Cats (0)

AMBER AND RUSSET - LATE COLOUR CHANGE GENES
Copyright 2014, Sarah Hartwell
The ancestors of the domestic cat were nondescript black / brown striped tabbies. Over the centuries , mutation produced a wide array of colours based on 2 different pigments. Eumelanin gives the blacks, browns and blues while phaeomelanin gives the reds, fawns and creams. A few other genes give further variations on those colours such silvers, colourpoints and solids/selfs. Mutations continue to occur and unexpected colours also turn up due to inbreeding where recessive genes, hidden for generations, start showing up.
AMBER AND LIGHT AMBER
During the 1990s , some purebred Norwegian Forest Cats in Sweden produced chocolate /lilac and cinnamon /fawn offspring. However , those colours are not found in the purebred Norwegian Forest Cat gene pool. Had the gene pool become polluted by someone , perhaps generations ago, breeding their Norwegian Forest Cat to another breed? Was it a spontaneous mutation? Crossing of those cats with known chocolate and cinnamon colour cats of other breeds ruled out chocolate/lilac and cinnamon/fawn genes. These cats were a totally new colour, peculiar to the Norwegian Forest Cat gene pool and dubbed the "X Colours". They are now called Amber and Light Amber. The Amber effect is due to the extension gene (also called red factor ) which controls the production of red and black pigment . The dominant version of the gene produces normal black pigment in the coat while the recessive version produces red pigment. The name comes from the effect of black or brown pigment not being extended throughout the whole coat, but being restricted to the skin of the extremities and to the eyes (for example in bay horses ).
This Norwegian Forest Cat was bred by Yve Hamilton Bruce from a silver mackerel tabby female (imported from Denmark) and a classic red tabby and white male . The result was 1 silver tabbies and 2 silver tabbies with white. At just over 3 months old, this silver and white tabby male developed a large patch of bright red hair on his back which continued to spread . Eventually the whole fur will become amber. The effect of amber during the colour-change stage depends on the original colour - solid black or blue , bicolour or tabby. The cat pictured is not a typical amber as it has the silver gene so the amber effect is overlaid on silver.
A non-agouti amber Norwegian forest Cat resembles a silver tabby, but has a distinctive black nose and black paw-pads instead of the pink /reddish nose with black outlining found on conventional silver tabbies. The photo below shows a non-agouti amber. According to Rui Pacheco, although this cat is clearly non-agouti because of the black nose, he has the phenotype of a silver shaded. Rui's theory is that the amber gene invalidates the action on the non-agouti gene; wide-banding will further lighten the base colour giving a very pale cat with black nose and paws. Amber cats are basically black cats with a colour modifier that affects the deposition of pigment on the hair shaft, but not the skin colour. That is why in non-agouti cats the nose remains black and in agouti cats it remains pink.
PLATINUM
The platinum effect is found in some lines of Silver Persian/Chinchilla Longhair. In 1986/7, Cheryl Bennett reported in The Silver and Golden Persian Newsletter (ACFA) that one of her shaded silver females (Kelley Lane Contessa of WeANDE) had changed from pure silver to pale golden. Contessa’s parents were a shaded golden and a shaded silver; these were full siblings. At 10 – 12 months of age, Contessa began to “tarnish” i.e. show cream /reddish patches and by 3 years of age she was entirely pale golden. She produced a number of Silver Persians without tarnish, but she failed to produce any offspring when mated to a golden sire . One of Contessa’s male offspring turned from silver to golden as an adult . An adult Silver Persian from England (Lynchard Silver Shadow) was exported to Australia and also turned golden. Shadow had a few golden hairs on his paw, but did not turn golden until he was a year old when his coat turned to pale beige. By 3 years old he was entirely pale golden. Shadow was bred to a genetically golden female, but the pairing only produced silver offspring. However, at least one of his silver offspring later turned pale golden. Other descendents of Contessa also went through the late colour change. During the 1980s, several other breeders of Shaded Silver and Chinchilla Persians came forward to report that their cats had developed reddish, brownish or golden-coloured fur along their spines as they aged . Many of the cats had no golden in their ancestry. At first it was dismissed as an unavoidable genetic fault where silver was incompletely dominant and did not hide the recessive golden colour.
A common ancestor of all the colour- changing Silver Persians was a stud cat called Kelly Lane Andromeda (in the UK) whose descendents were exported to the USA in the 1950s and 1960s . Some of those descendents were influential stud cats and would have spread the mutant gene far and wide. Controlled inbreeding (linebreeding) helped establish the gene, which would later double up to produce colour-changing Silver Persians. The effect of this gene on Golden Persians, if indeed it has any effect, is not known.
Except for red-silvers and cream-silvers, silver cats should entirely lack phaeomelanin (red or cream pigment) and only have eumelanin (black, brown, blue etc) present . Perhaps some other gene causes the eumelanin structure to change so that it is perceived as a golden colour. Chemical analysis proved that the late colour change Silver Persians did not have phaeomelanin pigment present.
AMBER-TYPE EFFECT IN MANX
An amber-type effect has also been seen in a Manx cat. Janet McArthur bred a male Manx (a longy i.e. with tail ) that was born an ordinary black/brown mackerel tabby without much rufousing, as was his sister . In addition , the male had white ticking at the tips of his hair. As the male grew, his stripes faded out and became spots while the black markings also changed from black to a burnt red-brown colour and his white ticking became smaller. The change from stripe to spots was probably an optical effect caused by the change from black to red-brown. His dorsal line became a burnt golden brown, and the background colour was similar to a washed out sorrel colour (sorrel is a colour found in Abyssinians).
There are a few oddities in his ancestry. His mother , Pearl , was a black Manx and his sire was a normal black/brown tabby from the same black Manx mother. The inbreeding was accidental , and may have brought out a recessive gene. Pearl was born to a black/red tortie (carrying dilute and colourpoint) and a red van-pattern sire; this pairing should not produce solid black females so, barring an anomaly in her sire's germ line, Pearl may be a tortie where the red has not been expressed in the coat. Pearl has also produced a kitten that changed from black with white ticking at the hair-tips, to black smoke and then to solid black without ticking.
BIMETALLIC / SUNSHINE
The informal term "bimetallic" (referring to a mix of silver and golden or silver-gilt) describes some Siberian silver tabbies that turn golden in a rather patchy fashion. Unlike the amber gene, the colour change does not start on the back and work downwards, but seems distributed throughout the coat. The formal name "sunshine" has been proposed for this emerging gene.
RUSSET
Similar to amber is russet, which turned up in a line of seal (brown) European-style Burmese in New Zealand in 2007. It has subsequently occurred found in the related Mandalay (similar to the Asian in Europe ) and appears to be a mutation of the extension gene. The first known russet was a pure-bred Burmese called “Molly” in 2007. There is now an experimental programme in NZ to breed Russet Burmese and to investigate dilute russet, russet tabby and solid russet (as opposed to the Burmese sepia form of russet).
The first russet kitten “Molly” was born an "odd-coloured lilac (lavender)" which gradually lightened as she grew, progressing through lilac-caramel and chocolate ticked tabby and then dramatically changing to red. Chocolate ticked tabby and red were both impossible from her pedigree, and in any case , reds are not born as chocolate tabbies! Several more unusually coloured kittens were born in different litters and all went through the same colour/pattern changes . The ancestors of these kittens were seal Burmese and had no silver, tabby or Mandalay (Asian) blood . The pedigree had both dilute and chocolate, there were no reds, creams or torties until 4 generations back. DNA testing showed some kittens to be genetically seal and other genetically chocolate. Hence the new colour is due to a different mutation currently known as russet. Russet appears to be due to a recessive mutation that causes black pigment (eumelanin) to gradually fade to a minimal amount while leaving red pigment (phaeomelanin)unaffected.
Russet kittens resemble tabbies at birth , but have pink noses and paw-pads, pale fur around the pads and genitalia and a pale tail-tip - all of which would be dark in tabbies. The muzzle and fur around the eye is ivory. The back is solidly dark rather than ticked, becoming pale ivory halfway down the flanks. The back becomes more ticked appearance, almost a saddle, as kittens undergo the colour change and the face becomes reddish. By age two, they may resemble a red Burmese. It differs from amber as ambers have dark noses and paw-pads. Non-agouti (solid) amber kittens are very dark with a dark face that is last to go red while russet kittens have off-white faces (possible due to Burmese sepia gene in the mix), which are the first part to go red (rather than the last as in ambers), and pale undersides. The russet colour change appears to be slower than the amber colour change. Russet kittens to date have been larger at birth than their siblings and somewhat on the large side as adults.
RECESSIVE BROWN - THE ENIGMATIC BARRINGTON BROWN GENE
Copyright 2010 Sarah Hartwell
This page pulls together what is known about Don Shaw 's Barrington Brown gene, a form of recessive brown/colour dilution only reliably recorded in a colony of laboratory cats, none of which are believed to have left the laboratory. To make this comprehensible to the non-genetics expert I have referred to "copies of genes" or "versions" of genes although the correct terminology is "alleles". There is also a brief guide to Shaw's terminology at the end as Shaw's writing pre-dated modern "standard" symbols and terminology.
Don Shaw was an early feline geneticist in the USA. During the 1950s and 1960s, there was no standard form of genetic coding and Shaw used his own system of genetic coding which can be difficult to read today . He also referred to chocolate, which is a mutation of the black gene, as "chocolate dilution". What modern fanciers call dilution, Shaw called "maltesing" (Maltese i.e. blue cats were a genuine dilution of black). Shaw viewed dilution as being due to the reducing amount of melanin present in the hair and not by the way pigment forms clumps in the hair. Chocolate had less melanin than black, therefore Shaw called this dilution. Blue has a the same amount of melanin, but arranged differently, so Shaw did not consider this dilution.
In simple terms , the black (eumelanin) colours in cats is due to a series of genes (alleles to be precise) with the following order of dominance/recessive (most dominant at the top, most recessive at the bottom ). The chart also shows how these combine with dilution genes and dilution modifier genes. A cat inherits 2 copies of the black gene and only the more dominant version will show up while the recessive gene will be hidden. The caramel form only shows up in cats that already have the dilute colour and it lightens and adds a brownish cast to the dilute colour.
ORIGINAL COLOUR
DILUTE VERSION
CARAMEL VERSION OF THE DILUTE VERSION (DILUTE MODIFIER, DOUBLE DILUTION)
Black
Blue
Caramel (UK: Blue-based caramel)
Chocolate
Lavender (Lilac)
Taupe (UK: Lilac-based caramel)
Cinnamon
Fawn
UK: Fawn-based Caramel
These colours will naturally look a little different on Burmese (due to the sepia gene), Tonkinese ( mink gene) and cats with the silver Inhibitor gene. For clarity, this article will ignore those additional genes and just look at the eumelanin (black) series of colours.
An additional type of recessively carried brown colour has been reported in laboratory cats, but not in the outside world. Termed "Barrington Brown", a cat with two copies of this gene had black pigment diluted as shown below. Because it hasn't (yet) been seen in the cat fancy , this gene is enigmatic to many fanciers.
Some have suggested it is the same as caramel (dilute modifier), however some reports indicate cats with 2 copies of the Barrington Brown gene were different in colour from cats with only one copy of the gene (which seems odd since the gene is recessive and cats with only one copy should have dilution at all). When 2 copies were present, Barrington Brown had an additive effect on black and chocolate, and presumably on cinnamon (which may have been unfamiliar to Shaw).
ORIGINAL COLOUR
BARRINGTON BROWN DILUTION
Black
Deep mahogany brown
Chocolate
Light brown
Cinnamon
Cafe -au- lait ( milk coffee colour)
The dilution only happens if 2 copies of Barrington Brown are present, although some sources suggest the genes had additive effect on each other to create the pale milk coffee coloured cats when there were 2 Barrington Brown genes. Since Barrington Brown is a recessive gene. If only one copy was present it should be masked by the dominant non-Barrington copy. This is part of the problem when interpreting reports that use an old, non-standard genetics notation!
ba - Barrington Brown Locus (unverified)
Genotype
Description
Phenotype
Ba/Ba
Homozygous - Non Barrington Brown
Cat unaffected - i.e. Black/Brown/Chocolate etc
Ba/ba
Heterozygous - Non Barrington Brown - carrying Barrington Brown
Cat unaffected - i.e. Black/Brown/Chocolate etc
ba/ba
Homozygous - Barrington Brown - liberty of renaming
Mahogany Brown/Light Brown/milk coffee in colour (depending on whether cat is black/chocolate/cinnamon)
In the 1960s, chocolate was described as a dilution of black and the third possible version of the gene, cinnamon, was apparently unfamiliar to Shaw (or not reflected in his terminology). Shaw’s Barrington Brown article was first written in the 1960’s in the Journal of Cat Genetics, and reprinted in the early 1970’s in Cats Magazine and has been subject to much reinterpretation, especially when a dilute modifier emerged in the cat fancy.
According to Shaw's breeding data, Barrington Brown (mahogany brown) dilution was inherited in much the same way as “standard chocolate dilution" (black/chocolate alleles) by reducing the amount of pigment in the hair and by producing elliptical pigment granules instead of round granules. Elliptical pigment granules refract light differently and give a reddish-brown colour instead of black/sepia. In Shaw's terminology which can confuse modern readers, chocolate is a dilute of black (while blue is "maltesing" of black).
It became apparent that Barrington dilution gene was in a different location to the ordinary black/chocolate genes and was inherited independently of black/chocolate. That means it wasn't the modern cinnamon. Genes in different locations can affect different enzymes involved in production of the same protein, in this case the production of eumelanin pigment. Shaw referred to the “standard chocolate dilution" as affecting enzyme D while the Barrington system affected enzyme B. He identified Barrington Brown as having 2 alleles; the dominant wild type and the recessive Barrington Brown dilution.
B+ = Wild type gene. Apparently responsible for normal Enzyme B production, giving full intensity of melanin.
b = Barrington Brown recessive gene. When 2 copies are present there is less Enzyme B produced which means less melanin produced; the pigment granules are elliptical (reddish-brown) instead of round (black).
He gave it the name Barrington Dilution or Barrington Brown because it was discovered at the Quaker Oats Nutritional Laboratories in Barrington, Illinois. The first cat known to be homozygous for Barrington Brown was born at the Barrington Laboratories. This was a strangely colored light brown ( rich caramel or perhaps cafe-au-lait) male kitten homozygous for Barrington Brown as well as having “standard chocolate dilution". The 2 gene systems were additive in nature and each expresses its effect as if the other were not present i.e. the end result is the sum of both Barrington Brown and "standard chocolate dilution" being expressed. Shaw's description of "rich caramel" (light tan) misled 1970s breeders to believe the emerging dilute modifier was the same as Barrington Brown.
Don Shaw and Wayne Durdle investigated this gene in depth. Cats carrying the Barrington gene were donated by the Quaker Oats Nutritional Laboratory to the Feline Research Laboratory at Tuskegee Institute, Alabama (maintained there from summer 1966 to mid-1969). No Barrington Brown carriers were believed to exist outside of the colonies and Barrington and Tuskegee. Studies established that the light tan or cafe-au-lait colouration was due to the combined effects of "standard chocolate dilution" and their Barrington Brown gene.
Shaw had left the project in July 1968 and Durdle reportedly terminated the colony in June 1969 (this suggests the cats were destroyed, neutered or used in other research). To their knowledge - and to the ongoing chagrin of many breeders - the Barrington Brown cats had no direct descendents in the cat fancy and the gene was lost . However, Shaw noted that some "doubly diluted" kittens were appearing in the cat fancy. These had no apparent connection to the Barrington or Tuskegee cats. He also mentioned unconfirmed reports of a dilution system similar to the “standard chocolate dilution" but not traceable to any known chocolate ancestry.
Shaw offered the following possibilities:

• a leak of the Barrington Brown gene from one or other of the laboratory colonies into the cat fancy population; perhaps some cats or kittens had been taken home as pets and bred.
  
• a new mutation in a completely different gene location
  
• a new mutation at the same location as his Barrington Brown gene
  
• an independent recurrence of the Barrington Brown mutation.
  

To identify the relationship , if any between the Barrington Brown cats and those with "double dilution" would require test-matings between the new "doubly diluted" cats and some known Barrington Brown cats. Since there were no known Barrington Brown cats outside of the laboratory and the lab colony had been terminated, this wasn't possible. In the 1970s, Pat Turner stated , without any supporting evidence , that caramel (the double dilution mentioned by Shaw) was the same as Barrington Brown. The dilute modifier (caramel) colours are not the same as the colours described by Shaw, though some websites continue to repeat Turner's claim that Shaw called the caramel colour Barrington Brown. Turner had simply jumped to conclusions. Barrington Brown cannot be the same as caramel because Barrington Brown affected black and chocolate, but caramel only affects blue, lilac and cream.
Shaw noted that recessive traits emerge in pedigree cats because of line-breeding and inbreeding which matches together genetically similar individuals. This means a better chance of cats inheriting 2 recessive versions of a gene and new traits showing up as result - just as they did in the Barrington colony of cats.
From a breeder viewpoint, it is sad that "recessive brown" has been lost. Shaw's descriptions indicate that it wasn't the same as caramel. Its effects on the wider palette of feline colours - torties, ticked and patterned tabbies, Burmese sepias, minks and colourpoints - can only be hypothesised. From time to time there are reports of odd colours in cats, including a tantalising "palomino" described as "the colour of a brown paper grocery bag" from the USA that might just have been the light tan noted by Shaw.
SHAW'S TERMINOLOGY
Modern feline geneticists use b/b for chocolate, but Shaw defined chocolate as d/d, because to him it was a dilution of black. Reading Shaw's work on black, chocolate and Barrington Brown without knowing his terminology is therefore confusing. In Shaw's defence, the standard gene symbols had not been defined in the 1950s and 1960s.
Light tan/cafe-au-lait in Shaw's terminology is d/d, b/b. To a modern reader, d/d, b/b indicates lilac. This, as much as any visual similarity, helps explain why his Barrington Brown is so often confused with the dilute modifier.
Shaw’s Chocolate alleles
D+- = Normal colour (black/sepia)
d/d = chocolate (in Shaw’ s words , Chocolate Dilution)
Shaw’s Maltese alleles
M+ = Dense colours e.g. black
m/m = Maltese (or in our terms dilution) e.g. blue is the maltese of black
Shaw's Barrington Brown allele
B+ = Normal coat
b/b = Barrington Brown
Lilac (in Shaw's terminology) would be d/d, m/m = Dilution allele (chocolate) + Maltese allele = lilac. 
Lilac in modern terminology is b/b, d/d = Chocolate allele + dilution allele = lilac.
Although there is no official modern symbol for the lost Barrington Brown, Ba is often used for convenience in modern discussion (since it isn't allocated to anything else ).
Brown Locus (modern symbols)
B/- = Black/Brown (Shaw's black/sepia)
b/b = chocolate (Shaw's "standard chocolate dilution")
bI/bl = cinnamon (probably not recognised by Shaw)
Barrington Brown Locus (modern symbols)
Ba/Ba - no Barrington Brown dilution
ba/ba - Barrington Brown dilution/recessive brown
THE BASIC SELF (SOLID) COLOURS OF CATS
Copyright 2002 - 2013 Sarah Hartwell
Genetically speaking, there are four basic self (or solid) colours of cats: black, chocolate, cinnamon and red. All other self colours are modifications of these. Although covered here as a self colour, red is a form of tabby and it is impossible to completely eliminate the tabby markings. Why are there not five basic colours? White is counted as an absence of colour rather than a colour.
Different countries, registries and breeds have different names for some of the same basic colours. Even where the same name is used, there may be different views on what is an acceptable or ideal version of that colour. Colours which appear identical to the human eye are caused by different genetic interactions.
The same colours are called by different names in different breeds. Even in the same breed, the colour may have different names depending on which country the cat comes from and which registry it is registered with. American registries like to add "mink" after the Tonkinese colours whereas British registries use the same name for that colour as is used in the equivalent Siamese or Burmese colour. Confused? Don't worry - there are some cross - reference tables later on!
The jet-black colour you known as "black" is called " ebony " and "ebony tabby" in Orientals, "black" in solid coloured domestic shorthairs, "brown" when it refers to brown tabby domestic shorthairs, "bronze" in Egyptian Maus, "tawny" in Ocicats and "ruddy" in Abyssinians. In colour-pointed cats, "black" is called "seal". In Burmese it is " sable " or "seal sepia" and in American Tonkinese it is "cinnamon" or "natural mink". In the Asian breed (self Burmese cats) it has a breed name to itself "Bombay". Shaded silvers, black smokes and chinchilla cats may look various shades of grey or silver, but they are black cats with silver roots to their fur. Add dilution and it becomes "blue". Modify the dilution and it becomes "caramel". Yet it is still basically a black cat.
WHY RED SELF CATS ARE STILL RED TABBIES
Strange as it may seem , all red cats are actually red tabby because the non-agouti gene (the gene that turns a tabby into a self/solid colour) does not affect the way red pigment is deposited. A variety of other genes, called polygenes or modifiers, control the intensity of colour and contrast between markings and background colour. Selective breeding has reduced the tabby markings to produce a cat that looks solid red by breeding from those cats with the least red markings (cats with "low contrast" between markings and background colour). Because the non-agouti gene does not work on the red pigment, red tabby ghost markings can never be completely eliminated and may be seen on the tail, legs and forehead and as a darker region along the spine . Even though red cats are registered as "red self" they are still red tabbies, albeit red tabbies with very reduced markings. Because they lack the polygenes for high contrast between markings and background colour, their offspring also appear to be red self. More information can be found in Robinson 's "Genetics for Cat Breeders".
DILUTION AND MODIFIED DILUTION - MALTESING AND CARAMELISING
The simplest modification of the 4 basic colours is dilution. As the name suggests, this "washes out" the original colour to something paler. This is also called maltesing since the it was first identified in black cats and blue cats - Maltese is term for blue-grey and some of the eary blue cats imported into Britain were known as Maltese cats.
A secondary type of dilution is called the dilute modifier. It only affects already diluted colours. Since it gave rise to the colour "caramel". I have referred to it here as "caramelising" purely to avoid confusing the layperson. Textbooks always refer to it as the dilute modifier.
ORIGINAL COLOUR
DILUTE VERSION
CARAMELISED VERSION OF THE DILUTE VERSION (DILUTE MODIFIER, DOUBLE DILUTION)
White
N/A
N/A
Black
Blue
Caramel (UK: Blue-based caramel)
Chocolate
Lavender (Lilac)
Taupe (UK: Lilac-based caramel)
Cinnamon
Fawn
UK: Fawn-based Caramel
Red
Cream
Apricot
Amber
Light Amber
 
Black, chocolate and cinnamon are all versions of the same gene. With the exception of "red" which is a special case, genes are inherited in pairs - one from each parent . Some genes are dominant over others and only the dominant one will be expressed (show up). The other gene (the recessive) will still be lurking in the background and can be passed on to offspring. Depending on which pairing a cat inherits, it will be one of those basic colours. The fact that at it may look different depends on many other genes which alter the way in which these three basic colours are expressed.
If it inherits black + black OR black + chocolate OR black + cinnamon it will be black.
If it inherits the dilution gene it will be blue.
If it inherits the dilution gene AND the caramelising gene it will be caramel.
If it inherits chocolate + chocolate OR chocolate + cinnamon it will be chocolate.
If it inherits the dilution gene it will be lavender.
If it inherits the dilution gene AND the caramelising gene it will be taupe.
If it inherits cinnamon + cinnamon it will be cinnamon in colour.
If it inherits the dilution gene it will be fawn.
If it inherits the dilution gene AND the caramelising gene it will be a pink-brown..
Many early caramels were probably registered as fawns or lilacs, probably as poor quality individuals. The colour differences are subtle enough that it may be necessary to check what is in a cat's pedigree to work out the exact colour. To the rest of us, such cats are simply "biscuit colour" and are no less attractive for it!
Red is a different gene to black, chocolate and cinnamon. It is a sex- linked gene which means a female must inherit 2 copies of the red gene in order to show up as a red cat. A male only needs one copy of the red gene to be a red cat. This is why ginger tomcats are more common than ginger females (though contrary to popular belief , ginger females are neither rare nor infertile!). If a female only inherits one red gene, she is a tortoiseshell.
If a female inherits red + red she will be red (ginger).
If she inherits 1 red gene she will be tortoiseshell (red/black, red/chocolate or red/cinnamon depending on the other genes present)
If she inherits red + red AND the dilution gene she will be cream.
If she inherits 1 red gene AND dilution gene she will be dilute tortoiseshell (blue/cream, lilac/cream or fawn/cream)
If she inherits red + red AND the dilution AND the caramelising gene she will be apricot.
If she inherits 1 red gene AND dilution gene AND the caramelising gene she will be dilute tortoiseshell in the caramelised range of colours (though this may not show up very well and may be impossible to identify.)
If a male inherits 1 red gene he will be red.
If he inherits 1 red gene AND dilution gene he will be cream.
If he inherits 1 red gene AND dilution gene AND caramelising gene he will be apricot.
In reds, there are genes for rufism i.e. for the depth of the red colour. This is why show-quality reds are a rich, deep red colour while alley cats are more often marmalade or ginger. Early reds (1880s) were known as yellows; the depth of colour was improved over many generations of selective breeding.
In 1924, a series of breeding experiments between a Siamese female and a tabby male resulted in black offspring that themselves produced tabby offspring. This suggests a gene for black that is dominant to tabby, the opposite of the known behaviour of tabby and black! It is possible that the Siamese female, one of a pair imported from Bangkok , had a mutation for black colour that was dominant instead of recessive. There have been no further reports of dominant black in the cat population. Dominant black is genetically different to the black colour described above , but unless it occurs again its interaction with other genes will remain unknown.
Note : The usual explanation of black (B), brown/chocolate (b) and cinnamon (bl) is that there are 3 alleles of the black locus gene. Chocolate carrying cinnamon can appear as light chocolate in some breeds, but basically there are 3 phenotypes: black, brown/chocolate and cinnamon. An alternative explanation suggests 2 gene loci, each with two alleles. One locus is black/brown. The other locus is a modifer that, when homozygous (2 copies are present), changes black to cinnamon and brown to pale cinnamon. This gives 4 phenotypes: black, brown/chocolate, dark cinnamon/light chocolate and light cinnamon.
An additional type of recessively carried brown colour was reported in laboratory cats in the USA in the 1960s, but not in the outside world. Termed "Barrington Brown", a cat with two copies of this gene has black diluted to deep mahogany brown, chocolate dilutes to light brown and cinnamon dilutes to a pale cafe-au-lait. There are suggestions it may have been the caramel gene, but the descriptions of the colours quite different. The confusion seems to stem from the discovering describing the light tan of the Barrington Brown cats as "caramel coloured". Barrington Brown cannot be the same as caramel because Barrington Brown affected black and chocolate, but caramel only affects blue, lilac and cream.
CARAMEL
The Dilute Modifier gene was posited by Patricia Turner who erroneously believed it to be the same as the Barrington Brown gene reported in a colony of laboratory cats. In essence, caramel lightens and gives a brownish cast to the underlying colour. How can adding brown make something lighter? Brown (in terms of colour, not genetics) comes in a variety of shades ranging from fawn to chocolate and adding a brownish hue is not the same as mixing paint . With the dilute modifier, blue becomes caramel, lilac becomes taupe (brownish grey) and red becomes apricot. In practice , these colours may only be visually distinguishable from each other by knowing the cat's genetics. Some find caramel cats resemble golden series cats and suggest that a theoretical hypostatic (hidden) silver/golden gene is showing through, especially in the lighter colours where there is nothing to mask a hidden silver/golden colour.
In the past, the Dilute Modifier has also been called “double dilution” and “caramelising”. Some people are still not convinced about caramel because almost identical colours can result from blue + caramel and from lavender/lilac + caramel. With a modifier gene, it is possible to get the same end colour (visually) from 2 different base colours (genetically). It depends on what the Dm gene is modifying (the expression of a protein) which is different from mixing paint. Two different original proteins (relating to melanin production or deposition) could be modified by such a gene to produce almost identical expression.
EXTENSION GENES (BLACK MODIFIER) - AMBER/LIGHT AMBER AND RUSSET
The Extension gene (formerly the Black Modifier gene) brightens black areas of the coat. At birth, kittens appear to be black or blue (it has not been found in combination with other genes recessive to black), but become brighter as they grow. The Black Modifier has so far only been observed in the Norwegian Forest Cat. Recognition of the colour by registries will allow the cats to be correctly registered and distinguish them from chocolate, lilac, cinnamon and fawn (not found in Norwegian Forest Cats).
COLOUR
WITH EXTENSION GENE/BLACK MODIFIER
Black
Amber
Blue
Light Amber
Chocolate
Lilac
Cinnamon
Fawn
 
During the 1990s, some purebred Norwegian Forest Cats in Sweden produced chocolate/lilac and cinnamon/fawn offspring. Because these colours are not recognised (they indicate outcrossing) they were called "x-colours". Crossing a "cinnamon spotted" x-colour with a fawn Somali produced blacks and blues, not fawns. Some x-colour cats were registered as "golden", but the x-colour can be found in combination with silver (x-colour silver tabbies), ruling out golden: a cat can be either silver or golden, but not both. The offspring of an x-colour "cinnamon spotted" and a chocolate point Birman were black and blue tabbies, ruling out the recessive chocolate, lilac, cinnamon or fawn and ruling out a recessive masking factor (i.e. that found in colourpoint cats). To produce black and blue offspring, the x-colour must be genetically black (dominant to chocolate). Further test-matings upheld these conclusions.
Self (solid) x-colour cats were born as poorly coloured black-silver or blue-silver tabbies. Their tabby ghost-markings faded as they matured and the colour became bright apricot to cinnamon colour with dark brown paw pads and nose leather with no black rim (a black rim is characteristic of silvers). Mating a self x-colour cat to a black-and-white and mating two self x-coloured cats together showed that the colours were not agouti (not ticked), but were new colours, now called Amber and Light Amber. Kittens are born dark and undergo a period of extreme brightening of the black/blue areas as they mature. Their original birth colour is often seen only on the back and tail, allowing amber and light amber to be distinguished from one another.
Amber is apricot-to-cinnamon colour with brown paw pads, nose leather and eye rims. Kittens are born dark or black, with ghost markings, and brighten as they mature. Light Amber is a pale beige colour. Kittens are born blue and brighten with age, becoming pink-beige to fawn at maturity. The nose leather, eye rims and paw pads are dark blue grey. (To complete the description of amber/light amber: Amber Tabbies are born apricot with black markings; the markings brighten to reddish-brown/cinnamon at maturity. The nose is pink and the paw pads and eye rims are brown. Light Amber Tabbies are born beige with blue tabby markings; the markings brighten to pink-beige/fawn at maturity. The nose is pink and the eye rims and paw pads are blue-grey. Amber/light amber replaces black/blue in torties. Amber also occurs with silver and in bicolours/tabbies -and-white.)
The provisionally named Russet Burmese is an experimental Burmese colour in New Zealand. In 2007, Nicki and Bob Mackenzie's line of seal (brown) Burmese (carrying dilute and chocolate) produced "odd-coloured lilac" kittens which gradually lightened as they grew, progressing through chocolate ticked tabby to red. The kittens traced to the same father and two related females (mother and daughter ) and all three shared a common ancestress only a few generations back. DNA testing showed the russet kittens to have the standard seal or standard chocolate genes, meaning there is an a different gene causing the colour change. Russet kittens also tended to be larger than their littermates, both at birth and as adults. Russet appears to be a recessive gene from a spontaneous mutation. It causes the black pigment (eumelanin) to gradually fade to almost nothing while leaving the red pigment (phaeomelanin) unaffected. Like the amber colour change, it might be an extension gene mutation.
Russet kittens have pink noses and paw-pads and a pale tail-tip. The muzzle and fur aroundthe eye is ivory. The back is solidly dark rather than ticked, becoming pale ivory halfway down the flanks. The back becomes more ticked appearance as kittens undergo the colour change and the face becomes reddish. By age two, they may resemble a red Burmese. It differes from amber (in Norwegian forest Cats) as ambers have dark noses and paw-pads. Non-agouti (solid) amber kittens are very dark with a dark face that is last to go red while russet kittens have off-white faces (possible due to Burmese sepia gene in the mix), which are the first part to go red, and pale undersides. The russet colour change appears to be slower than the amber colour change.
PINK- EYED DILUTION
The type of dilution seen in cats is blue dilution (it dilutes black to blue). A second type of dilution seen in many mammals is "pink-eyed dilution". Pink-eyed dilution is characterised by a pink or ruby glimmer to the eye (depigmentation). The coat colour is often diluted to bluish-fawn and the pink-eyed dilution factor is generally inherited as a recessive gene. A possible pink-eyed dilute female cat was reported in 1961. She was described as pink-eyed with a light tan coat. She was mated to a chocolate point Siamese and produced three tabby kittens 10 days premature. Sadly none of the kittens survived. The colour of the kittens implies that pink-eyed dilution in cats is inherited as a recessive trait and is independent of the colourpoint genes. The pigment granules in the hairs of pink-eyed dilute cats were very small and yellowish brown,instead of the normal dark-brown or black. ( Todd NB: A pink-eyed dilution in the cat. JHered 52:202, 1961.27.)
ALBINO, DOMINANT WHITE, WHITE SPOTTING
Albino is generally thought of as pure white, but the situation in cats is more complex . There are five known alleles for albinism : blue-eyed albino, pink-eyed albino, Burmese pattern, Siamese pattern and full colour (non-albino). Full colour is dominant to all of the other four alleles. Burmese pattern is incompletely dominant to Siamese pattern; cats that inherit one of each of those genes will be intermediate in pattern and is known as Tonkinese. A quirk of the Siamese form of albinism is that it is temperature dependent with warm areas of the body being paler than cooler areas. For this reason , it is often described as "colour restriction" rather than albinism. Pink-eyed albino appears to be recessive to all of the other albino mutations.
The albino cat reported in Europe and the USA seems to be intermediate between pink-eyed albino and blue-eyed albino. Although it has the white coat of a true albino, its eyes have ruby red pupils and pale blue irises. A true pink-eyed albino was reported in 1931 and again in 1980s in the USA. Albino kittens have turned up more recently in the Bengal breed, unsurprising since albinism is found in the Asian Leopard Cat (the wild parent of the Bengal).
Dominant white is the colour associated with deafness in cats. Dominant white masks all other colours and cats may have blue, orange or odd eyes. Those with blue eyes have a high chance of deafness. Those with one blue eye have a high chance of deafness on the blue-eyed side. Those with orange eyes are far less likely to be deaf . Some dominant white kittens are born with smudges of coloured fur on top of the head, this smudge of colour usually disappears by adulthood, but kittens with colour smudges are more likely to have normal hearing.
The gene for white spotting can also create the impression of a self white cat. This gene is semi-dominant and is variable in the way it is expressed - a cat may have no visible white spots or may be wholly white and all stages in between those two extremes. Unlike dominant white, white spotting is not linked to deafness.
COLOUR RESTRICTION, TICKING AND TIPPING
Another form of modification is colour restriction. This is seen in colourpointed cats where the colour is restricted to the head, tail and legs ( plus scrotum in males ). This is a form of partial albinism. As well as restricting where the colour will be exhibited, it tends to "bleach out" the colour to a greater or lesser degree depending on the type of colour restriction. It may seem odd to think of these as solid colour cats, but genetically they are; they have just had a "special effect" overlaid on them.
There are three colour-restriction patterns :
In the Siamese (Himalayan) pattern, there is maximum contrast between the points and the body colour.
In the Burmese, there is the least contrast but just enough to be able to see that the cat is not a single solid colour.
In the "Mink" Tonkinese, the effect is moderate.
The self colours may have different names in these cats. Alternatively a colour which appears identical to one of the solid colours may be genetically different because of the mild to moderate bleaching effect. A seal-point Siamese may appear to be brown, but genetically it is black!
In Abyssinian and Somali cats, the agouti gene causes the colour to be distributed in bands along each hair. This creates a ticked effect. The most confusing aspect is "red". The bright red colour in sorrel Abyssinians is really cinnamon. The dark red of the Usual Abyssinian is genetically black. Where registries recognise the sex-linked red and sex-linked cream colours the colour name is prefixed by the words "sex-linked". In the USA, the sex-linked colours are apparently not recognised to avoid confusion with the sorrel and fawn colours. To the naked eye (or without the benefit of a pedigree chart), the colours are practically indistinguishable.
It has been thought there were 2 different shades of sorrel red: a bright coppery red and a dark red characterised by an influential American Abyssinian sire called Champion Dhmahl's Diablo. Some pale kittens darken to red, while others become deep solid- looking red with minimum ticking. The dark red is more likely to be chocolate. Chocolate and lilac, along with the silver series, are considered alien to many North American breeders, but are familiar in Europe.
Full Expression
Sepia (Burmese)
Mink (Tonkinese)
Pointed (Siamese)
Abyssinian/Somali
Australian Mist
Black/Brown (in tabbies)/Ebony
Brown/Sable/Seal Sepia
Sable/Natural Mink
Sable/Seal
Usual/Tawny/Ruddy/Brown
 Brown
Blue
Blue/Blue Sepia
Blue
Blue
 Blue
 Blue
Chocolate/ Chestnut/Brown
Chocolate/ Champagne Sepia
Chocolate/ Champagne Mink
Chocolate
 Chocolate
 Chocolate
Lilac/Lavender/ Frost
Lilac/Platinum Sepia
Lilac/Platinum Mink
Lilac/Platinum
 Lavender
 Lilac
Cinnamon
Cinnamon/Cinnamon Sepia
Cinnamon/ Honey Mink
Cinnamon
Red/Sorrel
Gold
Fawn/Light Lilac
Fawn/Fawn Sepia
Fawn
Fawn
Fawn/Beige/Dilute Sorrel (non sex-linked)
Peach
Red
Red/Red Sepia
Red
Red
Sex-linked red
 Red
Cream
Cream/Cream Sepia
Cream
Cream
Sex-linked cream
 Cream
 
The colour can also be restricted to the ends of each hair. This is caused by a gene for silver. Chinchilla ( shell ) is the lightest form of tipping or shading - hair tip is coloured and hair shaft is silver, giving a sparkling appearance. Shaded is the next degree - the colour extends further along the hair shaft, darkest on the back (where fur is longer) to create a mantle of shading. Smoke is the heaviest tipping - the undercoat colour is reduced to a small band near the hair root so that the cat appears to be solid colour with a pale ruff until the coat is parted and you can see the silver roots. In goldens, the effect is similar except that the hair shaft is gold rather than white. These cats are still self colours, but the colour distribution has been modified!
There is another effect known as silver-tipping; this is the sparkling silver tips to the otherwise black fur of Chausies. It is an effect inherited from Jungle Cats (F chaus) and is seen in purebred melanistic Jungle Cats. Affected cats are self coloured, but with a modifier which is new to domestic cat genetics.
A WIDER PALETTE OF COLOURS
The previous sections dealt with solid colours and how they are affected by two types of dilution, three types of colour restriction, agouti ticking and silver/golden shading.
There are genes which control how evenly the hair is coloured i.e. the bands of colour on each hair. In agouti cats, it is easy to see different bands of colour using a magnifying glass . In cats with apparently solid-coloured fur, you would need a microscope. Even though the effect on an individual hair can't be detected, the overall effect on an area of fur is make the colour more or less dense i.e. darker or lighter.
In general the self colours (with plain English descriptions) are:
Colour name
 Plain English Description
Albino
White
Amber
Born black, brightens to bright apricot to cinnamon with age.
Apricot
Pink-brown or hot cream, with a metallic sheen,
Beige
Abyssinian/Somali: Non sex-linked cream, Fawn
Black
Jet-black
Blue
Blue-grey
Brown
Chocolate. Genetically black cats with tabby pattern are known as brown tabbies. (Burmese "Brown" is equivalent to black.)
Caramel
Caramelised blue, cafe-au-lait colour (biscuit colour), cool toned bluish fawn, metallic sheen
Champagne
Burmese/Tonkinese: equivalent to chocolate
Chestnut
Oriental: Medium -dark brown, equivalent to chocolate
Chocolate
Medium-dark brown
Cinnamon
Milk-chocolate (reddish) colour
Cream
Buff, dilute of sex-linked red
Ebony
Foreign : equivalent to black
Fawn
Abyssinian/Somali : hot cream (non sex-linked cream), equivalent to Light Lilac
Light Amber
Born blue, brightens to pink-beige or fawn with age.
Frost
Lilac/lavender
Gold
Australian Mist: equivalent to cinnamon
Honey
Burmese/Tonkinese : Equivalent to chocolate/chestnut
Indigo
 
Lavender
Pinkish grey (dove grey)
Lilac
Lavender
Light Brown
Equivalent to cinnamon
Light chocolate
Burmilla: Milk chocolate
Light Lilac
Equivalent to fawn
Natural
Tonkinese: equivalent to sable/seal
Peach
Australian Mist: Pink-brown, equivalent to light lilac/fawn. Peach is also seen as a dilute of Russian Blues and may be caramel.
Platinum
Burmese/Tonkinese: Equivalent to lilac/lavender
Red
Rich ginger red (poor reds are yellowish due to other genes)
Red
Abyssinian/Somali: Equivalent to cinnamon
Ruddy
Abyssinian/Somali: Equivalent to black/brown
Sable
Burmese: Dark brown (genetically black)
Seal
Siamese: Dark brown (genetically black)
Sorrel
Abyssinian/Somali: Equivalent to cinnamon, honey mink of Tonkinese
Taupe
Caramel dilution of lilac/lavender
Tawny
Abyssinian/Somali: Equivalent to black
Usual
Abyssinian/Somali: Equivalent to black
White
Non-albino white, this is the absence of colour.
 
In all likelihood, there are many other genes which subtly alter the colour e.g. by modifying the hair structure slightly to change the way it absorbs or reflects light or by affecting the distribution of the pigment granules in the hair shaft. The appearance of silver-tipped black cats suggests that hybridisation is going to introduce new colours and effects into domestic cats.
Some colours which have already appeared are still disputed. For example "caramel" is a subtle form of dilution. It does not affect non-dilute colours (black, chocolate, cinnamon), but it changes the appearance of already dilute colours (blue, lilac, fawn). Caramels which are genetically blue cats have a brownish cast while those which are genetically lilac are lighter in tone . So should they have separate colour names e.g. blue-caramel (caramel), lilac-caramel (taupe) and fawn-caramel or should all the slightly varying colours be lumped under the name "caramel". They jury is still out on this one, but it demonstrates the importance of knowing what colours the cat has in its pedigree!
Another new colour, indigo, is described as a richer, darker version of blue. This suggests some sort of intensifying effect acting on diluted colours. However since blues can be variable in hue, there is no confirmation that indigo is due to a specific gene or that it is genetically heritable. The only example I have seen depicted a tortoiseshell cat with blue-black and hot cream patches i.e. to richly coloured to be blue-cream tortoiseshell, but not a red-black tortoiseshell. This could mean a gene with the opposite effect to caramel i.e. it acts on dilute colours only, but it intensifies them. If so, it is probably invisible to the naked eye and only noticed in individuals with the most extreme effects of that gene. The same gene would have no discernible effect on non-dilute colours since there are already at their most intense . Note: since other breeders describe indigo as a darker version of blue, the "blue-black" may have been exaggerated by lighting.
If there is a colour intensifier turning blue into indigo, it could be expected to work on the other dilute colours as well e.g. turning lavender into deep lavender, fawn into rich fawn and cream into rich cream. The differences would probably be too slight to be visible - it is already hard enough to detect the caramels!
According to Inina Sadovnikova, "I believe every breeder has had cats of one and the same colour, but of a different shade . Take a black (brown) tabby, for instance. They range in shades from a light grey with a black pattern to a rich brown with a black pattern. Reds are also very different, from a ginger to an intensive orange. The same concerns blue, I have seen blues so dark that one doubts if they are really blue and not black. From my experience , I tend to think that there is a modifier, though I don't know of any studies of this factor. The litters of my female Konkordia always had at least one kitten coloured intensively, either a very dark black tabby or a very dark red tabby. She has a light shade of colour, but her grandmother is darker and also had several dark kittens in her litters. This looks like a recessive, but one needs a lot more of statistics to prove if and how it is inherited. A dark shade is desirable for red, but at present undesirable for blue. So I don't know if "indigo" has any future."
ODDITIES
Strange as it may seem, in some animals the mother's diet can affect coat colour. Recent work at Duke University in Durham, North Carolina, has shown that in a certain strain of mice , the "agouti" coat colour gene can be turned off through methylation. The extent of methylation was dependent on the mother's vitamin intake during pregnancy. (Methylation turns off genes by chemically modifying them). The effect has not been observed in cats, but in theory it could turn genetically agouti (ticked) cats into solid colours.
Some cat shelter workers have noticed apparently self red kittens turning into black adults. Solid black is known to be prone to rustiness through damp or sunlight and evidently, in some cases , to conditions in the womb. Self black kittens are often "rusty" at birth, though few cases are as striking as that of a ginger kitten called "Marmalade" who was entirely black at 6 months old (personal correspondence).
A better documented oddity is that of apparently self red or self black female cats that are genetically tortoiseshell cats. In rare cases, one colour predominates to such an extent that the other is represented by a few isolated hairs.
BEAUTIFUL BICOLOURS - TUXEDO AND MAGPIE CATS
Copyright 2002 - 2016 , 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 Varietäten" (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!
In January 2016 , it was announced that piebald patches form when cells fail to develop in the womb. The gene for white spotting affects the embryo cells (melanoblasts) which will become pigment-producing skin cells (melanocytes) which make the pigment for hair. Pigment cells move and multiply as an embryo grows and there aren't enough cells to cover all the skin, so the animal gets a white belly. The findings were published in the journal Nature Communications. The pigment cells 'fail to follow instructions' during early development . They move and multiply randomly as an embryo grows, without complex cell-to-cell communication sending them in one direction as once thought. The University of Edinburgh 's Dr Richard Mort said that the cells move and multiply at random which is not what was expected. The University of Bath's Dr Christian Yates said: that piebald patterns could be caused by a faulty version of a gene called kit. What the researchers found was counter -intuitive. Previously it was widely thought that the defective kit gene slowed cells down, but instead they found that it reduced the rate at which the pigment cells multiply. This did not surprise cat fanciers who had noticed that pigmented patches could fit together like a jigsaw and likened the process to “plate tectonics” where pigmented patches drift across the embryo’s skin leaving unpigmented areas inbetween the patches. Researchers at the Universities of Bath and Edinburgh said that in addition to kit, there are many other genes that can create piebald patterns, but their the mathematical model can explain piebald patterns regardless of the genes involved.
This wasn't entirely surprising to many cat fanciers with an interest in genetics or embryology! Independently of scientific researchers, and based on observation, fanciers had some ideas of their own, which have turned out to be close to the mark.
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.
Part of this theory was that the patterning of black and white was the result of the skin surface "cracking" during early embryo development. The skin of such bicolour cats at an early embryo stage would be basically pigmented, but the dominant S (white spotting) gene, causes the pigmented surface to crack into domains or islands . These islands drift apart over the embryonic surface as the embryo grows, but the pigment cells don't multiply fast enough to fill the spaces between the island . That leaves white areas in the regions between the islands of pigment. 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 not enough melanocytes available to fill the 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 - it is suspected that the belly area expands greatly during embryo growth . Megacolon in cats and exposed gut conditions in some rabbits upholds this theory. 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 .
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. This is also explained by the mechanism detailed in the 2016 report findings. The black pigmented cells that have moved across the embryo surface multiply to form black islands (clonal patches), while the red pigmented cells that moved across the surface multiply to form red patches.
OLDER / ALTERNATIVE THEORIES FOR BICOLOUR PATTERNS
Above: "Olga", an attractive brown bicolour owned by Sarah Richie
It used to be thought that when the melanoblasts arose from the "neural crest" - the area along the back of the embryo -they migrate all over the body during formation of the skin. Where cells failed to reach their allotted positions before the skin was fully formed, those areas lacked pigment. This seemed to explain why the white is most often found on the paws, belly and chest - those 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. It also explained 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.
Another theory involved either of two cellular mechanisms in white spotting that "turned off" pigment. One mechanism was 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 involved 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.
Some cat breeders believed there were 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.
Leukoderma ("white skin"), leukotrichia ("white hair") or vitiligo, is a cosmetic condition that 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"), 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 . "Leukotrichia" is a generic term. Acommon term is "piebaldism" because it causes white patches in the skin and fur. It is an "aquired depigmentation" that occurs during the cat's lifetime, is usually progressive and may be triggered by illness or environmental factors. Ultimately, a cat with leukoderma may become almost entirely white. Antibodies are formed against the pigment-producing melanocytes. The melanocytes are destroyed leading to the white areas. A type of leukoderma has been identified in some Persian cats and these are used as laboratory subjects in the study of depigmentation conditions. Periocular leukotrichia, causes the fur around the cat's eyes to become pale - as though the cat is wearing spectacles.
Vitiligo in a big cat - the "cobweb panthers" that turned progressively white.
Charva with feline vitiligo (Jason Reeves).
 
"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. 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. 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.
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.
"BIMETALLIC" SIBERIANS & PLATINUM PERSIANS
2013-2014, Sarah Hartwell
Many thanks to Silvia Perego and Lesley Morgan for allowing the use of their photographs on this page.
Note on Terminology: At present, terminology isn't standardised. "Bi-metallic" describes a visual effect, while "sunshine" is the (current) proposed genetic name. In Siberian Cats, sunshine-silver produces the bi-metallic pattern.
For some years, strangely patterned Siberian cats have been turning up. Some are covered in A Torbie Dynasty: A Torbie Dynasty. Caroline Sharp in Germany had several generations of apparently tortoiseshell-tabby male cats. This should be impossible as tortie tomcats are genetically abnormal in some way and even when fertile they should not produce further generations of tortie tomcats. They couldn’t all be chimeras, so there had to be some unidentified mechanism at work.
Other Siberian cats had both silver and golden areas of fur and were dubbed “bimetallic.” This X-colour (mystery colour) looked similar to the partway stages of the amber colour change seen in Norwegian Forest Cats. Siberian breeders had referred to their cats as "golden" but this term clashed with the wide band colour found in Persians, Exotics and British Shorthairs. The official term "sunshine" has been proposed to distinguish Siberian golden from "wide band golden".
At birth, Siberian "Sunshine" resembles the early stages of the amber (ee) colour change in Norwegian Forest Cats caused by a non-extension gene. However, the Siberian cats tested negative for the amber gene. There were no genetic tests for Wide-band or for silver inhibitor, let alone for Siberian sunshine, but it can be distinguished visually and by studying pedigrees. Bimetallic colouration has also been seen in the Kurilian Bobtail and in random-bred cats in the Ukraine .
Judge Lesley Morgan come across a number of strangely coloured cats that had been nicknamed “bimetallic” as they displayed a mix of silver tabby and golden tabby in their coats. The effect was beyond the rufousing or breakthrough colouring sometimes seen in silver tabbies and in some cats the golden areas increased as the kittens grew into adulthood. The only link she found was that every bimetallic Siberian she saw had close Polish ancestry. Other Siberians had been registered as black silver tortie tabby, but were more likely to be bimetallics (silver tabby + sunshine).
3 views of "bimetallic" including a close-up of the fur.
In 2013, Silvia Perego of Veselka Siberian Cattery in Italy described more such cats. Her stud male, Yankee, came from Poland and is a black silver tabby with white. When he arrived at her cattery at 4 months old, she noticed that he had a "strange" golden-brown spot on his neck and some little other spots on the back. At first this appeared to be rufism. His first litter included a silver tabby daughter, Lady , who had the same golden-brown spot on her neck. Another litter included a daughter, Lizzy, with the same spot. Then Lady, from the first litter, had her own litter with another silver-and-golden kitten like them. In summer 2013, Yankee had two kittens with the golden-brown patch. Although a judge suggested chimerism, there appeared to be a newly emerging gene at work.
The classic golden colour found in Persian cats is due to the dominant wide-band (Wb) gene. This widens the pale brown area at the base of each agouti (banded) hair and confines the darker colour to the ends of the hairs. The paw-pads are either white (i.e. pink) or match the colour at the hair tip. Combined with the silver inhibitors, this resulted in chinchilla and shaded silver Persians. A golden Persian has wide-band and the recessive form of the inhibitor gene.
Research by Eleonora Ruggiero into the pedigrees of Siberian Cats brought out some intriguing details . For example in “Onix Gloria” lines there are golden tabby Siberians born of non-golden parents. In the pedigrees of “Siberian golden” (provisionally denoted sg), golden offspring are born from two non-golden parents. This means the gene was recessive and only expressed when two copies are inherited. It is not the same as Persian golden (Wb with recessive inhibitor). Bimetallic females were often registered as tortoiseshell, but didn't breed as a tortoiseshell as they lacked the red (O) gene (evident when the paw-pads are examined and are never reddish). Their nose leather was dark pink, unlike the nose leather of tabby or genetic tortie cats.
When golden cats started turning up in silver Siberians, it resulted in cats expressing both colours in the coat; something impossible were it the same as Persian golden. The degree and intensity of golden colour (probably influenced by polygenes) rules out rufism or tarnish. These cats were initially referred to as “bi- metal ” and later as "sunshine". Siberian sunshine was established in a breeding programme that used much inbreeding to bring out this recessive trait. Early sunshine cats were dismissed as silvers with high degrees of rufism and not bred from. In 2013, all known sunshine Siberians are agouti (tabby markings) and the colour hadn’t been seen in non-agouti (solid) cats. The sunshine gene affects the agouti (ticked) hairs only".
Sunshine silver looks very different from a silver with rufism; the nose leather is pinkish unlike that of a tabby cat. The red colour of a sunshine red tabbies and sunshine torties is brighter and the paws are lighter. The absence of pigment in the sunshine tabbies extends beyond the edges of the nose leather so they have no nose-liner and have whitish fur at the bottom of the nose.
There is a link to one of the Snowknight cats, Joschka, around 7 generations back, so his torbie dynasty continues. The common ancestor from Poland is Snowknight Destiny. Either Destiny or her brother Snowknight Derringer had a very colourful torbie male offspring, but the breeder had him neutered rather than test-mating him. This was carried over from Joschka, who had an extra Chromosome. Caroline Sharp’s torbie male Bogomir Tschudovich of Misinza aka Meli is black-and-white, with a big flame of red over and around his ears and no sex-linked red parentage. Because those cats don't have the inhibitor (silver) gene, they looked even more like tortie-tabbies.
Michelangelo, a colourful torbie male (without silver) from Caroline Sharp's breeding line.
While silver-and-gold has turned up in Siberians, a silver-TO-gold effect has turned up in some lines of Silver Persian/Chinchilla Longhair where it has been dubbed "platinum". In 1986/7, Cheryl Bennett reported in The Silver and Golden Persian Newsletter (ACFA) that one of her shaded silver females (Kelley Lane Contessa of WeANDE) had changed from pure silver to pale golden. Contessa’s parents were a shaded golden and a shaded silver; these were full siblings. At 10 – 12 months of age, Contessa began to “tarnish” i.e. show cream/reddish patches and by 3 years of age she was entirely pale golden. She produced a number of Silver Persians without tarnish, but she failed to produce any offspring when mated to a golden sire. One of Contessa’s male offspring turned from silver to golden as an adult. An adult Silver Persian from England (Lynchard Silver Shadow) was exported to Australia and also turned golden. Shadow had a few golden hairs on his paw, but did not turn golden until he was a year old when his coat turned to pale beige. By 3 years old he was entirely pale golden. Shadow was bred to a genetically golden female, but the pairing only produced silver offspring. However, at least one of his silver offspring later turned pale golden. Other descendents of Contessa also went through the late colour change. During the 1980s, several other breeders of Shaded Silver and Chinchilla Persians came forward to report that their cats had developed reddish, brownish or golden-coloured fur along their spines as they aged. Many of the cats had no golden in their ancestry. At first it was dismissed as an unavoidable genetic fault where silver was incompletely dominant and did not hide the recessive golden colour.
A common ancestor of all the colour-changing Silver Persians was a stud cat called Kelly Lane Andromeda (in the UK) whose descendents were exported to the USA in the 1950s and 1960s. Some of those descendents were influential stud cats and would have spread the mutant gene far and wide. Controlled inbreeding (linebreeding) helped establish the gene, which would later double up to produce colour-changing Silver Persians. The effect of this gene on Golden Persians, if indeed it has any effect, is not known. Except for red-silvers and cream-silvers, silver cats should entirely lack phaeomelanin (red or cream pigment) and only have eumelanin (black, brown, blue etc) present. Perhaps some other gene causes the eumelanin structure to change so that it is perceived as a golden colour. Chemical analysis proved that the late colour change Silver Persians did not have phaeomelanin pigment present.
BIMETALLIC ORIENTAL SHORTHAIR
This is an Oriental Shorthair, genetically a chestnut silver spotted tabby and white, but with with non-silver patches resulting in a "merle" type effect. This looks similar to the bimetallic effect seen in the Siberians. Kotchera Trail Blazer of San-Toi was bred by Wendy Heidt, a Siberian breeder and is being investigated by Pat Decano (CasaDecano Orientals, Siamese and Colorpoint Shorthairs). Pat has also considered chimerism as a possible explanation.
COLOURPOINTED AND MASKED CATS
Copyright 2002-2015, Sarah Hartwell
The colourpoint pattern is a form of albinism and is known as the "Himalayan pattern" although it's more often called "Siamese" pattern by cat lovers. It is found in a number of mammals including rabbits and goats from the Himalayas region. The exact effect of the gene depends on the temperature since its effect is inhibit colour in warm areas of the body and allow colour to develop on the cooler areas such as the ears, face, legs, tail and (in males) scrotum. In warmer climes, there is less contrast between the body colour and the point colour. Colourpoint kittens are born with almost pure white coats on a pink skin because of the uniform warmth in the womb; only after birth does the point colour develop. In cooler regions, the contrast may be very pronounced. The characteristic blue eyes reflects reddish at night because albinism affects the tapetum (reflective layer) of the eye; non-albino cats' eyes reflect greenish.
The most familiar colourpoint breeds are the Siamese and Himalayan (Colourpoint Longhair/Colourpoint Persian), but there are a number of others. Most, if not all, colourpoint cats have Siamese somewhere in the history, or at least have Thai, Asian or Japanese cats in the family tree since this pattern occurs naturally throughout Asia and into Russia . The Burmese has a "low contrast" form of the colourpoint gene andthe Tonkinese is halfway between Burmese and Siamese. There are a series of gene variants and combinations controlling the contrast between body colour and point colour; these are shown in the charts below.
 
 
SIAMESE/COLORPOINT SHORTHAIR
Modern Siamese come in different varieties - one is the "Modern" Siamese with its exaggerated wedge-shaped head, bat ears, a muzzle sometimes referred to as a "banana nose" and an often excessively tubular body. Another is the "Classic" Siamese that resembles the 1950s-1960s Siamese and intermediate between the more robust "Old-Style" that resemble early Siamese imports and the extreme modern Siamese. There is a continuum between the old-style and the modern Siamese, with most people considering there to be two types: old and modern. Personally speaking, I find many of the modern showbench Siamese extremely ugly creatures and many owners, as opposed to breeders, prefer a less extreme style of cat. For this reason, there are also traditional (old) style Siamese (Thai Siamese) being bred, recreating the chunkier shape familiar in the late 1800s and early 1900s. Both originate from the same cats.
The naturally-occurring colourpoints I have seen in Malaysia and Thailand are closer in type to those being bred to the older (non- ultra -typed) Siamese standard. Of the conformation of Siamese cat, Phyllis Lauder wrote in "The British, European and American Shorthair Cat" (1981): "Cats of eastern provenance have not shown long heads: in the 1930s and 40s there was, at the Natural History Museum in London, a stuffed Siamese cat, and this animal’s head was ‘as round as an apple ’ to quote one of England’s prominent experimental breeders, the late B. A. Stirling- Webb . The taxidermist’s work showed a large cat of strictly ‘domestic’ type. "
There are many theories about the origin of the Siamese. The Seal Point Siamese was known as the Royal Cat of Siam because it was found in palaces. It seems to have been familiar in the ancient Siam city of Ayudha which was founded in 1350 and was Siam's capital until destroyed by fire during the Burmese invasion of 1767. In Bangkok's National Library there are manuscripts from Ayudha which describe native fauna. "The Cat Book of Poems " depicts pale coated Seal Point Siamese, describing them as having black tails , feet and ears with white hair and blue eyes. No one really knows what the cats of ancient Siam looked like or whether they were bred for body type or purely for colour and pattern with little regard for shape. Another problem is that artists often stylize their illustrations.
The extinct Annamese from Vietnam is also considered by some to be the ancestor of modern Siamese and that the Siamese cat of Thailand arose through interbreeding Birman and Annamese cats. An early description of the Seal Point Siamese in 1676 calls it "Vichien Mas" (meaning "diamond mouth") and drawings depict extreme expression of the colourpoint pattern - dark ears, nose, paws and whiskers. A cat with Siamese markings appeared on an old engraving discovered by the naturalist Pallas on his journey into Southern Russia between 1793 and 1794. It was the opinion of Sir Russell Gordon (who closely studied these cats) and Auguste Pavie ( French explorer and one time resident at Bangkok), that the Siamese cat derived from a cross between the Burmese Temple Cat (i.e. Burmese, not the Birman which was known as the " Sacred Burmese Temple Cat") and the Annamite Cat. The Annamite cat was described as a slender, small and gracefully built variety of cat with a short tail. Like so many varieties it was said to have disappeared due to interbreeding with introduced domestic cats.
Although apparently favoured as palace cats, there is no clear record of an distinct Siamese breed until the 1800s. The British became interested in Siamese cats and imported them from Siam. The earliest documented imports were during the 1870s, but these were apparently not bred. In 1884, the departing British Consul-General Gould was given a Siamese cat by the Siamese king . He brought the cat to England and its progeny were exhibited at Crystal Palace in 1885. The early Siamese cats were round-headed, solid and muscular, but even so, their appearance was so extraordinary that they were described as an "unnatural nightmare of a cat". In Ceylon, the Siamese cat was, for a while, known as "Gould's Cat", having been introduced there by Mr Gould. The Burmese Sacred Cat was known to early British cat fanciers as the "Gold Cat". A wild cat of the region was known as the "Golden Cat" (Temminck's Golden Cat) or "Bay Cat". HC Brooke believed these similarities of name to be the reason that Temminck's Golden Cat was claimed to be an ancestor of the Siamese. The first champion Siamese, "Wankee," was born in Hong Kong in 1895 and exhibited in 1898. He was relatively large and round-headed by modern standards, but had a more distinct muzzle and longer body than modern appleheaded (as in the rounded headed "applehead" dolls of the USA) Siamese - more of an intermediate type.
Frances Simpson , editor of "The Book of the Cat" (1903) included contributions from several early breeders of Siamese cats. While acknowledging the existence of blues, blacks, whites and tabbies in Siam, she stated that only the "Royal Siamese" and "Chocolate Siamese" were recognised in England at that time. These were sometimes bred to each other although opinions on the quality of the offspring were contradictory. The royal Siamese was sometimes bred to white short-hairs because the English type was preferred over the foreign type by judges. Because white is dominant and masks other colours, " sports " with "any other colour" points occurred a generation or two later.
Most early breeders considered the Siamese cat to be more delicate than the English cat, having delicate lungs and being prone to disease and other upsets. Many did not risk sending their precious cats to shows. One early breeder noted a rarity of female kittens in a litter, the average being 5 males to 2 females. The kittens were said to be difficult to rear , as they suffered from worms and teething, and it was common to foster Siamese kittens on English cats to make them more robust and healthier. Males were described as extremely powerful, great fighters, had terrible voices and would kill strange cats and fight dogs. "The males are, however, antagonistic to others of their sex, and fight with a terrible persistency. I have heard of a stalwart fellow who, being allowed his liberty, cleared the neighbourhood of all other wandering toms. When made neuter, Siamese become most charming home pets."
Miss Forestier- Walker and her sister, Mrs Vyvyan, had received a pair of Siamese cats from the Siamese Palace in 1884-5; Miss Forestier-Walker wrote "Siamese cats were first introduced into England about twenty -five years ago, but were not often seen until a few years later. Since then they have become fairly common. There are two distinct varieties in the present day. (1) The royal cat of Siam" by which she meant the seal point Siamese, "(2) The chocolate cats are deep brown in colour showing hardly any markings, and have blue eyes. The tails are sometimes straight , which is not a fault; but a knot or kink in the tail is a peculiarity of the breed, and therefore desirable. In England it has been asserted that this is a defect , but in Siam it is highly prized […] In the East a cat with a kinked tail fetches a higher price ."
"There is a legend that the light-coloured cats with blue eyes represent silver; the dark cats with yellow eyes, gold; and that the possessor of both will have plenty. This rather gives the idea that originally the eyes of the pure chocolate cat were yellow, and that the present variety has been crossed with the royal cat. It is a great mistake to mix the varieties, as the result after they become adult is a blurring of the markings and a patchy coat." What Miss Forestier-Walker was describing was the range of colours from the brown Burmese (the "Burmese cat" depicted in the 1903 book was an Oriental ticked tabby), through the "blurred" or less distinct markings of the mink range (Tonkinese) to the sharply defined colourpoint of the Siamese.
Champion Wankee was bred in Hong Kong in 1895, the offspring of a female kitten stolen from the Palace in Siam, and imported to England aged 6 months. His owner , Mrs Robinson, wrote "One of the most beautiful of the short-haired cats is undoubtedly the royal cat of Siam, and the breed is increasing in popularity ; but is never likely to be common, as the cats are delicate in this country. […] The [standard of] points of the chocolate Siamese are the same as the royal, with the exception of body colour, which is a dark rich brown all over, thus making the markings less noticeable. All Siamese darken with age, and when they get dark there is a tendency to call them chocolates. I know of only one real chocolate - Mr C Cooke's 'Zetland Wanzies' - so consider them more likely to be a freak than a distinct variety."
She also described the different body types: "Of the royals there seem to be two types in England: the one - rather a small, long-headed cat, with glossy , close lying coat and deep blue eyes, and with a decided tendency to darken with age - is generally the imported cat or having imported parents; the other is a larger cat, with a rounder head, a much thicker, longer and less close-lying coat, and the eyes a paler blue (these cats do not darken as much or as soon as the other type, and have generally been bred for several generations in England)."
According to another early breeder, Mrs Parker Brough, "There are distinct varieties of Siamese known to fanciers - the palace or royal cat, the temple cat (chocolate), and there is likewise the common cat of the country, which is also found within the palace. The points of the chocolate cat are identical for shows with those of the royal except body colour, but the imported chocolate is often dark chocolate, with blue eyes, stumpy tail with a marked kink, short legs, and heavy , thick body. There are not many chocolates exhibited, owing to the preference given to the royal variety. It must be understood that there is no definite royal breed as such, but the palace breed seems to have originated by selection ." Lady Marcus Beresford wrote that Siamese imported from a Bangkok temple "differed from the royal Siamese, being darker and having a more pointed head and face, and their eyes were larger and fuller."
According to Miss Forestier Walker "This breed is said to be kept very carefully in the palace in Bangkok - hence the title 'royal' - and is by no means the common cat of Siam. One gentleman (a missionary), who had lived there fifteen years, had during that time seen only three. A few years ago there was a pair of these cats in the Zoological Gardens at Bangkok, but they were very poor specimens. […] The first specimens were brought to England about twenty-five or thirty years ago, and Mr Harrison Weir says that among those who possessed them were Lady Dorothy Nevill, whose cats were 'imported and presented by Sir R Herbert of the Colonial Office. The late Duke of Wellington imported the breed, also Mr Scott of Rotherfield.' "
Mrs Parker Brough wrote "Until recently the Siamese was but little known in Europe, but occasionally was to be found in the various zoological gardens. At present there is a fine female specimen to be seen at the Zoo at Frankfort-on-the-Main, having been purchased from the King of Roumania. One or two are to be seen at the Hague. London has the first one it has had for six years, but it is not shown owing to its want of condition. […] A point on which the Siamese fancy is divided is where the ideal cat should have a kink in the tail or not.[…] There is a peculiarity in breeding the Siamese - i.e. the rarity of female kittens in a litter, the average seeming to be five males to two females. […] They have naturally rather delicate lungs."
Some of those early (1890- 1902 ) Siamese were evidently large, robust creatures, for example a neutered male called Attache was described as very large and powerful, with massive limbs, and an unconquerable antipathy to all other cats of any description. Frances Simpson summed up in 1903 by saying "I do not believe that Siamese will ever become common in England, for many reasons. These cats are expensive to purchase , difficult to rear, and fancier are afraid to risk them in the show pen; but in spite of these drawbacks, I think as time goes on and the Siamese Club extends its labours, we shall see and hear more of these really curious creatures, for what we call the royal Siamese bears no resemblance to any other cat, and the distinguishing difference , being so great, tend to make the breed one of our best show cats and a clear class to itself, for the Siamese of the purest blood should not be crossed with other cats. We have heard of 'any other colour' Siamese, but these cats of varied hue claiming to be Siamese are but the offspring of a cross. We have been told of black and blue and tabby Siamese; but the fanciers of Siamese look askance at these freaks, and feel that it is worse than useless to attempt to produce any other variety than that which we have learned by custom to designate the Royal cat of Siam."
A great deal has been written about the origins of the Siamese breed. Sydney W France , editor of "Cats and Kittens" magazine, wrote the book "Siamese Cats" in 1949. This was the first major publication on Siamese cats since 1936.
"The history of the Siamese in this country is a very short one, and it is true to say that they have only been here within living memory, and that the first ones actually were from The Royal Palace of Siam. Even on this point there is much controversy and it is interesting to note that the first Siamese of which there is any record were said to have been brought to England in 1884 by Mr. Gould who was then Consul General in Bangkok at that time. In 1886 a pair of cats and two kittens were brought to England by Mrs. Vyvyan, these had actually been procured from the King’s Palace in Bangkok. In the same year, Mrs. Walker, the General’s daughter, brought over one male and three females. There is no doubt that at that time the true Siamese were kept in the Royal Palaces and Temples , and that few of them ever found their way from there except as gifts, which were then considered as of great worth."
In direct contradiction, Ida M. Mellen, well-known American authority on cats, in her " Practical Cat Book" (1939) writes "Although this cat generally is referred to as the Royal, and even as the Sacred Siamese, it is the common cat of Siam, just as the Manx, equally an aristocrat, is the common cat of the Isle of Man."
France also reproduces a letter Mellen received from Dr. Hugh M. Smith, Adviser in Fisheries to His Siamese Majesty’s Government between 1923 and 1934. In addition to the Siamese cat, he mentions a "mauve" cat which is no doubt the Si-Sawat (Korat) and kinked- tailed cats which were, and still are, common throughout Thailand, Malaysia and Singapore (and in the modern Japanese Bobtail breed).
"I was well Acquainted with cats in Siam, but made no special study of them. There appear to be two races peculiar to the country: the common form with pale fawn colour, black or dark brown feet, tips of ears, tail and muzzle, and blue eyes, well-known to cat fanciers all over the world, and a form of uniform mauve or Maltese colour with yellow eyes. There are no "palace" cats in Siam. There are no "royal" cats, although the strikingly marked creatures would be the natural ones to be kept in palaces. Any person can have a Siamese cat, and as a matter of fact there are many people outside the palaces and many foreigners who keep such cats as household pets. There are no "temple" cats. The Buddhist priests, who do not live in the temples but in special buildings in the temple grounds, may keep cats, as they do dogs. A Siamese prince whom I know very well was visiting in London and was interviewed by one of the thousands of Siamese cat fanciers there. He told her there were more Siamese cats in London than in all Siam.
You probably know about the cat not peculiar to Siam but found over much of South Eastern Asia, which always has a sharp kink near the tip of its tail. It is of various colours but never of the special Siamese cat colour, and is of no interest except for its tail."
There is no doubt that Siamese cats came from Siam (modern day Thailand), but some disagreement in the 1940s as to their "royal" origins. France disagreed about Siamese not being Royal cats. As well as mentioning the various royal legends relating to the tail kink and squint (faults in show cats), he stated:
"This does not bear out my own information on this point, because whilst living in Jersey I had the good fortune to meet and become friendly with Major Walton of Verona House, Grouville, who was until recently, in Siam in connection with the Rice purchasing Commission, when he and his wife , both cat lovers, became friendly with the Prince Regent of Siam. Major Walton told the Prince Regent that he and his wife wanted to bring back some Siamese to England, but had not been able to find any at all in the country that were for sale. Before leaving Siam, Mrs. Walton was presented with a pair, male and female by the Prince Regent, and I actually went and saw them in quarantine in Jersey where they had a litter of five kittens, of which, later on, I bought two; a male and female.
These Siamese cats had coats of extremely fine quality texture, and colour, and very good head and body shape, splendid long whip-like tails, but with eyes which definitely failed in colour according to our standards here in England. They appear to be hazel , whereas there is no doubt that we have enormously improved the eye colour and have cats with beautiful deep blue eyes."
Major Walton’s remarks to France about the Siamese cats of the Royal Palaces were also mentioned in an issue of "Cats and Kittens" magazine, of which France was editor. This prompted the following letter from Mr. A. N. M. Garry of Minehead, Somerset:
"My wife and I are ardent cat lovers, and having spent most of my working life in Borneo, I feel I have some justification for writing to you about Siam and its cats. When I was in Siam in 1930, I was told that there were two distinct types of Siamese cats - apart from the Malay cat and crosses with it. The first is the one we see in England, but I think its points are a shade different, [chocolate] brown instead of seal. The second, which was said to be peculiar to the Royal family and palaces, had the body colour of the first - but not the points; and hazel eyes.
Having been a contemporary at Eton with the then King, I got a special permit to see the Bangkok Palace more thoroughly than the usual tourist does, and I saw one or two of these "Royal" cats, whose appearance was (to the best of my recollection after so long) as I have described. At that time, the export of the first type, except neuter ones, was absolutely forbidden, owing to the fear that they might become extinct in Siam, because so many had been exported.
The second type was absolutely unobtainable, far less exportable, for it was not to be seen outside the royal palaces. But owing to the war, and the various constitutional changes that Siam has undergone, the rules may well have been relaxed. It certainly looks as though Major Walton has been lucky enough to get hold of a pair of the scarcer "Royal type." Incidentally, I hope he is aware of the tendency of Siam born Siamese to chest troubles ~ damp or cool weather . I know of several people even out there, who have lost their pets from this cause.
I am afraid I am quite unable to agree with Mrs. Adney’s friend that a Siamese must have a kinked tail to be considered pure bred out East. In my experience, the .connoisseur out there, just as at home, demands the straight tail; but the fact remains that the majority of so-called Siamese cats in Malaya and Borneo have kinked tails, owing to one of their parents or forefathers having been a Malay cat. They still make lovely pets, and have the characteristics of pure-bred Siamese, such as the deep voice and the love of following their owners like dogs.
The Malay cat, like the British domestic cat, is of varied colours, ginger, black, black and white, tabby and tortoiseshell. Some having Persian forbears, are long haired. Practically all have kinks, and I had one, a beloved ginger, called Peter, who was considered a cat of particularly good omen , as he had two kinks quite close together. They were very tender, and he hated them being touched. He lived to the age of seventeen - very old indeed for an animal in the tropics. He died a few months before the Japanese invaded Borneo."
The first Siamese fanciers club was founded in Britain in 1902 at which time they were apparently variable in type. Possibly the conformation depended on which cats the early Siamese had been out-crossed to, there being few pure Siamese in the country at the time, or to the variability of imported cats and inbreeding from a limited gene pool.
Early photos show differences between the "compact" cats and the "lithe" cats, but the difference is nowhere near as extreme as that between modern "classic" and "old-style" Siamese. Early Siamese are more robust and have more rounded heads than modern classic Siamese, however, they were longer-bodied (less cobby) and more wedge-headed than the British Shorthairs and Persians of the time. The 1892 Siamese breed standard (Harrison Weir) described them as a marten -faced, Oriental type of cat distinctly different from the cobby, round British cats. Weir described the Siamese wedge as beginning at eye level, at the muzzle (in modern cats it begins from the ears downwards).
Based on only a few imported Siamese, early breeders believed that there to be 2 types of Siamese. Judge and Siamese-fancier, Mrs. Carew Cox, said (reported 1903) "There appear to be two distinct types - the compactly built, short in body, short on legs, and round in head; and the long-bodied, long-faced, lithe, sinuous, and peculiarly foreign-looking variety."
An early Siamese breeder, Mrs. Robinson said, "Of the royals there seem to be two types in England: the one - rather a small, long-headed cat, with glossy, close-lying coat and deep blue eyes, and with a decided tendency to darken with age - is generally the imported cat or having imported parents; the other is a larger cat, with a rounder head, a much thicker, longer, and less close-lying coat, and the eyes a paler blue (these cats do not darken as much or as soon as the other type, and have generally been bred for several generations in England)."
The Siamese Cat Society of America was founded in 1909 although the date of their arrival in the United States is not precisely known. Many early Siamese had kinked tails and cross-eyes or a squint; these faults have largely been bred out of modern Siamese.
In 1949, Kit Wilson (Vice-Chairman of GCCF) wrote in the Sydney W France's book "Siamese Cats"
"A few years before World War 2, interest was beginning to be shown in Blue Pointed Siamese. These had caused some considerable controversy among breeders, many of whom were of the opinion that they were "sports" and therefore could not be bred true, but a few, whose convictions based on research refuted these opinions, and "The Blue Pointed Siamese Cat Club" was formed.
Lately particular interest is being taken by some breeders in Chocolate Pointed Siamese. The greatest living authority on this fascinating variety is Miss Wentworth Fitz-William of Slingsby Yorks., one of the most valued breeders and judges of many years standing who has always owned some, and from whose stock most of those in existence to-day have sprung. [...] It is an interesting fact however that there are considerable numbers of chocolate points in France, but their Owners have no idea as to their origin, the strain however must have been good, as they are without exception the only Siamese worth showing over there (or at least they were at the January 1948 show, when I was judging. [...] For those who have never seen a Chocolate they are often smaller in build than the seal or blue point, and their points are of a rich milk chocolate colour.
Blue points are very popular in the U.S.A. many of them winning high awards at shows over there, but, although I am open to correction, I have not heard of any chocolates. I only had one of these in my classes at the Danish (Darak) show in 1946, and this was a very poor specimen and hardly worthy to carry on the breed, and was, in all probability a " sport ." It had been bought at quite a high price when a kitten as a seal point, and having met the vendor as well as the owner I am of the opinion that neither of them had the slightest idea that it was anything other than the usual seal.
While writing this an interesting fact has come to light with regard to chocolate points. Mr. Brian Stirling-Webb, the Hon. Treasurer of the Siamese Cat Club has been making a close study of this variety, and he states that practically every chocolate in the country can trace their pedigree back to an imported cat brought into this country in 1896, this may also account for those on the continent . Who knows, perhaps another from the litter was taken over there and that those now being shown are direct descendants."
In that book, Wilson wrote in disapproving tones of cross-breeding of Siamese. Luckily the experiments in breeding Siamese with other breeds (not species!) did continue in spite of early disapproval, leading to a wide variety of attractive Oriental and Foreign varieties.
"From America we have heard of the Black Siamese with orange eyes but to my knowledge no specimen has ever been seen in this country. Then there have been long haired specimens, described as Burmese, they have the same colouring as the seal point, and long fur, which although in no way comparable with the Persian yet is definitely more long than short. Experiments however have proved that in breeding Siamese to other species - varied forms can be made - this practise is not to be encouraged, as it may lead to definite malpractices as have occurred in other livestock."
Her comments show some confusion between Burmese and Birman. The "Black Siamese" is probably the Foreign Black; essentially a Seal Point Siamese without the colourpointing gene.
While they became more popular in the US and Europe, the purebred Siamese was dwindling in its native Thailand. In the 1950s, a breeder obtained 3 Siamese kittens of the robust type from Thailand. While the debate over "2 types" of early Siamese continues to this day, what is certain is that early Siamese cats were far more moderate in type than either of their modern counterparts. During the early 1960s, the "robust" Siamese lost out to the longer thinner type. While owners often preferred a moderate cat, exhibitors preferred an extremely slender cat with a very long, triangular head, almond-shaped eyes, and flaring ears. This look caught on with show-oriented Siamese breeders and judges. A cat called "Fan Tee Cee" was an early example of the increasingly extreme conformation that was impressing the judges. Fan Tee Cee appears many times in some pedigrees, demonstrating his contribution to the new-style Siamese. The growing number of new-style Siamese put pressure on cat fancies to change the breed standard to reflect the changing shape, putting old-style cats at a disadvantage.
The original Siamese cats were Seal Points ("black" points) although Blue Points and Chocolate Points also appeared in Siamese litters and were for many years considered "poorly marked" Seal Points. In the USA at the end of the 1960s, breeders in Britain and the USA were working with different colours of Siamese-type cats e.g. ebonies, reds, chestnuts and lavender. Lynx -points (tabby points) were being bred in Britain. Because blue-pointed Siamese appeared spontaneously in seal-point litters, they were known as "sports" or "freaks". In 1896 Louis Wain, father of the modern cat show, refused to judge a Siamese on grounds that it was blue rather than seal. When the first book on Siamese was written by Phil Wade she wrote, ‘"Even the best blue pointed cannot, I think, equal the beauty of our seal pointed cats and I can see no real object in trying to breed them. Their value at the moment is their scarcity, but I cannot believe there will ever be a great demand for them."
Tabby Point Siamese had been mentioned as early as 1902 in Britain. Between 1944 and 1949, they were bred in Scotland and known as Silverpoint Siamese. They were introduced to the cat fancy at a London cat show in the 1960s and in 1966, the Tabby Point Siamese was granted recognition by the Governing Council of the Cat Fancy (GCCF).
By 1986 the old-style Siamese had vanished from the showbench and many people assumed that Siamese cats had always had a long body, wedge-shaped head and disproportionately large ears. Siamese cats had become extreme parodies of the original imports. In Australia, some Siamese appear even more extreme than their American cousins - with larger ears and even more fragile bodies. In Britain they have also become more extreme - and ugly - in type, especially facially. It really is up to judges to do a sanity check and prevent the cat from being taken to ever greater extremes of type.
In the USA, the old-style cat was still being bred in a small way even though it could no longer compete against the modern style cats. It was popular with the general public. There is a separate Siamese breed known as the traditional and recognised as distinct from classic Siamese by some registries in the USA. The term "Old-Style Siamese" is often used.
In Britain, interest from the public wanting to buy old-style Siamese cats has led to a revival of the Traditional style Siamese during the late 1990s. Those would be owners expressed dismay at the wedge-headed Siamese seen on the showbench. On mainland Europe the Thai Pointed is a naturally occurring colourpointed (Blue-point, Lilac-point) Korat variant. The Thai Siamese is a European term for the traditional (round-headed) style of Siamese derived purely from non-ultra-typed Siamese cats. The term Thai Siamese has sometimes been used for colourpointed Korat variants.
In the USA, the Siamese is recognised in seal point, blue point, chocolate point and lilac point. Other colours (red point, cream point, tabby point, tortie point etc) are known as Colorpoint Shorthair (Colorpoint Oriental Shorthair). Elsewhere, these other colours are classed as Siamese. The additional colours were introduced through out-crossing, however they have been so extensively back-crossed with Siamese cats that only the most ardent purist would claim that they are not Siamese! The additional colours are not permitted in the traditional old-style of Siamese which seeks to breed only the traditional colours as well as the more moderate conformation.
Lynx point (UK) 1990.
Lynx point (UK) 1990.
 
Breakthrough Tabby Pattern in Colourpoints
Cats born of Siamese parents sometimes have a tabby/spotted pattern breaking through so strongly that the cat appears to be a tabby/spotted Oriental or a tabby/spotted sepia (Burmese) pattern. Viewed closely, the pattern colour is heavily ticked on a paler ticked background. This breakthrough pattern is evident from kittenhood and not the same as age-related darkening in Colourpoint cats. It is inherited, possibly due to modifier genes that prevent the colourpoint gene from properly inhibiting the colour/pattern on the normally pale areas of the body.
A stongly marked seal-tabby Siamese with a breakthrough spotted pattern (looked like a Spotted Tabby Oriental with blue eyes) mated to a Siamese produced a litter containing one strongly marked classic tabby that looked more like a Classic Tabby Oriental with blue eyes. The breakthrough pattern has become a problem in some Australian lines. Another example was a lynx-point Siamese female (old-style conformation) in England that looked like a slightly washed out tabby with blue eyes. Her colour raised questions as to whether she was genuinely colourpoint or was genetically mink or sepia pattern from Tonkinese lines.
BALINESE/JAVANESE AND SINGHALESE
As with the Siamese, there are 2 types of Balinese - Balinese and the traditional style of Balinese (Apple- or round-head). In both cases, they are semi-longhaired versions of the 2 different types of Siamese
Semi-longhaired variants have appeared now and again in purebred Siamese litters. There have been various explanations. The recessive nature of the longhair gene means it can be carried for many generations without coming to light - in which case the gene was present in one of the early imports of Siamese. Some say that at least one Chinese tapestry depicts pointed longhairs. Longhair can also occur as a simple mutation; again the gene may be carried hidden for many generations. An unlikely explanation is that it came from the domestication of the longhaired Pallas cat (F manul) of western China although genetic evidence is lacking. More feasibly early british breeders may have outcrossed early Siamese imports to longhaired cats or to shorthaired cats which had a recessive gene for longhair. Possibly they were crossed with Turkish Angoras, another breed popular at the time.
After many years of hiding their existence, a "Longhair Siamese" was registered with the Cat Fanciers Federation in 1928, however they were not bred in earnest until 1955 when Marion Dorsey began breeding and showing the longer-haired variety in the USA. Since long hair is a recessive trait, these long-haired Siamese bred true. To keep it separate from the conventional shorthaired Siamese it required a new name. "Balinese" was chosen to reflect its grace , like that of Balinese dancers, and also because Bali is close to Thailand and this reflects the breed's links with its Siamese ancestors.
In 1961 it was recognized in the USA and accepted for registration in the same point colours as Siamese: seal, chocolate, blue and lilac. Other colours were introduced from outcrossing: red, cream, tabby, tortie, cinnamon, fawn, smoke, silver and more. These were registed in the USA in 1979 as Javanese, but are considered Balinese in other countries since they have been so extensively back-crossed to Balinese that they can no longer be considered hybrids .
The development of the Balinese paralleled that of the Siamese. In the 1950s most Siamese and Balinese cats were less extreme than we are used to today. The more extreme look gained in popularity for the Siamese and its longhaired counterpart followed suit since Balinese were bred back to Siamese to improve the type of the Balinese. In addition, much of the Balinese cat's long fur was lost, leaving it as a more-or-less shorthaired cat with a plumy tail, a side-effect of breeding back to the Siamese. As with the Siamese, a few breeders preserved the older style and it is now becoming more popular in its own right.
In the 1960s, a cat very similar to the Balinese was bred in the USA and Canada . These resulted from crossing Siamese cats to red/red-tabby Angora (presumably the Turkish Angora) resulting in semi-longhairs with Siamese conformation and red or tortie points. The fur was shorter, but fluffier than the Balinese and the tail much fluffier. The temperament was also calmer than the Siamese due to the Angora influence. Essentially these were the red-series equivalents of the Balinese which US registries recognised only in seal, blue, lilac anc chocolate. The Singhalese was allowed to be bred to either Balinese or Himalayan and the progeny could be registered as Singhalese. It was judged to the Balinese standard with allowances made for its different coat type. Over time, it appears to have lost out to the Balinese.
BIRMAN
The Birman has the colourpoint pattern with the addition of white bootees/gauntlets. The Birman arrived in Europe around 1920 and has also been known as the Tibetan Temple Cat and the Sacred Cat of Burma . Modern Birmans are descended from a pair brought to Europe by Mr. Vanderbilt. It is also reported that 2 Englishmen, Major Gordon-Russell and August Pavie, then living in France, received a pair of Birmans from the Kittah people in 1919 as gratitude for their part in saving the temple from being overrun by invaders. The remainder of the tale is the same: the male unfortunately died during the voyage, but the female was pregnant. Of her kittens, only a female survived. This meant two female Birmans and no male. To re-establish the type, these cats were crossed to Siamese, Colourpoints and White Longhairs.
In 1927, judge Mrs Basnett reported on the Paris Cat Show held on 14th and 15th of January by the Cat Club de France and wrote "The Sacred Burmese Temple Cats interested me very much, with their long fur on the tail and coat resembling that of a poorly bred Persian; their colouring is exactly like that of the Siamese, but their feet sometimes have white toes. I was given to understand that they are very difficult to rear, only about one in ten survive . I do not think they possess the same quick movements as the Siamese, life to them seems much more dreamy and slow , but they are very loving and intelligent." This clearly referred to the Birman; confusingly the name Burmese Temple Cat was also used at that time for the gold-eyed brown Thai cats analogous to modern Burmese or brown Orientals.
As a result of its early near-loss and re-creation, some say the Birman comes from temples in Burma, while others claim it is a breed entirely manufactured in France using Siamese andblack-and-white longhairs. French breeders had to re-create the Birman not once, but twice since the Second World War caused pedigreed cats in Europe a great set back. Only two purebred Birmans survived and their offspring formed the new post-war foundation of the Birman breed in France. They were bred with other longhairs out of necessity, but by 1955, the Birman was once more recognised as pure bred and there was no more outcrossing. In the early 1960's Mrs Elsie Fisher and Mrs M. Richards imported the first Birmans into Britain from France.
While some claim the cats were created in France, it is interesting that in 1960, a pair of "Tibetan Temple kittens" was given to a North American cat lover and were identical to the Birman cats being bred in Europe. In recent years new point colours have been introduced, including chocolate, lilac point, red, cream, tabby and tortie point. Not all societies recognise the newer colours. It's worth noting that the older style of Birmans tended to develop "hood-type" head colouring.
Early in the breeding of Birmans in the USA, Gertrude Griswold attempted to keep the Birman breed under her control. It was possible to adopt a female Birman from her for breeding, but to actually breed the cat, there had to be a notarized agreement (a contract with the Griswolds) that the sire would be of Gertrude Griswold's choosing and the adoption of the resulting kittens into new homes would be under her jurisdiction. Later on, Ann Baker attempted a similar degree of control over Ragdolls, another colourpointed breed.
Birman variants are known e.g. the " Khmer " being a bootee-less Birman-type cat and the Tibetan being a Birman in solids colours and tabbies. Some of the early Birmans lacked the gauntlets.
TEMPLECAT (BIRMAN SHORTHAIR)
The Templecat is the shorthaired equivalent of the Birman. It is a Birman in every respect apart from the slightly springy short hair. It was developed in New Zealand by outcrossing Birmans to a cinnamon spotted tabby Oriental.
The Templecat began development in 1995 and was originally called the Birman Shorthair. It later became the Tsuncat (after a temple priest in mythology surrounding the Birman cat) and became the Templecat in 2001.
The early colours were seal point, blue point, chocolate point, lilac point, cinnamon point and fawn point It is now also in red point, cream point and in tabby and toritie points.
HIMALAYAN
The Himalayan is a Persian in a colourpoint coat and with blue eyes. In the UK it is known as the Colorpoint Longhair (or Colourpoint Persian). In the USA, Himalayans with tabby or tortie points are known as Colorpoint Longhair in some registries.
Experimental breeding of Siamese and Persians dates back as far as the 1920s . The first recorded deliberate cross between Siamese and longhaired cats was made by Swedish geneticist T Tjebbes in 1924 using white longhairs. However it was not his intention to create a new breed for the cat fancy. In 1931 in the USA, Virginia Cobb and Dr. Clyde Keeler began an experimental breeding programme with the purpose of learning the inheritance involved in producing a Colourpoint Longhair. In 1936, they produced the first Himalayan kitten. They published an article about colourpoint inheritance in the American Journal of Heredity. Having accomplished their aim, the breeding programme was abandoned.
Sydney W France's book "Siamese Cats" (1949) also mentioned the crossing of Siamese cats with Persians, a practice which eventually led to the now familiar Colourpoint Persians (Himalayans). In 1949, such breedings were regarded with a mix of curiosity and disapproval.
"Some curious experiments have been made from time to time in an effort to cross Siamese cats with Persians and even the tabby cat, and at least two treatises have been written on the subject. "Siamese-Persian Cats" by Clyde E. Keeler and Virginia Cobb, "Journal of Heredity" v. 27. No. 9. Sept. 1936, and "Crosses with Siamese Cats" by K. Tjebbes, Journal of Genetics, V. 14. p. 335, 1924. From this we find that Swedish Dr. K. Tjebbes in about 1924 crossed a white Persian female with a Siamese and the colour of the Persian dominated to the extent of seven white kittens and three coloured ones. Back crosses all gave 50 per cent . white. Like Siamese colouring, short hair dominated.
It took Mrs. Virginia R. Cobb five years of experiment working in conjunction with Dr. Clyde E. Keeler of the Harvard Medical School to produce the first successful experiment and breed long-haired Siamese kittens, using black Persians instead of white, as used by Dr. Tjebbes in his experiments. Only pedigree cats were used, and from each litter only the most perfect kittens were selected, to be mated up in due course to carry on the experiments. A black Persian male was mated to a Siamese female, and a Siamese male was mated to a black Persian female. The kittens in every case were black and short-haired. After a time a female of one of these litters was mated to a male of the other, and produced among her kittens a long-haired black female! this female being bred back to her short-haired black father. She subsequently had a litter containing two Siamese-Persian kittens which had the long hair of the Persian and the markings of the Siamese. This mating was repeated and of the eleven kittens three were long-haired Siamese, the other eight being black. These three long-haired Siamese kittens had the blue eyes of the Siamese, and the same voice, which they used just as often as the true Siamese do.
In 1939 when Ida M. Mellen reported these facts in her practical cat book the experiments were still proceeding."
In 1931, Virginia Cobb ( Newton Cattery) and Dr. Clyde Keeler (Harvard Medical School) conducted an experimental breeding programme to produce a Colourpoint Longhair. They wanted to study the inheritance of longhair and colourpoints to produce a "longhaired Siamese". They first crossed Siamese with Black Persians. This resulted in black shorthair kittens. They mated the offspring together and achieved the desired longhaired Siamese. In 1936, Keeler and Cobb produced the first Colourpoint Longhair (Himalayan) type kitten, "Newton's Debutante". They published an article and the " formula " for producing her in the American Journal of Heredity and then abandoned the breeding programme. They had accomplished the goal of studying how the factors were inherited (longhair and colourpoint are both recessive), but did not intend to create a new breed.
In Britain, at the Siamese Cat Club Show in 1947, Brian Stirling-Webb was approached by a cat owner, Mrs Barton-Wright, who wanted to mate her "long-haired Siamese queen " to one of his Siamese stud cats. She lived close to his Briarry cattery and was anxious to breed more long-haired Siamese, believing they would make a great addition to the Cat Fancy. Stirling-Webb disagreed and told her that she would get only short-haired kittens from mating her cat with a Siamese. Stirling-Webb had seen the results of Keeler and Cobb's experiments and described their longhairs as having Siamese type (far less extreme in type than modern Siamese). He had felt that Keeler and Cobb's experiments were the worst possible thing that could happen to the Siamese - the Siamese cats' beauty was due to their svelte outlines, which Stirling-Webb felt would be obscured by a long coat. Stirling-Webb therefore recommended either having the queen spayed or mating her to a Black Persian and then mating the offspring together. The owner was "averse to either course" and asked to take the queen to Stirling-Webb for him to see her for himself .
Stirling-Webb later wrote: "When I saw this queen, I was astonished at her beauty. Apart from her colouring, she possessed practically no Siamese characteristics, and was reasonably Persian in type. I was also amazed that nothing was known of her origin due to the fact that she had been a stray living in a country churchyard for about six weeks before being adopted by this lady. The police had been informed of her adoption, and advertisements were placed in the local papers in an attempt to contact the original owner, but without success." The stray queen later became known as "Bubastis Georgina". Unfortunately her habits meant she was "quite unfit to live in a civilised house" and in spite of persevering with her, her adopter offered her to Stirling-Webb as a cattery cat. By then, Stirling-Webb was enthusiastic about producing colourpoint Persians (but emphatically against longhaired Siamese!). He was joined by Mrs SM Harding (Mingchiu cattery) and began to develop the variety further. He repeated Cobb and Keeler's "formula" and also acquired a Black Longhair male who had one Siamese grandparent. This male was mated to the colourpoint queen and they produced kittens close to his ideal.
Getting GCCF recognition for Colourpoint Longhairs (Persians) was harder as he needed to produce 3 generations of Colourpoint Longhairs for it to gain breed status and his stock was becoming inbred. In 1955, the cats achieved breed recognition and to continue outcrossing his breeding stock. He recommended the use of "bad" Siamese (round-headed) in Colourpoint Longhair breeding programmes. From the outset, the variety was considered a colour variety of Longhair (Persian): Breed Number 13B. Side-effects of his efforts were the Self Chocolate Longhair (Briarry Bruno ) and Self Lilac Longhair (Minghiu Lilac).
In the 1940s and 1950s, Ben and Ann Borrett bred cattle at their Chestermere ranch in western Canada. They were also lovers of Siamese cats and became aware of Stirling-Webb's experimental breeding programme in England. The Borretts travelled to England to visit Stirling-Webb and to purchase several of his cats to start their own breeding program . Their experience as cattle breeders meant that the Borretts understood the amount of work required to establish the new breed and the need for multiple bloodlines. They built up a large cattery (Chestermere cattery) and developed several different and totally separate bloodlines. Only the very best cats from each generation were bred. In 1957, they exhibited two of their imported cats at an ACFA show in Calgary, Alberta, Canada and were asked to create the "Himalayan" breed standard (Himalayan is the name given to the colourpoint pattern in rabbits and goats from the Himalayan region).
Around the same time, California artist and cat breeder Marguerita Goforth (Goforth cattery) agreed to look after a friend's cat. The cat was a seal-point Longhair called "Princess Himalayan Hope" and had come from the San Diego Humane Society. Goforth gained permission to breed the cat to create a Persian-type cat with Siamese colouring. When seeking acceptance for the breed in the USA, she used the Borretts' breed standard. In the 1960s, Goforth wrote that she regretted the decision to recognise the Himalayan as a distinct breed: "I have felt for many years that the Himalayan cat should be recognized as a colour of Longhair". While the Himalayans were kept separate from Persians they began to diverge in type, becoming a long-nosed, colorpoint longhair.
In the 1950s, the Himalayan was recognised in the same 4 colours as the Siamese of the time: seal point, chocolate point, blue point, and lilac point. Red (flame) points and tortie points are recognised in the USA in 1964. Over the next several years these were joined by blue-cream point, cream point, lynx (tabby) points and tortie points. In 1984, the CFA made the Himalayan a division of the Persian breed.
Left: Flame-point (red-point) Himalayan
Right: Seal Point Himalayan
Bottom: Chocolate-point and Seal-point Himalayans
Only a few Himalayan breeders will ever breed bicolour Himalayans, using bicolour Persians to introduce the white spotting pattern. Paul Beall, Richmond, Texas is one such breeder. Paula's photo (left) 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. At present, Paula is working with the cinnamon gene in Himalayans; though the gene should have been present in Siamese cats used to develop the Himalayan, it does not seem to hve shown up in the Himalayn breed and breeders may have to introduce it using out-crosses to Persians.
For a while in the 1950s there was a French breed known as the Khmer. It resembled a semi-longhair colourpoint and was apparently overtaken by the Himalayan. "La Vie a Campagne", 1935 dated Khmers back to the 1920s and the photographs resembled early Colourpoint Persians. The story goes that a pair was taken to Paris and abandoned there by a returned serviceman from Indochina. By 1935, this strain of Khmer had evidently almost petered out. The French Khmer is also been described as a bootee-less Birman-type cat. The Khmer is described in an aside by Fernand Mery in 1966 when he describes the "Colourpoint Longhair" as being "remarkably like a breed recognised in France, the Khmer, which has its own standard in that country, though it is not recognised by La Federation Internationale Feline d'Europe". The term "Khmer" had actually been dropped in 1955.
COLOURPOINT EXOTIC SHORTHAIR
The Exotic Shorthair is a shorthaired version of the Persian. It was developed during the 1950s and recognised in the mid 1960s. The Colourpoint Exotic Shorthair is an Exotic Shorthair with colourpoint pattern and blue eyes.
RAGDOLL AND RAGAMUFFIN
Ragdolls are medium to large boned, fairly long cats and possess the Siamese colourpoint gene and the white spotting gene. The have broad modified wedge shaped head - best described as a large piece of pie, with the corners rounded. Their ears are wide set and slightly tilted forward and the eyes are very blue, but not round in shape.
The Ragdoll comes in seal point, chocolate point, blue point and lilac point although red points, cream points and tabby points are now being bred. The "colourpoints" are exactly that. The "mitted" versions have the colour points but have white mittens, white boots and a white blaze on the nose. The "bicoloured" has additional white on the stomach, chest and ruff. Genetics means that there are three other patterns: "High Mitted" where the mitts extend up legs; "Mid-High White" which is a Bi-colour with additional white in the "saddle" area; and "High White" which is a Bi-colour with even greater degree of white, the "saddle" may be absent.
The colourpoint and white spotting genes interact to produce the different patterns. The three recognised patterns are colourpoint, mitted and bicolour. All Ragdolls are colourpointed and some also have white. Due to the way genes interact, some cats Ragdolls which look identical actually have different genotypes (different genetic make-up). For example, High Mitted, Mid High White and "true" Bicolour can all appear identical, though the exact amount of white is variable (almost infinitely so!) and some High Mitteds display more white than a show quality Bicolour while some Mid High Whites may have so much white on them that they appear to be High Whites! At times like this, only a knowledge of what genes the cat has inherited allows breeders to know what colour their cat really is!
High White is also known as Van pattern or Harlequin in some associations and is a recognised colour in CFA (America).
Ragdolls also produce pet quality variants which are not shown (except as household pets), but which have the same excellent temperament. The variants include white toes on colourpoint Ragdolls, dark toes on mitted Ragdolls, odd white marks appearing in dark areas and dark marks appearing in light areas! The white spotting is hard to get perfectly right!
The Ragdoll originated in California in the early 1960s and is surrounded by myth thanks to their somewhat eccentric breeder, Ann Baker. Ann Baker bred Persian cats. Josephine was a white Persian-Angora cat belonging to Baker's neighbour ; she was a semi-feral who produced kittens as wild as herself . After Josephine was hit by a car she had to stay indoors while recovering and she became somewhat tamer.
While Josephine's earlier litters had been half wild like their mother, the litters born after the road accident were very relaxed, docile and social. The exact reason why is not known and later became the subject of wild claims. Since kittens inherit much of their personality from their sire, the most likely explanation is that she was breeding with different males, ones carrying genes for more sociable offspring. Judging by the kittens' appearance, Josephine's suitors were most likely Birman and Burmese. In addition, Josephine was less wild and perhaps did not train her kittens to be quite so fearful of humans.
Baker decided to create a new breed, one which retained the very sociable, relaxed traits, non-matting fur, large size and the property of going limp when handled. She acquired a black Burmese-looking female called Buckwheat and a bicolour female called Fugianna. Fugianna's sire was another of Josephine's kittens, a seal-point Birman-type cat called Daddy Warbucks. There were to be no more founding cats since Josephine and her next litter were destroyed by their owner. Based on the appearance of the two founding females, Baker split their progeny into two groups: Fugianna's kittens were termed "the Light Side"; Buckwheat's kittens were termed "the Dark Side".
Ann Baker formulated a strict breeding policy to preserve the purity of the Ragdoll and charged other breeders (franchisees) a royalty fee for every kitten they sold . She also made wild claims about the cats: their docility was due to changes in Josephine after she was hit by a car (a long-discredited Lamarckian theory), that while Josephine was being treated for injuries she was infused experimentally with skunk and/or racoon and/or human genes (even if done , these genes would not have entered the egg cells in the ovaries), that Ragdolls were immune to pain (they have normal pain thresholds but are remarkably tolerant cats) and even that they are a link between us and space aliens! A more plausible suggestion was that Josephine had a mutation in her ovaries and this mutation was present in the egg cells (a germ-line mutation) and was passed on to her offspring.
Her claims and methods alienated other breeders, some of whom broke away to develop the Ragdoll into a registrable breed using sound breeding practices. Depending on which history you read, the Ragdoll was recognised as a purebred cat in 1965, 1966 or 1967. Also according to many sources, Denny Dayton achieved US recognition for the Ragdoll in 1967, however he didn't buy his first Ragdolls until November 1969. In 1971 Baker created the International Ragdoll Cat Association ( IRCA ) and claimed that non-IRCA cats to be fakes, frauds, look-alike, half-bred, not authentic etc while promoting her cats as the only legitimate Ragdolls. In 1975 Baker patented the Ragdoll name; she died in 1997 but her patent remained valid until 2005 and allowed only IRCA breeders to use the name "Ragdoll". Earlier breakaway breeders felt that the restriction did not apply to them because they had purchased and bred their cats prior to the time of the patent. In 1981, Ragdolls were exported overseas for the first time.
The stories of the two types of Ragdoll ran parallel for many years. Baker's trademarked umbrella term for all of her IRCA varieties was Cherubim Cat meaning "Angels non-fighting cat". In 1985 and 1987, one UK breeder also advertised "Cherubinis" which appear to have been non-colourpointed offspring from outcrossing Ragdolls to other breeds. "Cherubini" was a legally safe equivalent of IRCA's trademarked "Cherubim" name. The IRCA Ragdolls were not recognised by any other registry and upon Baker's death, a number of breeders re-registered their Miracle Ragdolls, Honeybears and other IRCA varieties as foundation stock for the new RagaMuffin breed which is now recognised by many registries around the world. The RagaMuffin is not merely a non-pointed Ragdoll, it also has a different conformation from the Ragdoll. After Baker's death, some breeders tried to sell their IRCA Ragdolls as "rare mink-pointed Ragdolls" or "rare solid colour Ragdolls" to unwary owners/breeders who did not know that there had been two conflicting Ragdoll breeds. These "rare" colour cats, though attractive, were not registrable as Ragdolls in any other cat association.
Throughout her life, the eccentric Baker defended her breeds and claimed that only IRCA Ragdolls were the real article and guaranteed free from genetic defect, while other Ragdolls were half-bred or cross-bred inferior lookalikes trading fraudulently on the Ragdoll name. Quarter-page advertisements to this effect appeared in an American cat magazine in 1992. Periodically, she would send threatening mail to breeders non-IRCA Ragdolls, including a mailshot that contained photos of dozens of dead cats laid out side-by-side along with a claim that someone had broken in and killed her cats (some believe Baker killed her own cats and might not have known what she was doing due to a claimed history of head trauma ).
The RagaMuffin is another descendent of the original Ragdolls, but in a more extended colour range. Their similarity to the accepted Ragdoll meant that many registries would not accept them, though the conformation is not the same as the TICA Ragdoll (note: the NZCF accepts "Solid Ragdolls instead). In the days when Ann Baker and IRCA were active , a there were a number of other "breeds" or named breeding lines. IRCA's other named breeds/lines were the Honeybear/Honey Bear (resembled old style Persians), Ragdoll Hobby Cat, Miracle Ragdoll ("a highly upgraded Ragdoll" formerly known as the "Ragdoll Tu", "experimental Persian" or simply "Miracle"), Baby Dolls/ Doll Babies, Angels, Shu Schoo, Symoneese/Symonees, Manxees and Fuzz. The named IRCA Ragdoll/Miracle breeding lines included Maximillions (silver tabbies), Minks, Bears (thicker fur, shorter ears, rounder eyes, flatter face, cobbier body) and Catenoids (blue-eyed solids surrounded by claims they could be bred to any cat to produce an IRCA-type Ragdoll!). After the demise of Baker and of IRCA, many of these were absorbed into the RagaMuffin and some were apparently re-registered as TICA Ragdolls.
SNOWSHOE, SNOWSHOE SIAMESE AND SNOW -TOES
Originally, the Snowshoe (or Silver Laces Cat) resulted from crossing Siamese, or Oriental Shorthairs, with American Shorthairs. The white mitts occur natually in some lines of Siamese where they are considered a fault. Today Snowshoes generally result from Snowshoe to Snowshoe breedings although they may still be outcrossed to one of the founding breeds in order to expand their gene pool, introduce new colours, or improve type or colour/pattern. Officially the breed claims no Birman ancestry despite the superficial resemblance in pattern.
Snowshoes were developed in the USA by Dorothy Hinds in the late 1960s when her two Siamese cats produced kittens each with four white feet. White spotting sometimes showed up in early Siamese cats (white toes are occasionally still found). She dubbed them Snowshoes. The variant did not recur so she crossed a Siamese with a bicolour American Shorthair. In addition to the white mitts, this eventually produced the inverted white 'V' on the face.
The Snowshoe was recognised by various American registries in 1974, however in 1977 there was only one registered breeder of Snowshoes. Luckily, interest in the breed picked up and the Snowshoe has become a popular breed in the USA.
It has Siamese style points and blue eyes combined with white paws and white spotting and comes in blue, seal, lilac and chocolate points. The Snowshoe pattern is divided into two categories: Bicolour Snowshoes where up to two-thirds of the cat can be white and Mitted Snowshoes where the white ranges from just the toes to halfway up the leg. Snowshoes have a white inverted 'V' extending from between the eyes down over the nose.
Early on the breed was erroneously thought to be a Birman Shorthair, based purely on its appearance. The Birman Shorthair (Templecat) only came into existence in 1995 in New Zealand.
The Snowshoe conformation is intermediate between the parent breeds and is sometimes described as close to the older (or traditional) and more moderate style of Siamese cat although it is sometimes described as resembling the American Shorthair. The Snowshoe inherits the muscular body from the American Shorthair and the body length from the Siamese/Oriental. The conformation is, to some extent, affected by the outcross breeds permitted in countries where the American Shorthair is not found.
In the 21st Century , an experimental breed called the Snowshoe Siamese has also been announced. Its body type is more extreme and is closer to the modern Siamese, while the original Snowshoe maintains the more moderate body type. Some Siamese cats have appeared with white toes and this was considered a serious fault in the breed.
The Snow-Toes was bred in the late 1960s by crossing Birmans to Himalayans, seemingly with the aim of a mitted Himalayan. Whether this will see a revival alongside the Snowshoe Siamese has yet to be seen!
NEVA MASQUERADE
In 1990, the first Siberians were imported into the USA. However the colourpointed variety with blue eyes was not imported and colourpoint Siberians were not recognised by registries on in American breed standards. However Russian breeders have recognised the colourpointed variety since they began to keep records (in around 1987). According to anecdotes, colourpointed feral cats bred with feral Siberian cats along the banks of the Neva river at St Petersburg (then Stalingrad). The name Neva Masquerade reflects the origins of this masked cat.
The first Siberian cats were imported into the USA in 1990 by Elizabeth Terrell of Louisiana, in exchange for Himalayan cats sent to establish that breed in Russia. The first colourpoint Siberians were imported into the USA in 1997 and registries are have recently accepted it following early controversy.
TONKINESE
The Tonkinese is intermediate in type between Burmese and Siamese. Its pointed pattern is less pronounced than that of the Siamese and its shape is less extreme. Long known as a variety in Thailand, it has existed in the West for at least a hundred years. Towards the end of the 1800s, a cat described as a chocolate variety of Royal Siamese was exported from Singapore to England. It was a rich chocolate colour with a darker face, ears and tail; and its legs were darker than the body. It was not, however, recognised as anything other than a brown Siamese and interest in them had waned because it had been hard to compete against the striking coat and eye colour of the Siamese. These cats were either Burmese or Tonkinese, or most likely a mix of both types since some, when bred to the royal Siamese, produced offspring with "poor" markings (i.e. "mink") while others evidently produced offspring had royal Siamese markings. In 1903, the "Burmese Cat" (shown here) would be, in modern terms, an Oriental Ticked Tabby!
The Tonkinese is considered a hybrid of Siamese and Burmese and this has hindered its acceptance in some countries e.g. Australia because "as a hybrid it cannot breed true". The alternative standpoint is that it is that Tonkinese is the naturally occurring intermediate form of a colour range where the two extremes are represented by Siamese and Burmese. The two extreme forms have been selectively bred, but the intermediate form was ignored until relatively recently. Tonkinese type cats have been described for many years: the Chocolate Siamese (Britain, 1880s; generally considered to be Burmese), Golden Siamese (USA, 1950s, Siamese/Burmese hybrids),Golden Chechongs, Si-Burms and Zibelines (the French name for Burmese). These all had tan to brown bodies with seal or almost black masks and points.
The self-coloured Chocolate Siamese (as opposed to the Chocolate Point Siamese) was described by Harrison Weir in 1889 and by Frances Simpson in1903, however the "Royal Siamese" with its greater contrast between body and points was the form that predominated in Britain and the Chocolate Siamese was lost. The Chocolate Siamese is variously claimed to be an early Burmese or an early Havana Brown type cat. The founding female of the Burmese breed, Wong Mau, was a walnut-brown Tonkinese type cat imported into the USA from Rangoon in 1930. She was described as brown with darker points on her face, legs, feet and tail. When bred to a seal-point Siamese, she produced kittens identical to herself and some with a Siamese coat pattern. Over several generations and through mating her to her own sons , three varieties of offspring arose: those with a Siamese pattern, those which were dark brown all over (Burmese) and those with a dark brown body but identifiable darkening at the points much like Wong Mau herself. Wong Mau is considered to be the founding mother of the Burmese breed but was herself a Tonkinese and breeders chose to develop her Burmese offspring.
In the early 1940s, a Hawaiian breeder, Lelia Volk, bred a dark variety of Siamese: its colour was lighter than the Burmese, but darker than the Siamese. In a report in the Journal of Heredity, the colour was said to be close enough to the Siamese that it was unlikely to be perpetuated. Between the 1950s and early 1960s, Milan Greer in New York City bred Golden Siamese. These were Siamese/Burmese hybrids which had a rich mahogany body and dark points. Greer had been told that such cats were normally considered defective and destroyed because they did not breed true. Using Seal Point Siamese and Sable Burmese and cross-breeding their offspring, he produced chocolate brown cats with darker points and claimed to have bred pure "Golden Siamese" for 5 generations. Having proven his point, he stopped breeding, though his cats were popular with pet buyers.
In the 1960s and 1970s, the Tonkinese was revived in Canada and the USA by crossing Siamese with Burmese in spite of opposition from breeders of both Siamese and Burmese who viewed them as embodying everything they were trying to breed out of those two breeds. In the mid 1960s, Jane Barletta of New Jersey switched from the increasingly extreme Siamese to breeding the more moderate Tonkinese. At about the same time, Margaret Conroy in Canada bred a female Burmese to a Seal point Siamese as she was unable to find a suitable Burmese stud in her locality. Her tan-bodies cats bred true for 5 generations and were dubbed Tonkanese (with an "a"). The original cats were Sable (US: Natural Mink), but crosses between Blue Burmese and Blue Point Siamese introduced Blue Tonkinese. Later on, the other Siamese/Burmese colours were also bred together to expand the Tonkinese colour range: red, lilac (US: platinum), chocolate (US: champagne) and cinnamon (US: honey).
Though the Tonkinese breed in the USA has been recreated by crossing the Siamese and the Burmese, the variety has occurred naturally in south east Asia for hundreds of years. Rather than being a hybrid, it is a naturally occurring breed and the Burmese, Tonkinese and Siamese represent the two extremes and the mid-point of an overall genetic pattern caused by 2 mutant genes. Recognition in various cat fancying countries came from the 1980s onward. In the UK, it was recognised in 1990. In Australia it was grudgingly accepted in the 1990s despite being condemned as a hybrid and unable to breed true.
The colours of Tonkinese are (the term "mink" is not used by some registries) blue mink, champagne mink (light chocolate), honey mink (chocolate/chestnut), natural mink (seal), platinum mink (lavender/lilac), red mink, cinnamon mink, fawn mink, cream mink. In some registries Burmese are accepted in tortie patterns and this will be reflected in the Tonkinese. Its genetic make-up means that Tonkinese-to-Tonkinese breedings will always produce a mix of patterns: self (Burmese-type) variants, Tonkinese and colourpointed variants. The colourpointed Tonkinese find favour with those who seek a less extreme version of the Siamese. Although most registries don't accept the variants for exhibition, the variants are an important part of the breeding program; bred together they produce Tonkinese-pattern cats.
COLOURPOINT SHORTHAIR (COLOURPOINT BRITISH SHORTHAIR)
 
The Colourpoint British Shorthair is exactly what the name suggests - a British Shorthair in a colourpoint coat with blue eyes. It was developed in the 1980s by crossing the British with the Himalayan.
OPAL (COLORPOINT AMERICAN SHORTHAIR)
In the 1990s, breeders of appleheaded (roundheaded) Siamese were invited to register their cats as Opals. Little has been heard of the Opal since then although the traditional style of Siamese goes from strength to strength (despite the registry's rather bizarre refusal to allow its actual breed name to be used in articles or reviews in case it erodes, rather than promotes, the breed). The Opal appears to be a Colourpoint American Shorthair.
THAI POINTED AND THAI SIAMESE
The Thai Pointed is a naturally occurring colorpointed (Blue-point, Lilac-point) Korat variant. Korats are related to Siamese and have interbred naturally in their native Thailand.
The Thai Siamese is a European term for the traditional style of Siamese derived purely from non-ultra-typed Siamese cats.
The term Thai Siamese has also been used for colorpointed cats derived from naturally occurring colorpointed variants in Korat litters, but this usage is not encouraged.
Andrea and Beata Brunetti, in Italy, have a Thai (the European name for the less extreme, traditional or old style of Siamese cat) called Urban Hunter with a visible blotched tabby pattern and an interesting dorsal stripe. Urban Hunter has been mated to a red-point Thai called Kasia Ruda, resulting in one male and four female kittens. All of these have dorsal stripes similar to Hunter.
In some lines of old type Siamese, there is less contrast between body and points. The Siamese pattern is caused by an albinism gene, but is partly dependent on temperature, with a darker colour developing in cooler regions. This is unlikely as Urban Hunter lives in a climate where the temperature ranges between 25 and 30 celsius.
Polygenes can also help to allow or suppress the colour. Variable contrast between body and points is seen more often in the older style Siamese, probably due to greater genetic diversity compared to the modern extreme-type cats. It is also possible that a novel mutation has occurred as Urban Hunter has aqua , rather than Siamese blue, eyes. Because of the aqua-blue eyes, I initially wondered if Urban Hunter had Tonkinese in his ancestry (Tonkinese is intermediate between Siamese and Burmese in colourpoint). Genetic testing and test-mating would be required to determine whether Hunter’s colouring is due to a mutation of the colourpointing gene.
AUSTRALIAN CLOUD
 
 
The Cloud is an Australian shorthair/semi-longhair bred in mink colors. This does not appear to be a recognised breed, but is a cross-breed from Birmans, Ragdolls and British Shorthairs.
Some photos depict a Ragdoll-like cat, others depict a cat similar to the traditional style of Siamese or a colourpointed British Shorthair (as seen here). The accompanying photo is provided by an owner of a cat bought as a "Cloud".
 
KUCING MALAYSIA/PIAWAIAN KUCING
Kucing is Malaysian for "cat". The Kucing Malaysia is similar in type to the Tonkinese with a colourpointed pattern similar to Ragdoll and blue/blue- green eyes. A White blaze on face and muzzle is desirable.
 
FRENCH KHMER
A French breed known as the Khmer was described by Mery in 1966. He noted the similarity of emerging colourpoint Persians (Himalayans) to the existing French Khmer, although the photographs of the Khmer suggested a semi-longhair or even a a fluffy , cobby colourpoint shorthair. Although Mery described it in 1966 and noted that it had a breed standard albeit one not recognised by FIFe, the term Khmer had been dropped back in 1955 when Colourpoint Longhairs and Himalayans were being developed in America/Canada and Britain respectively. There are earlier descriptions of the Khmer dating from 1935 (by which time it was close to being lost) and photos from that time depict a longhaired cat similar to an early colourpoint Persian. "La Vie a Campagne" in the late 1920s showed a longer haired Khmer, so possibly the coat length was seasonal. A pair of Khmer cats were supposedly taken to Paris from Indochina by a French serviceman - a tale very similar to that of the Birman. Two Birman were reputedly obtained by Major Gordon Russell from Burmese priests whom he had helped escape to Tibet; when he went to live in France in 1919 he took his Birman cats with him. This suggests that the French Khmer was a Birman that lacked the characteristic white markings of the Birman. This corresponds to the use of the term Khmer to describe bootee-less Birmans - and at one time there were indeed unsuccessful attempts to create"Himalayans" from Birman stock.
 
MASKED SILVER
A variety described by Mery and others in the 1960s, this is not related to Siamese cats but was considered a form of shaded silver (agouti + silver) longhair with a black mask on its face. It is a poorly marked black smoke. Similar cats appear in black smoke (non-agouti + silver) individuals of curly -haired breeds due to the black colour being more intense on the straighter fur of the face and legs.
Martine Sansoucy of Butterpaws LaPerms has seen a number of masked silvers and notes that they appear to be smokes rather than shaded silvers. Butterpaws BC The Crow, known as "Cairo" is a black smoke that meets the general description of a masked silver. As can be seen from the photo, the different textures of fur on the face and the body give the impression of a masked cat.
Historical Masked Silver.
Masked Silver (black smoke) LaPerm.
Historical Masked Silver.
Some authors have written that the masked silver dates back to 1900, but may have been referring to shaded silvers, a variety not recognised in Britain at the time and considered to be a badly bred chinchilla or poor quality smoke. Chinchilla and shaded silver are genetically tabby cats, smokes are genetical solid colour cats; this difference was not understtod in the early 1900s. According to Milo Denlinger in 1947, "Masked silvers are a new variety and very few are bred." Denlinger went on to describe the variety: "The ideal masked silver is a very beautiful animal; in colouring or, I should say, marking, they should resemble the Siamese Cat; that is to say, they should have a black mask, or face, black feet, and legs. The body should be as pale a silver as posible." The eyes were to be deep golden or copper . Several authors have observed that the description of the masked silver resembles that of the Siamese.
 
OTHER COLOURPOINTS
In addition to the recognised breeds, the colourpoint pattern is recognised as a permissible colour/pattern in many other breeds. In most cases the cat is simply described as a "colourpoint 'breed-name' ". The colourpoint Japanese Bobtail has only recently found its way into the West although it has existed for decades in Japan and a similar breed known as the Mekong Bobtail (formerly the Thai Bobtail) is known in Russia. Mekong Bobtails also occur naturally among feral cats in Malaysia and Thailand.
In others the prefix "Si" is used e.g. the Si-Rex is a colourpointed Rex and is further identified as Cornish Si-Rex or Devon Si-Rex. It exists unofficially or experimentally in a few breeds e.g. Si-Manx (Manxamese). In the American "Cats Magazine" of June 1965 there was a photo of two Blue-Point rex-coated kittens born to a blue self (Cornish) Rex in August 1964. Their pedigree showed that four generations back, there had been a Blue Point Siamese outcross in the breeding program. In addition, one ancestor was an Oregon Rex mutation owned by Mrs Mildred Stringham of Warrenton, Oregon. A Si-Rex was being developed in England at the same time, by Mrs Ashford (Annelida prefix).
At a Feline Fantasy staged by the Silvergate Cat Club of San Diego, USA. in 1952, a "Sianx" cat was exhibited. This was a Siamese-Manx cross with Seal Point colour and type, sans tail. However, the Western province Cat Club of Cape Town claimed precedence in Sia-Manx breeding. The late Father Fowler's Sia-Manx was exhibited at their show during 1952, some months before the cat at the San Diego show, and resembled a Siamese Seal Point with definitely high hind quarters and complete absence of tail. He was described as a very charming person by Miss P. Ashby-Spilhaus, the Registrar of the South African Cat Union.
The colourpoint trait can be introduced into any breed by crossing it with an existing colourpoint breed, although the first generation kittens will not show the trait. The gene for colourpoint has also entered the gene pool of free-breeding cats and there are many pretty "look-alikes" to be found in shelters and humane societies. Although not registrable as Siamese, Himalayan etc, they are still good pets and can be shown in household pet classes.
An unexpected colourpoint (tabby point) kitten born among a litter of black-and-white bicolours. The mother was a non-pedigree and the father was suspected of being a Birman cat that lived in the area.
Below are some non-pedigree colourpoints. Capi ("Paws") is a male Siamese mix who resembles a Snowshoe. He was photographed by Koraljka Polack on Cherso island (north Adriatic sea) in 2007. There are several "Siamese mongrels" in the area due to a Siamese cat that lived in the area some years ago. This bred with stray and feral cats. The third photo is a random-bred male colourpoint.
COLOUR DILUTION IN CATS: BLUE DILUTION, PINK-EYED DILUTION, CHEDIAK-HIGASHI SYNDROME DILUTION
Information collated 2015, Sarah Hartwell
Broadly speaking, dilution refers to making a colour lighter – like mixing a coloured paint with white paint.
COMPARISON STUDIES
The colour of cat fur is due to the size, shape and distribution of the melanin (black pigment) or phaeomelanin (red pigment) granules in the hair. The hair has a central shaft (cortex) and outer sheath ( medulla ). Melanin granules may be distributed differently in cortex and medulla. Granules in the medulla can be a different size and orientation to those in the cortex. It is the combination of location, size, shape and density of melanin granules that gives different colours.
The Structure Of A Hair Shaft
 
Simplified Diagram Of Pigment Granules In Cat Hair - based on the microscopy examination photos
Prieur & Collier (1981) compared the melanin granules in hair of black, smoke, blue, Chediak-Higashi-smoke, and pink-eyed dilution cats. A portion of the preserved skin of the pink-eyed dilute cat was used in this study.

• Black cat hair (non-agouti) contains numerous small dark brown to black melanin granules uniformly distributed throughout all portions from base to tip of the hair.
  
• Smoke cat hair (non-agouti + inhibitor (silver) gene) has a scarcity of melanin granules at the base of the hair.
  
• Blue cat hair (non-agouti + dilution) has larger basic melanin granules, but still relatively regularly shaped, and a non-uniform distribution of granules.
  
• Chediak-Higashi Smoke cat hair trait has even larger melanin granules than the blue dilution, and they are relatively irregularly shaped. All of the cats were Persian and resembled blue smoke Persians.
  
• The pink-eyed dilute cat hair had very small yellowish-brown melanin granules and very few larger granules. Under the microscope, some lighter banding could be seen on the hairs, but the tip was coloured. The base of the hair was paler than the tip (agouti banding).
  

- Prieur DJ, Collier LL. Morphologic basis of inherited coat- color dilutions of cats. J Hered. 1981 May-Jun;72(3):178-82.
BLUE DILUTION
The normal dilution seen in cats is "blue dilution" that changes black to grey (blue), chocolate to lilac, and cinnamon to fawn. Blue dilution affects the skin and coat colour by altering the pigmentation granules, but leaves the eyes pigmented.
How Blue Dilution Affects Different Colours
Prieur DJ, Collier LL. Maltese dilution of domestic cats. A generalized cutaneous albinism lacking ocular involvement. J Hered. 1984 Jan-Feb;75(1):41-4. "The Maltese dilution is an autosomal recessive trait of cats that dilutes black cats to blue, and orange cats to cream. The pigmented cutaneous and ocular tissues of Maltese dilution and control cats were examined and compared by light microscopy. Most of the melanin granules in all of the pigmented cutaneous tissues of the Maltese dilution cats were aggregated together into large clumps. However, none of the intraocular tissues containing melanin producing cells of either neural crest or optic cup embryologic origin contained clumped melanin granules. It is concluded that the Maltese dilution trait is a unique generalized albinism without ocular involvement."
PINK EYED DILUTION
Pink-eyed dilution is found in a number of mammals but there is only one good account of it in cats. It changed black to a fawn/bluish-tan colour and the eyes had a red pupil and gold flecked red/pink irides (the coloured section around the pupil). There may have been occasions where pink-eyed dilution has been confused with pink-eyed albino which is reported sporadically in cats.
Analysis of hairs from the preserved skin of the female pink-eyed dilute showed very small yellowish-brown melanin granules and very few larger granules. The base of the hair was paler than the tip and some hairs had lighter banding (agouti banding) that was only visible under magnification. The tip of the hair was coloured.
For those with a scientific background, I’ve included details from the two known studies of the single confirmed cat with this mutation.
Pink-Eyed Cats /Albino Series
To untangle the different pink-eyed mutations, here is a short description of the albino series in cats showing where pink-eyed albino fits in. It is the most recessive of the albino alleles (gene variants). A pink-eyed albino has an unpigmented white coat. A pink-eyed dilute has a coloured coat (right now we don’t know how pink-eyed dilution interacts with dominant white (white masking)).

• C/C, C/cb, C/cs, C/ca, C/c - full colour
  
• cb/cb, cb/ca - Burmese colour restriction (temperature- sensitive )
  
• cb/cs - Tonkinese Colour restriction (temperature-sensitive)
  
• cs/cs, cs/ca, cs/c - Siamese Colour restriction (temperature-sensitive)
  
• ca/ca - Blue-eyed Albino (not temperature-sensitive)
  
• c/c - Pink-eyed Albino (not temperature-sensitive)
  

Temperature-sensitive albinism in cats is caused by tyrosinase (TYR) mutations. For the molecular genetically minded, it is a cytosine deletion in the TYR gene. Cat fanciers refer to it as the "C" locus (a sort of genetic "address"). Albino cats have been reported in scientific literature (Todd 1951; Turner et al. 1981) but their eye-colour has not always been well described. Blue-eyed albinos have not been properly distinguished from pink-eyed albino cats (Bamber & Herdman 1931; Todd 1951; Leventhal 1982; Leventhal et al. 1985). This made it unclear as to whether there were more than one "one non temperature-sensitive albinism alleles" in cats i.e. both blue-eyed and pink-eyed. Lyons believed the single report of a red-eyed albino cat to be erroneous (Imes et al. 2005), but since 2005 there have been more reports and photographs of pink-eyed albino cats (this mutation has entered the Bengal breed via the Asian Leopard Cat). Because of the cat's eye structure, pink-eyed albinos often appear lilac-eyed with a red pupil, however the coat is always white. Blue-eyed albinos have pale blue irides and must not be confused with "blue-eyed white" cats that have the white masking gene and vivid blue eyes.
How is this relevant? A test-mating of a female pink-eyed dilute to a colourpointed cat demonstrated that the pink-eyed dilution is recessive, but it's not on the "C" locus and therefore isn't related to the known albino series. This makes it interesting to cat fanciers as a potential new variety, providing it doesn't have any damaging side-effects. This was documented by Neil B Todd in 1961:
A Pink-Eyed Dilution in the Cat - Neil B Todd
September 1961, 52 (5) - Journal of Heredity
It was the author 's original intention to report in detail on the genetic basis, if any, for the cat to be described below. However, her untimely death due to an intestinal obstruction ended the work almost before it was started.
During the summer of 1958 three kittens from a litter were turned over to the SPCA in Springfield, Massachusetts. Their estimated date of birth was April 1958. Two of these animals were apparently quite ordinary, although not detailed description of them was available. The mother was presumably normal; the father was unknown. The third kitten, a female, had pink eyes (although a small amount of yellow pigment was present in the iris ) and a light tan coat. She was a blotched tabby (dominant) with moderate white spotting (dominant) and had a slight kink at the tip of her tail. Her hair was short and dense with a fine texture. In all other respects she conformed to the wild type. It is suspected that she had some visual impairment from the rather clumsy way she would run into things and from her hesitancy to jump from a height which would not disturb most cats.
The animal apparently never came into heat spontaneously during the first three years of her life, but this may have been due to her caged existence rather than to any inherited or congenital defect. She was treated with hormones in January, 1961, and subsequently mated to a chocolate point Siamese. This cross was made with the purpose of discovering if any relationship existed between her phenotype and the albino series of alleles. A chocolate point Siamese is homozygous recessive for a pigment-determining allele at the albino locus (cbcb), for brown (bb) and for non-tabby (aa). [Note "+" meant "wild type" in the genetic nottion of tht time.]
The cross produced a litter of three kittens, born 10 days prematurely with the result that none survived. However, all were fully pigmented tabbies and therefore of the genotype +acb+bb+aa. From the data it can be concluded that the phenotype of the mother was due (if indeed genetically determined ) to the homozygous recessive condition at some locus other than that of the albino series of the black-brown alternatives.
[Modern notation for the offspring: Aa Bb Ccs Dd oo T-       (I’ve used “T” on its own because the report doesn’t say if the tabby is classic (Tc) or mackerel ( TM )]
Neil B. Todd
The Biological Laboratories, Harvard University
 
And that seemed to be the end of it until 2015 when Kathryn Eden posted photos of Milkdud, a Donskoy (Don Sphynx kitten) whose colour defied analysis. Born to black parents, Milkdud’s genetic test results were:

• Agouti Result: a/a - Non-agouti (solid colour)
  
• Albino Result: N/N - No copies of albino allele are present.
  
• Amber Result: E/E - No copies of the mutation for Amber.
  
• Brown Result: B/B - Full color black, does not carry brown or cinnamon.
  
• Colourpoint Result: C/C - Full colour, cat does not carry Burmese or Siamese alleles.
  
• Dilute Result: D/D - Full (Dense) color. Cat does not have the dilute allele.
  

Milkdud's skin was closer to a fawn colour and his eyes had yellow irides with red pupils. There is no genetic test for pink-eyed dilution. Photos exist of the original pink-eyed dilutes hairs under high magnification, so that is one route to investigate (the Don Sphynx has slight furring on the tail). The other is to breed furred cats with the same colour and eye colout and to analyse their fur. While the genetic basis isn't yet known, visually Milkdud appears to be a pink-eyed dilute and hence a new opportunity for the cat fancy.
Anna Kholmska, in Ukraine, has bred 4 generations of these cats (over 7 years) to establish a breeding line. All have been healthy. The unidentified colour is known in discussion (during 2015) as "Ukraine chocolate." From the visual appearance, it closely resembles the description of pink-eyed dilution. What is exciting, is the possibility of a lost phenotype reappearing in cats. The colour also needs to be bred into furred cats for study. Anna notes that the chocolate colour (b at the TYRP1 (B) locus) is not accepted in the Donskoy, although it can be very attractive. For more than seven years, Donskoy Discovery has bred show-quality cats, including some that resembled chocolate colour. Unlike Chocolate Orientals, the Donskoys had an unusual eye colour that appeared pink in some lighting conditions. This was initially attributed to a combination of the chocolate gene with the dominant hairless gene, but the work of Kathryn Eden and Anna Kholmska and genetic testing has shown that the chocolate allele is absent. The cats need to be tested to confirm the presence of the dominant B allele (black). The coat and eye colour could be caused by a novel dilution of black (if B is present) or by some other mutation (if B is absent). All of the cats share a common ancestor. Their genotype (genetic makeup) shows they should be solid black but their phenotype (visual appearance) is chocolate. This indicates a diluted-colour but the cats do not have the dilution gene (d/d) at the D locus. In any case, “d” causes blue dilution i.e. black to grey. “d” is one of the three genes involved in melanosome transfer and translocation. The unexplained “Ukraine chocolate” seems due to a gene for which there is no test (at least in the cat). Possible candidates are the other two melanosome transfer and translocation genes: Myosin Va (MYO5A) and RAB27A. Genes involved in the synthesis of eumelanin other than TYRP1, for example, TYRP2, are also possibilities. The other known dilutions in the cat are the extinct Barrington Brown cats (none of which were bred outside of a laboratory colony) and the single example of a pink-eyed dilution cat.
UNDETERMINED DILUTION
There are 4 cats of Thai descent whose dilution factor hasn’t been genetically identified.
The first of these is Mai Thong. In January 2015, I received information and photos of a native Thai cat that TIMBA hoped to use in the Suphalak breeding programme. Nolan Betterley lives in Thailand and is working on the breeding project to resurrect some of the rare cats that are described in the ancient cat poem manuscripts. Breeders working on the Suphalak, which is a solid (i.e. no Burmese pointing) chocolate self, have searched all over Thailand to find cats suitable to be used in the breeding program. They found cat, Mai Thong, aged approx 7 months, on the street . Nolan hoped that she was either cinnamon or lilac (these are recessives). DNA was tested at UC Davis and came back as aa BB CC DD (Non-Agouti, Black, No Colourpoints, No Dilution), which should produce a black cat. However Mai Thong is very obviously not black, but has an orangey colour. The gene candidates are a non-extension gene (since Russet Burmese exist) or a different form of dilution.
The remaining 3 cats that failed the test for the Suphalak breeding programme were also street cats; two were brown and one was tortie. They are now in the USA with two different breeders. The plan was to mate them and test offspring, but this has not happened. Doug Schar has the tortie sister. The owner of the other 2 cats has not allowed access to a male sibling for test-mating. UC Davis found no mutation of the TYR gene for these cats. Unfortunately, no other labs appear interested in following up to see what mutation is present..
CHEDIAK-HIGASHI SYNDROME (CHS) DILUTION (CHEDIAK-HIGASHI SMOKE)
This has only been found in Persians and manifested as blue smoke fur with yellow to light-green eyes (normal blue smoke Persians have darker fur and orange/copper eyes). Their eyes reflected red and tended to be sensitive to light. The cats developed cataracts as early as 3 months old. Melanin pigment granules in all tissues, not just the fur, were abnormally in size and shape due to combining with abnormally large lysosomes (part of a cell's digestion mechanism). CHS cats tend to develop kidney lesions, and have impaired vision due to the abnormal pigmentation in the retina. The University of Sydney, Australia's Faculty of Veterinary Science defines CHS as a "rare, lethal disorder " but it better meets the criteria for an "impairing mutation" as its effects can be mitigated with proper care. Affected cats are highly susceptible to infection, due to the abnormally large lysosome in their white blood cells. Bacterial infections occur repeatedly because the white blood cells cannot respond normally to infection; antibiotic treatment is required to combat infection. Prolonged bleeding tends to occur because of abnormal blood platelets delaying clotting. In the laboratory, CHS cats had a longer period of eye-flicker, compared to non-CHS Persians, after being spun around.
A Blue Smoke Persian and a Chediak-Higashi Smoke Persian
Despite the many side effects, cats given appropriate care can have a good quality of life. Because it affects health, the trait should not be deliberately propagated as a colour curiosity. For those with a scientific background, here are several references and abstracts about the condition.
Collier LL, King EJ, Prieur DJ. Aberrant melanosome development in the retinal pigmented epithelium of cats with Chediak-Higashi syndrome. Exp Eye Res. 1985 Sep;41(3):305-11. "Chediak-Higashi syndrome is a genetic disorder characterized by greatly enlarged cytoplasmic granules, including lysosomes and melanosomes. Eyes of humans and animals with Chediak-Higashi syndrome are hypopigmented to various degrees. Intraocular melanin granules vary in size, with some being massively enlarged. Electron microscopic examination of retinal pigmented epithelium of kittens with Chediak-Higashi syndrome disclosed a number of abnormalities of premelanosomes and melanosomes. Few premelanosomes were present. Most of the melanin granules were giant sized , but their structures varied. Some of the giant granules were composed of several premelanosomes and melanosomes in different stages of maturation. Others contained randomly oriented melanofilaments between melanosomes. There were also complex giant granules consisting of both melanosomal and lysosomal components. Inappropriate fusion of cytoplasmic granules appears to be the most likely mechanism for formation of the giant granules. Fusion of premelanosomes with lysosomes and resultant destruction of the premelanosomes probably is a major cause of the ocular hypopigmentation of Chediak-Higashi syndrome."
Hargis AM, Prieur DJ. Animal model: renal lesions in cats with Chediak-Higashi-Steinbrinck syndrome. Am J Med Genet . 1987 Jan;26(1):169-79. "In this study designed to characterize the renal lesions in cats with the autosomal recessive Chediak-Higashi-Steinbrinck syndrome (CHS), the renal tubular epithelial cells of CHS cats were examined by light microscopy. The lesions observed were similar to those in the renal epithelial cells of other species with CHS and were consistent with those reported previously in other tissues of CHS cats."
Kramer JW, Davis WC, Prieur DJ. The Chediak-Higashi syndrome of cats. Lab Invest . 1977 May;36(5):554-62. "Three cats with Chediak-Higashi syndrome were found in a single line of 27 Persian cats, and three additional affected cats were produced from two prospective breedings of the original line. The disorder was characterized genetically as an autosomal recessive condition. All cats in the line with the combination of yellow eye color and "blue smoke" hair color exhibited the disorder. Four of the five cats examined had bilateral nuclear cataracts as early in life as 3 months of age. No increased susceptibility to infectious disease was observed. A bleeding tendency was noted. Light microscopic studies of hair and skin revealed enlarged melanin granules. These manifestations were similar to those in man, mink, cattle, mice, and the killer whale with Chediak-Higashi syndrome. Cats are the sixth species in which this genetic disease has been reported." [Note: later studies of CHS cats found the immune system to be impaired]
Prieur DJ, Collier LL. Morphologic basis of inherited coat-color dilutions of cats. J Hered. 1981 May-Jun;72(3):178-82. "The melanin granules in hair of black, smoke, blue, Chediak-Higashi-smoke, and pink-eyed dilution cats were studied. The hair of black cats contains numerous small dark brown to black melanin granules uniformly distributed throughout all portions. The basis for the dilution in smoke cats is a paucity of melanin granules in the basal portions of the hair. Blue cat hair has a larger basic melanin granule, some very large but relatively regularly shaped granules, and a non-uniform distribution of granules. The granules in the blue cat hair resemble those in the hair of dilute mice. The Chediak-Higashi trait causes even larger basic melanin granules than the blue dilution and enlarged and relatively irregularly shaped granules. The melanin granules in the hair of the pink-eyed dilution cat are very small, and are yellowish brown compared to the dark brown to black of those of black, moke, and blue cats."
Prieur DJ, Collier LL. Inheritance of the Chediak-Higashi syndrome in cats. J Hered. 1981 May-Jun;72(3):175-7. "The phenotypes with respect to the Chediak-Higashi syndrome (CHS) of 245 cats from 67 matings were analyzed. It was determined that the gene for feline CHS, like that in other species with CHS, is inherited in an autosomal recessive manner with complete penetrance. We propose the symbol ch for the gene for CHS in cats. "
EYE COLOURS
Copyright 2009, Sarah Hartwell
Wild cats in temperate regions generally have hazel eyes, but domestic cats' eye colours vary from blue, through green to yellow, orange and brown. Within each of those colours there is wide variation in hue and intensity. The colours are not discrete, but form a continuum, for example between blue and green there are "sea-green" and aqua while between green and yellow there are lemon and lime shades. The coloured part of the eye is called the iris, a word that means "rainbow".
Some eye colours are linked to coat colour, for example colourpoint (Siamese pattern) cats have blue eyes. In most breeds, various eye colours were possible so breeders set chose the eye colour that harmonised with the coat colour and wrote that into the breed standard. While breeders might prefer certain pedigree black cats to have vivid orange eyes, in the general cat population black cats can also have green or yellow eyes.
HOW EYES GET THEIR COLOUR
There are two major factors that influence feline eye colour: iris pigmentation and blue refraction.
The iris has 2 layers containing pigment-producing cells (melanocytes). The outer layer is the stroma, consisiting of consists of loosely arranged cells. Underneath the stroma is the epithelium which contains tightly packed cells. Both the stroma and epithelium produce pigment, but in different amounts. The pigmentation of the iris is caused by melanin and the colour ranges from lemon yellow to hazel to deep orange or brown. In addition to this, the "transparent" structures of cats' eyes absorb and refract light much like a sheet of glass. Viewed head on, glass appears colourless but viewed edge-on it is greenish or bluish. The size, spacing and density of the fibres in the stroma determines how it reflects and refracts the light, determining the shade of blue. Just like glass, the transparent structures in cats' eyes are transparent blue, ranging from practically colourless to deeper blues and violet-hued blues. The combination of the blue refraction and iris pigment produces the overall eye colour. Looking into a cat’s eye is like looking through a blue-tinted window; the blue tint affects how we see the other pigments present in the iris.
The iris pigmentation and the transparent structure pigmentation are both controlled by multiple genes in different places on the chromosomes (polygenes). Littermates can inherit different mixes of these genes from the same parents and have different eye colours. Different mixes of polygenes causes the continuum of shades.
The type of colour and the intensity depends on the number of melanocytes in the eye and how active they are. If there are no melanocytes, the eye appears blue (or in rare cases, pink which is due to the colour of the blood vessels). A low number of melanocytes gives green. A high number of melanocytes gives orange. A second factor is how much pigment those melonocytes produce ( activity ). Those that produce less pigment give a lighter shade, while those that are more active and produce a lot of pigment give the deeper shades. There is no single inactive/active switch , so there are varying shades inbetween. Eyes with a low numbers of melanocytes (green) can range from pale green (less active melanocytes) to strong green (industrious melanocytes). Similarly , the yellows range from light amber to deep copper. The activity levels of melanocytes is genetically determined so it is possible to selectively breed for either deeper or paler eye colours. The way that light passes through the various blue-tinted structures gives us the final visible colour.
Kittens are born with blue eyes. The adult eye colour develops around 6-7 weeks old, but may not reach its adult hue until 3-4 months old. During a cats' lifetime, illness and injury can cause variations in eye colour, but there is no truth in the old wives' tale that feeding a cat on fish will cause its eyes to change colour! That tale is due to old wives weaning kittens (probably onto boiled fish in the days before commercial cat foods) around the time the eye colour naturally changed.
There is also some pigmentation on the inside of the eye and a highly reflective area behind the retina called the Tapetum Lucidum. This is an iridescent layer of tissue that reflects light back through the eye. and helps cats see in low light - it also causes eye shine in flash photos. Most blue-eyed white cats lack the Tapetum Lucidum; their eyes reflect a dull red colour (caused by the blood vessels) rather than an iridescent eye-shine.
THE BLUES
Kittens are born with blue eyes and this is not a guide to their adult eye colour! In a few breeds, the blue colour is permanent , but in most it will be replaced by another colour starting around 6 weeks old. The blue colour depends on the intensity of blue refraction.
Blue eyes are a feature of colourpointed cats where the colour is linked to the temperature-dependent albinism that produces the colourpoint pattern. The blue colour varies in hue and intensity and breeders must selectively breed cats with the best eye colour to try to pass on that colour to the kittens.
Blue eyes also occur in white cats and are associated with the epistatic white (often called dominant white) gene which is quite different from albinism because it masks an underlying coat colour. The white spotting gene also affects eye pigment if there are white patches around the eyes. White cats can have one or both eyes blue. The non-blue eye will be whatever colour the breed standard requires for example in white Persians, the other eye is vivid orange. In the randombred odd-eye white cats, the non-white eye might be green or yellow or even a different shade of blue. For more on blue-eyed white cats White Cats, Eye Colours and Deafness.
The Ojos Azules is characterised by cornflower blue eyes, but the mutation is linked to lethal deformities if a kitten inherits 2 copies of the mutated gene. In this breed, the eye colour is independent of the coat colour. Other blue eye mutations that are independent of coat colour crop up from time to time. To date, most have been a pale blue rather than an intense blue and none of these other mutations have been deliberately bred.
A grey bicolour (tuxedo pattern) cat with intense sapphire blue eyes was found on a country road in Windellama, New South Wales, Australia by A M Schnieder. Since the Ojos Azules has not been imported into Australia, this represents a spontaneous mutation among the feral population. Whether it is the same as the American Ojos Azules mutation is not known.
GREEN AND AQUA
Green ranges from a yellowish-green through gooseberry-green to deeper greens and blue-toned greens. Some of the older standards spoke of grass-green eyes, indicating the green be as rich as possible. Green tints and flecks also appear in some hazel brown eyes.
The Chinchilla Longhair (Chinchilla Persian) and its shorthaired equivalents are notable for their black-rimmed sea-green eyes. Aqua (blue-green) is found in the Tonkinese which is intermediate in type and colour/pattern between Siamese and Burmese cats. The Siamese has blue eyes while the Burmese has orange eyes.
YELLOW, ORANGE/AMBER AND BROWN
Yellow varies from a pale lemon to more vivid hues. There is overlap between the yellows and browns (pale hazel or tan) and also between yellows and greens.
While the boundary between yellow and green may be fuzzy, there is no mistaking an orange-eyed cat. The colour is often described as coppery. Many of the breeds developed by early cat fanciers in the UK called for orange eyes as the pale greens and yellows were considered undistinguished, while bright orange complemented the coat colour.
Brown covers a wide range from hazel (the normal colour of temperate climate white cats) through to darker browns. Some browns appear tinted or flecked with green, orange or yellow.
ODD EYES (HETEROCHROMIA)
The scientific term for odd eyes is heterochromia iridium (different coloured irides). It can be genetic (inherited), congenital (development defect) or acquired (illness, injury, medication). In the cat fancy, odd-eyes means having one blue and one other-colour eye. Odd eyes are most common in epistatic white cats where one eye is blue and the other is orange, yellow, brown or green. Pedigree odd-eyed white cats have one blue and one orange/amber eye, but in randombreds the non-blue eye may be yellow, green or brown. Another form of odd eyes is found in white cats where one eye has a tapetum lucidum (green eye shine in the dark) and one eye lacks the tapetum (red eye shine). These different eye shines are seen in reflected headlights or flash photography. Bicolour cats with a high degree of white on the face may also have odd eyes.
Heterochromía from birth is common in cats (blue/amber), horses (blue/brown) and some breeds of dog (blue/brown). It is uncommon in humans. In cats, there is no specific gene for "odd eyes," but it is associated with the white spotting gene and the white masking gene (epistatic white/dominant white). These genes prevent pigment production in the embryonic stage, which includes the pigment of the iris. If some melanocytes (pigment cells) remain active in the eye area, this is sufficient to make the eye green, amber, brown or yellow etc. Depending on the activity level of the melanocytes, the eye colour ranges from blue with other tints through to fully coloured, and from pale through to intense colour. During the cat's lifetime, the effect may be incremental so that the intensity of eye colour deepens, gradually replacing the blue colour.
Heterochromia is mostly found in cats with white coats or white patches. Occasionally it is seen in coloured cats. When present from birth, it can be a result of congenital abnormality (birth defect), mosaicism (merged embryos), or a somatic mutation in some cells that form the skin and eye (birthmark). These are not hereditary and this form of heterochromia is not passed from parent to offspring.
Odd-eyed tortoiseshell cat.
Odd-eyed white cat photographed by Rodrigo Arancibia in Iquique, Chile. She shows no sign of deafness. her kittens are solid white, indicating the dominant white gene.
Less commonly , other colour cats may have odd eyes for example a tortie with one yellow and one blue eye. This can be due to the eyes developing differently in the embryo. Perhaps the cat has a white spotting gene that hasn't showed up on the fur, but has affected the eye. Below is another odd-eye tortie: a 10 year old tortie/tabby called Debby photographed by Marjan Boonen in The Netherlands. Since tortie cats' patched colouration is due to mosaicism, it's possible this has extended to the eyes resulting in two different colours
Occasionally the different colour eyes are due to an eye injury or illness that has damaged the iris or one eye. In this case the trait cannot be passed on.
DICHROIC (DICHROMATIC) EYES
Partial heterochromia (dichroic eyes) occurs when different amounts of melanocytes, or melanocytes with different activity levels, are present in distinct areas of one iris. This gives the “pie slice” or “coloured halo” effects. Dichroic/dichromatic eyes have two colours in one iris. For example there may be a yellow ring around a green iris. This is considered a fault in show cats, but can be very attractive.
Less commonly, there may be a distinct differently coloured section (like a slice of a pie) in the iris, for example a brown area in a blue or green eye. This has been reported in some white cats. Instead of affecting the pigmentation of the whole iris, the white gene affects only part of the iris. One or both eyes can be affected; if both are affected there may be a mirror image effect. These are sometimes called "weird-eyed" cats because they are not odd-eyed in the normal sense .
The photo is "Gilbert" who has one blue eye and one dichoric eye.
LILAC / BLUE-EYED ALBINO
Albinism is found in the Siamese to closely related Oriental breeds due to inbreeding. Emma Stothers (2013) provided photos of her albino Oriental Longhair, Pangur. Pangur's eyes look like faded lilac-blue. On close inspection, the pupil is a dull red, while what looks like lilac is just the pale blue refraction of the lens overlaid on a pinkish pupil. This makes Pangur very photo-sensitive. Pangur's parents are half-siblings with the same father.
PINK
Pink is not a normal eye colour in cats. It is found in pink-eyed albinos and is due to complete lack of pigment in the eye combined with very little blue refraction. True albinos entirely lack melanin and the iris is pale pinkish-white. Because of blue refraction, the eyes usually have a bluish hue, result in a blue-eyed albino (as with Pangur above). Very rarely the transparent structures are colourless and a pink-eyed albino results.
It was once believed impossible for a cat to have pink eyes due to the structure of the eye and true albinos had pale blue eyes, but there have now been several pink-eyed albinos reported in recent years. Albinism has been recorded in the Bengal breed and in its wild parent, the Asian Leopard Cat (shown here).
Pink-eyed dilution has been reported once in the cat (Todd NB. 1961. "A pink-eyed dilution in the cat" Journal of Heredity 52, pg 202). The type of dilution found in cats is blue dilution which turns black into grey. A second type of dilution seen in some mammals is "pink-eyed dilution" which gives a bluish-tan/fawn coat and also depigments the eye, giving a pink or ruby appearance. This has been reliably reported only once in cats when a pink-eyed female with a light tan coat was produced (none of her kittens survived so pink-eyed dilution in cats seems to have been lost).
Pale irises and underpigmented retinas also occur as part of Chediak-Higashi syndrome along with a pale coloured coat (seen in Blue Smoke Persians) .
EYE SHAPES
In addition to the different colours, there are several distinct eye shapes. These are shown in slightly exaggerated form in the chart below.
TORTOISESHELL AND TRI-COLOUR CATS
Copyright 2002 - 2013, Sarah Hartwell
Because of the way tortoiseshell is inherited, almost all tortoiseshell and calico cats are female. The very few male tortoiseshell cats are caused by genetic aberration or development abnormalities in the foetus. These are discussed in detail in "Mosaicism, Tortie Tomcats, Genetically Impossible Kittens and Gender Anomalies". This article only considers normal tortoiseshell cats.
Jean Bungartz referred to tortoiseshell-and-white cats as "Spanish cats" in his 1896 book "Die Hauskatze, ihre Rassen und Varietäten" (Housecats, Their Races and Varieties) in " Illustriertes Katzenbuch" (An Illustrated Book of Cats) though he said it was hard to explain why they got the name as they were not restricted to the Pyrenean peninsula.
HOW TORTOISESHELL PATTERNS OCCUR
The ginger colour of cats (known as "yellow", "orange" or "red" to cat breeders) is caused by the "O" gene. The O gene changes black pigment into a reddish pigment. The O gene is carried on the X chromosome. A normal male cat has XY genetic makeup; he only needs to inherit one O gene for him to be a ginger cat. A normal female is XX genetic makeup. She must inherit two O genes to be a ginger cat. If she inherits only one O gene, she will be tortoiseshell. The O gene is called a sex-linked gene because it is carried on a sex chromosome. Tortoiseshell cats are therefore heterozygous (not true-breeding) for red colour.
The formation of red and black patches in a female with only one O gene is through a process known as X-chromosome inactivation. Some cells randomly activate the O gene while others activate the gene in the equivalent place on the other X chromosome. This only shows up visibly in skin cells as these produce pigment. This occurs early on in the embryo and as skin cells multiply, they form patches. The skin is a mosaic of cells where some have the O gene active (making ginger pigment) and some do not (making black pigment). This can only happen in cats with two X chromosomes. Male cats only inherit one X chromosome so this is active in all skin cells as there is nothing equivalent on the Y chromosome which could "switch off" the O gene.
There are two main theories regarding brindled torties and patched torties. One (the "early/late deactivation theory") suggests that the time at which X chromosome deactivation occurs during foetal growth determines whether the cat has well defined patches or is brindled with intermixed black and orange hairs. Skin cells multiply during growth and spread out across the skin; as the embryo grows the skin cells multiply. If deactivation occurs early on each pigment cell has room to multiply into, a "red" cell will multiply into more red cells while a "black" cell will give rise to patches of black. If it occurs later, the patches are smaller as the cells have less room to multiply into; some "patches" will be no larger than a single hair! All "red" cats are red tabbies - where there are large red patches, the tabby pattern will usually be discernible.
All Red Cats Are Red Tabbies? Strange as it may seem, all red cats are actually red tabby because the non-agouti gene (the gene that turns a tabby into a self/solid colour) does not affect the way red pigment is deposited. A variety of other genes, called polygenes or modifiers, control the intensity of colour and contrast between markings and background colour. Selective breeding has reduced the tabby markings to produce a cat that looks solid red by breeding from those cats with the least red markings (cats with "low contrast" between markings and background colour). Because the non-agouti gene does not work on the red pigment, red tabby ghost markings can never be completely eliminated and may be seen on the tail, legs and forehead and as a darker region along the spine. Even though red cats are registered as "red self" they are still red tabbies, albeit red tabbies with very reduced markings. Because they lack the polygenes for high contrast between markings and background colour, their offspring also appear to be red self. More information can be found in Robinson's "Genetics for Cat Breeders".
The other theory regarding brindling and patching (the "migration theory") is that brindled torties occur when there are more pigment producing cells produced from the neural crest (which becomes the back and spine area). The cells are assumed to have undergone X chromosome deactivation before migration. The migrating cells carry either O (red) or o (black) and they migrate at the same rate into their final positions. Where there are many pigment producing cells, there is more competition ( imagine cells jockeying for position on the skin) and they become intermingled. Where there are fewer pigment producing cells, each cell has room to grow into patches of colour (imagine plants forming clumps in a flower border).
As an interesting side note, human women can also have X-inactivation. Although there are no tortoiseshell women, there is an uncommon condition called anhidrotic ectodermal dysplasia caused by a faulty gene on the X chromosome. The faulty gene reduces the number of sweat glands per square centimetre on the skin. In women who inherit one normal gene and one faulty gene; their skin may be divided into hundreds of small patches which only become visible when they sweat. The normal patches of skin sweat normally, but the patches derived from cells with a faulty gene don't.
To be completely accurate, O = orange and o = "non-orange" meaning that o allows whatever colour is carried on other chromosomes to be displayed (black, blue, chocolate etc). The O gene converts normal black/chocolate/cinnamon etc pigment into red pigment. For the sake of simplicity, this article refers to o as "black" since that is the non-orange colour most commonly seen on tortie and calico cats.
TORTOISESHELL-AND-WHITE (CALICO) CATS
The white patches in tortie-and-white (tricolour, calico) cats is caused by the piebald spotting gene discussed in Beautiful Bicolours. This is a semi-dominant gene with very variable expression ranging from nearly all white to nearly all coloured with only a few white hairs. The gene affects the embryo cells which will become pigment-producing skin cells. These cells are initially formed along the "neural crest" - the embryo's backbone area - and migrate to all over the body during formation of the skin. Where these pigment producing cells fail to get in position before the skin is fully formed, there will be areas of skin which lack pigment producing cells i.e. white areas. White areas are usually the areas furthest from the cat's backbone - paws, belly, chest and chin - these areas take longest to reach.
One effect of white spotting in tortoiseshell cats is to change the pattern from brindled to patched. Tortie cats with little or no white tend to have brindled coats. However, the more white there is, the more the black and white will also be separated out into patches instead of being intermingled. The phenomenon of tortie-and-white cats having better defined patches of colour is familiar to most cat lovers.
In the developing embryo, the pigment forming cells migrate from the neural crest. If the "migration theory" is right, cells which activate O (red) and those which activate o (black) appear to migrate at the same speed, leading to a brindled pattern. If the embryos also inherit the gene for white spotting, the fur develops as patches of colour. The bigger the white areas, the bigger the and better defined the separate patches of black and red. The presence of the white spotting means fewer pigment producing cells and less competition between them as they migrate into position. One or two cells reach an area and these multiply in situ to form a patch of colour (a clonal patch).
Red Tabby Markings in Tortie and Tortie-Tabby Cats. Where a tortie (or calico) cat has the non-agouti gene it will appear to have solid patches of colour. Due to the quirk of the red pigment being unaffected by the non-agouti gene, there may be tabby markings visible in the red areas. How much red tabby pattern is visible is due to other genes called polygenes that also affect the depth and contrast of the red colour (the illusion of solid red is due to low contrast between background and markings). Where a tortie (or calico) has the agouti gene instead, the tabby markings will be visible in all of the coloured patches. In a tortie cat, the non-red colour will be a solid colour. In a torbie cat, the non-red colour will also have tabby markings. More information can be found in Robinson's "Genetics for Cat Breeders".
TORTIE ABYSSINIANS AND TORTIE TICKED TABBIES
There is another gene which gives rise to the red colour. This is the non-sex-linked red ("sorrel") found in Abyssinian cats and other ticked breeds and it does not produce tortoiseshells. Genetically, the Abyssinian pattern is not a solid colour, but is a form of tabby, called ticked tabby, and is described in Striped and Spotted Cats. The ticked pattern changes the visual appearance of the cat's colour. Although some registries call this colour "red" ("non-sex-linked red" in Australia), many registries prefer to call it "sorrel" to avoid confusion with the sex-linked red gene described earlier. Genetically, Abyssinian "non-sex-linked red" is the equivalent of cinnamon in non-ticked cats. It appears as a reddish-brown due to the ticked pattern of the hair. Chocolate Abyssinians appear a deeper red which can be differentiated from sorrel by the chocolate's darker tail tip.
Strictly speaking, because Abyssinian is a tabby pattern, the tortie versions should be termed ticked torbies (tabby-torties) rather than ticked torties. This photo shows Harley (Harlequin) and Spot (provided by owner Lizzi) who were resuced as feral kittens. Harlequin is a ticked torbies, while littermates Spot is ticked tabby with white, and Blaze and Mr Grey are mackerel tabbies with white. The two mackerel tabbies are quite lightly marked on a ticked background and show some rufousing (red tints).
The true sex-linked red Abyssinian is bright orange with a red tail tip. It is important for breeders to know whether they have sex-linked red or non-sex-linked red as this will affect the breeding program. Where there is sex-linked red, there can also be tortoishells. Tortie Abyssinians occur, but since the breed does not permit white, these are always brindled. The combination of brindling and ticking can make it almost impossible to determine whether a female is tortie or not just from a visual inspection. Sometimes a female Abyssinian is only known to be a genetic tortie when she produces a mix of red and black kittens!
Ticked tabby also occurs in non-pedigree cats. In randombred cats with the ticked tabby pattern, white markings may be present as well, resulting in "ticked tortie tabby and white" (ticked calico). The photo shows part pedigree tortoiseshell ticked tabby kittens with white markings.
DIFFERENT TORTOISESHELL COLOURS
The basic tortoiseshell combination is black and red. Because red does not occur as a true solid colour, the "red" patches are actually red tabby and this will be more apparent on some cats than on others. The addition of a dilution gene gives rise to the other tortoiseshells combinations. In Burmese cats, where there is a different dilution gene at work, diluting black into brown.
 
The action of caramel is noted as theoretical, in reality the colours may be indistinguishable from other colours unless the presence of caramel is known from the cat's pedigree; the entries are based on data from solid colour cats. The colour intensifier is also theoretical, based on the possible "indigo" gene which turns blue into a deeper richer colour. This could equally be due to polygenes (multiple genes having a cumulative effect). I have included it after seeing one cat which was dark-blue/hot-cream tortoiseshell i.e. where both colours were intensified.
NON-DILUTE COLOUR
DILUTE VERSION
CARAMELISED VERSION OF THE DILUTE VERSION (DOUBLE DILUTION)
INTENSIFIED VERSION OF THE DILUTE VERSION
("INDIGO" GENE)
Tortoiseshell
Black/Orange Tortoiseshell
(Brown Tortoiseshell in
Burmese cats)
Blue-Cream Tortoiseshell
Caramel/Apricot
Indigo/Hot Cream (dark-blue and hot cream) (theoretical)
Cinnamon Tortoiseshell
Milk chocolate and cream
Fawn Tortoiseshell: 
Fawn and cream
Caramel/Apricot
Chocolate (Chestnut) Tortoiseshell
Warm milk chocolate, red, and cream
Lilac (Lavender) Tortoiseshell: 
Frosty lilac-grey and cream
Taupe/Apricot (theoretical)
Above are black (smoke), solid chocolate and chocolate tortie kittens. The black is shown for colour comparison. Chocolate and choc -tortie are more common in pedigree cats than in randombreds, but these kittens were born to non-pedigree parents. Photo courtesy of Leva Cygnet.
In Siamese cats, only the points (legs, tail and face) show as tortoiseshell and because the gene causing the Siamese pattern turns black into dark brown, the tortoiseshell pattern will contain dark brown rather than jet black.
TORTOISESHELL TABBY AND TORTOISESHELL TIPPED
Tortoiseshell combines with the tabby pattern to create tortoiseshell-tabbies, also known as patched tabbies or (in the US) torbies. The tabby pattern may be classic (blotched), mackerel or spotted.
 
In a usual tortoiseshell, the black patches become brown tabby (i.e.. black markings on brown background), the red patches become more obviously red tabby (red patches on paler red background). Patched tabbies occur in other tortie colours e.g. blue-cream patched tabby.
In combination with silver, this would produce tortoiseshell markings on a silvery background e.g. black-silver (black markings on pale grey) and red-silver patches; though such combinations are not common.
The tortoiseshell markings can also be combined with the genes for tipped, smoked or shaded in which case the tortoiseshell colours are restricted to the hair-tips rather than going all the way to the root. This gives the effect of a shimmering tortie pattern on a pale undercoat.
BREEDING TORTOISESHELLS
In order to produce tortoiseshell kittens, at least one of the parents must carry the O gene. Red male and red female only produce red kittens. The combinations which produce tortoiseshell kittens are red male and non-red female, red female and non-red male. The following can also produce tortoiseshell kittens: tortie female and non-red male, tortie female and red male. It also depends on some kittens being female.
Parents
Kittens
Female
Male
Males
Females
Red
Red
Red
Red
Black
Red
Black
Tortie
Tortie
Red
Red
Red
 
 
Black
Tortie
Tortie
Black
Red
Tortie
 
 
Black
Black
Red
Black
Red
Tortie
Black
Black
Black
Black
 
This is what is happening when a tortoiseshell female is mated to either a red or a black male. Because a tortie female is heterozygous, she can pass on either the gene for red or the gene for non-red (black). The colour of the kittens depends on whether they are male or female and what gene, if any, they inherit from the male.
GENES FROM RED MALE
GENES FROM TORTOISESHELL FEMALE
 
GENES FROM BLACK MALE
GENES FROM TORTOISESHELL FEMALE
 
O (red)
o (non-red [black])
 
 
O (red)
o (non-red [black])
O (red)
X chromosome
OO
Red female
Oo
Tortie female
 
o (non-red [black])
X chromosome
Oo
Tortie female
oo
Black female
No gene (Y chromosome)
O-
Red male
o-
Black male
 
No gene (Y chromosome)
O-
Red male
o-
Black male
 
Because other conditions can give the impression of being tortoiseshell, some matings which theoretically have a high probability of producing tortie kittens will only ever produce red or black kittens. At the other extreme, a genetically tortoiseshell cat may appear to be solid ginger or solid black because of the way the X chromosomes activated - these cats will produce unexpected tortie kittens in appropriate matings.
CLONING TORTOISESHELLS
This may seem an odd topic, but if you clone a tortoiseshell cat you will end up with a cat of one or other of the constituent colours and not a tortoiseshell clone. If you clone a red/black tortoiseshell, the clone will be either red (ginger) or black.
DNA tests on the tabby-and-white cloned kitten ("Cc") proved that she was a clone of her tortie-and-white genetic mother (the cat whose cell was used to create Cc). The pattern of pigmentation in multicoloured animals is the result of genetic factors combined with developmental factors within the womb. This means bad news for owners who want an exact replica of a tortoiseshell or calico cat. Why isn't Cc also tortie-and-white? The answer is due to ‘X-linked Inactivation'.
Tortoiseshell cats have two X chromosomes, one carrying the gene for orange coat colour and the other carrying the gene for black coat colour. As the embryo develops, a process called ‘X-linked inactivation’ occurs in its tissues. One or the other X-chromosome in every cell in a tortie cat embryo is randomly inactivated. This only shows up in pigment producing cells, producing the familiar tortie effect.
Regardless of which cell was used to produce Cc, because that cell is already an adult cells, one or other of the cell's X chromosomes would have been inactivated while the  donor cat was an embryo. Cc had an equal chance of being orange-tabby-and-white or black-tabby-and-white, but would never be tortie-tabby-and-white. Unless a way can be found to undo X-linked inactivation at the embryo stage, owners wishing to clone a tortoiseshell cat will have to settle for a cat of a different colour entirely. If the X-linked inactivation can be reset, the inactivation is a random process so the clone will have the right colours, but not in the same places as the donor cat - it may have well-defined patches of colour while the donor cat was thoroughly brindled.
TRI-COLOUR BENGALS
Confusingly for cat lovers, tri-colour Bengals are not tortoiseshell cats.  In the Bengal breed, tri-colour refers to a cat with black rosettes on a golden brown background. The centre of the rosette is generally a darker version of the background colour, hence the cat has three colours: pale background, black markings, dark rosette centre.
Tortoiseshell Bengal variants have occurred during breed development, but it is not an accepted colour in a breed which aims to re-create a wild look.
ANOMALOUS TRI-COLOURS
A number of anomaous tricolours are discussed in "Mosaicism, Tortie Tomcats, Genetically Impossible Kittens and Gender Anomalies". These include a white, black and grey tricolour and a white, red and grey tricolour. A further unusual tricolour is best described as a "white brindle".
The "white brindle" was one of the most striking "tortoiseshell" cats I have seen. Imagine a brindled tortoiseshell in which all the black areas have been converted to white. The result was a cat with intermixed fur of white, cream and ginger; the fur being brindled like a tortie rather than being patched like a red-and-white bicolour. This was seen while on holiday (either Turkey or Tunisia) and unfortunately I didn't have a camera with me. I have never seen anything like it since. A likely explanation is a genetic mutation in the fertilised egg had caused a pigment production fault in the areas which should have been black. It would have been interesting to see if the trait could have been perpetuated.
One reader suggestion for the "white brindle is a red-tabby-and-white smoke shorthair. This would give a similar effect, but the colour in a smoke cat does not go down to the roots and the brindled effect would not be so thorough. I am currently inclined to think that the cat was a conventional brindled tortie which was not producing black pigment.
WHITE CATS, EYE COLOURS AND DEAFNESS
2001 - 2016, Sarah Hartwell
This page has two linked topics - the different causes of blue eyes and white fur in cats is one, the other is the relationship between eye colour, coat colour and deafness. This means some information in one section might also be repeated or summarised in onther section (that way you can read a section and not have to read the whole page).
ALBINO OR JUST WHITE?
A true albino cat
The eyes are unpigmented and appear pink. The skin is pale pink.
A true albino kitten, age 7 weeks, owned by Michelle R. McGaha. The eyes are unpigmented and appear pink. The skin is pale pink. The mother is a tortie Manx, the father is unknown. He is very light sensitive and will need sun- protection on his ears to guard against skin cancer (all white-eared cats have a risk of skin cancer if they sunbathe too much).
Contrary to popular belief, white cats with blue, orange or green eyes are not albino. Albino cats, such as the one in this photo, have pink (unpigmented) or bluish-pink eyes and, like most albino animals, their eyes are sensitive to light. The white colour in cats is due to a gene that masks any other colour genes (this is why white cats can have non-white or bi-colour kittens). Albinism is a different mutation that causes the absence of colour, not the covering up of colour. When owners talk about having green-eyed or orange-eyed albino cats, they mean green-eyed or orange-eyed "dominant white" cats. In this context, dominant means a gene that masks other genes, it does not mean the cat has a dominant personality.
Recessive white is an alternate name for the blue-eyed/pink-eyed albino gene which is part of the colourpoint series of genes. The white spotting gene can sometimes produced solid white cats and is also sometimes called recessive white.
It is also a fallacy that "all blue-eyed white cats are born deaf". All kittens are born blue-eyed and with their ears folded down. Whether they will stay blue-eyed and whether they will be deaf can only be ascertained after a few weeks. Not all blue-eyed white cats will be deaf (as this article explains).
DOMINANT WHITE
Dominant white (more properly "epistatic white" since it occurs on a different gene to the black-based or red-based colours) denoted by the gene symbol "W", is the colour associated with deafness in cats. Dominant white masks all other colours and cats may have blue, orange or odd eyes. Those with blue eyes have a high chance of deafness. Those with one blue eye have a high chance of deafness on the blue-eyed side. Those with orange eyes are far less likely to be deaf. Some dominant white kittens are born with smudges of coloured fur on top of the head where the colour is incompletely masked, this smudge of colour usually disappears by adulthood, but kittens with colour smudges are more likely to have normal hearing. These cats are not albinos; genetically they can be any colour, but the white is dominant to those other colours (albinism is an absence of other colours).
Lacy has the dominant white gene associated with blue eyes and deafness (Lacy is deaf). Lacy owns Greg Schultz who refers to her as "The Queen".
Mikey has the dominant white gene, but has green eyes rather than blue. He is also deaf. Mikey is owned by Jennifer Moore (ibjennyjenny).
In some animals, the blue-eyed white trait is sex-linked (carried on the X chromosome) and is found in males more often than in females. In cats, the gene for white is carried on an autosome (a chromosome other than the X or Y sex chromosomes) and the trait occurs equally in male and female cats. Blue-eyed white is not sex-linked in cats.
WHITE SPOTTING
The gene for white spotting, denoted by the gene symbol "S", can also create the impression of a self white cat. This gene is semi-dominant and is variable in the way it is expressed - a cat may have no visible white spots or may be wholly white and all stages in between those two extremes. Unlike dominant white, white spotting is not linked to deafness.
 
Some white cats are due to the very variable expression of the gene for white spotting. The diagram above shows a typical progression from solid colour through to solid white caused by this gene. The number by each diagram is the "Grade" of spotting from Grade 0 (no white spotting) through to Grade 10 where white spotting has obscured all of the base colour. Grade 10 white spotted cats resemble solid white cats, but are rarely deaf. Again, small spots of colour may be discernible on cats that appear solid white due to the white spotting gene.
ALBINO CATS (RECESSIVE WHITE)
Albino is generally thought of as pure white, but the situation in cats is more complex. There are five known alleles for albinism: blue-eyed albino ("ca"), pink-eyed albino ("c"), Burmese pattern ("cb"), Siamese pattern ("cs")and full colour (non-albino, "C"). Full colour is dominant to all of the other four alleles. Burmese pattern is incompletely dominant to Siamese pattern; cats that inherit one of each of those genes will be intermediate in pattern and is known as Tonkinese. A quirk of the Siamese form of albinism is that it is temperature dependent with warm areas of the body being paler than cooler areas. For this reason, it is often described as "colour restriction" rather than albinism. Pink-eyed albino appears to be recessive to all of the other albino mutations. Albinism is not linked to deafness in cats ("dominant white" (W) is the gene linked to deafness).
Full albinism - also known as recessive white - is rare. When present it affects the structure of the tapetum of the eye so tht it reflects red/pink instead of reflecting green.
In 1927, judge Mrs Basnett reported on the Paris Cat Show held on 14th and 15th of January by the Cat Club de France and wrote "One lady brought for my inspection a Siamese Albino of about 4 months; I could not see a trace of any shading anywhere on the white coat, tail, legs, ears or mask; the eyes were a very beautiful blue, and their fiery centre seemed to be accentuated by the absence of all shading. In appearance it was a very typical Siamese with the long sleek body and whip tail, and a very beautiful wedge-shaped head and face." Albino Siamese have also been purposely bred; they are completely white Siamese-type cats with bluish-pink eyes (true "pink eyes" are uncommon due to the physical structure of a cats' eyes). The "European Albino" bred in Belgium is a European shorthair type white cat with ruby-red eyes which have pale translucent blue irises. The albino cats reported in Europe and the USA seem to be intermediate between pink-eyed albino and blue-eyed albino. A true pink-eyed albino was reported in 1931 and again in 1980s in the USA.
Albino Siamese had pinkish-blue eyes rather than the clear blue related to dominant white in the Foreign White breed. The mutation for "true albinism" was reported when a Chocolate Point female bred to her Chocolate Point son (this line became quite inbred in order to produce the albino cats). The recessive white gene has continued to lurk in the Chocolate and Cinnamon Oriental gene pool, where it somes manifests by affecting the eye structure and producing blue-eyed Orientals. When pink/red eye shine shows up in breeds of cat, or in randombreds, it is most likely due to chocolate or cinnamon in their ancestry. It has been observed in some oriental cats and in chocolate/cinnamon Devon Rex, Affected cats are not necessarily wholly or partly white and the only hint of this gene seems to be the red eye shine instead of green eye shine! Why does it only visibly affect eye structure? All multi -celled creatures are mosaics where genes can be switched on in some cells but not in others. For some reason, during the development of the embryo, the lurking albino gene got switched on in those cells that formed the eyes.
OTHER ALBINISM
An albino kitten was born at Chelmsford Cats Protection League shelter in the 1990s and required handrearing. It was described as white furred and having very pale pink ears, nose leather and paw pads though I have no information on its long term survival. Albino kittens have turned up more recently in the Bengal breed, unsurprising since albinism is found in the Asian Leopard Cat (the wild parent of the Bengal).
In the Ojos Azules cat, normally typified by bright blue eyes in combination with colours other than white or colourpoint, the homozygous form of the gene has caused dead albino kittens.
WHITE CATS AND DEAFNESS
A few years back I was asked three related questions on a newsgroup. This article is adapted from my answer.
- Are white cats, particularly blue eyed white cats, always deaf or is this an old wives' tale? 
- Is deafness linked only to odd-eyed white cats? 
- Some blue-eyed whites aren't deaf - why?
There is an established link between the white coat color, blue eyes and deafness. The tapetum lucidum is generated from the same stem cells as melanocytes (pigment cells). The blue eyes in a piebald or epistatic white cat indicates a lack of tapetum. Deafness is caused by an absence of a cell layer in the inner ear that originates from the same stem cells as well. In odd-eyed white cats, the ear on the blue-eyed side may be deaf, but the one on the orange-eyed side usually has normal hearing. Not all blue-eyed whites will be deaf since there are several different genes causing the same physical attributes (whiteness, blue-eyedness) so it all depends on the cat's genotype (its genetic make-up) not its phenotype (its physical appearance). Some people claim that 99% of blue-eyed white cats are deaf. This is inaccurate because blue-eyedness and whiteness can both be caused by different genes. It all depends on what genes the cat has inherited. These are the actual figures from scientific studies around the world. The percentages are given in ranges because results are different in different areas, partly because of the different genes found in the cat population. Where a cat is classed as deaf, the deafness may affect one or both ears.

• 95% of the general cat population is non-white cats (i.e. not pure white) and congenital deafness is extremely rare in non-white cats.
  
• 5% of the general cat population is white cats (i.e. pure white). 15-40% of these pure white cats have one or two blue-eyes.
  
• Of those white cats with one or two blue eyes, 60-80% are deaf; 20-40% have normal hearing; 30-40% had one blue eye and were deaf while 60-70% had one blue eye and normal hearing.
  
• Of the 5% of white cats in the overall population, 60-80% had eyes of other colors (e.g. orange, green). Of those 10- 20% were deaf and 80-90% had normal hearing.
  
• Deaf white cats with one or two blue eyes account for 0.25 - 1.5 of total cat population
  
• Total number of cats with white coat and blue eyes account for 0.75 - 2.0% of total cat population
  

It is evident from those studies that blue eyed whites exhibit a higher incidence of deafness than do orange/green eyed whites or non-white cats! But not all blue eyed whites are deaf and here's why:-
There is a known link between white coat color, blue eyes and deafness - but since the coat and eye color can be caused by different genes it means that only some blue eyed whites are deaf. There is a gene/gene complex which causes white coat, blue eyes and deafness, but not all cats get their white coat and blue eyes from that particular gene, so not all white cats will be deaf.
If the cat is a Foreign/Oriental White, it carries the gene for 'Siamese Blue Eyes' which is not linked to deafness (the gene for Siamese Blue Eyes is linked to cross-eyes instead). Siamese blue eyes have a reflective tapetum, but this is depigmented because the Siamese colour is caused by albinism. This depigmentation gives the red-eye with flash cameras. Random matings can mean that this gene sometimes appears in non Oriental-looking cats which have colorpoint cats in their ancestry.
Crystal (owned by Alana Harley, Vancouver Island, Canada) has two different types of blue eye.
As this photo shows, one eye has normal green eye-shine and the other has pink eye shine. Depigmentation gives the red-eye with flash cameras
Albino cats are also white. True albinism causes pinkish eyes, but some albinos or partial albinos have pale blue eyes. There are too few albino cats studied to draw firm conclusions, but this mutation is not necessarily linked with deafness. It is also hard for a cat owner to determine whether their cat is a blue-eyed albino cat rather than an ordinary blue-eyed white.
Jamie Linton, who volunteers at a shelter in California, came across this blue-eyed white young cat. The blue is restricted to an inner ring around the pupil, fading to white around the outside of the iris. It wasn't known if the cat was deaf. The most likely explanation seems to be that pigment remained concentrated at either end of the iris muscles as the eye grew, rather than being distributed evenly - a bit like pulling toffee where the middle section goes pale as it it stretched and the more intense colour remains at either end.
There is also a gene for blue eyes which is inherited separately from coat color. This is the gene responsible for the Ojos Azules breed. If the cat is white colored, there is no easy way of telling whether it has the blue-eyed-deaf-ear type gene or the Ojos-Azules-blue-eyes type gene. It's only possible to tell that a cat has this particular gene if the cat is non-white and has blue eyes. Genes for blue eyes independent of coat color may be more common than previously realised. I have encountered three blue-eyed random-bred cats (one ginger, one silver tabby, one brown tabby) in one UK town between 1989 and 1995. This may be the same gene as Ojos Azules, or it may be due to different gene mutations. Other types of blue eye are being discovered in all-white cats in Asia, the blue is different than Siamese Blue Eyes, but does not seem to cause deafness.
The Lady Chablis, belonging to Denise Parmentier, is an odd-eyed white. One eye is yellow-green and the other is blue. She has perfect hearing.
Odd-eyed white cat photographed by Rodrigo Arancibia in Iquique, Chile. She shows no sign of deafness. her kittens are solid white, indicating the dominant white gene.
The white coat can be caused either by a gene for white coloration or by a gene for 'white patching' - sometimes the white patching is so extensive that the cat appears solid white. If the white cat exhibited a few colored hairs or a smudge of color on its body (usually on the head) as a kitten, then it should have normal hearing even if it has blue eyes because it has inherited a non-deafness causing gene for white coat! However, blue-eyed bicolour cats occur and the more white they have in the region of their eyes and ears, the more likely they are to be deaf. This is because one of those genes for white affects the development of both the eyes and ears - it causes lack of eye pigmentation (i.e. blue eyes) and deafness - and since the eyes and ears are close together, if that gene affects that area of the body, it is likely to affect both sense organs (hence odd eyed whites may be deaf on the blue-eyed side).
Aries, a blue-eyed silver-tabby-and-white Norwegian Forest Cat bred by Raquel Ortega Cormenzana of MONTEGANCEDO*ES (Madrid). Sometimes bicolour cats with white faces have blue eyes. Aries has has perfect hearing because the white does not extend to the ears.
Congenital deafness can also be caused by a variety of hereditary factors, just as in humans. Due to random mutation or the wide gene pool, cats of any color can be born deaf (probably less than 1%) - including orange-eyed whites. Hence deaf white cats with non-blue eyes can occasionally appear and in these cases the deafness is not linked to coat color. Deafness can also be caused by illness or injury, so a person adopting a deaf cat may not know whether the cat was born that way or became deaf later on.
There are also green-eyed white cats, the Russian Angora is green-eyed and white is a favorite color. Green-eyed white cats have a lower incidence of deafness than blue-eyed white cats because the gene for white they carry does not normally affect their eye color. It is similar for orange eyed whites; they rarely have congenital deafness. Genetic expression is very variable and orange-eyed/odd-eyed/blue-eyed whites are interbred in many breeds - hence up to 20% of non-blue-eyed, white cats may have some degree of hearing impairment depending on what gene is causing them to have a white coat.
So overall, blue-eyed white cats stand a higher than usual chance of being deaf; but they are not guaranteed to be deaf. Odd-eyed white cats may be deaf on the blue-eyed side. If you have a deaf white cat, it is not advisable to breed from it as this would pass the trait along. Deaf white cats are banned from exhibition or breeding by some fancies in Europe and there is a move to reduce or eliminate this trait from British breeding lines of various breeds. Deafness can cause problems because a cat cannot hear danger approaching. It can cause problems to breeders because deaf female cats cannot hear their kittens crying out and may neglect them. Deaf kittens cannot hear their mother calling to them and may get lost. Deaf cats also seem to have no volume control when meowing. For more information see Living With a Deaf Cat.
NOTES 1. THE "ALBINO SIAMESE"
Feline geneticist Don Shaw, writing in Cats Magazine, September 1972 to February 1973, looked into the so-called Albino Siamese and wrote that not all so called Albino Siamese were true genetic albinos and that many are blue-eyed whites of Siamese type (what we now call Foreign Whites). However, some Albino Siamese had pinkish-blue eyes rather than the clear blue of other blue-eyed white breeds. His research found that early registrations of "Albino Siamese" mentioned "Chinese Whites" - reportedly cats of Asian origin with Malay type and apparently blue-eyed whites having the piebald (white spotting) gene. Unlike other albino animals, the cat's eye structure always gave bluish tone, even to the pink or ruby eyes normally associated with albino animals; this made true albinism in cats difficult to detect.
A mutation for "true albinism" was reported when a Chocolate Point female bred to her Chocolate Point son. Most of the early breedings to produce these reported "Albino Siamese" were within the same Chocolate Point line, meaining the cats were relatively inbred. It had always been suggested that the mother/son mating resulted in doubling up of a recessive allele for albinism, however Shaw challenged this theory.
A Chinese White, and thus white spotting, had been introduced into the Chocolate Point Siamese line and this Chinese White may have carried both white spotting and albinism. If so, the apparently true albinism in Albino Siamese would have been introduced from outside and not occurred as a mutation within the breed. Alternatively, he had noted that Chocolate Point and Lilac Point Siamese seemed to have a pink glow to their otherwise blue eye which wasn't seen in Seal Points or Blue Points.
Shaw believed the supposed Albino Siamese were blue-eyed whites resulting from the white spotting inherited from the Chinese Whites PLUS the pinkish tone due to being homozygous Chocolate Point Siamese (this would be masked by the white spotting) - this combination mimicked true albino. Since the alleged Albino Siamese resulted from inbreeding Chocolate Point Siamese, so Shaw reasonably expected the offspring to have the same pinkish-blue eyes he had observed in Chocolate Points along with a double dose of white spotting that produced a wholly white coat.
The possible alternatives presented by Shaw were that Albino Siamese might be true genetic albinos or they might simply be blue-eyed whites with pale blue, slightly pink-tinted, eye colour. He didn't have enough breeding data to determine which, but there was enough data to demonstrate the presence of white spotting genes in that line (white toes, lockets etc). Since Shaw’s time, true albinism has been seen in cats (the eyes being pink with only a slight blue tint). It has been recorded in the wild in the Asian Leopard Cat and also in the domestic descendants of Leopard Cat hybrids, the Bengal.
Albinism has spread from the Siamese to closely related Oriental breeds. Emma Stothers sent me this report of her albino cat in 2013. Pangur is an oriental longhair with very pale skin and nose. Her paw-pads and ears, are pink-tinted white threaded through with red veins. Her eyes look like faded lilac-blue (Emma describes it as an Elizabeth Taylor colour) and when you look closer,  you can see that the pupil is a dull red, while what looks like lilac is just the pale blue refraction of the lens overlaid on more red. You can very clearly see the blood vessels beneath the iris which can be a bit creepy, because when the light hits Pangur in the right way, her entire eye turns a muted pinkish red. In the dark, they reflect bright red.
Pangur is an indoor cat, and has only been into the backyard under supervision. In direct sunlight, she's unable to open her eyes, and has to squint through little slits. She also squints when looking out of windows . She's very sensitive to light, and hates being photographed using flash. In contrast to her poor daytime vision, she can see very well in the right lighting. Her hearing has been tested by the vet and is perfect. Emma has checked Pangur’s pedigree and found inbreeding. Pangur has two grandmothers, but only one grandfather , which means her parents are half-siblings. It seems likely that the grandfather passed on his recessive albinism gene to Pangur’s parents, and Pangur inherited two copies of the gene. One of her sisters appears to be albino, but her other siblings have neon-bright cyan eyes (i.e. the blue colour normal for foreign white cats). 
NOTES 2. OTHER BLUE-EYED CATS
Not all blue-eyed cats are white, nor are all blue-eyed non-white cats Ojos Azules. The blue-eyed trait turns up surprisingly often in random-breeding cats. Below: silver tabby female feral cat, black-and-white male, red tabby (with white bib) male, red tabby longhair male, odd-eyed tortie, black-and-white male.
CURLED, CURVED AND FOLDED EAR CATS
Copyright 2002-2011, S Hartwell
Cats generally have prick (upright ears) which are triangular in shape. The size, set and exact shape varies from the small ears of Persian cats to the "bat-like" flared ears of some Siamese cats. Some breeds require the base to be more widely flared than others. Some breeds have ears placed close together high on the head, others have wide-set ears with a broad expanse of forehead between. The tips may be rounded, pointed, tufted or fringed depending on the breed.
There is also a rare genetic condition causing the cat to have a second, smaller, pair of ears behind the normal ears. These are apparently non-functional and may be no more than ear flaps with no middle or inner ear section. Though affected cats are generally physically healthy, some have reportedly been lethargic or sluggish which suggests that the condition might affect the brain. This rare condition must not be confused with accounts of "four-eared cats" from China - these were Persian-type cats where the fur inside the ear gave the impression of extra ear-flaps.
In addition to variations on a general theme, there are two distinct ear shapes - folded forwards and curved backwards. Both mutations (or very similar mutations) have occurred more than once.
FOLDED EARS
The Sumxu (Chinese Lop) is now regarded as extinct, but was once found in the area around Peking , China. Descriptions of the Sumxu as longhaired cats with glossy black or yellow coats and pendulous ears derive from a series of mistranslations and the confusion of two entirely different animals! A Polish Jesuit missionary to south China, Michael Boym (ca. 1612-1659), first described the Sumxu in his illustrated book Flora Sinensis (1656) in which he also described Chinese fauna. "Sum Xu" was the Portuguese rendering of songshu, meaning " pine rat". Though Boym's illustration resembles a squirrel, the description indicates the Yellow Throated Marten (Martes flavigula) found in the region. Boym wrote that the Sumxu was a pretty yellow-and-black animal that was commonly tamed and wore a silver collar . They were valued as hunters of mice and sold for up to 9 silver coins (indicating their usefulness or rarity).
The cat-like Sumxu was described in early 1700s as a curiosity, and in 1796 when a droop-eared cat brought back from China. But how did the Sumxu name end up attached to a variety of cat?
This engraving is from German Jesuit Athanasius Kircher's book "China Monumentis, Qua Sacris qua Profanis" (1666). The book is written in Latin and describes the Sumxu as being like a cat. The engravings would have been done from descriptions or rough sketches rather than from life. Kircher was a scholar rather than a missionary and he relied on the expert knowledge of those who had travelled to China, such as Boym and one of Kircher's former students, Martino Martini . Kircher's description of the sumxu referred to it as a domestic animal similar to a cat (in Europe, martens were also known as "tree cats"). He described it as black and saffron coloured with splendid hair. Following Boym's description, Kircher also mentioned that the Chinese tamed this creature and put a silver collar around its neck. It was an avid hunter of mice and was rare and valuable. The creature depicted, while similar to Boym's illustration, was the size of a small bear and is shown chasing deer, rather than mice! In this it resembles a wolverine, rather than a marten.
Italian Jesuit Martino Martini had visited China in the 1650s and published Novus Atlas Sinensis in 1655. In the section on Peking Province, Martini described a race of white, long-haired and long-eared cats found in the region. He wrote that the cats in that province were white all over with long hair and long ears like lap-dogs and that these well-fed cats did not catch mice, but were ladies ' lap-cats. Martini became the definitive source of information about China and his description was used in 1673 by John Ogilby and by others who described the long-eared, milk-white cats as being companions and not hunters (there being other cats that were good mousers). The cats were likened to the Maltese lap-dog and to spaniels. It retrospect, it is likely that these white cats were also blue-eyed, a combination of traits associated with deafness in cats - quite likely they couldn't hear the mice in order to investigate mouse bolt-holes! If this was the same mutation as the Scottish Fold , it is associated with sometimes crippling changes to the tail, hind legs and vertebrae which would make an affected cat reluctant to move around.
In 1736-37, Martini's work was superseded by that of French Jesuit du Halde's works on the Chinese Empire. This compiled material from French Jesuits and re-used some of Martini's book, including the description of the long-eared white cats: "Among the animal species one finds singular (i.e. unique) cats which the Chinese ladies seek out to provide them with amusement and feed with plenty of delicacies: they have long fur and drooping ears." This remained the standard reference book for many years and was reused by other authors, including the passage about the lop-eared cats. For example John Green's book described the lop-eared cats of Peking Province as "a particular Sort of Cats, with long Hair, and hanging Ears, which the Chinese Ladies are very fond of." This was later translated back into French by abbé Prevôt as "among the animals one finds a singular species of long-haired cats with hanging ears, which the Chinese ladies greatly liked."
In volume 4 of his "Histoire Naturelle" (?1767), Georges Louis Leclerc, Comte de Buffon, wrote "The Natural History of The Cat". Buffon noticed that, unlike the many breeds of dog, there were few distinct races of domestic cat: Spanish, Syrian and Khorasan (Persian). He believed the long-haired lop-eared Chinese cat could be a fourth distinct type. "They (cats) are in nature much more constant , and their domesticity is not as complete nor as universal as that of the dog, so it is not surprising they are less varied in type. Our domestic cats, though different from each other in colour, do not form distinct and separate races. Only the climates of Spain , Syria or Chorazan ( Persia ) have produced constant varieties; to find another, one must join the climate of the province of Pe-chi-ly ( Northern Zhili) where there are cats with long hair and hanging ears, much liked by the Chinese ladies." Buffon was translated into English in 1781 by William Smellie. According to Buffon, "Our domestic cats, though they differ in colour, form no distinct races. The climates of Spain and Syria have alone produced permanent varieties: To these may be added the climate of Pe-chi-ly in China, where the cats have long hair and pendulous ears, and are the favourites of the ladies. These domestic cats with pendulous ears, of which we have full descriptions, are still farther removed from the wild and primitive race, than those whose ears are erect ."
In a supplement, Buffon added that there was some doubt as to whether the Sumxu was a cat or some other animal: "I formerly remarked, that, in China, there were cats with pendulous ears. This variety is not found any where else, and perhaps it is an animal of a different species; for travellers, when mentioning an animal called Sumxu, which is entirely domestic, say, that they can compare it to nothing but the cat, with which it has a great resemblance. Its colour is black or yellow, and its hair very bright and glittering. The Chinese put silver collars about the necks of these animals, and render them extremely familiar. As they are not common, they give a high price, both on account of their beauty, and because they destroy rats."
Buffon's source was Prevot, whose source was Green, whose source was de Halde, whose source was Martini's 1655 work. Buffon noticed a process we now call neoteny - domesticated animals retain juvenile features into adulthood. Thus wolves ahd pricked ears and domestic dogs often had softer, hanging ears. Buffon believed this could apply to cats as well and that the mild climate and ancient civilisation of China was conducive to domestic cats developing a similar trait to domestic dogs. In his work, Buffon also mentioned the sumxu as a pretty domestic animal of China, not unlike a cat. In his Vol VIII (published 1777), Buffon goes so far as to suggest the lop-eared cat of China is a different species from the ordinary domestic cat and that it might therefore be the cat-like sumxu:
"We said (volume VI, page 14) that China has a race of cats with hanging ears; perhaps this variety is found nowhere else and is a species to the (domestic) cat, because Travellers speak of an animal called the Sumxu, which is completely domestic in China and which is in many respects like a cat. In colour it is black and yellow with extremely glossy fur. The Chinese give them silver collars around their necks and make them very tame . As they are uncommon, they are expensive; not only because of their beauty, but because they most cruelly wage war upon rats."
This seems a big leap (and an erroneous one) from Boym's black-and-yellow squirrel-like creature to Martini's white lop-eared cat, however du Halde had omitted to mention in his compilation that the lop-eared cats were milk-white! While Boym's sumxu was valued as an avid mouser, Martini's lop-eared white cats were not interested in mice and were kept as lap-cats, another piece of information du Halde omitted in his compilation. Boym's illustration of the sumxu ignores its ears, while Martini described them as the defining feature of the white cats. Boym didn't liken the sumxu to cats and it is Kircher who made that comparison. The sumxu (probably the Yellow Throated Marten) was a creature of south China while the lop-eared cats were from Peking Province in the north. Buffon's leap was further misinterpreted by later authors such as French writer Anselme Gaëtan Desmarest who combined the name "sumxu" with the description of the lop-eared cat (its white colour being unknown to him) as "hanging-ear cat, fur long and fine, black or yellow, domesticated in China in Peking province (Pe-chi-ly) under the name of sumxu". This was perpetuated through the 19th and early 20th centuries, especially by cat fanciers looking for new and exotic cats to import .
When China reopened to foreigners after the Opium Wars, a number of missionaries, entrepreneurs, cat fanciers and researchers went looking in vain for the lop-eared cat. In his book "Variation of Animals and Plants Under Domestication" Charles Darwin referred briefly to a drooping eared race of cats in China. In "The Cat" by Lady Cust (1870) it states "Bosman relates that in the province of Pe-chily, in China, there are cats with long hair and drooping ears, which are in great favour with the Chinese ladies; others say this is not a cat but an animal called 'Samxces'"
In his 1885 book "The Cat: Natural History, Husbandry ["Hygiene"] and Illnesses" Gaston Percheron suggested the lop-eared cat might be a hybrid between the cat and a marten: "Certain respected naturalists even claim even that the [domestic cat] mates with the marten and produces among the hybrids those like the latter in colour and fur. In this way they explain the hanging-ear cats of China [...]" but instead of being fed delicacies by its mistress, the cat has become a delicacy to be eaten, "The Chinese Cat. It has long, silky hair and hanging ears, like those of a badger. Its flesh is highly esteemed by the inhabitants of the Celestial Empire. As with the dog, it is treated with great solicitude by feeders and fatteners of this country and when it is fattened, it appears alongside swallows' nests on well-served tables. " As a side-note, in 1926, Siamese cat fancier Lilian J Veley remained adamant that Siamese cats were not merely a form of domestic cat, but resembled an unknown type of viverrine that lived in the region (the most likely contender again being the Yellow Throated Marten) in their sable-and-dun colour and "marten-like" face! Maybe "Sumxu" and "Siam" sounded similar to her ears.
The Chinese, or Hanging Ear, Cat according to "Die Hauskatze, ihre Rassen und Varietäten" (Housecats, Their Races and Varieties) from "Illustriertes Katzenbuch" (An Illustrated Book of Cats) written and illustrated by Jean Bungartz, published in Berlin in 1896 (There is no published English translation of this book, this gives the gist of the text)
The Chinese or Hanging-Ear cat is most interesting, because it provides proof that by continual disuse of an organ, the organ atrophies. So with the Chinese cat the hearing and/or the ears have deteriorated. Michel says the Chinese, not only admire the cat in porcelain, but also value it for culinary reasons. The cats are regarded as special titbits and enjoyed particularly with chains (noodles?), with rice". This cat is bred particularly for the purpose of meat production, and is a preferred Chinese titbit; this is not unusual if one considers that the Chinese consume much the sight of which turns the stomachs of Europeans. The poor creature is locked up in small bamboo cages and much like a kind of geese fattened with plentiful portions. Extensive trade is carried on with other parts of Asia and the Chinese allow no tomcats to be exported so there is no interference in this lucrative source of income.
Due to the restrictive conditions that have deprived the cat of its actual use, its hearing decreased because it was no longer needed as for hunting its own food. With no need for watchfulness, it was useless to have sharp hearing to listen for hidden things so the hearing became blunt and in natural consequence the ear lost its upright nature, gradually become lower and becoming the hanging ear that is the characteristic feature of the Chinese cat. At first impression this is a surprising and amusing look, but is impression is lost with closer examination. If one ignores the characteristic of the ears, one sees a beauty similar to the Angora cat: a long, close coat of hair, albeit less rich, covers the body. The hair is silky- soft and shining and the colour is usually a light yellow (isabelline) or a dirty white yellow, although some have the usual colouring of the common house-cat. In size it considerably exceeds house-cats and is stronger. The ears hang completely, as with our hunting dogs and are large in relation to cats.
Although the Chinese cat is found in considerable numbers in its homeland, it rarely arrives at European animal markets. Only one such cat has reached us in the flesh; we acquired this years ago when a sailor returning from China brought it into Hamburg . The accompanying illustration is based on this. In character it is like the Angora cat and somewhat inactive. It also prefer to live by a warm fire, is a little sensitive to attention , hears badly and is at its most animated when it sees the milk or food. Apart from its unusual ears, it does not have any really attractive characteristics and is a strange representative of the house cat.
Though reports refer to the Chinese Lop having pendent or pendulous ears (suggesting abnormally long or floppy ears e.g. like a labrador dog) this is probably an exaggeration. In all likelihood, and in the absence of any current examples or pictorial evidence, the ears were folded in a manner similar to the Scottish Fold. In Frances Simpson's "The Book of the Cat" (1903), contributor H C Brooke wrote "There is said to be a variety of Chinese cat which is remarkable for its pendent ears. We have never been able to ascertain anything definite with regard to this variety. Some years back a class was provided for them at a certain Continental cat show, and we went across in the hope of seeing, and if possible acquiring, some specimens; but alas the class was empty! We have seen a stuffed specimen in a Continental museum, which was a half long-haired cat, the ears being pendent down the sides of the head instead of erect; but do not attach much value to this."
In 1926, Brooke wrote that "for donkey's years" Continental cat shows had offered prizes for the Drop -eared Chinese Cat. On each occasion, the cat failed to materialise and Brooke considered it to be mythical. Other writers suggested it was the result of haematomas causing the ears to fold or crumple. Brooke noted that although no-one ever saw the cat itself, one always met "someone who knows someone whose friends has often seen them". He had been assured by a Chinese gentleman he had met only once that "he knew them well". HC Brooke, and other fanciers, made enquiries of the Chinese Embassy, of Hagenbeck's (a major Hamburg animal dealer at the time) and of a "certain well known author, who has lived for years in China and knows that country well", but to no avail. The American Express Company had instructed their representatives at Shanghai and Peking to make enquiries, again without success. None of the wild animal dealers knew of the Chinese Lop.
The German naturalist, Brehm, had given a very detailed description of the cat in the 1700s. Brehm was usually very accurate. In 1882, Brooke had seen a stuffed specimen in a Continental museum. The specimen was "half-coated with yellowish fur". He admitted that it might have been a fake or a cat with its ears deformed by canker (i.e. cauliflower ears) that had been presented in all good faith. All avenues of enquiry exhausted, Brooke declared the Chinese Drop-eared cat extinct. The last reported sighting of the Chinese Lop seems to have been in 1938 when a droop-eared cat was imported from China. On that last occasion the mutation was thought to be restricted to white longhaired cats. It is hard say for certain whether these were isolated cases or whether the Chinese Lop was a genuine variety. It cannot even be said with certainty that the trait was an inherited one.
Cats with folded ears have been reported in the Hebrides, Germany and Belgium, but were apparently regarded as no more than isolated curiosities and not bred.
The only widely recognised fold-eared cat is the Scottish Fold and its longhaired version, the Highland Fold (Coupari). These have ears which form a close-lying cap. Also known as Scottish Lop, the Scottish Fold traces back to a female white shorthair farm kitten discovered near Coupar Angus , Perthshire, Scotland in 1961. This cat, Susie, produced some fold-eared kittens. One of these, Snooks, became the founding mother of the breed after Susie's unfortunate death in 1963. By 1967, numerous "Lop-eared Cats" had been born however the UK cat registries declared the trait to be a deformity and refused to accept the "Scottish Fold" for breed status because of potential ear problems (an inconsistency considering they recognised the tailless Manx). Interest in Scottish Folds in the USA led to breeding being continued there and it was recognised as a breed in 1973. The cat does not suffer undue ear problems and one British registry accepted it in 1983.
The Coupari is the name given to Longhair Folds in the UK, although these are known elsewhere as Highland Folds. The argument is that Coupar Angus is not in the Highlands, however Highland Fold is no more inaccurate than other breed names e.g. Balinese, Tonkinese. Longhaired cats were present in the original Scottish Fold, but the shorthaired variety was preferred because the ear shape was more visible. It was developed in the 1980s and recognised in 1986. It is known by various names: Coupari (in UK), Highland Fold, Longhair Fold and Scottish Fold Longhair.
Kittens are born prick-eared and start to develop the folding at around 4 weeks old. The fold is fully developed at around 3 months of age. The gene causing the folded ear trait is a dominant gene which causes skeletal problems if the cat inherits two copies of the gene. Scottish Folds are therefore always bred to prick-eared to keep the incidence of problems to a minimum. The problems are a thickened tail caused by tail vertebrae fusing and thickened legs with swollen feet due to overgrowth of cartilage around the paws. These side-effects cause problems with walking.
Prick-eared Scottish Folds gave rise to the Scottish Shorthair in Australia. This breed (with longer tail and legs, and different coat texture to the British Shorthair)was recognized and named by the Queensland Independent Cat Council ( QICC ) as a result of a submission by Rona Sandilands at a meeting at then QICC- president Pat Mercer’s home attended by prominent Scottish Fold and British breeders, including Lillian Carter founding breeder in Queensland. The first Scottish Shorthair exhibited was Ptah Dunstan Gold, bred by Lillian Carter and owned by Rona Sandilands. During the 1980s he was exhibited at the Queensland Longhair Cat Club show, achieving a Reserve in Group 3 under Victorian Judge Marie Orchard. There is also a Scottish Longhair.
The Poodle Cat (Pudelkatze) is essentially a breed developed from the Scottish Fold and Devon Rex to create a curly coated fold-eared cat - a curly-coated Scottish Fold or fold-eared Devon Rex. It was developed initially in Germany where its future is threatened by rulings prohibiting the breeding of cats with harmful defects. This ruling affects Scottish Fold cats because of the skeletal abnormalities which can occur. The breed is attractive and if breeders are careful to breed only from healthy cats, there is no reason it should not be accepted. Astonishingly, there appears to be the intention of adding Manx into the mix so that the cats are tailless as well. This would create a more dangerous mix of semi-lethal genes since the Manx condition can cause other skeletal abnormalities.
Back in 1981, Phyllis Lauder wrote in "The British, European and American Shorthair Cat" wrote of news from Australia of "interference not beneficial to the domestic shorthairs". A correspondent, Mrs Batten, had been asked for her views on the idea of crossing Manx with Scottish Folds. Scottish Folds had not yet been recognised. Lauder wrote that the idea was born of "love of change for its own sake and by the desire to meddle" since a Manx/Fold cross would not only produce a freakish-looking cat, it would be a tragedy for the cats - the two breeds had enough to contend with in refuting charges of deformity.
The unrecognised Oriental Fold is a Siamese/Oriental type cat with folded ears (I believe it came from attempts to breed colourpoint Scottish Folds). The Hemingway Fold is an unrecognised variety of fold-eared cat with extra toes. The term "Hemingway" is often used in America to describe polydactyl cats (mitten cats). It results either from a spontaneous folded ear mutation in a polydactyl cat or from accidental or deliberate crossed of Scottish Folds and polydactyls, possibly for curiosity or as attractive pets.
A fold-eared cat aiming for recognition is the Foldex or Exotic Fold. The Foldex is a cat of Exotic (shorthaired Persian) body type developed through crossing Scottish Folds with Exotics. It has a shorter nose than the Scottish Fold but not as short as the Exotic Shorthair. In other respects it is a fold-eared version of the Exotic Shorthair. Facially, the Foldex looks like "a little furry owl". Like the Scottish Fold, it is prone to the same bone deformities and produces some non-folded offspring.
In 2005, the Ukrainian Levkoy Cat, a fold-eared naked cat, was created using the Don Sphinx and Scottish Fold. The Ukrainian Levkoy is less extreme in body and face type than the Don Sphinx (the face is wider and ronder) and the ears do not fold tightly to the skull as in the Scottish Fold, but stand out from the head and fold closer to the tips. It also occurs in velour and prick-eared forms.
In 1986 there was a report of a "Hebridean Cat". The owner said that the tiny ears were a feature of Hebridean and in addition the ears were folded like those of a Scottish Fold. No more was heard of the Hebridean breed.
In the mid-1990s, a fold-eared black cat turned up in a feral cat colony in Essex, England. It was not possible to determine whether the trait was inherited or was a birth defect. It is extremely unlikely to be due to a free-ranging Scottish Fold as these are uncommon in Britain - there would certainly be no free-ranging unneutered Scottish Fold males around.
In 2007, Vasilis Lekkas from Athens (who is deeply involved in the recently formed Greek network for the preservation of domestic livestock including cats) reported possible folded ears on the Greek island of Myconos, but these may have been due to injury.
Lizzie Ellis (The Feline Rescue Association Inc, Maryland, USA) provided this photo of a folded ear apparently caused by frostbite. The ears had to be surgically removed. This shows how inury, including haematoma ("cauliflower" or collapsed ears from fighting) and frostbite can mimic folded ears at a casual glance.
CURL -EARED CATS
The first recognised curl-eared breed was the American Curl. The ears are curled or swept back on themselves and feel rigid to the touch . The originate from a stray kitten, Shulamith, in 1981. Shulamith was a black longhair female with strangely curled ears. She went on to produce kittens, some of which also had curled ears. These attracted attention when shown at a cat show in 1983 and in 1985/6 the American Curl breed was recognised.
The mutation is a dominant gene so litters will often contain a mix of curl-eared and prick-eared cats. It is impossible to tell which kittens will develop a good curl - they are all born prick eared. The ears curl up tightly over the next few days. During the next four months, the tight curl starts to relax until the final semi-curled state is reached. As far as is known, the gene does not cause detrimental side-effects.
The Hemingway Curl is a localised variety rather than a breed. The first Hemingway Curl appeared as a spontaneous curl-eared mutation in a colony of polydactyl cats known as Hemingway cats (after the polydactyl cats of Ernest Hemingway) on Key West. Hemingway Curls have since been crossed with African Jungle cat hybrids to produce the Jungle Curl breed. It occurs in shorthair and longhair versions and may attract sufficient interest for a breed to be established.
Having established the curl-eared trait, it became possible to cross American Curls with other breeds to create new varieties e.g. the Kinkalow which is a mix of American Curl and Munchkin to produce a short-legged curl-eared cat.
The Ruffle was, according to one source, an accidental side-effect resulting from attempts to improve the ear size of American Curls by introducing the Cornish Rex into a breeding line. Whether the Cornish Rex is an allowable outcross for the American Curl is another matter entirely. According to another source, the original mating was a deliberate attempt to produce a new breed. Progeny from this breeding line was apparently acquired by a breeder unaware of the Curl-to-Rex matings which had the side-effect of introducing the recessive gene for curled fur. The gene remained hidden until two carriers were mated and produced curly coated American Curls some generations later. The rippled effect of the fur reminded the breeder of a brand of potato snack hence the name Ruffle. Unfortunately the breeder's circumstances changed and further development of the Ruffle breed could not be funded. Interestingly, the Rex coatd in American Curls was mentioned as early as 1991 by feline geneticist.
 An experimental breed called the Jaguarundi Curl (shorthaired and longhaired) was reported in 2001, but its existence is unconfirmed. The Jaguarundi is a South American felid which is very un-cat-like in shape. It is suspected that the reporter had confused it with the Jungle Curl. The Jungle Curl does exist and is a hybrid of Jungle Cat (F chaus) and American Curl or Hemingway Curl with Serengetis, Jungle cat hybrids, pure Jungle Cats, Bengals, Egyptian Maus, Abyssinians and other domestic shorthairs contributing their genes to the mix. It is still in the developmental stage to produce wild looking cats, preferably with a spotted or rosetted pattern, with curled ears. Several breeders in Illinois are working with a curl-eared polydactyl cat called Impians (formerly called Tulips). Impians were originally developed in the 1990s by crossing American Curls with polydactyls, creating a harlequin patterned semi-longhaired breed. The markings, which can be any colour, are restricted to the head, down the spine, shoulders, hips and tail.
The Australian Curl was a breed that never was, rather it was a single female cat, Matilda , which was discovered in 1996 and who failed to pass on the trait. The curl of the ears apparently differed from that of the American Curl though no precise details were provided. She was mated to a normal eared cat of similar type, but none of the kittens were curl-eared. She suffered severe illness after the birth (1997) and could not be bred from again. No back-crossing could be attempted to determine whether the gene was carried as a recessive. Although none of the offspring had curled ears, they were not bred to each other (or it was not possible to do so) to determine whether they carried the trait was carried in recessive form. It is possible that a breed opportunity has been missed due to Matilda's unfortunate illness.
In 2007, Vasilis Lekkas from Athens (who is deeply involved in the recently formed Greek network for the preservation of domestic livestock including cats) reported "In the islands of Paxoi in the Ioania Sea in western Greece, an otherwise conventional cat with curled ears is common. Locals consider it their "normal" cat and view as unusual cats of the mainland or other islands. These cats tend to be white bodied with restricted coloured patches. It appears the mutation may be similar to American Curl. There are two Paxoi islands, but is unclear, which one the report relates to." A young cat with a moderate curl has also been reported.
The Elf breed was initiated in 2006 with the first full hairless, curl-eared Elf being born in 2008. It combines the Sphynx hairlessness with the American Curl's ear conformation. Permissible out-crosses include non-pedigree domestic shorthairs. They have a sturdy, athletic build, are sociable, intelligent, inquisitive and people centred. Half-Elf cats e.g. furred Elf variants result from outcrosses to diversify the gene pool.
The Dwelf also combines curl ears with hairlessness, but adds Munchkin and Highlander (a bobtailed, curl-eared polydactyl breed) to the mix. These combine hairlessness, curled ears and short legs. Non-standard (long legged) Dwelfs may resemble the Elf breed.
IRISH SHORT-EARED CAT
Described in a light-hearted book depicting cats which had been "painted" with vegetable dye (in fact all painted cats were photo-manipulations), the Irish Shortear is a mutant-eared cat that never was. This footnote has been included because people have asked where to find an Irish Shortear!
The book in question was the spoof "Why Paint Cats" and the fake breed among genuine breeds was the Irish Shortear. It was described as Burmilla x Scottish Fold with large protuberant eyes, short ears and very relaxed nature due its excellent vision. The photo was that of a Brown Burmilla which was been edited to give the cat larger eyes, narrower chin and short ears (the original ear tip had been photographically enlarged and grafted back onto the face). The face was then grafted back onto the image of the cat's body. One giveaway was that the facial area had a "floating effect", which is common in photo-composites. A Scottish Fold would have given folded not shortened ears. In addition, the partial-dominant "macro-retinal" gene mentioned in the text is fictional (and it should have been a macropthalmia gene since the retina is the back of the inside of the eye).
BOBTAILED AND CURLY TAILED CATS
Copyright 2002 - 2013, S Hartwell
THE TAIL
The normal tail has 21 - 23 vertebrae on average with the normal range being 18 - 28 vertebrae. It is on average 25 cm (10 inches) long, but can range from 20 cm (8 inches) to 30 cm (12 inches) with a few exceptional specimens having 35 cm (14 inch) tails
There are various mutations affecting the tail and in 1940 American zoologist Ida Mellen wrote of oddities in cats' tails including kinked, bobbed, curled and even double tails. Curly tailed cats were known in China in the 12th Century (probably bobtails) and ringtail cats were known in the USA sometime prior to 1940. In 1868, Darwin wrote in The Variation of Animals and Plants Under Domestication "Thropughout an immense area, namely the Malayan archipelago, Siam, Pegu, and Burmah, all the cats have truncated tails about half the proper length, often with a sort of knot at the end."
Tailless and cats have occurred periodically through random mutation. They have been reported in Bosnia , Burma, China, Crimea, Java , Malaya, Denmark, Nova Scotia and Thailand though some of these will have been bobtails. Bobtail cats occur throughout Asia and into Russia, with more recent mutations occurring in the USA. In a 1949 study, Searle found no kinked tails in London cats. He studied Singapore's cats in 1959 and eventually concluded that stubby-tailed cats were common in Singapore, but in general was rare in Europe except for the Manx breed where the tail was often completely absent. The Manx and the Asian bobtails are due to different mutations. Research indicates that the “tailless gene” has 4 alleles i.e. there are four different versions of the same gene. The Manx breed is based on one of those mutations, and that mutation can have side effects such as spina bifida. The PixieBob is based on a different mutation of the same gene; that mutation has a different appearance from the Manx mutation. The American Bobtail, which occurred spontaneously in a different geographic area, may be based on the same, or a different mutation of the same gene. The Kurilian Bobtail is due to an incomplete dominant gene. However, the similarly named Karelian’s bobbed tail is due to a recessive gene, as is the Japanese Bobtail.
BOBTAILED CATS
The bobtailed mutation is not related to the Manx mutation. It has occurred independently in various geographic areas. It is widespread in Asia and parts of Russia. An Abyssinian Bobtail has been reported in non-pedigree Abyssinian-type cats (location not specified). A Spanish bobtail has also been reported. Cats with knotted, shortened, kinked and pom-pom tails are relatively common in Tenerife and the Canary Islands, the mutation apparently having become established due to the isolation of the gene pool (tourists frequently mistake the trait for breakage). Similar mutation have occurred spontaneously in the USA. There are also purported Bobcat hybrids that have inherited the short tail from the wild parent. According to leading feline geneticists, the genes governing tail conformation are located on a mutation hotspot.
According to an earlir anatomist, Sir Richard Owen, and to Professor H N Moseley, the kink in the Siamese's tail (a mild form of the bobtailed trait) was the relic of a prehensile tail, possibly inherited from civet ancestors (though neither civets nor genets had prehensile tails)! Others asserted that the kink was due to intercrossing the Siamese with the "common strain" however Lilian J Veley wrote in 1926 that this could be discounted since there was no other cat known in Siam, "common" or otherwise that had ever possessed an original kink, making it a folly to try to eradicate the trait. Presumably Ms Veley had not encountered the numerous bobtailed and kink-tailed street cats in Thailand. In "Our Cats" of May 1901 there was information on the Siamese's tail from interviews with the King of Siam and his Private Secretary. While the cats in the Royal Palace apparently had no kinks in their tails, occasionally a "tramp cat of Malay origin" strayed in and the resultant crossbred cats had kinked tails.
In 1783, Willian Marsden, Fellow of the Royal Society and late Secretary to the President and Council of Fort Marlborough wrote in "The History of Sumatra" of the Malay Cat: "All their tails imperfect and knobbed at the end." In 'The Variation of Animals and Plants under Domestication", Darwin wrote "throughout an immense area, namely, the Malayan Archipelago, Siam, Pequan, and Burmah, all the cats have truncated tails about half the proper length, often with a sort of knob at the end. […] The Madagascar cat is said to have a twisted tail." Another writer and traveller, Mivart, had corroborated the statement regarding the Malay cat, of which he said the tail "is only half the ordinary length, and often contorted into a sort of knot, so that it cannot be straightened […] Its contortion is due to deformity of the bones of the tail". Joseph Train had also mentioned th Malayan cats, comparing them with the Manx: "The Manks rumpy resembles some what in appearance the cats said by Sir Stamford Raffles to be peculiar to the Malayan Archipelago." Sir Stamford Raffles' name is closely associated with Singapore.
HC Brooke reported seeing three spotted tabby Malay cats in Holland in the 1880s and some very similar specimens at the Jardin d'Acclimatation in Paris around the same time. "The tails of these [Paris] cats, about three or four inches long, were tightly screwed, or at least the tail formed three complete revolutions. The ' screw ' tail, as also the spotted type of colouration, appear to be becoming very rare." In 1889, Mr O Gould had apparently taken four Siamese to Ceylon and soon Ceylon was "overspread" with their progeny, known as "Gould's Cats" and many had kinked tails in spite of there being no Malay cats for them to interbreed with. Lilian J Veley wondered if the Malay cat's kink had come from the Siamese rather than the other way round (she considered the Siamese to be the older type). She also implicated the Korat "a blue variant of the Siamese, to which our 'blue-pointed' freaks are due."
Jean Bungartz described several bobtailed cats in his 1896 book "Die Hauskatze, ihre Rassen und Varietäten" (Housecats, Their Races and Varieties) in " Illustriertes Katzenbuch" (An Illustrated Book of Cats). He noted that according to Brehm Martens, on the Sunda Isles and in Japan there were cats with different lengths of tail. Kessel had told Weinland that there were short-tailed cats on Sumatra. Bungartz added that the cats of Cochinchina had only a short, curled tail and the Madagascar cat had a turned, knotted tail.
R Shelford, former Curator of the Sarawak Museum wrote in his book "A Naturalist in Borneo" "It may be mentioned here that the domestic cat of the Malays is quite a distinct variety [...] it is a very small tabby with large ears and a body and hind-legs so long that it lacks all grace. The tail is either an absurd twisted knot or else very short and terminating in a knob; this knotting of the tail is caused by a natural dislocation of the vertebrae so that they join onto each other at all sorts of angles." The length of hind-leg was a trait shared by the Manx, leading some cat-fanciers to believe that the two were related.
Miss Lowndes, daughter of the novelist Mrs Belloc Lowndes described a Malay kitten that she had acquired. It had recently arrived, along with its mother, from the Straits Settlements. "It has a triple-kinked tail. It is, unfortunately, not of the spotted kind, but these seem to be very rare nowadays." More information was provided by Mr Boden Kloss, Director of the Raffles Museum and Library at Singapore "The tail which distinguishes these cats may be clubbed or kinked, very short or of medium length, and the animals themselves of many colours - plain, piebald, or patterned." He also wrote "A fair proportion of the cats of Singapore seen in native villages are short-tailed animals with a kinked tail. There would [be], I should say, three or four kinks. In colour they may be tabby, or boldly black and white. As a point of interest it may be noted that Felis planiceps [ Flat -Headed Cat], one of the wild species of the peninsula, tends to resemble the domestic Malay cat in the matter of tail." The cat writer HC Brooke, who had an interest in the Malay cat, wrote that F planiceps and the domestic cat were unlikely to be inter -fertile.
Mr H O Forbes had exhibited a bobtailed Malay cat to the Liverpool Biological Society and shown the cause of the knotting to be the development of wedge-shaped cartilages between the tail vertebrae. Forbes attempted to link the Malay bobtails to the bobtailed cats found in part of Portugal . In the 1920s, Forbes wrote "My remarks referred to the interest I had in exhibiting the creature's skin from the occurrence in the East of what I had noted as extremely common in the cats of Portugal when I lived there about 1876. The kink, I was told was then believed to have become hereditary, from a custom long practised by the Portuguese of pinching or breaking the tails of the new-born kittens, and it would be of special interest if it could be established that the kink in the Malayan cats' tails had been communicated to them through those imported by the early Portuguese into the East. If I can trust my memory the tail of this cat, though short and kinked had the full number of vertebrae, some of them reduced and wedge shaped." Others disputed his theory as the trait had been reported in the Malay cat since at least 1783 and no amount of tail-pinching would cause the trait to become hereditary!
Kuantan, Malaysia. Feral male with perfect pom-pom tail.
From Lake Chini, Kuantan area, local domestic male with bobtail and extra toes (not visible). His offspring ranged from bobtail, kinked tail to two-thirds normal length tail.
 
The bobtail trait ranges from a normal-length tail with a distinct kink, through to a short twisted pom-pom and just about anything between those two extremes. The degree of kink is variable and the vertebrae are affected so that the tail cannot be straightened. It is often possible to feel a bony knot inside the kink where vertebrae have fused. The bobtail mutation is widespread throughout Asia, extending as far as Russia. It is well know that early Siamese cats had kinked full-length tails and this is still seen in Siamese-type cats in Thailand, but has been bred out of pedigree Siamese cats. Colourpointed cats were kept by Thai royalty and legend has it that a princess entrusted her rings to a palace cat while she bathed. She threaded them on the cat's tail and the cat knotted its tail so the rings did not fall off. The kink therefore marks where it knotted its tail. Unlike the Manx mutation, there appear to be no detrimental effects.
The Madura or Buso cats are a closed colony of bobtailed blue cats with green eyes which live on the tiny and isolated island of Ra'as off the coast of the Indonesian island of Madura (it is a 6 hour ferry ride from Madura to Ra'as). There are reckoned to be less than 100 of these cats and they are in danger of extinction, in part due to local beliefs that prevent the exportation of fertile cats. The Madura breed and its decline have been documented by Dr Ronny Rachman Noor (Faculty of Animal Science, Bogor Agricultural Universtity). The cats are known to the islanders as "Kucing Buso" (grey cat) and the true Madura cat is blue self, suggesting they may have come from Korat stock taken to the island on trading ships. There is some mythology surrounding the breed, just as there was about the "Royal Siamese". The true Madura cat, originating from Ra'as, may only be kept by high-ranking people such as religious leaders, high rank government officials and informal leaders. Anyone attempting to smuggle a cat from Ra'as will find their boat sinking. The cats' posture and the triangular facial shape are similar to the wild cat and leopard. The cats are large with a medium length tail with a visible bend or kink at the end. The fur is relatively thick. The true Madura cat is solid grey known as "buso" often with lavender nose leather. A few so-called Madura cats on Madura can be found with the brown sepia, mink and colourpoint patterns as well as blue bicolours due to matings with local cats. A brown mink "Madura cat" exhibited at a cat show in Surabaya was of Japanese Bobtail type. The declining numbers appear due to people trying to export the cat out of the island and to a high mortality rate among breeding cats, both on Ra'as and on Madura, indicating either a lethal gene or a high degree of inbreeding. Cats brought out from Ra'as are castrated due a local belief that the fertile Buso cat may only be kept at the Ra'as island (castration in itself may pose dangers of infection). To preserve this cat as a distinct type and avoid extinction, breeders would need to establish breeding lines which would mean obtaining unneutered cats. The restriction on who may own a true Madura cat also limits ownership and controlled breeding.
There are various tales associated with the Asian bobtailed cats. The Siamese tail kink, for example, was supposedly due to a princess threading her rings on the cats' tails for safekeeping and the loyal cats kinked their tails to keep the rings in place. Siamese were also believed to be royal palace cats. Likewise Madura cats may only be owned by people of high status. A Malaysia belief is that if a kittens tail is cut off and buried under the doorstep, the cat will not stray from home. Another is that monks cut the tails off of cats so that the cats do not go to heaven. A cat with a stumpy tail is not perfect and imperfect creatures cannot go to heaven. I examined a tabby cat brought back from Japan whose owner claimed that its tail had been cut off by monks during kittenhood. I found evidence of knots and kinks in the remnant of tail which told me it was a natural bobtail. Also, the tail ended quite normally in a black tip. This was a perfectly normal genetic bobtail, though the owner preferred to think she had rescued an abused cat.
In 1988, the Cat Association of Britain finalised the standard for the "Oriental Bobtail"; a cat of oriental (or foreign) conformation and coat, but with a bobbed tail. Since then, little or nothing has been heard of this breed.
The most famous bobtailed breed is the Japanese Bobtail found in both shorthair and semi-longhair varieties. This is due to a recessive gene. It has appeared in ancient Japanese art and has evidently existed back as far as the 6th century. It is claimed that it was introduced into Japan from China at that time, which corresponds to Ida Mellen's account of curly tailed cats in China. Once the pet of Japanese nobles, it eventually spread to the general population but was not considered anything more than a common moggy. It attracted the interest of American breeders in 1968 and was recognised in the US in 1978. The longhaired version had always existed (longhair is due to a recessive gene) but was not given breed status until 1991.
In a study comparing the genetics of several breeds (Lipinski MJ, et al., The ascent of cat breeds: genetic evaluations of breeds and worldwide random-bred populations, Genomics (2007), the Japanese Bobtail was found to be more closely related to Western cats than to Asian cats, though the breed showed some Asian genetic influence. Cats are not indigenous to Japan, but reached the islands along Asian trade routes hundreds of years ago. However, the Japanese Bobtail was imported into the USA in 1968 and developed as a breed in the USA. As a result, their gene pool was largely influenced by European/American cats rather than by Asian cats. It would have been interesting to compare the Japanese Bobtail to the indigenous Russian Kuril and Karel bobtails which (visually) are of Western European, rather than Asian, type.
Distinguished by its bobtail and high-cheekboned triangular face. Close-lying, silky short hair. The tail is 10-13 cm in length if fully extended, but due to the kinked structure it appears only 4-7 cm long. The tail hair often grows straight out in all directions, giving it a rabbit -like fluff-ball or pom-pom appearance. In Japan it is found in all colours including agouti (Abyssinian pattern) and colourpoint though these are not accepted in the Western fancy.
In 2001 a Singapura Bobtail was reported in a breeding programme. The bobtail kitten appeared in a Singapura litter, tracing back to a part-tailed foundation cat. Inbreeding can cause recessive genes to reappear. The kitten initially appeared to be tailless, but proved to have a stumpy tail similar to that of the Japanese Bobtail. This is not surprising since there are numerous bobtails in Singapore.
Street cats from Singapore and Johor Bahru (the town at Malaysian end of causeway to Singapore) showing the typical bobbed or kinked tail of local felines.
 
The bobtail trait is also widespread in parts of Russia and following the break with communism, a cat fancy developed in Russia and several bobtailed breeds are under development. The Kurilian’s bobbed tail is due to an incomplete dominant gene, while the similarly named Karelian’s bobbed tail is a recessive gene.
The Karel Bobtail (Karellian) is a shorthaired/semi-longhaired breed which occurs naturally along the coasts and islands of Lake Ladoga. The mutation is claimed to be identical to the Japanese Bobtail. These are elegant, svelte cats with lifted rumps and short pompom tails (4-13 cm). The Kuril Bobtail (Curilsk) is smallish, compact and cobby with a short (5-13 cm) "bob" or "pompon" tail . It has shorthair and semi-longhair forms. The Mekong Bobtail (formerly the Thai-Bob or Thai Bobtail) is a medium-sized Russian breed resembling the Traditional style (Apple- or Round-head) Siamese in all non-mitted colorpoint varieties. The tail is short (3-11 cm), bobbed and its outline is smoothed by the coat. The Toy-Bob or Toy Bobtail (1986) is a Russian miniature breed, no larger than a normal 3-4 month kitten. Toybobs have short, solid bodies and excellent muscles, with short straight or curved tail-remnant (3-7 cm), straight or corkscrewed, covered with fur in a "pompon" or "brush" effect. All of these were recognised in the 1990s, but apart from the Toy-Bob have existed for far longer.
Confusingly, there is another variety known as a Thai Bobtail. It is a naturally occurring bobtailed variety of Oriental/Burmese type. These are mostly, but not exclusively, colourpointed and are found in Thailand and Malaysia. The Malay Cat reported in the Malaysian peninsula in 1881 and found throughout Malaysia is similar to Japanese Bobtail, but has not been adopted as a formal breed.
The second officially recognised bobtail breed is the American Bobtail bred from a foundation cat discovered in the 1960s in Arizona. This is a dominant gene mutation. It occurs in shorthair and semi-longhair varieties and has a powder puff tail up to one third normal length. It was reputed to have bobcat blood (based on purely circumstantial evidence), but it may be a Manx-type mutation since some lines produce a range of tailless, rumpy, stumpy, longy and kink-tail cats. To avoid mixing up different mutations, the American Bobtail is never bred with either the Manx or the Japanese Bobtail. This is the official American Bobtail breed, however the trait has evidently occurred independently several times. The Sno-Bob is a colour variety of American Bobtail. It apparently resembles the Alaskan Bobcat being pale in colour with darker ear tips and darker bob-tail.
In 1940, American zoologist Ida Mellen wrote "The American Domestic Bobtail Cat of the New England and Middle Atlantic States (called the Rabbit Cat) traces its ancestry to the Manx cat, but the distribution of tailless cats is wide, covering the Crimea and other parts of Russia, the Bismarck Archipelago, the Malayan Archipelago, Burma and Siam" However nothing more has been said about the purported American Domestic Bobtail and it may have been no more than a few isolated Manx cats. The cats from Malaya, Burma and Siam are related to the Japanese Bobtail rather than the Manx.
There have been claims of an American Bobtail cat breed which resulted from bobcat/domestic cat crosses in the late 1970s. The breeder indicated that bobcats will mate with female Siamese cats because the scent of a Siamese female in season resembles that of a sexually receptive female bobcat. Bobcats can and do mate with domestic cats to produce at least some fertile offspring.
The Si-Bob (Si-Bobtail) is not a formal breed. It is the colloquial name for colourpointed cats of Japanese Bobtail or American Bobtail type. These are not recognised as part of the main breeds and their breeding is not actively encouraged, but they are attractive crossbreds not unlike the Thai Bobtails. Si-Bobtails occur naturally in Japan.
The PixieBob (1995) was, and sometimes still is, reputed to be derived from natural bobcat/domestic hybrids and to have inherited its tail from the bobcat. It is a polydactyl (extra-toed) breed derived from "Legend Cats" and in some respects it resembles a bobcat. The presence of bobcat genes has not been confirmed though the cat certainly resembles the wild species in several respects. On the one hand there is a persistent tale of "Legend Cats" - cats that are the result of natural cross-breeding between the two species. On the other hand there are cat fancies that refuse to recognise hybrid breeds. This leaves a few PixieBob enthusiasts both promoting the Bobcat influence (some claiming 25% bobcat blood) and simultaneously playing it down! Apparently genetic markers for the Bobcat do not show up in modern PixieBobs, although this may be due to the genes being diluted by several generations of backcrossing to domestic cats.
There is an unconfirmed version known as the Munch -Bob which is a Munchkin/PixieBob cross to produce a short-legged bobtail. In theory, any of the bobtailed or tailless breeds could be used to produce bobtailed/tailless Munchkins.
Not to be confused with the PixieBob is the Poly -Bob (1998) which is linked to the infamous Twisty Cats (cats bred for detrimental deformities). Poly-Bobs are not a recognised breed. They are bobtailed polydactyls which carry the harmful form of polydactyly which also causes gross deformities of the foreleg and front paws, including vestigial, absent or deformed leg bones and flipper-like forelimbs. No attempt has yet been made to eliminate these harmful effects and they represent the darker side of cat breeding (See Polydactyl Cats). Occasional tailless cats suggest a Manx-type mutation in which case the breeding line contains semi-lethal genes as well as genes for gross deformity.
The current trend to developing hybrid domestic breeds by crossing domestic cats to wild species may have contributed the bobtailed trait to one or two American cat breeds. In the American Bobtail the evidence is anecdotal and based purely on the cat's appearance, in the PixieBob it is unconfirmed. In the following, the hybridization is deliberate.
The next group of bobtails are the "Lynx" group of bobcat hybrids developed during the 1990s. Although it is speculated that the Desert Lynx breed group (American Lynx, Desert Lynx, Highland Lynx, Alpine Lynx and Mohave Bob) is derived from bobcat crosses DNA testing has not confirmed bobcat ancestry. The American Lynx is a bobtailed spotted shorthair originally claimed to be derived from bobcat/domestic crosses. The Desert Lynx is speculatively a Manx/bobcat hybrid (some early advertisements stated Abyssinian/bobcat) which once claimed 12.5% bobcat (i.e. bobcat within last 3 generations) and the tail types range from absent, through rumpy-riser to hock length. The Desert Lynx comes in shorthaired and semi-longhaired varieties. The Highland Lynx was said to be a hybrid of Jungle cat hybrids and bobcats. The Alpine Lynx is a white version of the Highland Lynx. The Mohave Bob breed is a rexed Desert Lynx developed by crossing Desert Lynx with Selkirk Rex to introduced the rexed coat type. Mohave Bobs, Highland Lynx, Desert Lynx, and Alpine Lynx may be bred together, with offspring registered according to their appearance. Solid white offspring with either straight or curled coats and either straight or curled ears are considered Alpine Lynx. Non-white kittens with rexed coats are considered Mohave Bobs. Non-white kittens with straight coats and straight ears are considered Desert Lynx. Non-white kittens with straight coats and curled ears are considered Highland Lynx.
In spite of their bobcatty appearance, DNA testing of cats within the Desert Lynx grouping has not found bobcat markers and these cats are considered to be wholly domestic cats by registries.
The Jungle Bob is a mix of F chaus (Jungle Cat) and PixieBob producing Jungle Cat type cat with a bobbed tail. There are also alleged Bengal/Bobcat hybrids in existence.
As well as the main groupings of bobtailed cats, the trait crops up elsewhere through random mutation. In the 1980s a "Spanish Bobtail" was reported, though the exact location in Spain was not given. In 2004 I received information on a localised strain of bobtailed cats in one part of Spain. According to Russell Meyers, a bobtail mutation has become fixed in cats around the Spanish villages of La Drova and Barx in the mountains in the Gandia area near Valencia. There he encountered two kittens with deformed tails. one, a male called Stitch, had a half-length tail that bent back upon itself. The other, Milo, had a knobbly bit at the end of her tail. The trait is known in the area as "Barx tail", Barx being a neighbouring village . Meyers came across a large number of cats in the area with similar tails. The most likely scenario is that a tomcat carrying the mutation fathered most of the kittens during one or more breeding seasons. He was not necessarily the cat in whom the mutation originally occurred, but by fathering the majority of kittens his genes - including the mutation - would have become widespread and the trait would have shown up in later generations through inbreeding.
In 2006, TICA proposed to clamp down on certain breeding trends including new bobtailed breeds created by crossing existing breeds together. Their Genetics Committee report stated: "The Committee proposes that TICA does not accept any proposed breeds for Registration Only status that do not exhibit novel mutations. The current mutations would be reserved for currently recognized breeds exclusively. This would end the seemingly endless applications for "munchkinized" new breeds, and then deter the inevitable introduction of "rexed", "Bob-tailed" and Poly-ed" everything else."
A bobtail breed in development is the Tennessee Bobtail. A description of its origins and type is provided by Patty Shane. Scattered throughout the Southern USA are a variety of bobtailed cats. The more common type is the Manx-type with tails ranging from rumpy to full and bodies ranging from the cobbier high-rumped Manx-type to a more svelte build as a result of random breeding. Some have ear tufts. The less common type is the Oriental (Japanese) Bobtail type. These lose the distinctive bobbed tails when out-crossed to other cats, although the kittens may have kinks or curves in the tail. The bobtail shows up when they are bred to each other. They are long, tall cats with more Oriental features and conformation and level backs in spite of their long back legs. When bred together, these create a long, tall cat, with some individuals having a broader, heavier frame and others leaning towards one or other of the parental types. The goal is a long, tall 6lb - 15 lb cat with the high, angled hind-legs of the Oriental type. The head should be apple or pear shaped, the ears straight and moderate sized (with tufts permitted) and the eyes almond (preferred) or lemon shaped in any colour. Tails range from tailless to full tailed and included bobs and single or multiple kinks or curves. A natural bobtail with kink(s) and/or curve(s) is preferred over tailless or long-tailed cats (the standard prohibits the docking of long tailed kittens). They come in any fur-length, colour and pattern with tabby/spotted preferred. Hairless, curly or wirehair is not allowed.
Two Tennessee Bobtails - Siam and Miss Hissy. Although tabby/spotted is preferred, the colourpoint pattern also occurs. Photos copyright Patty Shane.
At present, foundation Tennessee Bobtails are being registered with REFR. These are random-bred natural bobtail cats that meet the desired type. Cats meeting the Tennessee Bobtail type and belonging to a known bobtailed breed are being used to help establish bloodlines. Where a Manx-type cat is used, it is outcrossed to another type of bobtail to prevent Manx syndrome (skeletal and neural tube defects) as a result of Manx-Manx matings. I don't yet know how this breed will be visibly distinct from the American Bobtail.
In 2007, Vasilis Lekkas from Athens (who is deeply involved in the recently formed Greek network for the preservation of domestic livestock including cats) reported One complete bobtail and four cases of shortened tails, about half the normal size, where the tail ends were blunt and thick rather than tapering as in regular cats.
In 2011 a colony in Canada was report to have a short-tailed gene not passed on as a dominant. Some lines of Bengal have had issues with both spina bifida and with spontaneously producing bobtailed/corkscrew-tailed kittens accompanied by other defects including still-born kittens with intestines outside the body. Less affected cats tended to have a lump or bump at the base of the tail. Whether this defect is akin to recessive gene found in the colony in Canada, or to an incomplete dominant,had not been established.
CURLY TAILED CATS
Curly tailed cats have occurred throughout history and around the world though only recently has any interest been shown in perpetuating the trait in the form of the Ringtailed Sing -a- Ling (which became the American Ringtail in 2004). Many of the 1990s reports of curious tails confuse curly tails with bobtails. Some suggested that the cats are "longtailed bobtails" i.e. have a kinked section of tail close to the body, combined with a full-length tail with normal vertebrae beyond the kinked section; the kink would lift the tail upwards and the normal section would hang over the back or flank. In "Variation of Animals and Plants Under Domestication" Charles Darwin wrote: " I have seen a cat which always carried its tail flat on its back when pleased."
Although judges examining the early American Ringtails had reported the trait to be very rare, this is not strictly correct. Judges see pedigree cats and pedigree cats are bred for uniformity and adherence to a standard. This means rogue genes have been eliminated generations ago. The genes for curly tails would have been weeded out of pedigree lines. In addition, a curly tail would be a deviation from the standard (and might be considered a sign of genetic impurity in the breed) hence the cats would not be entered in shows, except possibly in household pet classes. The trait is found with greater frequency in random breeding cats but until recently was seen as no more than a charming oddity. I was surprised to be told that the trait is so rare as I had already encountered several curly tailed cats and kittens (nicknamed pigtailed kittens) in Chelmsford, Essex, England. Their curly tails were quite distinct from the kink found in bobtails.
In the Jan/Feb 1998 issue of the Cats Protection League magazine "The Cat" there is an account of further curly tailed kittens from Basildon, a town only about 13 miles from Chelmsford:- "At the time of writing, our latest little problems are four nine-week old kittens who have curly tails. Our vet has advised us that their tails are deformed and not growing properly and will have to be removed when the kittens are a little bit older. Our vets have said that once their tails are removed, they will be able to live perfectly normal lives, although they will look like little Manx cats."
The report would have been written in November/December 1997 to make the Jan/Feb 1998 edition. The accompanying photo showed the kittens sitting on a cushion; their curled tails were not visible. A follow up report in a later issue merely noted that the kittens' tails had been removed and the kittens had found homes. The only other recorded case of a whole litter of curly tailed kittens was a case reported to Sue Manley (who is establishing a curly tailed breed) and was a litter born to a barn cat in Oregon.
Ida Mellen's 1940 discourse on oddities in cats included a picture of a curly tailed cat from Jenkintown, Pennsylvania. This ginger and white cat's long tail clearly forms a tight spiral. Phyllis Lauder, writing in 1981 about Shorthaired cats, noted that an ACA directive stressed that the tail shall not be carried over the back: "This last is not a common fault, but it certainly looks strange when it occurs; there is a Siamese neutered pet who carries his tail as would a Pekinese dog, and the effect is to make people laugh!" Several curly tailed cats have been reported in random-bred cats in Britain in the 1990s and early 2000s. A curly-tailed tabby and white male cat was reported in Perthshire, Scotland in 1986. The degree of curl ranges from loose through to a tight corkscrew. In one case, the tail curled at the base and for the rest of its length lay flat along the spine.
In the 1980's, Katrina Lee of Washington, DC, vacationed on the island of Guadeloupe . In a small restaurant on the lower island she saw about half a dozen various colored cats with corkscrew tails. The tails were not looped over, but stuck out from the body like a normal cat's tail while looking like they'd been wrapped around something to make them spiral!
In Britain, one curly-tailed cat has achieved fame with South Ribble Pet Cat Club. Raffles is a white semi-longhair born in June 1998 and homed via a cat shelter in Oldham. When sitting, his tail forms a curl behind him. Another British curly tail is Sprocket found at Spaghetti Junction (Gravelly Hill motorway interchange in the Midlands). She was taken to Erdington Cat Rescue and it seemed that some prospective adopters didn't want her because she had a curly tail and it was perceived as a deformity or health problem.
Since the breeding programme for the American Ringtail began, reports of curly tailed cats have come in from around the world, indicating that it is not as rare as previously claimed and that there might be several different genes involved. It is now believed to be a polygenic trait i.e. several genes interact to dictate the type of curl and the degree of curling . Though the trait seems to be polygenic, some curly tailed males pass the trait onto more than 50% of the offspring while females seem to pass it on to a lesser degree. This suggests sex-linkage. There might be several different gene mutations producing a visually similar effect.
Unlike the bobtail where the vertebrae are fused into permanent kink and motion is limited, curly tailed cats have mobile tails and no fusion of vertebrae. The trait is hereditary, possibly as a result of several interacting genes which govern the type and degree of curl, and not a birth defect as once thought. Generally, the tail curls up and over in an arc or full circle, coming to lie against one or other flank as the cat walks. Owners of curly-tailed cats are often intrigued as to whether similar cats are "left-handed" (the tail tip on the left flank) or "right-handed". Sue Manley, breeder of American Ringtails (formerly Ringtailed Sing-a-Lings) produced another unusual tail alongside the normal ringtails. She described it as "baboon tail". The cat holds it curled up in a compete ring underneath him while he stalks around or hunts his toys. He then brings it up over his back when he is petted.
One peculiar curly tail was seen in a cat brought to a cat shelter in Chelmsford, Essex, England in 1999 or 2000. This was a black female cat whose tail curled up over her back and then lay flat along her spine. It gave her the appearance of a Rhodesian Ridgeback dog. The tail was mobile, but returned to that position when she was at rest. Over the years several "pig-tailed" kittens have turned up in the area, either as spontaneous mutations or due to genes in local feral colonies. The occurrence of several curly-tailed cats and pig-tailed kittens around the Chelmsford and Basildon areas in a six year time frame suggests a curly-tail hotspot in that part of Essex.
Between 2000 and 2002 there were intermittent reports of curly-tailed cats and also of people seeking curly-tailed cats as companions for, to replace , an earlier curly-tailed cat. Most reports came from the USA (sadly from unidentified locations). These included a grey tabby female with a curly tail, and a grey shorthair female with a tail that curled over to rest on her back. The latter cat's tail was described as spring-loaded, quickly returning to its curled position if the tail is straightened by hand during petting. This led to a suggestion that the trait was associated with grey cats.
A red tabby male from Missouri was reported to have a cute pigtail generally held curled up and over his back in a large perfect loop. His littermates had normal tails. His tail sometimes unwound to lie flat on his back or was held in a more conventional unwound posture during play. As the cat aged, the tail was held curled more often, suggesting a gradual loss of flexibility. The owner stressed that this caused the cat no discomfort. A cream tabby female developed her curly tail relatively late in life. She apparently had a near normal tail until she reached 8 or 9 months old, after which it became progressively more curled, forming an up-and-over curl with the tip resting on her left flank. This again suggested a loss of flexibility as the cat aged.
The majority of curly-tailed cats are random-bred cats since purebred cats have necessarily lost a degree of genetic diversity in order to produce consistent conformation. In 1997/98 a curly-tailed Turkish Van was reported. A Singapura with an unusual tail was reported in 2002. The tail was described as very flexible and he could curl it round. It is not like American Ringtails, as it does not curl as much, nor does it curl up and over. It is held low and curled in a loop at his side. An American Keuda with a similar tail ("Curlietail") was reported at around the same time. She double curled the tail when she sat down, but at 5 months old she does not curl it as much. Again, it would appear that flexibility is lost as the cat matures. Since the establishment of a breeding programme, it seems that curly tails of varying degrees are more frequent that initially suspected. What was once seen as a fault now has a serious following. Curled tails have also turned up in Russian Blues, Siamese/Orientals, Ocicats, Persians, Ragdolls, Scottish Fold, Devon Rex and Bengals (interestingly, several of those breeds have Siamese blood in their ancestry).
The photos above, very clearly showing the curly tail trait, are of Barbara Clark's 4 year old curly-tailed cat "Bunzy". His brother has a normal straight tail. Bunzy and his brother were adopted from a cat shelter in Utah, USA at 8 weeks old so nothing is known of their family history. Bunzy's tail clearly loops over to rest on his left flank with the tail tip pointing outwards.
A gallery of curly-tailed cats sent in by visitors to Messybeast can be seen at Curly Tailed Cats (a separate page has been created to improve loading times).
WHITE RINGTAIL
It should be noted that historical reports of the "White Ringtail" cat refer to the Turkish Van with its ringed or banded tail, not to a race of curly tailed cats. Some early mentions of curly tailed cats will have referred to the bobtailed cats common in parts of Asia. Also, very obese cats may hold their tails in a curled tail position for the simple fact that rolls of fat at the base of the tail prevent it from reaching any point below horizontal.
LONGHAIRED CATS
Copyright 2002-2015, Sarah Hartwell
The gene for longhair in cats is recessive i.e. a needs 2 copies of the gene in order to have longhair. A cat with one copy of the gene is shorthaired, but can pass the longhaired trait on to its offspring - the carried gene will only come to light if the cat is bred to another carrier of the gene or to a longhaired cat.
Remains of early cats can't tell us about fur length, but cats depicted in early Egyptian art are shorthairs and the parent species of the modern domestic cat, the African Wild Cat, is shorthaired. The shorthair coat ranges from plush and dense as in British Shorthairs and sleek and close-lying as in the Oriental breeds, with a variety of intermediate coat types depending on actual fur length and whether or not there is a woolly undercoat. There are a number of theories as to how the longhaired domestic cat arose.
Hybrid Origins?
In the 1800s, Pallas suggested that Angora and Persian cats were descended not from Felis silvestris but from Pallas’s cat (Felis manul) which he discovered and named. This suggestion was quoted by Darwin in 1868 who wrote, "The large Angora or Persian cat is the most distinct in structure and habits of all the domestic breeds; and is believed by Pallas, but on no distinct evidence, to be descended from the Felis manul of middle Asia.". There is anecdotal evidence that Pallas cats can interbreed with domestic cats to produce offspring, but to introduce the gene, the hybrid offspring would have to be fertile and be bred back to domestic cats. In 1907, Pocock described the various English domestic cats for Royal Zoological Society and strongly refuted the Pallas cat theory since the skull of Pallas’s cat differed from that of the Angora or Persian of Pocock's time. Modern genetic studies have also shown that the Pallas cat did not contriubte to the gener pool of the modern domestic cat.
There are claims that the Persian is descended from the Sand cat (F margarita) based on that fact that both have long hair covering the paws which forms a pad over the soles. Measurements of the length of the hair on the body and feet of both Persians and Sand cats does not uphold this claim. In addition, the long hair on the feet of modern Persians is due to their long coat; in the Sand cat it is a feature since the cats are otherwise shorthaired. Again, there is no genetic evidence of Sand Cats contributing to the domestic cat gene pool (although hybrids have been bred more recently.
Single Origin or Multiple Mutations?
A more plausible explanation is a gene mutation in a group of cats with the foreign conformation. Inbreeding would have allowed the trait to become fixed. This mutation may have originated in Asia Minor and in a book published in 1876, long-haired cats were referred to as Asiatic cats.
From historical writings, the longhair mutation either arose in three separate areas (Russia, Persia ( Iran ) and Turkey), or that the longhair mutation originally occurred in Russia and that Russian Longhairs (Siberian, Russian Angora) spread from Russia into Turkey (becoming Angoras), Persia (becoming Persians) and surrounding countries and south east Asia (introducing the gene into native cats to create Longhair Japanese Bobtails) by land and sea trade routes, retaining many of their cold -climate adaptations: a heavier body, dense undercoat and coarser hair. If so, all longhairs are derived from the Siberian cat. Yet another view is that long hair arose and was developed in Turkey and transported on land and sea trade routes to Europe, the Middle East and Far East. Much of this is based on the supposed origins of the modern Persian breed which was derived from Turkish Angoras, Russian Longhairs and purported Persian cats. Long fur in domestic cats appears to be an adaptation to cold as the paws do not have long fur underneath; in desert species of wildcat long hair is found on the paws and protects against hot surfaces.
It is possible that the longhair mutation has occurred not once, but many times and that it is still occurs. Some genes are mutational hot-spots e.g. identical mutations for curly hair and for bobbed tails have occurred (and still do) independently in differently locations. Parallel evolution means that similar cold environments could lead unrelated cat populations to evolve similar traits through natural selection in a relatively short period of time. Long hair mutations are seen in other cat species e.g. the cheetah has produced semi-longhair "woolly cheetah" variants in the past.
Longhairs range from "woolly" usually due to the undercoat) to fine and silky and the length itself varies from the flowing coat of the modern Persian to the semi-long coat of Turkish breeds. Semi-longhaired variants in normally short-haired breeds are generally blamed on recessive genes, but could be due to more recent mutations.
The first longhairs recognised by the western cat fancies were Angoras and Persians whose histories are intertwined. The Persian group is defined as all varieties developed from cats imported from Turkey, Persia (now Iran), Afghanistan and Russia at the turn of the 18th and 19th centuries. Brian Vesey-Fitzgerald "Cats" (1958) mentioned the belief, held by some, that a female Black Persian was imported direct from the Shah of Persia himself.
EARLY LONGHAIRED CATS: ANGORAS AND PERSIANS
Longhaired cats were first seen in Europe in the 1500s, first in Italy (1521) and then in France. They were named after the Turkish city of Angora (Ankhara). The first documented ancestors of the Persian were imported from Persia into Italy in 1620 by Pietro della Valle, and from Turkey into France by Nicholas-Claude Fabri de Peiresc at around the same time. From France they soon reached Britain. The longhaired cats from Persia were interbred with Turkish Angoras. A tabby Angora cat is depicted in Buffon’s "Natural History" (1756). Its head length and conformation are generally similar to the Turkish Angora except that its ears are set lower on the head.
The first longhairs in Britain were variously described as Angoras or as French cats (the latter generally being white). They were also known as Chinese (some longhairs were imported from China where they were known as four-ear cats due to the furnishings of the ear), Russian (the Russian Longhair or Russian Angora was known) and Indian. It was the existence of longhairs in China which led some to speculate that the longhair trait entered the domestic cat population through hybridisation with the Pallas cat. It is suggested that longhairs reached China from Iran (Persia), as a gift of the king of Persia.
The eighteenth-century French naturalist the Comte de Buffon quoted the 16th/17th Century Italian traveller Pietro della Valle: ‘In Europe there is a species of cats which properly belong to the province of Chorazan the USSR and Afghanistan. Their beauty consists in the colour of their hair, which is grey, and uniformly the same over the whole body, except that it is darker on the back and head, and clearer on the breast and belly, where it approaches to whiteness. Besides, the hair is fine, shining, soft as silk , and so long, that, though not frizzled, it forms ringlets in some parts, and particularly under the throat. The most beautiful part of the body is the tail, which is very long, and covered with hair five or six inches in length. They extend and turn it upon their back, like the squirrel, the point resembling a plume of feathers. They are very tame; and the Portuguese have brought them from Persia into India.’ From this description it appears, that the Persian cats resemble, in colour, those we call Chartreux cats, and that, except in colour, have a perfect resemblance to the cat of Angora.
Buffon had not seen these "Persian" cats himself, but he was familiar with the Angora in France and he apparently believed there was no difference between an Angora and a Persian except in colour (the cat was possibly a black smoke or blue smoke Angora). He named them "Catus Angorensis".
An apparently now extinct longhaired cat from China was known as the Sumxu, a fold-eared cat which was reported from the area around Peking, China. It was a longhaired cat with white fur and pendulous ears. It was described (mostly with regard to its folded ears) several times in the 1700s, with the final report being in 1938; though evidently extinct it indicates the presence of longhairs in China.
In the late 18th century, longhair cats with coarser, denser coats, and a stockier build were imported into Britain from Persia, Afghanistan and Russia. Cats imported from Turkey were mostly whites with a short, soft, silky top coat and little undercoat (and no woolliness in the undercoat). Those from Russia, Afghanistan and Persia were mostly black or blue and less foreign in type. The Angora has a ruff, breeches and a plumy tail, but unlike the later Persian, its coat followed the lines of its body due to the absence of woolly "padding" beneath. The Russian Angoras had green rather than blue, eyes.
A letter from M Lottin de la Val, President of the Imperial Acclimatation Society, to the President of the French Zoological Society in 1856 stated, "When you recently did me the honour of calling on me, you imparted the recently held view that the so called 'Angora' cat does not exist or could not exist except in the vicinity of ancient Ancyra. I hasten to dispel this illusion. I myself came upon specimens of that lovely feline species in the great Armenian plateau, at Erzerum, where the climate is greatly different from that of Angora. The species is very numerous at Mourch in Kurdistan, where it is the dominant variety. I also found it at Billis and in the pashalik of Bayazit.. The finest specimens, however, which I saw belonged to the Archbishop of Van, a town in the east of Kurdistan, on the frontier of Azerbaidjan. He had three of them, one pearl grey, one orange-hued with black and white flecks, and a third, which was completely white. Their fur was magnificent, though there was thought to be nothing to be surprised at in them, as such cats are common in Kurdistan. I also saw some at the residence of Khan Mahmoud, Prince of Hekiars, at Alpeit. I can not recall having seen any in Persia, though, had I thought that scientists might have been interested, I would have taken care to seek them out, busy as I was. But what will surprise you most of all is that despite the high temperatures prevailing, one should find Angora cats at Baghdad, though certainly these are not so fine as those to be found on the northern slopes of the medique and Taurus mountains, though whether the difference is due to the hot atmosphere or the hostility of the people of Baghdad, I cannot say. You will no doubt settle that point better than I could, all I can say is that the people of Baghdad are in constant warfare with their cats, maintaining, not without good reason, in my opinion, that they bring the plague, because of their fur coats and their habits."
While these longhairs might not have been appreciated in their native land, their charms were not lost on British cat fanciers. In "Our Cats" in 1889, Harrison Weir wrote of the various longhaired cats: "There are several varieties - the Russian, the Angora, the Persian, and Indian. Forty or fifty years ago they used to be called French cats, as they were mostly imported from Paris - more particularly the white, which were then very much in fashion."
 
Angoras
It was not until the mid-l9th century that cat enthusiasts distinguished between Turkish Angoras and other long-coated cats coming from Persia and Russia. According to the “Illustrated Natural History” by the Rev JG Wood (1853, 1874 ): There are many varieties of the Domestic Cat, of which the most conspicuous are the MANX CAT and the ANGOLA . [...] the Angola Cat being gorgeous in its superb clothing of long silky hair and bushy tail [...] A fine Angola Cat is as handsome an animal as can be imagined, and seems quite conscious of its own magnificence. It is a very dignified animal, and moves about with a grave solemnity that bears a great resemblance to the stately march of a full-plumed peacock conscious of admiring spectators. It is one of the largest of domestic Cats, and in its own superb manner will consume a considerable amount of food. Wood, a naturalist rather than a cat-fancier, evidently meant the Angora.
In 1868, an English writer described the Angora as "a beautiful variety with silvery hair of fine texture generally longest on the neck but also on the tail". In the early days of the cat fancy, longhaired breeds competed against each other; the Persian was preferred over the Turkish Angora and the Russian Longhair (Russian Angora), so much so that any resemblance to the Angora was later frowned upon by cat fanciers.
In 1870, the Honourable Lady Cust produced a book called "The Cat" (in which she perceptively wrote "In the present day, a love for cats appears chiefly permitted to 'elderly spinsters,' and is often even ridiculed") in which she described the longhair thus: "Of the wild cat, there is supposed to be only one species, which extends with trifling variety of colour over all parts of the world, the difference from the tame variety being more in the internal than in the external structures, its intestines being the shortest and smalles of all the quadrupeds. […] Hiertro dello Valli evidently means the Angora kind when he says 'There is in Persia a cat (particularly in the province of Choragan) of the figure and form of our ordinary ones, but infinitely more beautiful in the lustre and colour of its skin. It is of a grey blue, without mixture, and as soft and shining as silk. The tail is of great length and covered with hair six inches long, which the animal throws on its back like a squirrel.'"
Jean Bungartz described the Angora (Felis maniculata domesticus angorensis) in his 1896 book "Die Hauskatze, ihre Rassen und Varietäten" (Housecats, Their Races and Varieties) in " Illustriertes Katzenbuch" (An Illustrated Book of Cats) as the most beautiful and best known of the foreign cats and originating from high Asia. He summed up the debate over whether the longhair meant it was related to Pallas's Manul or was a housecat adapted to a cold, mountainous climate. He added that there were bluish-grey Angoras in the south Siberians. Bungartz wrote that it could not be proven whether it really came from Angora or just got that reputation because of Angora goats and rabbits from that region. He noted that white and silver (chinchilla?) were most popular, followed by blue, black, "grey-touched" (shaded silver?) and isabelline (cream) while other colour occurred through addition of other blood. He noted that crosses between Angoras and usual housecats lacked the full, rich, silky hair. It was a favourite , and very indulged, salon-cat in Bungartz's time and he described it as calm, aristocratic, attached to humans and a coddled lap- child ! He also noted the need to groom its hair to prevent matting ("a horrible, inextricable felt ball") and the need to wipe the eyes clean.
In a footnote to his lengthy notes on the Angora, Bungartz added a short statement about the Persian: "The Khorassan or Persian cat seems to be a modification of the Angora cat, their hair is somewhat more woollier and curlier, but nevertheless still especially long. The colour is dark bluish gray. In terms of beauty, she is quite close to the Angora cat, but is far rarer. "
By 1903 the Persian had replaced the Angora and by the 20th century the Turkish Angora was virtually unknown outside of Turkey. In Turkey, they were seen in many colours e.g. sarman (red tabby), teku (silver tabby),  Ankara kedi/Van kedi (odd-eyed white), brown tabby, selfs (solids), torties and tortie-and-white although some cat fanciers still claim that the white Angora is the only pure native Turkish Angora. There is a true-breeding colour variant found near Lake Van originally known in Britain as the Turkish Cat (or Turkish Swimming Cat) and later as the Turkish Van. In Turkey the name "Van kedi" (Van cat) refers to the highly prized odd-eyed white. Turkish Vans are auburn (red) and white with the colour restricted to around the ears and on the tail. They are also known to swim in shallow pools and streams.
In their study comparing the genetics of several breeds (Lipinski MJ, et al., The ascent of cat breeds: genetic evaluations of breeds and worldwide random-bred populations, Genomics (2007), the American Turkish Angora breed and the American Turkish Van breed were found to be distinct from each other. The American Turkish Van was related to Egyptian random-bred cats while the American Turkish Angora was closer to random-bred cats from Tunisia and Turkey. Lipinski suggested that cats may have spread into different regions of Turkey from different directions: into the Lake Van region from Egypt via land trade routes and into the Ankara (Angora) region from Tunisia via sea routes. Cat fancy history also refers to "Russian Angoras" suggesting a historical link between Russian and Turkish longhaired cats. Studies of Turkish Angoras maintained as a pure breed at the Ankara Zoo found that the American "Turkish Angora" is not closely related to the authentic Turkish cats. The American version was bred from Persians and other cats to "resemble" the Angora and is physically and genetically different from true Turkish cats.
 
Angoras were extinct in Britain until after World War II. In the 1950s and 1960s, Turkish cats were taken to North America, Sweden and Britain, but only the Van cats were bred and recognised in Britain. In the US, white Turkish Angoras were recognised in the early 1970s and other colours in 1978. They are now recognised in a full range of colours including black, blue, tabbies, tortie-and-white and smoke colours. Turkish Angoras have silky coats and gentle dispositions.
Meanwhile , in Britain the name "Angora" was given to an impostor! In 1977, a breed confusingly known as the Angora was recognised in Britain. Although similar to the Turkish Angora, it is bred in a wider range of colours (including chocolate and lilac, which come from Siamese ancestry), its voice is similar to that of the Siamese and it is more fecund than the Turkish Angora. The British Angora is a Foreign Longhair bred to recreate or resemble the Turkish breed rather than reintroduce genuine Turkish cats. It is related to the Oriental and Balinese. Elsewhere in Europe it is known as the Javanese or Mandarin ; in the USA it is a Foreign Longhair. The British Angora is more foreign in type with a longer, narrower head and larger ears than the Turkish Angora.
This situation was rectified in 2002 when Britain's GCCF came into line with other registries and called the British variety Oriental Longhairs, thus removing confusion with the original Turkish Angora cats.
The possibility of importing Turkish Angoras was apparently considered, but bureaucracy prevailed since the British GCCF would apparently not accept the documentary evidence (verification of breed) supplied by the Ankhara Zoo. If true, then the British cat fancy places more importance of paperwork than on actual physical felines; a serious drawback where naturally occurring varieties are concerned! Importation of either Turkish or American lines of Turkish Angora would mean 6 months quarantine.
Breeders of the British Angora have attempted to recreate the characteristics of the original Turkish cats. Those working with Turkish Angoras/Vans have worked to preserve characteristics of those naturally occurring breeds and the Turkish Angora has prior claim to the Angora name. Luckily the genuine Turkish Angora has since returned to Britain.
The most obvious differences between the Turkish Angora, British Angora and Persian are the coat, head shape and general conformation. The British Angora is an oriental cat - long-bodied with a longer, wedgier head and larger ears. The Turkish Angora is less foreign in build, with a wider, shorter head and smaller ears. The Persian has a wide head, flat face, small-ears and extremely cobby build. In addition, the Persian has longer fur with a woolly undercoat and a tendency to tangle or knot. The Angora coat is silkier without the woolly undercoat; the full Angora coat is seen in the winter - in summer, they moult and may look like shorthairs with fluffy tails.
In an article in the Colourpoint, Rex-coated and AOV Club's Journal (reported in 1981 by cat breeder Phyllis Lauder), Robin Sims made the point that though the terms "Persian" and "Angora" were often used interchangeably, the cats concerned were not the same and that the flowing coats of the cats imported into Europe from Iran were not governed by the same gene as that concerned with the cats imported from Ankara. Sims made it clear that the Turkish Van were Angoras, and held the opinion that it was unfortunate that in the early 1900s the Angora was used to improve the coats of the Persians.
Persians
In 1868 Charles Ross described the Persian as "a variety with hair very long and very silky, perhaps more than the Cat of Angora; it is however differently coloured." The cats most likely to win prizes at early cat shows were those which combined the Angora's longer fur with the Persian's shorter faces. The Turkish blue-eyed white cats were crossed with shorter-nosed blacks, blues and fawns (creams). The whites became shorter-faced, but the distinctive blue eyes were often replaced by orange eyes or odd eyes. Another reason for cross-breeding might have been to tackle the problem of deafness in blue-eyed white Angoras.
In the 19th century, Harrison Weir described Persians in white, black, blue, grey, red and "any other" self colour; tabbies were recognised in brown, blue (black markings on blue background, possibly black-silver tabbies), silver and light grey-and-white (probably blue-silver tabbies). In 1872 Weir described smoke Persians, "a beauty was shown at Brighton, which was white with black tips to the hair, the white being scarcely visible unless the hair was parted". The following year these were numerous enough to have their own classes. The British National Cat Show of 1879 included a Persian with a coat of "strangely graduated grey" (possibly chinchilla or shaded silver). Chocolates, lilacs and tortoiseshells were not described. Creams, then called fawns, or derisively as "spoiled oranges", were first recorded in 1890 and were darker in colour than they are today. The first Smoke Persian Champion (1893) was Mrs H. V. James's Backwell Jogram and it is worth noting that he is completely different to the modern Smoke Persians and (judging from photos) bears a striking resemblance of type to the modern Siberian Longhair.
In her 1903 "The Book of the Cat", Frances Simpson did not distinguish between the Persian and Angora and did not mention the Russian Longhair. She wrote in "In classing all long-haired cats as Persians I may be wrong, but the distinctions, apparently with hardly any difference, between Angoras and Persians are of so fine a nature that I must be pardoned if I ignore the class of cat commonly called Angora, which seems gradually to have disappeared from our midst. Certainly there is no special classification given for Angoras, and in response to many inquiries from animal fanciers I have never been able to obtain any definite information as to the difference between a Persian and an Angora cat. Mr Harrison Weir, in his book on cats, states that the Angora differs somewhat from the Persian in that the head is rather smaller and ears larger, fur more silky with a tendency to woolliness."
She considered the Persian to be less reliable in temperament than the English short-hairs, but considered them more intelligent and also keen hunters. However, they were less healthy than short-hairs and the longest haired kittens were the most difficult to rear. She attributed these health problems to in-breeding. Persians were disadvantaged by the fact that cat shows were held in the summer months (many were held in marquees), at a time when long-haired cats were moulting and tended to look moth -eaten and unkempt. This, she felt, held them back - not that the numbers shown upheld this opinion, long-hairs outnumbered English short-hairs by about four to one!
According to John Jennings "Domestic or Fancy Cats", published a little earlier than Simpson's work, "Of the many varieties or breeds of the cat with which we are now familiar, it must be remembered that, however crossed, selected, re-crossed, domesticated, or what not, we have but two breeds on which the super -structure of what is known today as the 'classification of varieties' has been reared - viz, the long-hair or Eastern cat, and the short-hair or European." Jennings found little difference in the skull length between those long-hairs and short-hairs; indicating that the Victorian long-hairs had not yet become the snub-nosed cats familiar to 21st Century cat lovers.
When Weir drew up his "points of excellence" for judging cats in 1889 he defined differences between the Angora and the Persian. Pocock later stated (1907) that Persians had a marked shortening and widening of the face i.e. it had diverged from the Angora. In addition, the Angora had sleeker fur, lacking a woolly undercoat. By 1903, Simpson had effectively dismissed the Angora cat in favour of the Persian type. Only in very recent times has the true Turkish Angora come back from being thus relegated in Britain.
 
By 1901 the colours were black, white, blue, orange (red selfs, red tabbies), cream or fawn, sable, smoke, tabby, spotted, chinchilla, tortoiseshell, bicolour and tricolour (tortie-and-white). True spotted Persians were described as being rare or non-existent; almost all had some tabby lines and rings. In fact the long flowing coat of the Persian does not lend itself well to spotted or mackerel tabby patterns. The sables were described in a 1903 book as "a kind of brown tabby ... These cats have not the regular tabby markings, but the two colours are blended one with another, the lighter sable tone predominating" which suggests shaded goldens. The orange (red self) was always popular and by 1915 there were classes for red self or shaded and for red tabbies. Blue-creams were recognised in 1929.
Persians arrived in North America from Europe towards the end of the 19th century and by the turn of the century they were eclipsing the native Maine Coon. Frances Simpson was an early champion of Persians and devoted considerable space in her 1903 book "The Book Of The Cat" to Persians "distinguished by unusually long coats, round heads, tiny ears, and wonderful toe tufts" (she also included the Maine Cat in her book as we shall see later on).
Simpson wrote "A gentleman who has lived for ten years in Assam says that he never saw in that part of India any long-haired cats except blue-eyed whites" although these seemed mainly to belong to English colonials, rather than being native cats. At around the same time, Mrs. Clinton Locke , president of the Beresford Cat Club, wrote in a letter to "Our Cats" "The first white Persian I ever owned was brought to me many years ago from Persia by a distinguished traveller, and its eyes were amber".
The White Persian was especially popular: "There is always a keen demand for white kittens, either as pretty pets or, if with correct-coloured eyes, for breeding purposes, and, doubtless, when more encouragement is given to this beautiful variety, there will be an increase of fanciers of the white cat, whose praises have been sung in fairy tales, nursery rhymes, and by novelists who have a weakness for describing interiors with a beautiful white Persian cat reclining on the hearthrug."
"Whereas formerly blue eyes were considered quite a rarity, now it is seldom we see any yellow-eyed white cats exhibited at our principal shows [...] It is easy to tell whether the baby blue eyes are likely to retain their colour or turn yellow. If at about three weeks or a month old the blue becomes tinted with green, then surely but sadly may we make up our minds that these kittens have not a distinguished career before them, for they will see and be seen with yellow eyes." A well-known authority on cats, wrote in a letter to one of the cat papers, "A few years ago white cats with green or yellow eyes frequently were prize-winners, and a blue-eyed white was looked upon as a rarity. Now blue eyes have it all their own way." Occasionally green-eyed White Persians turned up, hinting at Russian Angora ancestry.
Imported longhairs (probably then of Angora type) were considered better quality than British-bred cats: "the most perfect type of a white Persian is assuredly to be found amongst the imported cats; there is a certain beauty of form and silkiness of fur which is not possessed by the specimens bred in this country [...] These imported cats are often of a rather savage disposition, and, although they can be sweet -tempered enough with human beings, they are extremely fiery with their fellows."
Angora cat by J B Huet, 1808 " Collection des Mammiferes de Museum d'Histoire Naturelle" (Paris)
Weir's idealised longhair, 1889
In 1906 a Canadian writer described the conformation of one Persian thus: "His head is magnificent, and he is short on the leg, has plenty of bone" and the best Persians were reckoned to come from Britain. In the 1960s, Fernand Mery wrote that the development of longhairs into extremely cobby cats was due to either the British climate or was a British preoccupation since the British had done the same thing with the Chow dog breed.
Persians were also favourite photographic subjects as Sydney W France's noted (with some dismay) in his book "Siamese Cats" (1949) though he was good enough to list the various colours of Persian: "Take a look at any cat magazine, or at any cat photographs you may see in the papers or periodicals. Mostly of the long-haired cats, are they not? Chinchillas, Blue Persians, Biscuit, Smoke, White Persians." The "Biscuit" colour sounds to be cream or fawn. Grace Cox-Ife, author of "Questions Answered About Cats" (1947) lists the official Persian colours as Black, White (blue-eyed, orange-eyed), Blue, Red Self, Cream, Smoke, Silver Tabby, Brown Tabby, Red Tabby, Chinchilla, Tortoiseshell, Tortoiseshell-and-White and Blue-Cream.
In a study comparing the genetics of several breeds (Lipinski MJ, et al., The ascent of cat breeds: genetic evaluations of breeds and worldwide random-bred populations, Genomics (2007), the Persian breed was more closely related to random bred cats of Western Europe/America than to random-bred cats from the Near East. Their study also found the Persian to be derived from multiple lineages. This accords with breed histories describing it as derived from Turkish, Russian and British longhaired cats. Although Lipinski's team were surprised by the result, it did not come as a surprise to those versed in Persian breed history!
The modern Persian has evolved to have longer hair, a denser undercoat, a wider, flatter face, shorter muzzle, smaller wide-set ears, shorter body and tail and large round eyes. The lighter-bodied type of the early Angoras was lost and the modern Persian bears little or no resemblance to the original imports. However, the early ancestors of the Persian are once more being recognised in the form of the Turkish Angora and the Siberian.
In Britain, the cats became known as Longhairs, though most still call them Persians or Persian Longhairs to distinguish them from other long-haired breeds. In the USA, Persian is the official breed name although some American associations do not recognise the chocolate or lilac solid (self) colours introduced via Himalayan (Colourpoint Longhair) breeding programmes; these are known as Kashmirs. In some American registries, non-colourpointed cats of Himalayan origin are called the Himalayan Reflections or Self Himalayan. Thankfully some have seen fit to sort out the muddle so that Persians and Himalayans can be interbred and some offspring are known as Colourpoint Carriers (CPCs). The history of the Himalayan is in Colourpointed and Masked Cats.
A 1950s colourpoint longhair and the 1990s equivalent
The British system had its own perversities - each variety of Longhair being regarded as a separate breed, not as a colour variation of a single breed. The Chinchilla is therefore seen as a different breed to the Blue Longhair! Conformation does differ across the Longhair "breeds" and some of the Chinchillas remained less ultra-typed than the selfs. Chinchillas were reckoned to be several years behind other Persians in terms of extreme-typing.
During the mid 1970s and early 1980s, American Persians changed dramatically. The pre-1980s look had heavy brows, flat-topped heads rather than the domed heads. The "sweet, open-expression" was lost as fanciers pursued the extreme head type (ultra-type). These were dubbed "pigs" or as having a "piggy expression." The nose became narrow and ultra-high; the breaks above the eyes were moving upwards into the foreheads; the eyes were tiny and the jaws often maloccluded to produce a frowning mouth. It is suggested that Peke-Face reds Persians were bred to other colour Persians to produce the piggy look and the teardrop-shaped eye of modern ultras is attriubted to the Peke-face cats. The new look was deemed an exciting development and open-ended standards favoured these cats on the showbench. During the 1990s, both open-face and Persian pigs were advertised, but the trend is now back to the healthier open-face cats.
True Peke-face Persians (solid red and red tabby) generally conformed to the red Persian standard, but only 98 had been registered with the CFA beteen 1958 and 1993 (the 3 registered between 2000 and 2002 may not have been genuine Peke-face Persians). They had higher ears and a different skull structure that produced a very round head with a strong chin and very wide-set eyes. The nose was depressed and indented between the eyes. The muzzle was wrinkled and there was a horizontal break located between the usual nose break and the top dome of the head. This second break created half-moon boning above the eyes and an additional horizontal indentation in the center of the forehead. The Standard for the Peke-Face called for a brow ridge, dimple, and a double dome. The term peke-face is often incorrectly used to describe extreme-type Persians.
 
While American breeders were the first to breed the Persian to extremes, practically eliminating the cat's muzzle, British breeders are now following suit with ultra-typed Persians which find favour on the showbench but less favour with the general public who prefer the older, less extreme facial conformation. Personally I find the ultra-typed Persian ugly and they are more prone to breathing problems and tear duct problems than those of more moderate type. In the USA, the older type is now preserved as a traditional style Persian (bizarrely the breeder does not like the actual breed name used in reviews in case it erodes, rather than promotes, the breed). For a short while, American breeders developed the Peke-Face Persian. This arose as a mutation in red/red-tabby Persians and suffered from breathing problems, tear duct problems, high palate (making suckling difficult) and sometimes the head was too massive for kittens to be born naturally (Caesarian section needed). These have now vanished, but the ultra-types are becoming almost as extreme.
In 1872, Dr Gordon Stables, in his review of cat shows, had noted that "urbanity of countenance" should not be overlooked when judging Red Tabbies; the modern equivalent would be "a pleasing expression". In 1981, Phyllis Lauder noted that the Peke-faced Persian (bred only in the USA) was a solid red or red tabby longhair (there being no Peke-faced Exotic Shorthair equivalent) with a very different expression to that of a regular Persian. Like Pekinese dogs, they had prominent eyes, sometimes described as "owl-eyed". It was debatable whether these cats (or indeed the modern ultra-types) could be said to have a pleasing expression.
The Persian is still evolving . The Mink Longhair (Mink Persian) is a Persian Longhair in the Tonkinese (mink) colour series. Similar cats have been experimentally in different countries under a variety of names and since about the 1960s. They are attractive cats, but have yet to attract a serious following or achieve formal recognition. Other names used for this variety includeBurmalayan, Himbur, Iranese, Layanese, Silkanese, Tibetane/Tibetaan and Tonkalayan (some resemble Persians, other resemble Tonkinese). Mink Persians would seem a logical and attractive development. The Napolean Cat Longhair is a short-legged Persian bred through outcrossing Persians to Munchkins in the 2000s.
To add confusion to the Persian story, there is also the Exotic Longhair. Longhaired individuals are sometimes born to Exotic Shorthair parents due to the recessive nature of the gene for longhair. In many registries, they are not accepted in the Persian class due to their non-Persian parentage. In addition they may have a relatively poor coat compared to Persians born of Persian parents. However, they are accepted as Exotic Longhairs by some registries in the USA/ The original proposed name was Tiffany , due to the original conception that the Exotic Longhair was to be allowed in silver colurs only.
Russian Longhairs
Harrison Weir, founding father of the cat fancy and organiser of the first formal cat show (also writer of the first standards for pedigree cats), wrote about the Russian Longhairs in "Our Cats" in 1889. The cat he described differs from the modern Siberian cat, for example it was woollier, although cats of the old Russian Longhair type apparently still exist among domestic longhairs in parts of Russia.
The Russian Longhair male described by Weir differed from the Angora and the Persian in many respects, being "larger in body with shorter legs. The mane or frill was very large, long, and dense, and more of a woolly texture, with coarse hairs among it; the colour was of dark tabby, though the markings were not a decided black, nor clear and distinct; the ground colour was wanting in that depth and richness possessed by the Persian, having a somewhat dull appearance. The eyes were large and prominent, of a bright orange, slightly tinted with green, the ears large by comparison, with small tufts, full of long, woolly hair, the limbs stout and short, the tail being very dissimilar, as it was short, very woolly, and thickly covered with hair the same length from the base to the tip, and much resembled in form that of the English wild cat."
According to Weir, its motion was less agile than other cats and it did not care for warmth, preferring to be outdoors in the coldest weather. Another peculiarity was that it seemed uninterested in hunting birds . Its habits were not like those of its companion shorthaired cats. It attached itself to no person, but was inseparable from a shorthaired, silver-gray tabby female. This pair produced a single black-and-white kitten. The kitten inherited the woolly coat, had somewhat of a mane and a short bushy tail. It also seemed uninterested in birds, but would attack rats!
"I have seen several Russian cats, yet never but on this occasion had the opportunity of comparing their habits and mode of life with those of the other varieties; neither have I seen any but those of a tabby colour, and they mostly of a dark brown. I am fully aware that many cross-bred cats are sold as Russian, Angora, and Persian, either between these or the shorthaired, and some of these, of course, retain in large degree the distinctive peculiarities of each breed." However to the practised eye, it was possible to distinguish whether the cat was Angora, Persian of Russian. Unfortunately, these were all judged together in a longhaired class and not as distinct breeds; something which ultimately led to the loss of the Angora and Russian types.
I have seen some 'first-cross cats' that have possessed all, or nearly all, the points requisite for that of the Angora, Persian, or Russian, while others so bred have been very deficient, perhaps showing the Angora cross only by the tail and a slight and small frill. At the same time it must be noted, that, although from time to time some excellent specimens may be so bred, it is by no means desirable to buy and use such for stock purposes, for they will in all probability 'throw back' - that is, after several generations, although allied with thoroughbred, they will possibly have a little family of quite 'short-hairs'. I have known this with rabbits, who, after breeding short-haired varieties for some time, suddenly reverted to a litter of 'longhairs'; but have not carried out the experiment with cats."
Weir noted that he had never seen tabbies among the shorthaired Russian cats, but the Russian Longhairs were all brown tabbies except for two black Russian Longhairs, which he supposed were the offspring of tabby or grey parents. He did not recall having seen any white Russian Longhairs and wrote "I should feel particularly obliged to any of my readers who could supply me with further information on this subject".
In 1926, Dr Jumaud's book "Les Races des Chats" (The Breeds of Cats), which was based largely on the works of Professor Cornevin of Lyons, described the Carthusian cat (felis catus carthusianorum) and Tobolsk cat. The Carthusian was apparently the "Maltese cat" known the the Americans, though Jumaud's description referred to a large head with large, full eyes, short nose and small, erect ears. Its coat, he said, was half long and woolly and the colour was grey with bluish reflections. However, there was another variety of Russian cat known as the Tobolsk variety: "This variety, described by Gmelin, exists in Siberia , and is sometimes called the Tobolsk cat. It is larger than our common cat, and somewhat resembles the Carthusian in shape. The head is large, with big eyes, short nose, and small erect ears. Coat: as is fitting for an animal of a cold country, the Tobolsk cat has long fur, longer than that of the Chartreuse cat. Its texture is woolly, and in colour, uniformly reddish."
In 1927, Mrs Amy Lawrence wrote "In the Natural History Museum [South Kensington, London] there is an enormous cat which is said to be a 'Russo-Persian' cat. It has an immense coat, and is similar in every way to a Persian long-hair, except that it is larger than any specimen I have ever seen. An old uncle of mine possessed what HE called a Russian cat, also a long-hair with immense coat and very large." However the only "Russian" cats Mrs Lawrence had seen at cat shows was the small short-haired Russian Blue that looked like a blue Siamese cat! Her uncle's huge Russian cat had been a tabby.She wondered "Do Blue Russians really come from Russia, and if so, then where do those immense long-hairs come from, and why were they called Russians even by Museum authorities?"
NATURAL LONGHAIRS: MAINE COON, NORWEGIAN FOREST CAT, SIBERIAN, GERMAN LONGHAIR
In hotter climes, a short sleek coat is an advantage e.g. Siamese, Burmese. In harsh conditions, longhair cats fare better - they are insulated against the cold and their outer coat may have water repellent qualities. "Refrigerator Cats" were a strain allegedly developed in 19th Century Pittsburgh to control vermin in refrigeration plants, but actually the product of a newspaper story. Natural selection supposedly produced a race of "Eskimo cats" which were at home in the cold, having heavily furred coats, thick tails like Persians and tufted, lynx-like ears. Although the idea of natural selection favouring thick fur for a harsh climate is sound, a little investigation shows that no such race of cats existed.
The American Maine Coon arose through natural selection in conditions which favoured robust, longhaired cats. Accounts suggest that some of the cats taken to America were longhairs e.g. Turkish Angoras i.e. the longhair gene was imported into America. These would have interbred with the various other cats taken there and over many generations the harsh winters would have selected in favour of longer, thicker, more protective coats.
In Europe, the same natural selection process gave rise to the Norwegian Forest Cat, the Siberian and the Rugkatt. To the casual observer these resemble the Maine Coon, but the conformation and fur type differ. A "Russian Angora" cat has been described as similar to the Turkish Angora, but with green eyes i.e. naturally occurring semi-longhaired version of Russian Shorthairs (the group comprising Russian Blues, Whites, Blacks etc).
Lipinski et al found the Norwegian Forest Cat, Persian, and Siberian each showed subdivisions within each breed, indicating multiple lineages. This is not surprising in the Norwegian and Siberian as these come from a broad base of random-bred cats that have relatively recently been recognised as a breed.
Maine Coon
The Maine Coon, or Maine Cat, is one of the oldest natural breeds of North America, and is regarded as originating from the state of Maine. Is has also been known as American Longhair, American Shag, American Forest Cat, American Snughead and Maine Trick Cat.
There are various explanations of its origin. One is that it is a racoon/cat hybrid. Another is that Marie Antoinette sent her Angora cats to safety in the USA and that these cats interbred with the shorthaired domestics. Another is that New England sailors took home Angoras from Turkey in the late 17th century. Due to its tufted ears and large size (though not as large as some media reports would have us believe), others believe that the cats descend from North American bobcats or bobcat/domestic cat hybrids or, even more implausibly, as a hybrid between domestic cats and lynx. The misconception that it is a lynx hybrid is unfortunately still perpetuated by some credulous cryptozoologists.
Most likely, it derives from a mix of longhaired and shorthaired cats taken to New England by colonists and as ships' ratters. The rugged longhaired cats of Scotland, Norway and Russia are good candidates for some of its ancestry with the addition of Persians and Angoras. In the late 18th century Maine was a major ship - building , sailing and trading state. Trading ships would have carried a variety of animals including European cats, both as pets and as ships’ ratters and mousers. The Maine Coon would have evolved from these.
Maine Coons were well-established by the early 1800s and had evolved into a hardy, handsome breed of domestic cats and excellent hunters. They had a rugged coat and build and were tough enough to withstand the harsh winters. They were also large - both tall and long-bodied. The long, flowing fur is relatively heavy and shaggy, shorter on the shoulders and longer on the belly and tail. Maine Coons also have a well-developed ruff, broad muscular chest, strongly boned legs and relatively long, square-muzzled head with slightly concave profile.
They were first recorded in cat literature in 1861 and became popular competitors at early cat shows in Boston and New York. A Maine Coon won the 1895 Madison Square Gardens show. It was described in "The Book of the Cat" in 1903. And by 1906 there were 28 registered Maine Coons, but interest was waning. The Maine Coon's popularity as show cats declined when Persians arrived and though they remained popular as pets, they were largely ignored by cat fanciers and breeders until the early 1950s. In 1967 they were recognised as a breed (1976 by the CFA).
The first Maine Coon in Europe was a pregnant female taken to Austria from Canada in 1953 or 1954 and her progeny were known in Germany as American Forest Cats. They reached Britain in 1984 and are now popular in Britain, Europe and Australia.
The are found in almost all patterns and colours; the only ones not permitted are those indicating hybridisation with colourpointed cats i.e. chocolate, lavender or Siamese-pattern. Brown tabby is the most popular colour. Polydactyly was found in early Maine Coons but was discouraged. In the late 1990s, some breeders became interested in reinstating the trait.
The early Maine Coons were documented by Frances Simpson who was an early champion of longhaired cats. In "The Book Of The Cat" (1903) she included a chapter about Maine Cats from American breeder F. R. Pierce . In childhood (1861) Pierce had owned a longhaired black-and-white, " Captain Jenks of the Horse Marines" and, like Simpson, was evidently a longhair enthusiast. Pierce did not know Jenks' ancestry, but assumed that longhairs entered Maine (a major ship-building and seafaring region) much in the same way, and at about the same time, that they reached England. The major difference was that the Maine cats were largely left to their own devices while the British cats were being selectively bred.
In 1869, Pierce saw a pair of blue-eyed white Persian kittens that landed from a foreign vessel which had put into a seaport town for repairs. These had been acquired by a sailmaker making repairs to the ship from the cook who owned a Persian female which had produced kittens. The two cats were both kept for 2 or 3 in the hope of getting a good male for neutering (as a pet); all the female kittens being destroyed! When the desired male arrived, the original pair were sent to a relative in the country. However, during those 3 years they had evidently met up with local cats since longhaired blue-eyed white kittens began to appear in unexpected places. Lack of selective breeding meant the strain generally vanished, only to reappear later on. Pierce owned ones such cat, Dot, said to be as good a specimen of Persian as the one that came from the original kittens eleven years previously.
One Mrs Thomas , also of Maine, wrote that her cat was descended from a blue-eyed white brought to Rockport, Maine on a ship from France. That line of whites, while in the same locality, was quite distinct and unrelated to Pierce's white longhairs. Pierce wrote on a little island well off the coast and inhabited by only three families, there were pure white blue-eyed Persian cats, but was apparently unable to obtain one of these cats.
Another early champion longhair was Richelieu, owned by Mr. Robinson, of Bangor, Maine in 1884. Richlieu was described as a silver or bluish tabby, very lightly marked, but rather a coarse-grained variety - "a drug store cat" (moggy). At that time Maine, near the coast, had many fine specimens of the longhaired cats, particularly brown tabbies. The Maine cats were not considered valuable at the time. From the coastal towns and cities, the longhaired cats spread inland. Around 1895/6 the "cat fad" struck the Middle West and cats from Maine were being acquired by enthusiasts inland of Maine, with considerable sums being paid even for poorer quality or mongrel kittens.
Many of the prize-winning Maine cats of the mid-to-late 1890s were described as being of Persian type. At the turn of the 20th century, smokes, silvers and chinchillas were uncommon. The most common colours were whites, blacks, blues, oranges and creams, plus tabbies. A line of creams was founded by a fine cream apparently brought from an unspecified Mediterranean port by one Captain Condon in the 1880s. The Maine cat was distributed along the coast, and for about 60 miles inland, but were not then common in the less populous northern portion of that State.
Norwegian Forest Cat
Although in some ways it resembles the Maine Coon, the Norwegian Forest Cat (Norsk Skaukatt or Skogkatt) is a Scandinavian breed which evolved in the cold northern climate of Norway. Generations of living in the cold and wet gave rise to a cat with a heavy, weather-resistant coat and full ruff. The woolly undercoat provides warmth while a medium-long, glossy outer coat resists rain and snow. It differs from the Maine Coon in several respects - including back legs slightly longer than the front. It is an excellent climber.
Longhaired cats are mentioned in Norse mythology and in books of Norwegian fairy tales written 1837 and 1852 which describe it as having a long, bushy tail. Due to its resemblance to the Maine Coon and Scottish Wildcat some have suggested the Vikings took cats to Scotland (where they interbred with wild cats) and to North America on Viking longships. This theory is unlikely. Domestic cats did not arrive in America until European colonists arrived; there is no archaeological evidence of domestic cats in supposed early Viking sites in America and reports of native cats almost certainly referred to the racoon. The Scottish Wildcat has an intractable temperament and cannot be reliably domesticated, hybrid offspring inherit this wildness.
Other theories suggest it derived from Angora cats which arrived at Norwegian ports as ships’ cats and interbred with native Norwegian shorthairs, or that Crusaders took British shorthair domestic cats and longhaired cats to Norway. There is a misconception, fostered by credulous cryptozoologists, that it is a cross between domesticated longhairs and Scottish Wildcats. Norwegian longhairs would have to swim a long way to meet up with Scottish Wildcats! TIt is not a hybrid with European Wildcats nor with European Lynxes. Crosses between domestic cats and European Wildcats are closer in type to the wildcats - wild in temperament and shorthaired. Most likely, it evolved naturally as an adaptation to harsh wintry conditions.
Breeding of the pedigree Norwegian Forest Cat from semi-wild outdoor cats and farm cats began as early as the 1930s, with the cats being exhibited in Oslo before the Second World War. In 1963 it was shown under the name Skogkatt. The breed was revived in the early 1970s; in 1972 it was formally recognised in its home country, serious breeding began in 1973.
Many of the foundation cats came from near the Swedish border. Swedish breeders declared that the breed belonged to Sweden as much as to Norway. But the Norwegians disputed this and it was agreed that foundation cats must come "straight from the Norwegian forests". While single Skogkatts might cross the Swedish/Danish border while hunting in the woods no entire (unneutered) registered cats were allowed out of Norway until the breed was recognised. It was eventually allowed to be exported and was recognised in Europe in 1977 and became popular in the USA during 1985 and Britain in 1987.
Siberian and other Russian Longhairs
In the 19th Century, Russian Longhairs were described as distinct from the distinct from the Persian or Turkish Angora; and these are described earlier in this article as they contributed to the Persian type. The Russian cats had a larger body with shorter legs, a woolly coat with coarse hairs among it, a large mane and short thickly furred tail. In Victorian times, it was extensively crossed with the Persian and the Angora and was lost as a distinct type. It continued to breed naturally in Russia as the Siberian cat. The Russian Angora is described as similar to the Turkish Angora, but with green eyes instead of blue, orange or odd eyes.
The first Siberian cats (Russian Longhairs) were imported into the USA in 1990 by Elizabeth Terrell of Louisiana, in exchange for Himalayan cats sent to establish that breed in Russia. Before 1990, it was almost unknown outside of Russia. The first colourpoint Siberians (Neva Masquerades) were imported into the USA in 1997 and registries are have recently accepted it following early controversy. The history of the Neva Masquerade is in Colourpointed and Masked Cats.
Cats would have entered Russia from Europe, by land and sea, from Persia (Iran) by overland trade routes and from the Far East on board trading ships. The existence of bobtailed Russian breeds demonstrates that cats from south east Asia reached Russia one way or another. The cats of Russia and Siberia are little changed from mediaeval times, perhaps earlier.
It is suggested that the longhair mutation occurred in Russia and spread from there to Turkey (Angoras) and Persia (Persians) or even that all long-haired breeds ultimately have their origins in Russian cats. The similarities between the Siberian and the Norwegian Forest Cat and Maine Coon suggests a common origin and some researchers suggest that the Siberian is the ancestor of both the Norwegian Forest Cat and the Maine Coon. Equally, it could be parallel evolution where similar environments have led to unrelated cat populations evolving similar traits through natural selection.
The Karel Bobtail (Karelian, Karellian) and the Kuril (Kurilian, Curilsk) are both shorthaired/semi-longhaired Russian breeds with bobtails. The American-bred Nebelung is longhaired version of the Russian Blue, developed through outcrossing Russian Blues to other cats. Longhaired Russian cats other than the Siberian exist naturally in their own country. In 1993/4, the first Nebelung in the Netherlands arrived from Russia. A purebred Russian Blue stud cat called "Timofeus" turned out to be semi-longhaired. Although not found in modern Nebelung pedigrees, Timofeus confirmed that the recessive longhair gene was present among Russian Blues. In the following years, it turned out that semi-longhaired Russian Blues were not uncommon in Russia and some have become part of Nebelung breeding programmes in other countries.
German Longhair
The German Longhair has waited a long time for international recognition although a breed standard and scale of points has existed since 1929. All that was missing until a few years ago was a registered breed that corresponded to this standard. Following the 2nd World War, which interrupted cat breeding in Germany, the only native German longhaired cat breed was considered extinct.
Longhaired cats in Germany were generically called Angoras and bred for colour, not conformation. Biologist and zoologist Professor Dr Friedrich Schwangart (1874-1958) criticized them as generally not meeting the "Hochzuchtperser" ("high-bred Persian") standard seen in British Persians, hence he created separate standards for the Persian and the German Longhairs in 1929, describing the differences between the two types. From that point, breeders of "Angoras" had to decide whether to breed British-style Persians or more natural-looking German Longhairs. Schwangart hoped the German Longhair, with its silkier "wash and wear" hair would take its place as a more natural counterpart to the Persian that had been bred in Great Britain for decades. The German Longhair was first exhibited and acknowledged nationally at the Exhibition of the Federation for Cat Breeding and Protection in 1930 in Berlin. In the following years it was frequently seen at cat shows and in 1932, German Longhair "Fox of the Rhine Castle", owned by Dr Heine in Leipzig, became a Federal winner.
With the standards laid down in 1929, the longhaired cat types fell into 2 categories in the "Classification, Pedigrees and Systems of House Cats" and these were not to be interbred in order to maintain their distinct types (in Britain, early Persians, Angoras and the native British Longhairs had been interbred to create a single Persian Longhair). There was more information in Schwangart's 1932 publication "Formation and Breeding of House Cats (Results and Problems)" which noted further longhair breeds being the German Longhair founded by Schwangart himself and, in the previous 2 years, the Burma breed that appeared in Paris (i.e. the Khmer/Birman). By the day's standards, both Persians and German Longhairs had a compact conformation, short sturdy legs, a broad head with relatively short, broad muzzle and moderately small ears (akin to the European Wildcat). Both had rather short, beautifully carried tails, level back and long fur (with age differences, seasonal coat and pregnancy to be taken into consideration). And in both cases a "half-Angora" type with svelte body or narrow, pointed face were undesirable.
However, the Persian was described as thicker-set with a rounded head and a prominent forehead that fell abruptly to a broad, short muzzle giving an "angry" expression (i.e. a shorter face). The Persian's fur was denser and woollier with a well-developed ruff and the cats were bred with cobbiness and size in mind. In contrast to the Persian, the German Longhair had a more moderate head: a less prominent, tapered forehead that curved gently up from a longer nose with a more gentle slope. The conformation was less compact, the movement more fluid and the tail longer than the Persian. In essence, German Longhair did not permit the short face and prominent forehead of the Persian and in profile the face resembled the Tabby Shorthair. The German Longhair was found in the same colours and patterns as the Persian: single/self colours (black, blue, cream, red and white), bicolours, tortoisehsells (with or without white), "masks" (colourpoints), smokes, Chinchilla (tipped), peach (goldens?), silvers and both "tiger" (mackerel) and "marble" (classic) tabbies.
In "The Formation and Breeding of House Cats (Results and Problems)" (1932) Schwangart suggested the head and face of the German Longhair showed the influence of the large Nordic form of F silvestris (European Wildcat) resulting in a native Longhair that was distinct from the Persian or Angora. The tiger pattern completed the image of a German Longhair that might trace its ancestry, in part, to a wild cat. It was already known that domestic cats and wildcats could interbreed and some still believed that local races of domestic cat had arisen independently from local wildcat species. In a last work " Overview and Description of Domestic Cat Breeds" (1954) Schwangart described the German Longhair in detail, noting the existence of intermediate forms between Persian and German Longhair which were found in some of the colours, and the need to eliminate the intermediates in order to restore the 2 breeds as distinct form each other. It's clear that the Persian had been bred together, perhaps due to the difficulties of maintaining breeds during wartime, perhaps to improve the traits of one or other breed or perhaps through ignorance that they had originally been separate breeds.
He elaborated on the breed standard, though by then he may have felt it a losing battle due to the increasing popularity of the Persian. In the solid-colour German Longhairs, amber/yellow was the preferred eye-colour, except in solid white cats where amber, blue or odd-eyes were permitted. Deafness was a disqualifying fault in white cats which were to be tested using a whistle out of the cat's sight. He also mentioned the potential for degenerative problems, such as deafness, related to "albinism" (blue-eyed white was mistaken for albinism) so some indication that the cat wasn't albino, such as a dark membrane, was desirable. The bicolour and tricolour cats were to be more colour than white. The "masked" cats were allowed to be less symmetrically marked than bi- or tri-colours. This group included the "black and yellow" tortoiseshell and the "Spanish" (tortoiseshell and white). The tortoiseshells ideally were to have large patches of colour, but Schwangart admitted that this was rare. In parti -colour cats, the eye colour was to reflect the predominating fur colour. The eye colours of the Chinchilla (black-tipped), peach colours (goldens?), smokes and silvery ones related to their coat colour (i.e. paralleling shorthairs and Persians).
In May 1935 the German Longhair was officially allowed to be bred under the auspices of the "Katzenverein des Deutschen Reiches" (Cat Club of the German Reich ), which was the only breeding club at this time. It was grouped in the longhair class together with the Persian and Birman. In October 1939 it was recognised by the Confédération Internationale Féline (CIF) as "Borealis" or "Boreali" ("Northern"). The CIF. was the predecessor of the Fédération Internationale Feline (FIFe) and had been founded by the Societa Felina Italiana, the Cat Club of Paris and the Fédération Suisse. The Second World War interrupted the breeding programme and the German Longhair stagnated for several years before apparently dying out. Dagmar Thies reported in 1979 that Mrs R Aschemeier had managed to locate German Longhairs from original bloodlines and had bred them at Blasheimer mill since 1968. These cats were considered very typy representatives of the breed and their descendents were useful in re-establishing the breed. By 2005 there were a growing number of breeders interested in preserving or recreating the German Longhair. They found foundation cats among free-ranging farm cats that were close to Schwangart's German Longhair standard. The foundation cats 5 of the remaining German Longhairs descended from Mrs Aschemeier's cats from 1968 (and thus preserving some of the genetic make-up of the original breed).
A provisional German Longhair standard was registered with the World Cat Federation in 2008 and based on the 1929 and 1954 standards. It is the only longhaired cat developed on German ground and is the longhaired "sister" of the European Shorthair breed which it resembles in general conformation. it does not have the wide muzzle of the Maine Coon or the straight nose line of the Norwegian Forest Cat. The modern standard calls for a medium-size cat with a long, rectangular, robust and supple figure. It differs from the European Shorthair/Celtic Shorthair in having a deeper chest and medium-length bushy tail that tapers to a round tip. The sturdy legs are short to medium-long with large firm paws. The head is rounded, but is longer than it is broad with medium-long and sloping nose with slight stop (a pronounced stop is a fault). Strong chin and cheeks, the latter suggesting the Nordic race of European Wildcat (F silvestris silvestris). Ears are small to medium size, upright and broad at the base with a rounded point. Eyes are round to oval, large and slightly diagonally set; the colour relates to the fur colour/pattern (or to predominant colour in parti-colour cats).
The coat is medium long at the shoulders and shorter on the head. It is longer on the flanks, back and belly and is particularly long at the ruff, hind legs (britches) and tail. However the fur is easy-maintenance, shining and not as woolly as the Persian. All colours are accepted except chocolate, cinnamon and their dilutes lilac and fawn (in both solids and in patterned cats). The colours/patterns otherwise include self/solid, bicolour, tortoiseshell, tortie-and-white, "masked", tipped, cream, red, smokes, shaded, silvers and both mackerel and classic tabbies. The personality is human-oriented.
At the end of April 2012, it was formally recognized as the " Deutsch Langhaar" (German Longhair) by the World Cat Federation (WCF) at the general assembly with a revised standard that still is nevertheless still based on Schwangart's description. The recognition comes into effect at the start of 2013.
A second longhaired German cat is the German Angora bred from semi-longhair housecats and originally bred by Mrs Aschemeier who retired from breeding in 2010 (Aschemeier always referred to her cats as German Longhair, not as Angoras, stating "Being experienced in keeping studbooks for hunting dogs from now on I kept records on the progeny of my German Longhairs, so I could provide them with pedigrees at any time"). The German Angora, bred since 2000 (breed club founded 2005), is not recognised by any cat association and is trademarked instead. Breeders of the German Angora claim there are no genuine German Longhair cats because they became extinct. They describe the German Angora as a naturally occurring longhair in the same way that the "Forest Cat" breeds are natural longhairs. Until 2007, when German Longhairs were recognised, German Angora and German Longhair were synonyms for one breed; the longhairs that began in 1929. In 2007 the Board of Directors of the first German Angora Cat Club dismissed this idea, resulting in a dispute that divided the breeders' group. The German Angora became trademarked and may only be bred in the 1st German Angora Cat Club. It has not sought international recognition. Meanwhile, the German Longhair is bred to conform to the old image. Both breeds are described as very similar and both have been crossed to old-style Persians to improve the conformation and coat.
THE IRCA LONGHAIRS - RAGDOLLS, HONEYBEARS AND CHERUBIM CATS (RAGAMUFFINS)
Baker's Cherubim Cats included Ragdolls, Miracle Ragdolls, Honey Bears, Doll Babies, Baby Dolls, Shu Schoos, Catenoids and Little Americans among the fanciful names. The origins included fanciful claims of secret government laboratories, infusions of skunk genes, human genes and/or alien DNA while a Catenoid would supposedly produce Ragdolls regardless of what you mated it to. Baker seemed increasingly unstable; she distributed photos of herself amidst a hundred dead kittens that she claimed were killed by rival breeders who broke into her home. Disturbed cat fanciers were certain she had killed them as part of a publicity bid and tried to figure how to get other cats safely away from her (I recall the pleas for information circulating on mailing lists at the time). Perhaps mercifully for the cats, Baker died in 1997. Her IRCA organisation limped along for a few years, but many breeders quit or defected to conventional registries (which had cat shows) and the IRCA trademark on Ragdolls lapsed in 2005.
Former IRCA Ragdolls, Honey Bears (IRCA's version of the Persian) and Miracle Ragdolls, not being accepted by other registries, were merged and renamed RagaMuffins and registered under that name. Although bred with the Ragdoll to begin with, the modern RagaMuffin has a shorter nose with an obvious nose dip, and the eyes are walnut-shaped rather than oval. RagaMuffin breeders have worked to produce a rounded, more heavily boned cat by outcrossing to Persians/Himalayans and have used domestic longhairs to give the breed a sound genetic footing. The plushy coat is shorter and thicker than the Ragdoll's medium-long, silky coat. Some registries only recognise non-pointed RagaMuffins (feeling pointed varieties are to similar to Ragdolls), others also recognise the pointed varieties.
Honey Bears had been bred from a pair of Persian cats and could be regidtered as Persians with the CFA. Ann Baker claimed skunk genes were infused into the female and her kittens then resembled young skunks , being born silver before turning black with stripes along the head, back or underside (which to the rest of the cat world sounds like shaded silvers developing their colour). The cats supposedly has tails flatted on the sides (actually an illusion caused by long fur!) that they held over their backs like skunks. Baker claimed the Honey Bear looked like a Persian, but did not have a cat skeleton. Essentially the cats were Persians and were even marketed as non-matting Persian lookalikes, but Baker's breeding philosophies and increasingly wild claims, led to them becoming unregistrable as Persians with conventional registries. For all practical purposes, Honey Bears were Persians, albeit less extreme in type than those seen on the showbench. A few breeders persevered with them, but otherwise they were absorbed into the RagaMuffin.
Miracle Ragdolls ("highly upgraded Ragdolls") were mentioned in some of Baker's IRCA advertisements and were also merged into RagaMuffins after her death. Her press releases became increasingly vitriolic towards the breakaway Ragdoll breeders and their "half-bred" or "overbred" copies of her cats. Catenoids and Little Americans also appeared in IRCA adverts close to the end of Baker's life. Most of the other "breeds" seemed to be products of a fertile imagination rather than distinct types of cat.
SEMI-LONGHAIRS (INTERMEDIATES)
Semi-longhair means intermediate between the thick, flowing coat of the Persian/Longhair group and the short fur of the various Shorthair breeds. By modern standards the parent of modern Persian longhairs, the Turkish Angora is a semi-longhair. Semi-Longhair cats have coats slightly shorter than the Persian, often lacking the undercoat of the Persian. They range in type from the Oriental (e.g. Balinese) to more robust types (Norwegian Forest Cat, Siberian) though all are more moderate in type than the modern Persian.
According to Phyllis Lauder, writing in "The British, European and American Shorthair Cat" (1981), longhaired cats had long existed among the non-pedigree population in the West - before Angoras and Persians had been imported from the East - though they lacked the tremendous length of the imported and carefully bred pedigree cats. The prized "fluffy cat" found in the native population was referred to by fanciers as an "intermediate" and it was the considered opinion of breeders that the exhibition longhairs were derived through crossing "intermediates" with imported Persians and Angoras. Australian geneticist Mary Batten believed that the fluffy moggy got his pretty coat from either the indigenous Scottish wildcat or from cats imported from the Middle East by the Romans (it is more likely to have come from later Viking imports). She wrote "Almost certainly the factor which has produced the 'fluffy' coat is common to both sources". In fact the Scottish Wildcat is not an ancestor of the modern domestic cat. Fluffy cats had been present in the west for far too long to owe their existence to relatively recent imports. Mrs Batten added that, in 1967, a Tabby-point male bred by her was mated to a Chinchilla Longhair, and produced kittens whose fur was of intermediate length, rather than the expected shorthairs.
Lauder noted that while pet classes were dominated by shorthaired cats, the majority of the "longhairs" in pet classes were "intermediates". By this she meant that they had "Persian-type" fur as opposed to short, but the fur bore no resemblance to the tremendous pelage of the exhibition quality longhair. She again noted that it was an unsolved question whether the show longhairs were originally bred from cats imported from Ankara or Iran (Persia), or whether they were the result of selective breeding from the native "intermediates" that exhibited a mutation for fur longer than that of the predominant shorthairs. Lauder wrote "probably simply a variation - with a coat of different length occurring in a predominantly short-haired population. These 'intermediates' were much prized by their owners in the early days of the century; people would say with pride 'He's got a fluffy coat!" Lauder added that though there had been surveys into cat colour distribution, the provenance of the "fluffy" cats did not seem to have been researched. According to Mary Batten: "It is known that there are polygenes which influence shorthair coat-length. These may also be present in longhair lines [...] It is arguable as to whether the 'fluffy' cat received its coat-length from the Persian or Iranian aristocrats or from the blending of cats brought from these areas by Romans, and the indigenous Scottish wild cat. However, almost certainly the factor which has produced the 'fluffy' coat is common to both sources. It is not an allele of either longhair or shorthair, but is a separate gene or polygenic series".
As well as those domestic longhairs, "intermediates" or "Semi-longhairs" arose in a number of breeds either by spontaneous mutation or due to recessive genes introduced generations previously. At first breeders sold the longhaired "sports" as pets, or worse they destroyed the kittens so that other breeders did not suspect any impurity in breeding lines. However, longhairs continued to occur in these breeds and many have achieved recognition in their own right. In other breeds, the longhair trait has been deliberately introduced. A British Longhair known as the Lowlander is now being bred (albeit not in Britain) which is less cobby and less extreme than Persian Longhairs.
Some have been bred to retain their natural characteristics (e.g. Birman, Turkish Angora) while others are longhaired variants of shorthair breeds e.g. Cymric (longhaired Manx), Scottish Fold Longhair, Longhair Japanese Bobtail etc etc. Some are bred by crossing different breeds: Tibetane/Tibetaan (longhair Tonkinese), Nebelung (from Russian Blues and domestic longhairs).
In some recently established shorthair breeds, recessive longhair genes are now coming to light. The Bengal Longhair arose due to recessive genes from foundation Abyssinian cats. It has a fluffier coat than the Bengal and a plumy tail and distinct ruff. Although frowned upon by Bengal breeders, some breeders are working with this as a distinct variety. For the same reason, longhaired Ocicats appear in Ocicat litters. The Australian Mist Longhair has also arisen due to recessive longhair genes, again probably from Abyssinian foundation cats. A Singapura Longhair has occurred, but the only known example was neutered. It was identical to the Singapura in all respects apart from the semi-longhair coat which may have been due to recessive genes or to a spontaneous mutation.
There are many others, with new breeds appearing. Only a few are described in any detail.
British Longhair, Longhair Scottish Folds and Scottish Longhair
Longhair Scottish Folds are known as Coupari (after their place of origin, Coupar Angus) in the UK and Highland Fold in the USA (although Coupar Angus is not in the Highlands). It arose through matings with British Shorthairs that carried a recessive longhair gene ("fluffies" are sometimes born to British Shorthairs). It produces both prick-eared and fold-eared variants. Prick-eared Scottish Folds gave rise to the Scottish Shorthair in Queensland, Australia. It has longer tail and legs, and different coat texture to the British Shorthair. It also gave rise to the Scottish Longhair.
The British Longhair (Britanica, Lowlander) is a cat with the British Shorthair's conformation but a semi-long coat and arose from the recessive longhair gene carried by some British Shorthairs and which sometimes gives rise to "fluffies".
Colourpoint Semi-Longhairs
Descriptions and histories of the Balinese, Birman and Ragdoll are in Colourpointed and Masked Cats.
Identical in conformation to the Birman, but in different colours, is the Tibetan, derived from Birman/Persian crosses when introducing new colours into the Birman. It was recognised in Britain in 1986 and should not be confused with the Dutch Tibetane (Tibetaan), a longhaired Tonkinese cat bred in the late 1990s.
Various mink-patterned longhairs and semi-longhairs have been bred during the past few decades from Balinese/Burmese, Himalayan/Burmese, Tonkinese/Persian or Tonkinese/Himalayan crosses . Some resemble Persians, others resemble Tonkinese. The names used included Burmalayan, Himbur, Iranese, Layanese, Silkanese, Tibetane/Tibetaan and Tonkalayan. Although attractive, there has been insufficient interest in perpetuating these.
Turkish and Greek Semi-Longhairs
Apart from the Turkish Angora, there are other longhairs originating from Turkey and neighbouring Greece.
The Turkish Van has existed near Lake Van in Turkey for centuries. In 1955 it was discovered by two British photographers who were given two kittens and acquired three more later on. Recognition was problematical at first since Turkey did not have a cat fancy and there were no pedigree records for this naturally occurring breed. While American cat fancies have mechanisms for accepting foundation cats and developing naturally occurring varieties the British GCCF appeared unable to cope with the physical reality of a cat unless it was accompanied by a four generation pedigree! The Turkish embassy provided documents stating that the cats represented a natural Turkish breed, but this was not acceptable (even today it is a wonder that new cat breeds ever get recognised in Britain). After being bred by enthusiasts for the required number of generations, the "Turkish" was recognised in Britain in 1969 and is now known as the Turkish Van. A politically correct name for this cat is the Kurdish Van.
The Van Kedi is an all-white Turkish Van originating from eastern Turkey and should not be confused with the Turkish Angora. "Van Kedi" is Turkish for "Van cat". In Turkey the self white Van Kedi is prized and the auburn/white variety is held in less regard. The most sought after is odd eyed white although blue eyed cats are also considered special while amber eyed whites are the least sought after. In Britain most matings are between an all-white cat and an auburn/white to produce a mix of all-white and auburn/white offspring (plus occasional cream/white offspring) with a mix of all three eye colours. Outside of Turkey, the Van Kedi may be recognised as a colour variant of Turkish Van rather than distinct breed.
It is worth mentioning the Anatolian (Turkish Shorthair, Anadolu Kedisi) which is a naturally occurring cat similar in type to the Turkish Van with which it is allowed to breed. The Anatolian is found in all natural colours, with and without Van markings. In the past, many Anatolian cats were exported and registered as Vans or Angoras although Dutch and German breeders are striving for purebred Anatolians. The mistaken identity of Turkish Van suggests that they produce semi-longhaired variants.
The Aegean Cat is derived from naturally occurring cats of the Greek Cycladic Islands. It is being developed by members of the fledgling Greek Cat Fancy and is currently the only native Greek breed. Selective breeding started in the early 1990s using native semi-longhaired. These are of a light European/Continental type i.e. neither cobby, nor oriental. The semi-longhaired is than that of Turkish Angoras. All colours are found, with bi-colours (colour-and-white) predominating.
Tiffany (Chantilly), Tiffanie and Asian Longhairs
The Asian Longhair (Tiffany, Longhair Burmese) is a longhair variety of Burmese type and colour and was recognised in 1986. The British breeding programme for Asian Shorthairs such as the Burmilla had the side effect of bringing together the genes for long hair (from Chinchilla Persians) and Burmese coat colour. This ultimately gave rise to longhaired Burmese cats and to longhaired Burmese-type cats in non-Burmese colours. At first the variety was known as the Tiffany, but the Asian Longhair group encompasses a wider variety of colours, paralleling that of Asian Shorthairs.
The similarly named American Tiffany (Tiffany/Chantilly) is not related to the British Tiffanie. It was developed in North America (late 1970s, early 1980s) from non-pedigree cats. It has a silky, semi-longhaired coat in chocolate colour and superficially resembling the Burmese, but is unrelated. At first they were thought to be longhair Burmese so the name Tiffany was chosen in line with the British cats. However Burmese kittens are born with lighter coats and dark paw pads while Chantilly kittens are born dark with pink paw pads. At first they were known as Foreign Longhairs, later as Mahoganies, then Tiffany (in line with British Tiffanie) and later Chantilly or Tiffany/Chantilly to reflect its non-Burmese origin. Its exact history is not known, but it may have been a by-product of the breeding program which gave rise to the British Angora Foreign Longhair). They are recognised only in chocolate and lavender colours.
The Australian Tiffanie derives from the Burmilla breeding program in Australia. Burmillas are Chinchilla /Burmese crosses. Subsequent Burmilla-to-Burmilla matings may produce longhair kittens due to a recessive gene. These are known as the Australian Tiffanie.
Oriental-Type and Abyssinian-Type Semi-Longhairs
The British Angora is known elsewhere as Javanese (reflecting its links to the Balinese), Oriental Longhair and Mandarin. It may also have been the ancestor of the American Tiffany/Chantilly. It was developed from Abyssinian/Siamese crossings in 1973 which produced a chocolate brown longhair with white roots to his fur although he was not genetically a smoke. In 1974 it was remarked that he resembled the old Angora cats. The breeding programme progressed slowly until it was recognised in 1983. In 1984 it was decided that the Turkish Angora and the British "Angora" were 2 distinct breeds. The British "Angora" was improved to have true Oriental type and in 1989 the Cat Association decided that "Angora" was confusing since it was basically a longhaired Oriental. The name Javanese was chosen since this name was already used in Europe for Oriental Longhairs. It does, however. Clash with the American Javanese which refers to red, tortie and tabby Balinese cats.
A British breeder who had read reports on cat gene surveys in the late 1970s was inspired to recreate the coat patterns found in some cats of the Seychelles. The breeding programme began in 1984 using two tortie-and-white Persians and Siamese and Oriental cats. A breed society was formed in 1989, but these cats remain rare. They are essentially Oriental Longhairs with a pattern known to geneticists as the Seychelles pattern and to cat-lovers as the Van pattern: white body, coloured tail and splashes of colour on the head. Some have small splashes of colour elsewhere on the body.
The York Chocolate is an American breed developed in the 1980s/90s from domestic, non-pedigree cats. It is distinguished by its semi-longhair soft, silky hair and chocolate colouring. It has a long lean body reflecting some Siamese ancestry, but is a large cat. In addition to solid chocolate, it comes in chocolate and white bicolour, lavender (dilute of chocolate) and lavender and white bicolour. It is rare in its home country, in part because it is not considered distinctive enough.
The Somali is a longhaired Abyssinian; the name reflects its link to that breed. Longhaired "sports" sometimes occurred in Abyssinian litters and one was exported to the USA as early as 1952. A longhaired Abyssinian was exhibited in Australia as far back as 1965. It was not until 1967 that they were bred in the USA in a deliberate manner. In Britain, the longhaired variants were generally hushed up until a breeder went public at a cat show in 1971. Always blinkered in its outlook, the British cat fancy made a great effort to eradicate the rogue longhair gene by clamping down on pedigrees in which it had shown up. Somalis from the USA were imported into continental Europe in 1977 although longhaired Abyssinians had cropped up in litters prior to that. They were already being bred and shown in Australia and New Zealand. Finally in 1982 (when the Somali has achieved respectability overseas), the British cat fancy woke up to the attractiveness of the Somali and cats from the USA were imported.
The Suqutranese was first shown in Britain in March 1990 and is a sparkling white cat with Somali conformation, but nothing has been heard of this variety since.
In America, the Snow Cat (Alaskan Snow Cat) is similar to the silver series of Somali recognised in the UK. It derives from crossed between Silver Persians and Somalis and is intended to have heavier boning, thicker fur and a rounder head than the Somali. The silver Somali series of Abyssinian and Somali are both rare in the USA which is very liberal when it comes to new conformations, but very conservative when it comes to extending the colour range of existing breeds. It seems that American cat fancies would rather recognise an entirely new breed than allow new colours into an existing one!
Semi-Longhair Rexes
To avoid unnecessary duplication, see Hairless Cats and Curly Coated Cats for information on longhaired cats which have curly hair. Some are due to recessive genes for longhair in existing Rex breeds, others are due to the Rex mutation spontaneously occurring in a longhaired breed. These include the Angora German Rex, Bohemia Rex (Rexed Persian), LaPerm Longhair, Longhair Devon Rex, Rexed Himalayan, Rexed Maine Coon. Combined with Persian-type longhair and undercoat, the Rex mutation can be unruly and unattractive - a Persian with a "bad hair day". With semi-longhair or longhairs without a woolly undercoat, the fur is soft, ringletted or wavy. In many cases, the curling of the hair makes it appear shorter and denser than it really is.