STRIPED, SPOTTED AND TICKED CATS
Copyright 2002 - 2013, S Hartwell

For most people, two of the most familiar "types" of cat are the "blotched tabby" and the "ginger tom" (or "marmalade cat"). These are two examples of tabby, or striped, cat. In the early days of cat exhibitions, tabbies were divided into banded and spotted. Their name was said to have been derived from a street in Baghdad celebrated for the manufacture of its watered or moire silks. When sold in England, this silk was called atabi or - by those who misheard - taffety.

There are four basic types of tabby (ticked, mackerel, classic, spotted) with variations on each of these types. For example the "marbled tabby" is a variation on the "classic tabby".

 

MACKEREL AND BLOTCHED TABBIES

The two most common tabby patterns are "mackerel tabby" and "classic tabby" (blotched tabby, oyster tabby). These two patterns are common in random-breeding pet and feral populations. In the mackerel tabby, the vertical stripes are thin like fishbones and may break up into bars or vertically aligned spots. In the "classic tabby" (blotched tabby) there are broad bands, whorls and spirals of dark colour on a paler background usually with a "bulls eye" (or "oyster") pattern on the flank. Sometimes the markings are extremely broad and fuse together, especially on the back and flanks. Viewed from above, the classic tabby pattern resembles butterfly wings, giving it the nickname "Butterfly Tabby" - a term better known in grandmother's day than in the modern day.

Andrea and Beata Brunetti, in Italy, have a pedigree 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 as shown above. 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, allowing the underlying pattern to show through (in spite of the low contrast and aqua-blue eyes, Hunter does not have Tonkinese ancestry). Dorsal striping in tabbies is very variable and in this instance has combined with the low-contrast Siamese pattern to give an even more distinctive appearance.

Classic Tabby pattern (top view and side view). This pattern is variable, but the "bulls eyes" on the flank are distinctive.

 

Both the mackerel tabby and the classic tabby have stripes and bars on the legs and tail and the classic "M" (or fleur-de-lys) marking on the forehead and the belly is usually spotted. The background colour is an agouti pattern, meaning that each individual hair has several bands of colour along its length. The foreground colour is the solid colour (non-agouti) of the markings.

Left: Silver Classic Tabby

Above: Brown Mackerel Mabby (with white)

Right: Ginger mackerel tabby - the archetypal "ginger tom" or "marmalade cat"

 

The tabby pattern occurs in various colours where the markings are a darker version of the background colour. There are also versions where the markings occur on a silvery (grey) background colour. It can also be mixed with the tortoiseshell pattern to produced torbies or patched tabbies.

Above: Patched classic tabby on black-red tortie.

Left: Patched mackerel tabby on blue-cream tortie.

 

In Britain, the classic brown/black tabby is most common especially in towns, to the point of being considered the quintessential "British Cat". Because the pattern is recessive to mackerel tabby, classic tabbies breed true for that pattern. The bold blotched markings are considered to be more attractive or desirable than the mackerel tabby. Meanwhile, the typical Australian domestic pattern is described as neither truly spotted nor striped, but having a pattern broken into bars. This can be seen especially in feral cats which have reverted to wilder-looking mackerel tabby which may offer them better camouflage. In the California Toyger breed, the mackerel tabby pattern has been selectively refined into a more tiger-like pattern of thin but solid stripes which do not break up. In red self-coloured Persians, the classic tabby pattern has been refined to coalesce and to mask the background colour as far as possible in order to create a "solid red" cat.

CANDLE-FLAME PATTERN/BRAIDED TABBY

There are attempts to produce a "candle-flame" or braided pattern of striping in domestic cats to mimic the pattern found in tigers. Ideally the stripes should be evenly spaced and close to each other. Rather than break up into spots, they should break up into elongated hollow stripes or braids.

TICKED TABBIES

The third tabby pattern is the "ticked". This is due to another gene which masks any other tabby pattern. It is seen in the Abyssinian and Somali breeds, the Singapura and the Asian and sometimes in random-breeding cats. There is almost no striping except possibly for thin stripes on the legs, face and tail. Instead, only the agouti pattern can be seen, giving a "ticked" effect.

The Abyssinian breed was also known as the Ethiopian and the Algerian and was believed to trace back to the Egyptian Middle Kingdom. Its agouti fur resembles that of the African wildcat, an ancestor of modern cats. The first documented "Abyssinian" was "Zula", brought back from Abyssinia (Ethiopia) in 1868. A stuffed Abyssinian (possibly also from Ethiopia) resides at Leiden University's Natural History Museum (Netherlands) and dates to between 1833 and 1882. Apart from that, there is little evidence of Ethiopian origins. Ticked tabbies occur in India and Ceylon (Sri Lanka) and might have been imported from there (perhaps by British governors of India). Frances Simpson described it as a "small cat of delicate colouring", a description now more appropriate to the Celonese or Singapura.

Ticked cats occured naturally in Britain and the Abyssinian breed might have been selectively bred from these. The rabbit-like agouti colouring led to early ticked cats being called Bunny Cat, Hare Cat, Rabbit Cat or Cunny (the latter somewhat naively - this being the slang term for female genitalia!). The most popular was known as the British Ticked and perhaps deserves to be recreated as a variety of British Shorthair.

H C Brooke (writing in Frances Simpson's "The Book of the Cat") wrote in the section on "Some Foreign Cats" of the Egyptian or Caffre Cat and identified it as "the progenitor of the majority of the domestic cats […] it is found over the whole of Africa, and it is quite easy to understand how, with its eminently tameable disposition, it gradually spread over Europe. Our so-called Abyssinian cats […] bear a very striking resemblance to this handsome variety of cat." Similar ticked cats came from India were they were believed to have been derived from the Jungle cat (F chaus) while spotted varieties supposedly derived from the Leopard cat, Desert cat and Rusty Spotted cat.

Brooke's own wife owned an "Indian cat" and Brooke wrote, "Very curious and handsome is the Indian cat 'Indischer Fürst' exhibited by Mrs H C Brooke. His most striking peculiarities are the length and slenderness of his limbs, the extreme shortness of his coat, and his thin and tapering tail […] His ears are small, but as a kitten they were of enormous size, and with his long and pointed head gave him a most weird experience. The voice of this cat is very variable, and far more resembles the raucous call of the Siamese than the voice of any European cat." This cat, and his sister, were apparently stolen from a hotel in Bombay by an English sailor. He was given to a shoemaker in Leytonstone and later sold to Mrs Brooke for a considerable sum plus part-exchange of a kitten with seven toes on each of its paws (these being lucky mascots for sailors). In retrospect it seems that the cat was probably an Oriental (of the older style), but in Brooke's times only the Siamese was known. The Siamese of that time being far less extreme than the modern variety; the illustration shows the Indian cat as similar to an Abyssinian.

"A very taking variety is the Abyssinian. A good specimen should very strongly resemble what one might well expect the Egyptian cat to become after generations of domestication. […] The colour of an Abyssinian should be a sort of reddish-fawn, each individual hair being 'ticked' like that of a wild rabbit - hence the popular name of 'bunny cat. The great difficulty in breeding these cats is their tendency to come too dark and too heavily striped on the limbs; the face should be rather long, the tail short and thick and the ears large. […] The Abyssinian should not be a large coarse cat. A small cat of delicate colouring and with the above-mentioned body properties is by far to be preferred to the large, coarse, dark specimens one sees winning under some all-round judges because of their size."

Breeders favoured the lighter coloured cats as these showed off the ticked pattern to better effect. However, some judges favoured very dark, almost sooty Abyssinians over better marked fairer specimens, penalising the latter because of a "dozen white hairs on the throat". However, some of the earliest Abyssinians in England were Silver Abyssinians and 1889, Harrison Weir's 1889 standard for the Abyssinian accommodated these: "The Abyssinian Silver Grey or Chinchilla is the same in all points, with the exception of the ground colour being silver instead of brown. This is a new and beautiful variety."

The opinions of early cat fanciers were firmly divided into those who liked the silvers and those who despised them. Brooke was opposed to them and in 1903 marked one down with the comment "It is a ticked cat but not the proper Abyssinian colour." Elsewhere, Brooke wrote "I regard silver as an absolute alien colour to the breed, and though there would have been no harm done if these silvers had been kept to themselves, I cannot but think that they did an infinity of harm to the breed, by introducing a grey tinge into the coat, with the result that the beautiful "ruddy" tinge which we used to see in the cats of long ago, is now apparently lost to us. How they originated, or whether any cross was made use of to obtain them, I do not know." A few years later, in 1908, W Johnson Wood commented that "the silver specimens are even nicer than the brown". (Silver Abyssinians remained and are popular in Britain, though less so in the USA)

A strain of apparently albinistic Abyssinians had been bred by Sir William Cooke, of Newbury, but in 1927, his last male died, thus ending a very remarkable strain of albinistic Abyssinians. HC Brooke noted that a lady in Yorkshire owned a pair, but had never shown them and that she was contemplating having the male neutered. It was suggested that the strain derived from a cross with Siamese cat, but Sir William was confident that this was not the case and that the colouration did not bear out this theory. These cats were creamy white, with rabbit-coloured fur on their ears and an "eelstripe" or dorsal line down the back. Their eyes were blue suggesting a form of albinism. At around the same time, another mutation of Abyssinian was reported from Vienna by Herr Lesti. In his second litter by Ras Tafari, one kitten was fawn with a pinkish tint, though the previous litter had all resembled their sire.

In random-bred tabbies, the number, width and placement of bands on hairs in the agouti areas varies from cat to cat. While the Agouti gene permits the hairs to have bands of colour, there are probably minor genes which influence the way in which the hair is banded. Further polygenes may reduce or remove residual leg-barring, tail markings and facial markings on ticked tabbies. Hence the showbench Abyssinian has far fewer markings than ticked tabbies in the random breeding population.

Demonstrating that the ticked tabby pattern can occur in the random breeding population; this British semi-feral cat has ticked fur in a cream colour with some tail rings and a darker cream face with very few facial markings. He does not have leg stripes or necklace markings.

 

According to Albert C Jude in his book "Cat Genetics" (1955) ticking increases with age so that the adult tabby has more pronounced ticking than a kitten stage. In his investigation into the inheritance of tabby coat pattern, PW Whiting (1915) had tentatively considered ticking factors as a series of three alleles (3 alternative versions of the same gene) which he called "much ticking" "little ticking" and "non-ticked". Whiting also considered 3 alleles of the banding factors (number of bands on each hair), calling them "lined" "striped" and "blotched". Both sets of alleles above are named in the order of dominance suggested by Whiting.

American Shorthair breeder Carol W Johnson noticed that unpatterned tabbies, closely resembling ticked tabbies, sometimes occur in purebred where the conventional ticked tabby pattern is not present and where outcrosses to ticked (Abyssinian type) cats are not acceptable. They occurred when a Shaded cat was crossed with either a Classic tabby or with a solid/self carrying classic tabby. She described this phenomenon and formed a hypothesis based on observations and test matings.

At the time, the Abyssinian-type ticked tabby was believed to be an alternative to mackerel tabby or classic tabby i.e. there were three versions of the same tabby gene. What she described was that mackerel and classic tabby were two versions of the same gene while "Unpatterned" tabby (i.e. Ticked Tabby or Abyssinian-type) was an entirely different gene carried elsewhere on the chromosome. It masked out any other tabby pattern present in the cat's genes. In addition, a cat with two copies of the Unpatterned (tabby gene (ticked tabby gene) had fewer residual markings (barring on legs, chest and head) than a cat with only one copy of the gene. A cat with no copies of the Unpatterned gene allowed any other tabby pattern present to be expressed. She also suggested other genes which remove residual markings, noting that some ticked tabbies show "ghost" striping which superficially resembles a mackerel tabby, but is genetically different, perhaps being caused by presence or absence of an as yet unidentified gene.

Johnson considered that the this Unpatterned Tabby gene was present in Shaded Silver American Shorthairs as mating Shaded Silver to Classic Tabby sometime produced ticked tabbies, which were not recognised by the breed society. It is possible that it did enter the Shaded colours at some point in history, but it is masked by the Shaded pattern and hence only manifests in kittens born from matings with non-Shaded cats.

 

Non-agouti (Self)

Agouti Mackerel/Classic

Heterozygous Agouti Unpatterned (Ticked)

Homozygous Agouti Unpatterned (Ticked)

Inhibitor

Smoke

Silver Tabby

Shaded/Chinchilla

Shaded/Chinchilla (with no breakthrough pattern)

No Inhibitor

Self

Patterned Tabby

Ticked (Abyssinian) Tabby

Ticked (Abyssinian) Tabby

 

According to Johnson, the occurrence of unpatterned Chinchilla kittens suggests that the ticked tabby gene may be present in Persians although I have seen no data regarding ticked tabby offspring from matings of Chinchilla/Shaded Silver to Classic Tabby.

It is possible to introduce the ticked tabby pattern into other breeds. For example red ticked tabby could be introduced into the Persian breed to create a red self (solid) Persian which has no tabby markings. All red self cats display the red tabby pattern. Ticked tabby and white cats occur in the random-breeding cat population, but are less popular than striped/blotched tabby and white cats.

The Abyssinian was recognised in the 1860s but the pattern itself is far older. It is the "wild type" of tabby pattern and there have been suggestions that it is related to the ticked pattern of the Jungle cat (F chaus) which may have interbred with domestic cats at an early stage of their evolution. Recent crosses with the jungle cat have introduced a striking new variation of the ticked pattern - the silver tipped black -into the domestic gene pool.

Abyssinian ticked tabbies.

 

The Somali is a longhaired version of the Abyssinian. Long hair is a recessive gene and can be carried hidden by cats, resurfacing when two carriers meet and produce offspring. The Somali was recognised in 1967 though longhaired Abyssinians had been cropping up from time to time before that, much to the embarrassment of breeders who regarded it as a sign of impurity in the bloodlines. About a century ago, Louis Wain wrote "The 'bunnies' [British Ticks] throw both long- and short-haired kittens" and he described them as very big cats, both shorthaired and longhaired, who were born black and later lightened to an unbarred agouti coat. Some of those "British Ticks" were almost certainly bred with Abyssinians and because they carried (and produced) longhair are implicated in the origin of the modern Somali. The recently recognised "British Tick" (a ticked shorthair of British conformation) recreates these Bunny Cats. Below is a Somali-type cat and her kittens. The kittens are a mix of tortoiseshell ticked and mackerel tabby.

In CFA and GCCF, the Somali is a standalone breed (CFA 1979, GCCF 1991). Due to the small gene pool, outcrosses are made to Abyssinians and the shorthaired Abyssinian x Somali progeny are registered as Somali (or Somali variant) in these associations. They are not registered as Abyssinian because they carry the recessive shorthair gene. Some other registries allow the breeds to be crossed and the progeny registered according to hair length. In contrast, FIFe in Europe (and many European registries) register the shorthaired Aby/Somali offspring as Abyssinian. Hence registries that strictly segregate the breeds may require an 8-generation pedigree to prove that an Abyssinian being transferred from another registry has no Aby/Somali crosses in its ancestry. This is not foolproof as there could still be longhaired carriers. In the Canadian Cat Association (CCA), cats with Aby/Somali ancestry are registered as either Longhair Somali or Shorthair Somali. A Shorthair Somali is therefore an Aby/Somali outcross, but can't be registered as Abyssinian in some registries because it is a longhair carrier.

The Singapura (1971) apparently originated from Singapore where the ticked tabby pattern is common in free-living cats. It is generally accepted to be a mix of Burmese and Abyssinian to produce a ticked ivory colouration. Other ticked tabby cats imported from Singapore around the same time are the Wild Abyssinian (black-ticked tabbies, larger and darker than the Abyssinian) and the Limau Kohlum (red ticked tabbies).

The Wild Abyssinian was imported into the USA and bred for a while, but was not recognised as a breed. The use of the term Abyssinian caused dissent among breeders of the modern pedigree Abyssinian. As a breed, it appears to have died out, however this type of cat is still found in Singapore. Reid and Cindi Morgan lived in Singapore from 1992-1999. While there, they found an inured Wild Abyssinian kitten in a large storm gutter. The Morgans note that feral cats of this type are far from extinct and are easily found living in the storm drains of Singapore.

Wild Abyssinian cat from Singapore. Photo copyright Reid and Cindi Morgan

Before World War I, British Abyssinians were bred and shown in several colours: usual (dark brown with black ticking), silver, chocolate, grey (blue) and fawn, but only the usual remained by 1929. These colours have since been re-introduced into the Abyssinian and Somali, and occur naturally in non-pedigree ticked cats around the world. Tortoiseshell ticked cats occur, but the ticking makes the tortie pattern hard to identify visually. The ticked pattern can be combined with the silver gene to create a cat with ticking on a silver background. Silver Abyssinians and Silver Somalis are bred in Europe and a similarly patterned cat, called the (Alaskan) Snow Cat has been developed in the USA.

The Asian group of breeds began in the 1980s with the Burmilla and include the Asian Ticked Tabby. This is a Burmese-type cat with the Abyssinian-type ticked coat. A Burmese/Somali hybrid has also been bred and was known as the Burmali. The Celonese (1984) is a small ticked tabby which originated in Sri Lanka and is now being bred in Italy.

It would be interesting to see the effect of ticked fur on the full-length Persian coat, though there seems to be little interest in this.

A slightly different form of ticking is found in the Chausie, a breed derived from hybridising F chaus (Jungle Cat) with the domestic cat. Black Chausies with silver tipped fur occur and this is belived to be a form of black agouti rather than smoke/shaded silver or chinchilla.

SUQUTRANESE

Although superficially solid white, the Suqutranese (Socotranese) is essentially a ticked cat. It is a Somali-type semi longhair with white fur with glistening silver banding. UK registries and Somali breed societies frown upon anyone advertising "White Somalis" so the Suqutranese is not well known. Somali societies distance themselves from the breed.

Caroline Garrard and Charles and Betty Barrett were holidaying on the island of Suqutra (a.k.a. Socotra) off the coast of Somalia where they noticed a pure white cat with the conformation and temperament of a Somali. Its white fur had well-defined bands of silvery white clearly visible on the individual hairs. Ms Garrard had a pedigree usual silver Somali male (Clyde) and two white shorthair females. She bred an odd-eyed "White Somali" kitten (Fanny) in 1988 from white queen Fifi Farouche and Clyde. First generation kittens were a mixture of Somali colours and pure whites, mostly semi-longhairs of Somali type. In 1989, Fanny was mated back to her father, producing five White Somali kittens. Fanny’s second litter, also in 1989, comprised four White Somalis and one usual silver semi-longhair. Ms Garrard and the Barretts set up a breeding programme to seek recognition for white semi-longhairs of Somali type. They named the breed "Suqutranese" since "White Somali" was unacceptable to registries. Fanny and three of her offspring (2 females and one male) were shown at the CA of Great Britain show in March 1990.

The Suqutranese standard is nearly identical to that of the Somali except the coat must be completely white with silver-white ticking and the nose-leather and paw-pads to be pink. The breeders were hopeful that approval for the breed would be granted by the Cat Association of Great Britain. However, in August 1995 the Somali Cat Breed Advisory Committee took offence at advertisement in various publications for "so-called White Somalis" and issued a statement that Somalis are cats with ticked coats i.e. each hair has several alternating bands of two colours. Any cat with a coat of only one colour is not a Somali and should not be described or sold as such. Since the Suqutranese has a silvery ticked coat and has its own breed name (White Somali being a descriptive term only), this seemed to be a breed society concerned about purity of coloured Somalis. Since then, nothing has been heard of the Suqutranese which is a great pity. A number of Somali breeders overseas have shown interest in re-creating the Suqutranese and it would make a glamourous new addition to the showbench. It would also not be too difficult to reproduce the results (under a less political cat registry) by using a silver Somali male on suitable white females.

Note: In a near parallel of this, during the mid/late 1990s a Somali breeder experimentally crossed Somalis with black shorthairs to create a cat of Somali type but with black fur with a distinct "shimmer".

SPOTTED TABBIES

The fourth type of tabby is the spotted tabby. In some mackerel tabbies, the stripes have a natural tendency to break up to form bars or vertically aligned spots. At first it was suggested that the spotted tabby was simply another mutation of the same gene which caused ticked, mackerel and classic tabby. Spotted tabbies were therefore "broken tabbies". In 2009, more research was done and showed that a separate modifier gene is responsible for turning stripes into spots although the location of the gene hasn't been determined.


Harrison Weir Tabby (1889)


Harrison Weir Spotted Cat (1889)


Broken Tabby

There are a number of spotted breeds in existence. The oldest recognised is probably the Spotted British Shorthair. British Shorthairs have been recognised since the 1870s while American Shorthairs have been recognised (as "Domestic Shorthairs") since the 1900s. The Egyptian Mau may be older, but was not recognised as a breed until the 1950s. The British and American Shorthairs and the Egyptian Mau are naturally occurring breeds which have been refined by selective breeding. In 1872, Dr Gordon Stables described the classes at cat shows at the Crystal Palace and at Birmingham. He wrote "Class VII is for Spotted Tabby [...] A broad black band ran along his back and down his fine tail; and, diverging from this band came dark stripes of colour down the sides, converging round the thighs, and swirling round his chest in two Lord mayor's chains; but the stripes had this peculiarity, they were all broken up into spots."

The Egyptian Mau is sometimes claimed to be the oldest or only naturally occurring spotted breed. This, however, overlooks the Bahraini Dilmun Cat, a spotted variety which is barely known in its own country, let alone internationally. It is also in danger of being lost due to breeding with abandoned Persian cats in Bahrain (Persians ar popular pets, but once the novelty wears off they are frequently turned out to fend for themselves). Dilmun cats are semi-foreign in conformation and have evolved to survive in Bahrain's extremely high summer temperature. Only recently has there been any interest in maintaining this as a breed. It is being bred by the Cat Club of Bahrain and few a Dilmun Cats have apparently reached America.


Male spotted Dilmun (Photo: Adele O'Shea)


Female Spotted Dilmun (Photo: Adele O'Shea)


Male red spotted Dilmun (Photo: Adele O'Shea)

The Ocicat (1964) and was derived from crossing Siamese with Abyssinian cats to create a ticked-point Siamese. The ocelot-like spotted cat was an attractive surprise result. The Spotted Mist (1976) arose originally from a Burmese/Abyssinian mating and again, the creation of a spotted variety was accidental. The addition of domestic tabby shorthairs refined the type and the pattern and some Spotted Mists show a tendency to a rosetted pattern.

The California Spangled Cat was an early "designer breed" bred in the 1980s to resemble spotted wild cats, but without hybridisation with wild species. The body is spotted and lower legs are striped. A "King Spangled" version had marbled markings like a King Cheetah. These cats never achieved the popularity of the Bengal or Ocicat. The breed was founded using mainly Abyssinians, American Shorthairs and British Shorthairs. However, there are suspicions that wild blood Cat blood entered the mix when breeder Paul Casey acquired a Jungle Cat and also purchased two whole litters of Asian Leopard Cat/Siamese crosses (from old-style Siamese) from a their breeder in Hollywood. The breed had a high profile launch in the 1980s and though still listed by TICA it has almost disappeared.

The Kanaani (Canaan Cat) being developed in Germany is being bred to resemble the spotted wildcat subspecies Felis lybica gordonii, but with domestic temperament. Early photographs suggest an Oriental-type or Ocicat-type spotted breed.

Those were the "first four" of the non-hybrid spotted cats - Spotted Shorthair, Egyptian Mau, Ocicat, Australian Mist (six non-hybrid spotted breeds when the Bahraini Dilmun and Kanaani are counted). There are also spotted breeds where the pattern has been inherited from a wild species of small cat which has been interbred with domestic cats.

The Ussuri is a natural breed of from along the Amur river, Russia. It has reputedly crossed with small wild cats in the area - "Amur Forest Cats" or "Amur Leopard Cats" (Asian Leopard cat subspecies) which may make it a hybrid breed. It is a more robust variety which occurs semi-wild and has also interbred with semi-wild cats of Siberian and domestic shorthair type. The pattern is said to be distinctive - vertical solid or merged spots, which may have been inherited from the wildcat. Whether it is a hybrid is not proven.



A superbly rosetted Cashmere (semi-longhaired equivalent of the Bengal). Photo courtesy Corine Kooy, Cashmere Cattery.

In 1963 the Bengal was developed from crossing domestic cats with the spotted Asian Leopard Cat. The original intent was to study the Asian Leopard Cat's supposed natural immunity to certain feline diseases and whether this could be transferred to domestic cats. The end result has been an attractive spotted breed that also includes some margay genes from the Bristol breed developed and abandoned around the same time (the Bristol cats are believed to have introduced the best rosette patterns). In the 1980s, the Geoffroy's Cat was crossed with domestic cats in order to produce the Safari breed with the wild-type rosetted pattern seen in big cats such as leopards and jaguars. Since then. Domestic cats have been deliberately crossed with bobcats (American Lynx and related "Lynx" breeds), jungle cats (Chausie), servals (Savannah, Ashera) and fishing cats (Viverral, Machbagral) in attempts to create wild-looking spotted breeds of domestic cat. In Belgium, Bengals have been crossed to the Indian Desert Cat (F lybica subspecies) to creat the Punjabi (Punjabi Desert Cat). Punjabis are large, elegant cats resembling the wild parent. The colours are grey-black low contrast spots on ticked ivory to chocolate spots on ticked sandy colour. The spots are randomly aligned and low contrast like the wild parent.

The Mojave Spotted (formerly Hemingway Spotted) is a breed being developed from Bengal x Polydactyl crosses.

Mojave Spotted (formerly Hemingway Spotted): Developed from Bengal x Polydactyl crosses.

 

MARBLE TABBIES AND SOKOKE TABBY

The marbled or clouded pattern is thought to be a variation of the classic tabby pattern.

The Sokoke is a natural breed found in the Sokoke Forest area of Kenya. It is now being developed in Denmark. The pattern of the Sokoke is a modified form of classic tabby peculiar to that breed. Agouti (ticked) hairs appear in the centre of the blotched markings to create "hollow" tabby markings.

Marble is the term given to horizontal swirling of the tabby pattern. This type of pattern occurs in the "Marbled Mist", the marbled variety of Australian Mist (a wholly domestic breed derived from Burmese, Abyssinians and domestic tabbies) and occurs in a different form in the Bengal through the interaction of wild-type spotting (from the Leopard Cat ancestor) and domestic-type classic tabby. Marbled patterns may be found in other wild/domestic hybrids as more and more of these are developed.

The marbled tabby pattern is occasionally been seen in cats with no known Bengal (or other wild-type) ancestry. It should be noted that in some countries with a long indoor/outdoor tradition of cat-keeping, the random-breeding moggy population contains a hotch-potch of genes including those from unneutered pedigree pets (demonstrated by the appearance of "pedigree lookalikes" in feral populations).

In the example shown (a random-bred cat), the pattern does not form the swirls and bulls-eye of the classic tabby. The centre of the dark markings contains areas of agouti i.e. the markings are "hollow" or "outlined" (as found in the Sokoke).

In the Highland Lynx breed, the marbled tabby pattern is known as "clouded leopard".

Interestingly, while researching the relationship between classic, mackerel and ticked tabbies and their relationship to silver shaded cats, Carol W Johnson suggested the existence of a "Chaos" gene which disrupts the normal striped pattern.

In ticked tabbies, Chaos (along with Johnson's theorised "Confusion" gene and Cathy Galfro's suggested "Erase" gene) might eradicate residual markings. In a classic tabby, Chaos causes an intermixing of the light striped areas into the black and vice-versa. The banding of each hair is normal, but the hairs themselves occur in the wrong places. The effect she describes is more clearly seen in the Sokoke, which has a modified tabby pattern. An alternative mechanism for this was theorised by Australian Mist breeder Truda Straede with what she referred to as the "finely divided tabby pattern".

Currently, genes such as Chaos, Confusion and Erase are theoretical. It is plausible that some of the many polygenes affecting hair ticking and banding do indeed have the effects attributed to these genes.

PASTEL TABBIES, MIST TABBIES AND SIAMESE "BREAKTHROUGH" TABBY PATTERNS

Pastel tabbies are an attractive version of the blue cream tortie tabby. Instead of forming distinct patches, the tabby pattern is overlaid on a brindled tortoiseshell (the blue and cream hairs are intermixed) giving a hazy effect as shown below.

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 similar effected of dark ticked markings on pale ticked bckground is found in the Australian Mist breed.

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.

AMBER TABBIES

Amber is a colour peculiar to Norwegian Forest Cats caused by a black modifier gene. It was discovered during the 1990s and initially thought to be due to the genes for chocolate, lilac, cinnamon or fawn; these genes are not permissible in the breed as they indicate outcrossing to other breeds. Some cats were registered as goldens. Testing matings with chocolate point Birmans and between the so-called "x-colour" cats demonstrated it to be a new colour (due to a black modifier gene). The two new colours are called Amber and Light Amber; amber is a modifed form of black and light amber is a modified form of blue. 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 (resembling poor quality silver tabbies), 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. 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/tabby-torties.

THE GENETICS OF SILVER TABBIES AND GOLDEN TABBIES

Silver tabby is caused by the presence of the dominant Inhibitor gene. Silver tabbies have a silvery background colour with a tabby pattern overlaid on it. Silver tabbies in the random breeding population are prone to tarnishing - the appearance of a yellowish or rusty hue. The Inhibitor gene seems eliminate the yellow pigment in the ticked hairs of the background colour, but does not affect the solid coloured areas (the markings).

 

The best known silver tabbies are those with black markings as shown on this page. Silver tabbies with other colour markings also occur e.g. Blue Silver Tabby, Chocolate Silver Tabby, Lilac Silver Tabby, Cinnamon Silver Tabby, Fawn Silver Tabby, Red Silver Tabby (Cameo Tabbies) and Cream Silver Tabby (Cream Cameo Tabbies).

The Inhibitor gene has also been introduced into Abyssinians and Somalis to create silver-undercoated varieties of ticked tabby.

The Golden Tabbies have a brighter more golden background colour gained from inheriting the "non-Inhibitor" gene from Chinchilla parents. The Inhibitor gene causes silver tabbies, but since most genes occur in pairs the Inhibitor gene has a partner gene, known as the inverse of Inhibitor or "non-Inhibitor". It has been suggested that the brighter background colour of Golden tabbies is due to a "Wide Band" gene which determine the width of the hair shaft colour between the pigmented tip and the follicle. Golden Shadeds lack the Inhibitor gene, but have a shading pattern comparable to Silver Shaded cats, which must be caused by "something"!. More likely, there are polygenes (Wide Band genes, plural) which affect the undercoat width rather than a single Wideband gene.

MOLECULAR GENETICS OF SPOTTED CATS

In September 2012 researchers announced they had identified the gene that switches between mackerel and classic tabby patterns. This is the first "pattern gene" to be identified. In inheritance terms, mackerel pattern is the dominant "wild type" gene while classic pattern is due to a recessive gene. Cats with narrow stripes (mackerel tabby) have a working copy of the gene. The blotched pattern (classic tabby) occurs when a mutation turns that gene off.

The researchers noted that cells in the black stripes (or spots in the case of cheetahs) know they are in a black stripe and remember that fact throughout the animal's life so that the pattern grows as the cat grows. The researchers looked at cells along the boundaries between light and dark stripes/spots. They analysed DNA samples from feral cats in northern California and the DNA of cheetahs, including samples from the blotched "king cheetah" (a recessively inherited "classic tabby" version of the cheetah).

The research pointed to a gene they called Taqpep (Transmembrane aminopeptidase Q). Blotched house cats had mutations in this gene, while striped tabbies did not. The king cheetah also had a Taqpep mutation while the spotted cheetah had a normal version of the gene. The Taqpep gene produces an enzyme that diffuses outside of cells, interacting with other molecules. Another gene, called Edn3 (Endothelin3), led to the growth of dark fur rather than light fur and coordinated localized colour differences. Together, Taqpep and Edn3 produced the patterning: Taqpep establishes the type of pattern of stripes or spots in early development while Edn3 appeared to carry this on during growth.

Other genes, not yet identified, would determine further variations in the basic tabby patterns in domestic cats, for example broken stripes, round spots, marbled tabby (Bengal) or pale-centred blotches (Sokoke).

* Science 21 September 2012: Vol. 337 no. 6101 pp. 1536-1541
"Specifying and Sustaining Pigmentation Patterns in Domestic and Wild Cats"
Christopher B. Kaelin; Xiao Xu; Lewis Z. Hong; Victor A. David; Kelly A. McGowan; Anne Schmidt-Küntzel; Melody E. Roelke; Javier Pino; Joan Pontius;, Gregory M. Cooper;, Hermogenes Manuel;, William F. Swanson;, Laurie Marker;, Cindy K. Harper;, Ann van Dyk;, Bisong Yue;, James C. Mullikin; Wesley C. Warren;, Eduardo Eizirik; Lidia Kos;, Stephen J. O’Brien; Gregory S. Barsh;, Marilyn Menotti-Raymond.

INHERITANCE GENETICS OF SPOTTED CATS

Early on when pattern inheritance was studied, it was believed that three alleles (gene variants) at the same locus (position on a chromosome) controlled the tabby pattern. Different pairings of these genes produced the following order of dominance in tabby patterns: ticked (Abyssinian) (Ta), Spotted (Ts), Mackerel (TM), and Blotched (Classic) (tb). However data from breeders indicated that mackerel and blotched were on one locus while ticked/non-ticked were at a different locus. Two copies of ticked completely masked the tabby pattern, one copy of ticked and one of non-ticked allowed stripes to show up on the extremeties while 2 copies of non-ticked allowed the tabby pattern to show through. Some other mechanism seemed to break up stripes into spots. More recent research (Eizirik, E, VA David et al) confirmed the 2 loci for ticked/non-ticked and mackerel/blotched and confirmed that a modifier at a 3rd locus can turn the mackerel pattern into well-defined spots.

Three independent breeding lines in controlled environments were used. The first showed the inheritance of mackerel and blotched Tabby patterns. This breeding line confirmed that mackerel and blotched were alleles at one locus with mackerel being dominant to blotched.

The second breeding line focused on the inheritance of the spotted (Ts) variant and was founded using an Egyptian Mau male mated to 3 unrelated blotched tabby females. Egyptian Maus are homozygous for the spotted pattern although the gene involved hasn't been identified. Blotched tabby is recessive to mackerel and ensured there were no hidden recessives to confuse the results. This breeding line produced "broken mackerel" pattern offspring with a mixture of spots and broken stripes. When bred to a blotched tabby, the broken mackerel pattern cats produced a mix of blotched, mackerel-striped and spotted offspring. As there were no mackerel pattern cats in their pedigree, this meant the spotted pattern was caused by interaction of the mackerel tabby gene with an unknown modifier. The modifier masked the mackerel pattern by turning it into a spotted pattern. In fact a tabby pattern modifier was posited by Truda Straede in Australia's "National Cat" magazine based on her observations of small and large spot patterns in the Spotted Mist (now called Australian Mist). It is possible that the variation between broken stripes and well defined spots depends on whether a cat has one or two copies of the spotted modifier.

The third breeding line focussed on the ticked tabby pattern which is semi-dominant (incompletely dominant) to the other tabby patterns. Cats with 2 copies of the ticked gene have no body markings, while cats with only one copy of the gene have tabby striping breaking through on the legs, head and chest. The founders were 2 half-Abyssinian/half European Shorthair males: one had one copy of the ticked gene and carried blotched tabby, the other had one copy of the ticked gene and carried the mackerel pattern. Those 2 males were crossed to mackerel tabby females and blotched tabby females and the results noted. This breeding line demonstrated that the ticked pattern is on a different locus to the mackerel/blotched locus.

Eizirik, E, VA David et al's paper proposed the symbol Ta with alleles TaM (mackerel, dominant) and tab (blotched, recessive) for the pattern-type locus; TiA (Abyssinian ticked) and Ti1 (non-ticked) for the other locus which can fully or partly mask the pattern. A name was not posted for the spotted modifier locus though Straede has previously posited Pmf" (breaks the pattern into spots) and recessive "pmu" (leaves the pattern unbroken).

A Férard categorised tabbies as ranging from tabby (classic pattern), tiger, self (non-tabby), oceloid and spotted. Self indicates a solid colour. Agouti is not included. The oceloid pattern is intriguing as it suggests rosetting or clouding, a pattern only recently introduced into cats via hybridization with wild species.

In 1992, Dr Truda Straede, originator of the Spotted Mist breed (now the Australian Mist, comprising Spotted and Marbled varieties) had concluded that there were only two tabby patterns - abyssinian and classic - and an independently inherited "pattern modifier" gene which she called "pm". The dominant "Pmf" form breaks the pattern into spots, the recessive "pmu" form leaves it intact (i.e. tabby). Pmf would interact with mackerel striping to give small spots or bars. It would interact with classic tabby to give larger, often oddly shaped, spots and bars. Since the Spotted Mist pattern size fell into large and small spots, she suggested that there was a second form of the classic gene which divided the large pattern concentrically into narrower bands. When mated together, "small pattern" cats never produced "large pattern" kittens. Since she had never observed very finely divided blotched patterns, either in purebreds on in random-bred cats, this suggested that the a gene for large pattern classic tabby (thl) was dominant to that for small pattern classic tabby (ths) and that the gene for the small form was only activated if the pattern modifier gene was active.

Although Truda Straede had never seen any finely divided blotched patterns, such a pattern does exist. In the Sokoke, a natural Kenyan breed, the dark classic tabby markings do have a paler central area. The leading feline geneticist at the time, Roy Robinson, found Dr Straede's hypothesis interesting but disagreed with the overall conclusions regarding large pattern tabbies and small pattern tabbies.

There does appear to be a modifier which creates a spotted pattern out of an otherwise striped one. In classic blotched tabbies, the markings sometimes break up into larger round spots as found in the Ocicat and the Egyptian Mau. This suggests that spotted cats are not simply "broken tabbies" but have a separate gene for a spotted pattern or which interacts with the other genes to modify the pattern. This is especially the case where the spots are not vertically aligned as they would be in broken tabbies and where they are round rather than elongated.

Other genes for creating or modifying spotted patterns have been introduced into the domestic cat from wild species. The Bengal breed (derived from hybrids with the Asian Leopard Cat) exhibits a more random type of spotting as well as the coveted rosette pattern (groups of spots) and a swirled marble pattern where domestic cat tabby and wild type spotting interact. The inclusion of the Bristol breed (a margay hybrid) into some Bengal lines may have added other genes for spots.

FRECKLED CATS

While the distinct spots found on Egyptian Mau, Ocicat and several other breeds are due to a spotted modifier gene acting on mackerel tabby, there is evidently more to learn about spotted patterns in cats. For example this India street cat (submitted by Vasilis Lekka) shows a distinct "freckled pattern" of very small spots. A small spot pattern is also showing up in some Savannah cats and may be due to a gene found in the Serval species (one of the parent species of the Savannah breed) which produces the "Servaline" pattern.

No doubt there is much more to be discovered about the striped, spotted and marbled patterns with the introduction of genes from the African serval, the Jungle cat, Geoffroy's Cat and the Fishing Cat into the genetic melting pot.

In 2011, the journal "Science" reported that the Agouti gene not only affects depth how deeply coloured the fur is, but is also involved in the formation of patterns. Researchers at Harvard University studied deer mice, which have dark backs and pale bellies. They showed that small changes in the activity of the Agouti gene during embryo development affects the distribution of pigment. Where the gene is highly active, it delays the maturation of the pigment-producing cells; this results in paler fur in adulthood, for example the pale belly found in many vertebrates. The researchers hope to find out how more complex patterns, such as tiger-striping and leopard-spotting, are formed and which genese are involved. In cats, the agouti gene determines the ticked (banded) pattern on the hair-shafts. Agouti areas of the body have ticked hair-shafts due to alternating bands of heavy and light pigment and are considered the "background colour". The solid patterns, such as tabby stripes, are non-agouti areas where the hair-shafts are not ticked.

Eizirik, E, VA David et al "Defining and Mapping Mammalian Coat Pattern Genes: Multiple Genomic Regions Implicated in Domestic Cat Stripes and Spots", Genetics Society of America, 2010.

Lomax, T. D., and R. Robinson, 1988 Tabby pattern alleles of the domestic cat. J. Hered. 79: 21-23.

MESSYBEAST : COLOURS, CONFORMATION & FUR TYPES