Inbreeding
and Diversity
by John
Armstrong
Disclaimer
This 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 Chocolate
Low inbreeding does not mean giving up all hope of
winning championships.
In
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 Tail
Bringing down the COI.
Another
notable poodle with a low inbreeding coefficient was Connie Rodgers'
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 Heritage
In 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
Dilemma
Jean Lyle's extremely successfull
Wycliffe kennel began with the aquisition of a puppy
from the famous Carillon kennel of
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 ancestors
Most 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.
Copyright 1997,
2000 John B. Armstrong. The Canine Diversity Project. All rights reserved. Reprinted with permission.
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