EARLY REPORTS & STUDIES OF TORTIE TOMCATS

HOW COMMON ARE TORTIE TOMCATS?
Over the years and from region to region, the figures given have ranged from 1-in-1000 to 1-in-many-thousands. Some of these would have been mis-dentified poorly defined classic tabbies, especially where white patches obscure the tabby pattern. Some breeds may be more prone to genetic gender anomalies than others (this has been noted in dogs, but there is currently no comparable data for cats).
According to Roy Robinson's 'Genetics for Cat Breeders' out of a total of 125 kittens produced by black female x orange male there was 1 tortoiseshell male. The majority of calico or tortoiseshell males studied in recent years are from tortoiseshell (with or without white) females and are XXY (Klinefelter) or XXXY (chimera) genetic make-up. Further studies in the early 2000s on the chromosome complement and histology (structure) of tortoiseshell males in the UK and USA indicated around 0.43% of chromosomally abnormal (XY/XY, XX/XY or XXY) tortoiseshell male cats in the UK and 0.033% in the USA.
EARLY REPORTS OF TORTIE TOMCATS
In 1872. Doctor Gordon Stables listed the classes at cat shows at the Crystal Palace and at Birmingham. Regarding the shorthairs of Class 1 he noted "First on the list comes Tortoiseshell Tom". Stables considered Tortoiseshell Tom an ugly cat and expressed surprise that he only seen one tortie tomcat, and that one died at three months old. According to Rose Tenent in her book "Pedigree Cats" (1955): "Both the Short-hair Tortoiseshell and the Tortoiseshell-and-White varieties are very handsome animals, and, like their Long-haired counterparts, most useful for breeding purposes. Both types consist almost entirely of females. In fact, the largest number of males which have ever been seen together was probably in 1871, when four were exhibited at the Crystal Palace Show. One of these was a Tortoiseshell, and the others were Tortoiseshell-and-White. During the next few years several Tortoiseshell-and-White males appeared at the various shows, including the famous prize-winning cat, Totty, which was valued at £100. Then, as now, however, most of the males were sterile." Helen Winslow wrote of the same cats in "Concerning Cats" in 1900.
Jean Bungartz mentioned tortoiseshell-and-white cats in his 1896 book "Die Hauskatze, ihre Rassen und Varietäten" (Housecats, Their Races and Varieties) in " Illustriertes Katzenbuch" (An Illustrated Book of Cats): "One generally says that this pattern occurs only with the female cat, and that three-coloured tomcats would be very rare. Since the female sex predominates in this cat, this may not be surprising, all the more so as the three-coloured variety is not strictly limited is and can appear in each litter (i.e. mixed with other colours)."
In 1897 a tortoiseshell tomcat in Matlock, Derbyshire became famous not for its colour, but for having wings (the genetic condition Feline Cutaneous Asthenia). It was described in a local newspaper as ‘an extraordinary large tortoiseshell tom cat with fully grown pheasant’s wings projecting from each side of its fourth ribs'. The story was reported in the High Peak News of Saturday 26 June 1897: Extraordinary Capture at Winster: A Tomcat With Wings. [...]Mr Roper of Winster, while on Brown Edge near that village, shot what he thought to be a fox, which had been seen in the locality some time previously, on Mr Foxlow’s land. Thinking he had missed his aim, Mr Roper gave up the quest, but returning later he found he had killed the animal. It proved to be an extraordinarily large tomcat, tortoiseshell in colour with fur two and a half inches long, with the remarkable addition of fully-grown pheasant wings projecting from each side of its fourth rib."
In "Concerning Cats" (1900), Helen Winslow wrote There has been a newspaper paragraph floating about stating that a prize of several thousand dollars had been offered in England for a male tortoise-shell cat. This is probably not true, as a Mr. Smith exhibited a tortoise-shell he-cat at the Crystal Palace Show of 1871. Several tortoise-shell and white toms have been exhibited since, and one of these has taken nine first prizes at the Crystal Palace Show; but the tortoise-shell he-cat is extremely rare However, there may have been some confusion over what a tortoiseshell cat looked like as she went on to note that a real tortoiseshell "is not a striped tiger nor a tabby. It has three colors usually, black, yellow, and red or brown; but these appear in patches rather than stripes."
Another early report of a male tortoiseshell cats is a British Shorthair exhibited in early English cat shows (1900s). Lady Alexander's winning cats included Champion Ballochmyle Samson (left), an apparently tortoiseshell male. There is also a reference to Champion Tortie Man of Gallus. Frances Simpson, in "The Book of the Cat" (1903) describes the black-and-orange tortoiseshell and states "At the present time we have Lady Alexander's and Mrs Herring's males - 'Champion Samson' and 'Champion King Saul'." In the chapter on short-haired cats, a cat judge, Mr Jung, elaborates on these tortoiseshell toms: "One of the singularities of the breed is the nearly entire absence of males in every litter; in fact, I remember the say was that a tortoiseshell tom was as scarce as the dodo. At the present time however, we hve two good toms - viz. 'Champion Ballochmyle Samson,' winner of no fewer than twelve first prizes and championships, the property of Lady Alexander, and 'Champion King Saul,' winner of numerous championships and first prizes, owned by Mrs herring. Both these males are very good, and whenever they have been penned together it has always been a difficult matter for me to decide the winner. In females, 'Ballochmyle Bountiful Bertie' (sire, 'Ballochmyle Samson') ..." which means 'Ballochmyle Samson' was a fertile tortoiseshell male. The photograph shows him to be very definitely a brindled tortoiseshell.
Elsewhere, Simpson noted that tortoiseshell toms were rare, but stated "A number of clever fanciers and breeders have used their best endeavours and patiently persevered in the fruitless attempt to breed tortoiseshell male cats. In my long experience I have never known anyone who has succeeded, and those specimens that have been exhibited from time to time have been picked up quite by chance. I recollect, many years ago, at the Crystal Palace show, seeing the pen of a short-haired cat smothered with prize cards, and the owner of the puss standing proudly by, informing inquirers that it was a tortoiseshell tom that lay hidden behind his awards. This man had been paid a shilling by a London cook to take away the troublesome beast out of her area! He had taken it away to some purpose, and his surprise at finding himself and his cat famous was amusing to behold."
Interestingly, in 1912, a male Tortoiseshell and White was registered with the GCCF, being "Ballochmyle Bachelor", owned by Lady Alexander, bred by C Adkin, born 11th March, 1905, sire "Jabez", dam "Kit". He won 1st Prizes at the National CC Show in 1909, 1910 and 1911 and at Westminster in 1911 and 1912. There seem to have been several tortie males from the Ballochmyle line suggesting either some very interesting genetics in the Ballochmyle line!
In 1927, Signor de Southoff, from Florence wrote of a local race cats seen at Viege, above Aigle, Canton Vaud, in Switzerland. In addition to red longhairs ("Carthusian cats") tri-coloured male cats were apparently not uncommon in the village, although they were not specially bred or selected. The strain was apparently restricted to the village.
Among pedigree cats, another early and famous tortoiseshell male was Poldhu in the 1950s, a male Cornish Rex sired by founding Cornish Rex stud Kallibunker. Poldhu was a blue-cream-and-white male who sired a number of kittens. There is controversy over whether Poldhu was truly a tricolour or whether he was a blue tabby (all photos from that time are black and white and show tabby markings). Poldhu was owned by the noted cat breeder Brian Sterling-Webb and had been examined by several geneticists who attested to the fact that he was a blue-cream. Because Poldhu was both tortoiseshell (blue-cream) and fertile, veterinarian Dr A G Searle took a tissue biopsy for research, something which would not have been done for a blue-tabby. This had the unfortunate side-effected of rendering Poldhu sterile. To add insult to injury, not only was Poldhu rendered sterile, the tissue sample was lost.
Some contest that Poldhu was actually blue-tabby-and-white (which contradicts the veterinarian's interest in getting a tissue sample). According to "The Rex Cat" by Phyllis Lauder, Poldhu sired red and cream females out of four different queens. He could only do this if his X chromosome carried the gene for red (diluted to cream by other genes). His owner, Mrs Ennismore, recorded that Poldhu sired red or cream females, but records apparently show that he sired blue tabbies out of a solid blue female. It seems most likely that Poldhu was an XY/XY chimaera (XY/XY chimaeras are described in detail later on).
Poldhu was not the only male tortie Rex. Mrs Roma Lund of Maidenhead owned a blue tortoiseshell Devon Rex called Ben. Though there were no details on his fertility, Ben was valued at £1,000 for his rarity. The account was undated, but appears to be 1970s when Devon Rexes were rare and expensive, whatever their colour.
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Blue-cream (dilute tortie) |
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There has also been a fertile tortoiseshell-point Siamese male in England who has sired a number of litters before being neutered. Marilane Harlequin or "Hal" (1980 - 1994) bred as if a seal point i.e. passed on the gene for black. In his show career he was a Champion and a Premier (equivalent of a neutered Champion) although there is no show class for tortie males. He was bred from a tortie-point female and a cream-point male. In the 1960s or 1970s a Van-patterned male longhair turned up in the USA. This cat had a blue spot on his head mingled with cream, a cream spot on his body and a blue tail. Blood and skin samples were sent to Pennsylvania University for chromosomal investigation. The University concluded that the tortoiseshell Van male would be fertile, which meant the blood and skin cells contained the normal XY complement. His sister was black, indicating a black father and tortie mother. In the 1980s there was a Dutch-bred dilute calico (blue-cream-and-white) British shorthair male called Benjamin's Jos Brink. This cat sired litters between 1982 - 1987.
One of the most interesting cases surely has to be the "Tripple" line of Norwegian Forest Cats. This line apparently produces fertile male calicos at least every third generation. The foundation male of this line was a fertile calico male called Tripple. Tripple had a black patch on his flank which may be a somatic mutation, but otherwise appears to be a red/white bicolour. Some of these calico toms could be seen at Dutch cat shows. They look like ordinary red or red bicolour males, but have small black spots - these spots are very definitely black, not merely dark red. Some also had a mix of red and black in their coat, with the red always predominating. Studies of pedigrees from Tripple's breeding line indicated that calico males occurred every third generation and sometimes as often as every second generation. There was apparently a distinct ratio of calico males which certainly merits a more detailed study of the phenomenon. Initial studies, based on phenotype rather than genotype, showed no evidence of XXY (Klinefelter) in the line.
A fairly casual surf of the internet and usenet (newsgroups) turns up a number of cases of tortoiseshell male pet cats, some identified as XXY make-up, including adverts offering rare male calicos for sale. Some reports must be taken with a large pinch of salt. For example a supposedly calico male called Bentley was described in 1992. The owner had no idea that Bentley was unusual until someone described Bentley as a "she" and then said that cats with tricolour noses were always female. Bentley's nose was white, black and beige. The fur on Bentley's side was a mix of black tipped with beige and solid beige. However, Bentley's owner added "the beige forms circles like bulls eyes on his sides" meaning that Bentley was not a genetic anomaly at all, but an ordinary classic tabby cat with an ordinary mix of agouti and non-agouti fur! One wonders how many "tortie toms" are nothing of the sort, except in their owner's imagination or because the owner is unfamiliar with the tabby pattern?
In 1992, Mindy Machanic reported a 12 week old male calico called Huggy Bear. The vet was initially dubious that the cat was male. Huggy Bear was white and black and red with a Van pattern: white body with one large black spot striped with light tan, white legs with a tiny black spot, a fluffy black tail, striped with light tan and a white head with black and red/reddish brown triangular patches in front of his ears. He was healthy and not at all retarded as initially feared; he was also developing some typically male behavioural traits. In 1994, Paul Atchley bred a litter of calico kittens including two male calicos which were mostly white with orange, black, and grey patches. Several other tortie males were mentioned in 1994, including one called Smudge that went missing for six weeks before being found in a ventilation pipe and a a 7 year old tortoiseshell male barn cat called Injun.
In 1996, a newsgroup posting referred to a calico male called Camille who was, quite reasonably, assumed to be a female when adopted as a kitten. Camille has wide hips, walks with a wide gait, is healthy and has a sweet temperament; traits suggestive of XXY chromosomes (Klinefelter). He sounds similar to Brody Baker whose case study is towards the end of this article. By contrast, in 2001, a very macho dilute tortoiseshell male, Toby, was reported on Usenet -his fertility was unknown, but even after neutering he tried to mate with other cats. In 2001, a large, sweet-natured dilute calico (blue/cream tabby + white) male called Calvin was reported. The owner was tempted to breed him due to his rarity. In 1999 another writer said that her mother had bred numerous tortoiseshells including 2 male torties, but to her disappointment both males were sterile. In 2002 a family of four stray cats were reported including a red longhair, a black shorthair, a tortie female and tortie (predominantly red) male - this covered most of the normal colour possibilities for the "O" gene plus the anomalous colour! In 2003, another posting reported an orange tabby female called Carrot who produced a male calico kitten in her first litter. Other reports sound like cases of mistaken identity e.g. on Sept 20th 2002, a shorthair tortoiseshell male cat coloured grey, black and white was impounded in Bismarck, ND, USA (though this could be a silver classic tabby and white).
Tortoiseshell males from pedigree breeds are perhaps the most interesting as these have complete genetic histories detailing the colours of parents and related cats. Riëtte van Beek has collected information on a number of tortoiseshell male cats, their parentage and their offspring (if they are fertile). In addition to the cases detailed above, these enigmas include Kapamba's Home Miracle (tortie-and-white Norwegian Forest Cat, fertility and genetic make-up unknown), Dar es Salaam's Varsha (male Seal-Tortoiseshell Point Birman who passes on red), and Norwegian Forest Cats from Abelia cattery (France): Patchouli (tortie and white, breeds as a red-and-white) and his full brother Santal's Neroli (blue-cream tabby and white, breeds as a blue tabby and white), both being fertile male tortoiseshells born in 2001.
EARLY STUDIES OF TORTIE AND CALICO TOMS
One of the first scientific studies was that of the fertile blue-cream male Cornish Rex Poldhu in the 1950s. Veterinarian and researcher Dr A G Searle took a tissue biopsy for research, but this had the unfortunate side-effected of rendering Poldhu sterile. Worse, the tissue sample was lost. Mating records indicate that Poldhu sired red and cream females out of four different queens and blue tabbies out of a solid blue female.
In his book "Cat genetics" (1955) Albert C Jude discussed the anomaly of tortoiseshell mails and included a photograph of a fertile brindled tortoiseshell tomcat. He noted that tortie males occurred only as rare exceptions and were almost always sterile. At the time, the genetic constitution of the tortoiseshell male was unclear; one explanation was that a tortoiseshell male was in fact a genetically-determined (XX) female which, under the influence of male sex hormone from another embryo of the same litter, developed as a sterile intersex animal like the freemartin seen of cattle (discussed later in this article). He added that where the male sex hormone had been unusually concentrated, a fertile XX tomcat might result. Cutler and Doncaster (1915) had found that sterile tortie males entirely lacked normal reproductive cells.
During investigations carried out by Jude himself during 1966 in connection with a new theory put forward by Professor Komai of Japan, six tortoiseshell-and-white tomcats were reported in Great Britain. Professor Komai’s theory proposed that the mothers of all these males were themselves tortoiseshell or tortoiseshell-and-white. Only one of the reported cats, Mrs Cotterell's fertile male, had been traced and Jude appealed to readers for more instances - with full data - of fertile tortie or tortie-and-white male cats. The tortoiseshell-and-white male belonging to Mrs Cotterell of Luton was described as having "a large patch on forehead and over eye of ginger and white. It has white under throat and belly. It has golden necklace, and from then on to the tail, the coat is broken with gold color on base black." That particular tortie male sired kittens on a tortoiseshell female also belonging to Mrs Cotterell. The kittens were described as "one very like the mother, one something like the mother, only darker, and a third, black with a ginger tint." Unfortunately, the "black with a ginger tint" kitten died before 4 weeks of age. It was thought to have been a male while the other 2 kittens were females. It was hoped to obtain test matings if the sire remained unsold.
In 1967, Cats Magazine reported on a show of five male calico cats at the previous October's Totem Cat Club Show, Wacoma, Washington. Four were displayed by arrangement of the Research Department of Rainier School, a Washington State Institution for the mentally retarded (a term politically correct at the time). One was owned by Mrs Mahaffie and had won ribbons in the previous year's show. A second belonged to Mr and Mrs RG Parrett of Bellevue, Washington. The third belonged to Mr and Mrs B Moon of Vashon, Washington. The fourth was owned by the Research Department of Rainier School, and had been given to them by Mrs Waddell of McGehee, Arkansas. The fifth was a kitten owned by Miss Erickson, of Tacoma.
For the previous 8 years, Dr Horace Thuline and his staff at the Research Dept of Rainier School had conducted a research project into chromosomal abnormalities. The main interest was Down's Syndrome in children, but male calico cats were of interest because their colouration was due to a chromosome abnormality. The report noted "It is the fortunate circumstance of black and orange being sex-linked in a cat that allows males with two x-chromosomes (a male should have only one x-chromosome) to be identified by sight rather than with expensive laboratory procedures. The rarity of Calico or Tortoiseshell male cats is evident from the fact that, in seven years of looking for them, assisted by many people all over the United States, only eleven had become available for studies at Rainier - including the five at the show."
The display included posters showing chromosome patterns for each of the cats, though amidst so many different breeds and colours of cat their unusual nature was easily overlooked by show visitors. It was not obvious to the casual passer-by that "these normal-appearing males are rarely born, and rarely survive. Because of the general casual attitude concerning pet cat litters, it is impossible to give an accurate estimate of how frequently Calico or Tortoiseshell males are born." Miss Erickson and Mrs Mahaffie were present at the display while scientific explanations about the cats, about "Project Calico" and its importance to the Rainier School studies were available from Dr Thuline and his associate Dr Darwin E Norby. The Totem Cat Club planned to donate space again in 1968 to Rainier School for "Project Calico" and with more advance planning, hoped for a better display set apart from other feline exhibits and with colour posters.
Unfortunately for "Project Calico" and its applicability to human chromosomal abnormalities, modern research indicates that tortoiseshell and calico tomcats are more often due to chimerism than to the XXY chromosome abnormality. This also means that most tortie/calico males are fertile. Dr Thuline and his associates would not have been aware of the chimera condition in 1967.
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