bicolour cats go by various names and come in many patterns. They range from almost solid colour cats with a white throat locket or white tail-tip, through to almost solid white cats with black smudges on the nose or between the ears. You may have heard of them as tuxedo cats (white mitts, white belly and white chin with an optional white tail-tip) or patched, pied, particoloured, harlequin or magpie cats (usually white with coloured splashes on the back and top of the head). The term covering all bicolour cats is "piebald" or "white spotted" with some variants of the pattern being called the Seychelles pattern.

DIFFERENT TYPES OF PIEBALD

 

 

The diagram above shows a typical progression from solid colour through to solid white. The number by each diagram is the "Grade" of spotting from Grade 0 (no spotting) through to Grade 10 (white spotting has obscured all of the base colour). The "solid colour" can be one of the true solid colours e.g. black or grey, or a tabby colour or tortoiseshell (the bicolour is then known as a calico and is actually a tricolour). Non-pedigree cats have a diverse range of combinations not recognised in the pedigree world - the "solid" area can be Abyssinian ticked (Aby-and-white), smoke, shaded or tipped. Black-smoke-and white is an attractive combination if not actually encouraged!

The Van pattern seen in pedigree Turkish Van cats and in the Seychellois (a Van-patterned Oriental) is Grade 8 - Grade 9 Piebald and represents the most extreme of the 'Seychelles' patterns. In the Turkish Van, the colour is restricted to auburn (red) or cream patches on the head at the base of each ear; the tail is the same colour as patches, often with darker rings because red and cream are not true solid colours. Although auburn has long been the traditional colour of Turkish Van markings, other colours are being developed within different cat fancies, but in its native Turkey, the Turkish Van actually refers to an all-white cat (Van Kedi).

 

(Almost) Van Pattern (Grade 8/9 Piebald)

Grade 7 Piebald Littermates

Grade 7 Piebald

 

The Van pattern is also found in Persian Longhairs in the form of Van Bi-Colours & Harlequins. These often have additional markings on legs, and one or two small splashes of colour on the body. The Seychellois is an Oriental type cat, occurring in both longhair and shorthair forms, exhibiting varying degrees of the Van pattern:-

Seychellois Neuvieme is a white cat with coloured tail and head splashes
Seychellois Huitieme is white with coloured tail and head splashes plus additional splashes of colour on the legs
Seychellois Septieme has splashes of colour on the legs and body in addition to those on the head and the coloured tail.

British cat fancier Pat Turner defined the different levels of white spotting as: Grade 2 - "White Trim"; Grade 3 - "Mitted"; Grade 4 - "Irish"; Grade 5 - "Saddle"; Grade 6 - "Pied"; Grade 7 - "Chinese"; Grade 8 - "Harlequin"; Grade 9 - "Van". In exhibition-quality bicolour cats, symmetrical markings were preferred.

The placement of the colour patches is variable and related to how the embryo develops and expresses the genes it has inherited. As a result, even more extreme colour restrictions can turn up, for example Wanda below (photos provided by Peggy Alden-Clapper) is an eyecatching variation on Grade 9 Piebald - all white apart from a black tail, but no splashes of colour on the head or body.

Variation Grade 9 (Extreme Grade 9)

"Moorish-headed" Cat (Variation on Grade 9)

In 1896, Jean Bungartz wrote in his "Illustrated Book of Cats" (in German only): The black-headed or Mohrenkopf (Moorish-headed) cat must be clean white, with contrasting colour on the head and tail. Consistent specimens of this variety are extremely rare and it can probably be regarded as one of the most peculiar colour patterns of the domestic cat. The colour of the head and tail can be either black, grey, blue or yellow with no white hairs except those regularly showing up on the head. As previously mentioned, cats with good and correct markings are highly valued.

THE TYPICAL RANGE OF BICOLOUR PATTERNS

Piebald cats are extremely common and varied in appearance. The white spotting can occur with any colour/pattern and varies from minimal white through to almost all white. The smallest amount of white may be no more than a white locket and a small white belly patch, each no more than a few white hairs. Though this pattern doesn't have a descriptive namem the term "locket cat" has been suggested. The image at the top of this page and the description below is a typical progression through the different grades of piebald. Due to developmental effects at the embryo stage, there are further variations on this general theme!

 

Tuxedo Pattern

White facial markings

 

At one end of the scale is the white locket, followed by the familiar tuxedo pattern where the belly is largely white. As the degree of white spotting increases, the white spreads up the neck and onto the chin (as shown in the three-quarter Persian above). The cheeks may be white or have white spots and there may be a white blaze (an inverted V) from the muzzle up between the eyes or a white "teardrop" on the nose (as shown in the cat above right). If the neck and chin are white, the front paws are also usually white.

 

Mask-and-mantle pattern

Saddle pattern

Grade 7 Piebald (with white ringed tail)

 

As the degree of white spotting increases further, the white extends up the sides of the cat, up the forelegs (stockings) and onto the hind paws. The next step is all white hind legs, white fully around the neck and the white blaze may extend right between the ears. Some cats also have white ear tips. This pattern is sometimes known as "mantled" because the cat appears to be wearing a coloured cape with a hood or mask. The mask-and-mantle is a common bicolour pattern as shown above.

As the amount of white increases further, it simply shrinks the mantle into a saddle. The mask may also shrink back to a "cap" giving the cap-and-saddle pattern (as with the cat in the above centre photo). Further white spotting breaks the saddle up into smaller patches as shown in the black-and-white Grade 7 piebald cat. Sometimes one or more white rings encircle the tail - for some reason this ringed effect seems more common in tabbies.

With any degree of spotting, there may be coloured smudges on the cheeks, chin or nose. Some cats also have black toes on otherwise white paws.

 

Jean Bungartz described two bicolour patterns in his 1896 book "Die Hauskatze, ihre Rassen und Varietšten" (Housecats, Their Races and Varieties) in " Illustriertes Katzenbuch" (An Illustrated Book of Cats). His description of the "masked cat" is the tuxedo pattern, which he describes as "sometimes the tail-tip is white also. The eyes of this variety are bright yellow with black rims. Regular and sharply defined white patches create the most beautiful Mask Cats which have many admirers."

Bungartz describes a less familiar bicolour he calls the black-headed or Moor-headed cat (Mohrenkopf) which was clean white, except for the head and tail, which were black, grey, blue or yellow with no white interspersed "except those showing up regularly on the head". Consistent specimens of this variety were extremely rare and valuable. His illustration depicts a cat with wholly black head, but it is possible he was describing the Van pattern.

It is possible to combine piebald with bicolour. Bicolour Siamese have been bred in Europe to a mixed reception. A few Himalayan breeders breed bicolour Himalayans, using bicolour Persians to introduce the white spotting pattern. Paul Beall, Richmond, Texas is one such breeder. Paula's photo (below) shows a bicolour (piebald) Himalayan, but unfortunately this cat has not produced piebald-point kittens and Paula has decided not to reintroduce the gene from Persians.

GENETICS OF PIEBALD CATS

When the amount of white is small (under 40%), it is called low-grade spotting and comprises the white belly patch, mitts, locket and blaze. In medium grade spotting, the amount of white varies from 40% to 60% of the coat and is the typical pattern of a show quality bicolour (a mask and mantle effect). When it is extensive (more than 60%), it is called high-grade spotting - the Seychellois or Van pattern.

Piebald spotting is a semi-dominant gene with very variable expression. Low grade spotted cats and some medium grade, spotted cats are heterozygotes i.e. they have only one copy of the white spotting gene. Other medium grade spotted cats and all high grade spotted cats are homozygotes i.e. they have 2 copies of the gene. Where cats are in the medium spotted range it is generally impossible to know whether they are heterozygotes or homozygotes unless they are bred to a cat known to be either homozygous or heterozygous. To further complicate matters for breeders, some apparently non-piebald cats are really piebald cats whose white bits are so minimal they might be no more than a few hairs in the groin or at the tip of the tail!

The gene for white spotting affects the embryo cells (melanoblasts) which will become pigment-producing skin cells (melanocytes) which make the pigment for hair. One theory is that the melanoblasts arise from the "neural crest" - the area along the back of the embryo - and they migrate to all over the body during formation of the skin. Where these cells fail to reach their allotted positions before the skin is fully formed, those areas of skin will lack pigment producing cells i.e. be a white spot.

This explains why the white is most often found on the paws, belly and chest - these areas are the most remote from the neural crest and take longest to reach. The slower the migration of cells, the less colour there will be. This also explains why the back and the tail may be coloured in an otherwise all white cat - those areas are closest to the neural crest and the pigment cells didn't have to migrate very far at all!.

Where the white spotting occurs over the eyes, it may affect the eye colour. Thus a few bicolour cats have blue eyes. Another interesting effect of white spotting is in tortoiseshell cats. Tortie cats with little or no white tend to have brindled coats with intermingled black and orange hairs. However, the more white there is, the more the black and white will also be separated out into patches instead of being intermingled.

ALTERNATIVE THEORIES FOR BICOLOUR PATTERNS

Above: "Olga", an attractive brown bicolour owned by Sarah Richie

In cats there appear to be two cellular mechanisms involved in white spotting. One involves apoptosis (programmed cell death) reducing the melanoblast (pigment granule) population. In other words, the colour cells migrate over the whole surface of the embryo, but then selectively die out. The other cellular mechanism involves intracellular chemical communication whereby colour cells are biochemically turned off in certain areas. Both of these suggest a chemical gradient so that cells at the extremities tend to be first affected and stop producing colour.

Another hypothesis for the distribution of white was originally presented in papers published by T Terada and T Watanabe in a 1930s Japanese journal. This hypothesis is now being investigated using computer simulation. There are several "unknowns" with the current hypotheses: the relative frequencies of the 10 grades of white spotting given that white spotted cats may be either SS or Ss; the relationship, if any, between birth order and patterning; and whether the "swirled" pattern is related to any birth defects or any difficulties during pregnancy. The melanocyte migration hypothesis may not contain the whole story, especially where swirled patterns, black feet on white socks and skunk stripes are concerned.

The alternative hypothesis is that the patterning of black and white is the result of the skin surface "cracking" during early embryo development. The skin of such bicolour cats would be basically black (or whatever other colour is involved), but the dominant S (white spotting) gene, causes cracks to appear, breaking up the black surface into black domains. These black regions drift apart over the embryonic surface as the embryo grows. The white areas form in the regions between the black domains. It may help to think of it as similar to the movement of the earth's continents with areas of sea inbetween.

It is just as though the white areas were scar tissue produced by the cracking. There are no more melanocytes available to migrate into the scar/white areas as their surface expands. This means that the black domains can end up pushed together to form a single tuxedo style black area (albeit there may be some small white markings in this single black area). The white belly area might be a ventral (belly) seam from a ventral crack earlier on or it could represent the ventral limit of melanocyte migration. It is suspected that the belly expands greatly (megacolon in cats and exposed gut conditions in some rabbits upholds this). Black feet could indicate a black domain that has been pushed to the foot extremity by the expansion of the ventral region at the same time that the limbs are being formed.

Many cats have patches which look as though they could be fitted together like jigsaw pieces; for example a coloured spot on a leg might correspond to an inlet on a coloured patch on the flank, exactly as though a piece of coloured area has broken off and migrated elsewhere. Spots could end up almost anywhere depending on the timing of cracking, the size of the coloured domains, and the trajectories they take over the embryonic surface as it grows. Terada and Watanabe found it possible to fit all the coloured areas of cat coat together. The seams can then be projected onto the surface of a sphere (the embryo). As the sphere expands, the coloured area cracks apart. Think of a balloon covered in solidified chocolate: the balloon can expand, but the chocolate can't. The cracked solid surfaces then gave patterns just like those found on bicolour cats. The swirled patterns in particular fit this hypothesis.

Some cat breeders also believe there are flaws in the conventional melanocyte migration theory. For example a breeder working with American Shorthairs has been crossing Van pattern American Shorthairs to bicolour and spotted Van pattern cats since the mid 1990s. She found that the melanocyte migration theory could not account for some of the spotted patterns these breedings produced. Alternative theories from other white spotting genes and mechanisms in other species (dogs, horse, pigs and goats) also could not account for what was appearing in her cats, particularly with 60%, 70%, 80% and 90% white on a cat. The terminology used differs from the Grades 1 - 9 of the cat fancy; 90% white with a lot of random spots is referred to as Level 15.

Selective breeding has isolated most of the white on the cats to a single layer of white. She can now produce solid coloured cats from the mating of two bi-colours/pied/vanish cats. In addition, all of her best breeding cats have black feet. Breeding experiments involved mating a level 15 stud to a solid black, two Level 7 piebalds, and a Level 9 cat. A cat with a large degree of white is most probably homozygous for the white spotting gene and mating it to other white-marked cats which also carry at least one white spotting gene should not, in theory, produce solid-coloured cats. An alternative hypothesis is that the cats have the dominant white gene (the one related to deafness) and that a second gene is causing this to break down so that spotting appears. It is possible to produce coloured cats from mating two all-white cats since each parent needs only one copy of the dominant white gene to make them all white and can carry masked genes for other colours.

OTHER PIEBALD CATS

There are quite possibly several genes which influence white spotting. For example, the patterns above might not be due to a single gene - there may be several other genes which modify its effects. There are also other piebald patterns which appear to be due to different genes or to cats being homozygous for a modifier gene since the effect is relatively uniform and predictable.

The gloves (mitts) on Birman and Snowshoe cats appear to be due to an incompletely dominant gene. The gloves may vary in length but they are restricted to feet and/or legs. The white markings on the forehead and chest of breeds such as the Snowshoe may also be due to a gene which limits the amount of white. Lockets - those small spots on the throat, chest, stomach and groin of otherwise solid coloured cats also seem to be due to a different gene - one which is normally hidden by the more extensive white spotting of piebald cats. There is also the phenomenon of white toes occurring on otherwise coloured cats.

In the York Chocolate breed, there is a particular white displacement in bicoloured individuals. The placement of white maintains the same configuration in all successive generations. This has been termed "Spotting Particolour" and may be due to an allele (variant) of the White Spotting gene. According to this theory, the White Spotting gene appears to have 4 variants: non-spotted, spotted, particolor, and Birman mitted. Spotted is the dominant form and is variable expressed. Non-spotted is the recessive wild-type and produces a coat without white. The hypothetical Birman allele (or Birman mitted) is also variable, but confines the white spotting to the legs and feet. The hypothetical Particolour allele produces an inverted white "V" with the apex in the centre of the forehead and passing through the centres of the eyes plus a white chin, chest, belly, legs and feet are white. Particolour is also variable and its least expression may be a white locket or white spot on the forehead. The existence of breeds such as the York Chocolate (consistent particolour pattern) and Birman and Snowshoe (consistent mitted pattern) appear to support the existence of a white particolour gene and a white mitted gene.

 

A pattern which has only rarely been reported in cats is the belt, blanket or sheet marking. This is common in pigs and cattle and in Dutch rabbits where a band of white encircles the animal's body like a belt.

Another cause of white spotting is a somatic mutation i.e. some skin cells have a chance mutation which prevents them from producing pigment. This is not hereditary.


Vitiligo in a big cat - the "cobweb panthers" that turned progressively white.


Charva with feline vitiligo (Jason Reeves).

 

There is a rare condition which produces a "cobweb" or "snowflake" effect and is most easily seen on black cats. White spots appear on the coat; these become more extensive with age until the cat has a white lace pattern on the black fur. Ultimately the cat may go completely white or be left with diminishing isolated patches of colour. This condition has been seen in black leopards ("cobweb panthers") and similar conditions are found in humans, dogs and other animals. This must not be confused with the normal sprinkling of isolated white hairs which appear in a cat's fur during its lifetime. It is a rare condition; I come across only 4 cases - 2 relating to black leopards and 2 relating to domestic black cats.

"Leukoderma" is a generic term meaning white skin, or more precisely "piebaldism" because it leads to the formation of white patches in the skin and fur. It is an "aquired depigmentation" i.e. a loss of pigment which occurs during the cat's lifetime, is usually progressive and in some cases is triggered by illness or environmental factors. Ultimately, a cat with leukoderma may become almost entirely white. A type of leukoderma has been identified in some Persian cats and these are used as laboratory subjects in the study of depigmentation conditions.

Vitiligo is one form of leukoderma. Only a few cases have been recorded in cats and it usually causes loss of pigment on the face and feet. Antibodies are formed against the pigment-producing melanocytes. The melanocytes are destroyed leading to the white areas. Another rare condition, periocular leukotrichia, causes the fur around the cat's eyes to become pale - as though the cat is wearing spectacles. "Leukotrichia" means white hair. "Charva" (Jason Reeves) has vitiligo; she was a black cat with patches of slightly longer/softer black hair. In 2009 she turned 4 and developed a patch of white hair. Since then the patches of white have turned into stripes, spots, and patches. Her skin is turning pink as well due to the loss of pigmentation.

In 2011, Kimberly Sexton of New York City sent these photos of her cat Frankie. Kimberly's 14 year old cat Miko (white with grey markings on his head and tail) had recently died and in his last 3 years had formed a close bond with her younger cat, Frankie, a mackerel tabby. The two cats often slept intertwined. Two years before Miko passed away, Frankie developed a white spot of fur under his neck, while his chin and nose became paler. As time passed, Frankie's white areas spread and new ones developed around his neck and behind his shoulder blades. Kimberly joked to friends that Miko's white colour was rubbing off on Frankie as they slept! Frankie's white areas are likely to continue spreading, but the effect appears entirely cosmetic.

On coloured cats, hair regrowth around scars is often white. Some dark coloured cats show white hairs in their coat as they grow older - some follicles stop producing pigment, just as humans get "grey" hairs. Unlike the cobweb effect (where pigment loss occurs in patches, like snowflakes or strands of gossamer), age-related white hairs are evenly sprinkled across the body giving a salt-and-pepper effect; there may also be greying around the muzzle. It is a normal part of ageing although cats seem far less prone to getting grey hairs than do dogs. It should not be confused with the white patches found in vitiligo.

Lizzie Ellis (The Feline Rescue Association Inc, Maryland, USA) provided these photos of a grey male cat with white patches due to scarring. Where the skin was injured or burned, the fur has grown back white, not grey. The cat had suffered frostbite injuries.

SWIRLED PATTERNS AND SKUNK STRIPES

While the piebald pattern is generally fairly symmetrical until the white exceeds 60% of the body surface, some cats exhibit a variety of swirled patterns. This may simply be due to the way the embryo developed or it may be the interaction between different genes which affect white spotting. The two cats below are unrelated, but both have swirled black and markings and black markings on the face. Both also have black toes. Coloured toes sometimes occur on bicolours - these can be individual toes or multiple toes. It may even look as though a white-footed cat has paddled through coloured paint, in which case the colour usually extends up the back of the leg. Coloured toes may possibly be another gene (modifier) interacting with the white spotting gene.Note: The cat shown top left has a mild form of radial hypoplasia which has caused her deformed forelegs; she also has a slightly deformed skull.

Swirled Patterns

 

More unusual is the "skunk marking" - a white dorsal stripe. Because pigment producing cells migrate away from the neural crest, the dorsal area itself usually remains coloured in low to medium grade spotting. The appearance of a white dorsal stripe is unusual and appears to be hereditary.

 

Photos courtesy of Magnificent Munchkins

BRINDLED BICOLOURS AND "TWEED" MUTATION

Pandora (owned by Bill B, Granby, MA, USA) is a brindled cat with what appears to be an unusual mutation. She has the brindled pattern normally seen on tortoiseshell cats, but the patches which should be red are white! The brindling and facial pattern is typical of tortoiseshell cats, but it seems that 15 year old Pandora has a mutation that prevents her producing red pigment. An alternative explanation is that Pandora is chimera formed when a black embryo and a white embryo fused in the womb; however chimeras tend to have a patched appearance rather than being thoroughly brindled, this makes chimerism a less likely explanation. A third possibility is that Pandora has additional X chromosomes (XXX or XXXX instead of the normal XX female), however this genetic anomaly is associated with mental retardation and physical anomalies/deformities (this is not Klinefelter syndrome, Klinefelter syndrome is only found in males e.g. XXY or XXXY). These two conditions can be identified using tissue samples, but this isn't recommended in an older cat as it involves sedation or anaesthesia. It seems likely that the gene that should produce red pigment is faulty or that some other gene is masking the red pigment out. Since Pandora was spayed before Bill obtained her, it isn't possible to breed her to see if it can be inherited. I have seen the opposite mutation - where a tortoiseshell cat produced red pigment, but not black, resulting in a red, cream and white brindled cat - but this is the first time I have seen a black-and-white brindle where the red is absent.

More brindled bicolours are explained in the separate page on Roan, Tweed and "Salt and Pepper" Colours

THE CHINESE HARLEQUIN

During the 1980s, there were attempts to breed shorthaired cats with a black tail, black patches on the head and small black (or other solid colour) patches or spots on the body. Named Chinese Harlequin, it was to resemble cats found in ancient Chinese art. Although still listed by some registries, it appears to now be extinct due to the difficulty of breeding bicolours with consistent spotted markings. A similar "strikingly spotted" cat, the Gao Taem, is seen in ancient Thai art and is described as having, black marked forepaws, black and white ears, black shoulders, two black spots on the back and black shoulders. In total there should be nine horse-like black spots on an all white background. If it existed as a breed, the mutation creating these features has been lost. In all probability it was a form of Seychelles pattern. Some bicolours have black toes or paws and and some breeders have attempted to fix this trait to create a black-footed Van-type bicolour.

Also depicted in Thai art (1676) is the Vichiens Mas which is shown as white with dark ears, nose, paws and whiskers. Although depicted as black and white, it seems to be a stylised or idealised depiction of the seal-point Siamese.

"BELTED" OR "SHEETED" PATTERN

"Belted" means a solid coloured animal with a belt of white around its middle, such as is seen in Dutch Rabbits. This pattern has occasionally been seen in Spanish feral cats. "Sheeted" means a wider the band of white i.e. from shoulders to haunches.

THE "DALMATIAN" OR "APPALOOSA" CAT

The concept of a white cat with coloured spots, akin to a Dalmatian dog or Appaloosa horse, has intrigued numerous people. In some feral colonies, white cats with numerous small black splashes have been found alongside bicolour cats with the conventional Grades 1 - 9 of white spotting. This suggests the presence of modifier genes which affect the distribution of colour and white. Inbreeding causes genes to double up so that recessive genes and traits controlled by multiple genes become visible. It would be possible to have genes which interact with the semi-dominant white spotting gene such that small patches of colour break through the white or which cause a breakdown in white spotting.

Some breeders report a consistently high incidence of numerous small splashes of colour on the body in certain breeding lines which suggests to them that some other gene(s) is being inherited alongside the white spotting gene. This may see a repetition of the Chinese Harlequin programme.

MESSYBEAST : COLOURS, CONFORMATION & FUR TYPES