The Canine Diversity Project
Inbreeding and Diversity
by John Armstrong
DisclaimerThis is a story about Standard Poodles. In the Poodle, we are fortunate to have a breed with ancient and diverse origins. They are intelligent and versatile, and a recent German study has suggested that they are among the longest lived. My own study, though not complete, suggests that a Standard Poodle should live 14 years or more, given reasonable luck, and a Miniature about 2 years longer. (The averages are actually closer to 12 and 14 years, respectively.)
I would like to believe that you can have healthy, long-lived, championship-quality dogs in any breed. However, this may be a function of their inherent diversity. Many breeds have been established with a much more limited number of founders and, therefore, may never have included the best alleles for certain genes, or may have lost them as a result of random events.
Is Inbreeding Necessary?Many breeders still cling to the idea that inbreeding is the only route to success, and that they can use it as a tool to identify and weed out genetic problems in their line. They will cite the success of certain breeders who inbred extensively, unaware (or conveniently ignoring) that the most successful litters from these kennels were often the least inbred. They also seem to be unaware that many studies on a wide variety of species have demonstrated that highly inbred individuals frequently live shorter lives and have fewer progeny. This is called inbreeding depression.
Inbreeding depression results, in part, from the bringing together of deleterious recessive alleles inherited via both parents from a common ancestor. In humans, where genetic diseases of this type are relatively rare, the frequency of affected individuals is often higher in small populations that are culturally or geographically isolated. In dogs, man has created similar isolated populations by restricting genetic exchange between pure breeds. However, given a sufficiently large and diverse group of founders, there is no reason why the average purebred should not lead a long, healthy life - if responsibly bred.
English ChocolateLow inbreeding does not mean giving up all hope of winning championships.
In England, Julia Taylor (Greekmyth) bred brown Standard Poodles for many years, taking particular care to avoid introducing "contamination" from the North American lines. In 1976, Roy Flowers imported Greekmyth Olympia and later bred her to "DJ" - Am. Can. Bmda. Mex. Dom. (FCI) Int. Ch. Stormy Lane To Sir With Love - an outstanding brown primarily of Bel Tor heritage. The two shared only 20 ancestors over 10 generations.
One of the puppies was Stormy Lane Send In The Clowns, CD (Abby). Abby had an inbreeding coefficient (COI) of only 1.2%*, and lived to 16 years of age! In 1979, Abby was bred to Greekmyth Hercules (Lute). From the all brown litter came Fran Fischer's foundation bitch, Cadbury's Chocolate Clown (Caddy). Fran is now 7 generations past Caddy, and has carefully chosen mates as she went that she believed would both complement her dogs and, at the same time, minimize common ancestry.
In 1995, she bred Cadbury's Kate Hepburn (COI 2.7%), the great granddaughter of Caddy, to a black male known to carry brown - El's Total Package (Bucky). Fran later admitted to me that she had some misgivings about the match. Bucky's origins are predominantly Wycliffe black. However, she saw many things in him she liked, and because Bucky and Kate shared little common ancestry, the inbreeding would still be low (5.4%). The combination seems to have been a spectacular success. The first of the pups to achieve recognition was all breed BIS Ch. Cadbury's Ruby Tuesday (brown), but she has since been joined in the winner's cirle by both black and brown siblings and progeny.
Fran Fischer's task has not been easy. The remarkable success of the Wycliffe blacks sent almost every breeder in search of Wycliffe and related dogs to breed to, and created an artificial genetic bottleneck - so that the vast majority of the current population are more closely related than first cousins. There are standard poodles with distinctively different heritage out there, but there are not a large number. If they are indiscriminately bred to the mainstream, it will be like adding a spoonful of white paint to a gallon of black - the effect will not be noticed.
Danny's TailBringing down the COI.
Another notable poodle with a low inbreeding coefficient was Connie Rodgers' Ch. DeNevillette Dapper Dan (1981-95), COI 3.3%. Danny's sire was the black Ch. Gervais Tabu; his dam the white Ch. Alekai DeNevillette Wahine. Though many would likely have predicted disaster from the line and color mixing in Danny's pedigree, he was not only a long-lived and respected champion, but has over 180 titled descendants.
Danny's male "tail line" (the sire of the sire of the sire...) goes straight back to the great German black Champion Anderl von Hugelberg (b. 1923), as do most black pedigrees. Inbreeding and linebreeding, particularly on Sir Gay (b.1949) has been common, and in this line reached a high of just over 40% at Country Gentleman (Gentry).
Male line COI (%) Bred to COI (%) Ch Annsown Sir Gay, CD 7.3 Clairedge Cinderella, CD 9.9 Ch Annsown Gay Knight of Arhill 11.0 Ch Wycliffe Jacqueline, UD 18.6 Ch Wycliffe Thomas 9.5 Yolanda of Wycliffe 22.6 Ch Wycliffe Ian 38.6 Ch Haus Sachse's Rebecca 25.4 Haus Brau Aladin 25.4 Haus Brau Cheri Beri Ben 37.5 Ch Winshire's Country Gentleman 40.8 Ch Jocelyene Marjorie 4.4 Ch Dassin's Broadway Joe 16.6 Ch Apiele Nominee 12.1 Apiele Lustig of Gervais 16.5 Ch Stylistic's Emerald of Juel 8.3 Ch Gervais Tabu 13.9 Ch Alekai DeNevillette Wahine 29.0 Ch DeNevillette Dapper Dan 3.2
Following that high, the inbreeding coefficient was been reduced to a more reasonable level through a series of partial outcrosses, or what I would prefer to consider intelligently-planned assortative mating (see Breeding Schemes).
One of the most notable was the mating of Gentry to Jocelyene Marjorie. Marjorie's sire was Wycliffe Virgil, who shares common ancestry with Gentry (which is why I consider it a partial outcross), but her dam is only distantly related. Thus Marjorie has a low inbreeding coefficient, which again illustrates my point that you don't have to inbreed to get a good dog. She was Best Puppy in the 1965 Poodle Club of America Specialty and Best in Breed in 1968, and was the mother of 17 champions.
Exploiting the HeritageIn the early part of the century, before a distinction was made between large and small poodles (not including Toys, which were then a separate breed), the sizes probably ranged from around 14" to 24". The English dogs were generally smaller than the German. As a result, today's Standard is about 60% German whereas the Miniature (or European Moyen) carries a much higher proportion of English heritage.
Though they are now recognized as different varieties, the Miniature and Standard poodles are, with the exception of size, bred to exactly the same standard. If there is nothing to give scale, in theory you should not be able to tell which is which. What happens if you breed the two? You will get puppies that grow up to be small Standards because there are few Standards close to the minimum height and many Minis are near the upper limit. Though such crosses are not common, they have allowed a limited gene flow between the two varieties. (There has also been exchange between the Toys and Minis.)
Because of the constriction in the black Standard gene pool caused by the popularity of the Wycliffe dogs (see The Influence of Wycliffe on the Black Standard Poodle) it is nearly impossible to find a black mate that is not a cousin. After careful consideration of the options, Finnish breeder Pirkko Ranta-aho successfully petitioned the Finnish Kennel Club to allow her to breed her 21" black Standard bitch, Helen, to a 17" Moyen male. Six black pups were born in late October, 1998. At 2 years, they 19 inches at the shoulder. All have good temperament, structure and movement. We will be following their progress closely.
The Beginnings of Wycliffe: Michelle and DilemmaJean Lyle's extremely successfull Wycliffe kennel began with the aquisition of a puppy from the famous Carillon kennel of New York in 1952 - Carillon Michelle. Michelle was the daughter of Santo-Labory of Carillon, a Swiss black (part Moyen) imported around 1950, and Carillon Jestina. Though some of Jestina's ancestors came from Switzerland in the 1930s, Michelle's parents share few common ancestors, and her inbreeding coefficient is only 1.9%.
Michelle was bred four times. Her first litter, sired by Petitcote Domino in 1954, produced Wycliffe Jacqueline, the all-time top-producing Standard bitch (21 champions from 41 progeny). She was bred to Domino again in 1955 and 1956, producing 4 more champions, and finally to Carillon Dilemma in 1957.
Dilemma was the son of Michelle's 1/2 sister, Robin Hill of Carillon (COI 1.4%). Dilemma (COI 7.6%) was the most successful son of Annsown Sir Gay, producing 33 champions from 347 progeny. In addition to Michelle, he was bred to Jacqueline and to two of Jacqueline's daughters.
Unique ancestorsMost of the dogs described above have a low COI due to their parents being from different lines (despite the parents themselves being moderately to highly inbred). There are also some whose parents are not highly inbred, but I presently have too few examples to tell whether there is any significant difference and health and longevity between the two groups.
The number of possible unique ancestors doubles in each generation. The following table gives the expected number in each generation and the cumulative number up to 10 generations.
Generation No. in generation Cumulative No. Best achieved 1 (parents) 2 2 2 2 4 6 6 3 8 14 14 4 16 30 30 5 32 62 59 6 64 126 115 7 128 254 218 8 256 510 385 9 512 1022 624 10 1024 2046 938
Most of the Standard Poodle pedigrees have 350-400 unique ancestors in a 10-generation pedigree. The lowest I have on record has 79, and an COI of approximately 70%. The highest have 700-800 and are all below 5%.
Among all the pedigrees I have examined, that of Dorothy Dehn's Cafe Rubio de Beau Raccoon has the greatest number of unique ancestors, as shown in the final column of the table. This is truly remarkable pedigree. (His COI is 3.34%)
Cafe, and Dorothy's other poodles, may be seen at http://www.geocities.com/Athens/Acropolis/5975/poodles.html
Thanks to Dorothy Dehn, Fran Fischer, Roy Flowers, Connie Rodgers, and Pirkko Ranta-aho for giving me permission to discuss their dogs.
* 10-generation inbreeding coefficients were calculated by Wright's method with CompuPed software.
© John B. Armstrong, 1997, 2000.