(Part I - History, Literature Search, Coat Color
by Ann Dresselhaus
Reprinted with permission of the
Review of: Journal of Genetics, Vol 42, No. 3, pages 359-427, 1941,
Wolf-Dog Genetics, Prof. Dr. N. A. Iljin
From the same country that brought us the 40 year Fox Domestication Project
comes an 8 year venture into the genetical analysis of 101 wolfdogs.
In 1923, about thirty years before the Russian scientist, Dmitry K.
Belyaev was to begin his project which would show that wild fox could be
domesticated in as little as 40 years, another Russian scientist, Dr. N.A.
Iljin who was the Director of the Institute of General Biology at the First
Institute of Medicine in Moscow began his 8 year breeding experiment to
facilitate the collection of data on the genetics and morphology of wolfdogs.
Improving local dog races:
Dr. Iljin did a literature search before he began breeding and found quite
an impressive record of wolves being crossed with dogs down thru time
beginning as far back as Aristotle in the fourth century B.C. Pliny in
A.D. 23 - 79 reported that the Gauls attached their bitches to trees in
order to have them mated with wolves. These practices continued into the
18th and 19th centuries. Iljin found that the breedings described were not
investigated in a scientific way but were either occasional experiments or
were directed toward the improvement of local dog races including the Eskimo
dog, the Indian dog, and the Hungarian dog.
Many wolf-dog crossings were carried out in zoological gardens and parks
such as the Moscow Zoo Park, Jardin des Plantes in Paris, Hanover,
Stockholm, Halle, Brehm, and the Marseilles Zoological Gardens, where they
were regarded in a "well-respected manner" according to Iljin.
The German Shepherd Dog Mores Plieningen, SZ #159, who was bred to the
first Stud dog, Horand Von Grafath (previously known as Hektor
Liksrhein) and whose blood is said to be in the pedigree of every
German Shepherd Dog in the world today, was the granddaughter of a
wolf at the Stuttgart Zoological Gardens. Their son, Hektor Von
Schwaben, SZ #13, figured heavily in the early German Shepherd Dog
A wild caught 'zonar-grey' (black phase with grey highlights) male wolf was
mated to a black female sheepdog mix resulting in an F1 generation of 13
puppies. Half of the F1 generation had the mother's solid black coat color
or were black and tan and half had the father's zonar-grey coat color.
There were 61 puppies in the F2 generation, 24 in the F3, and 3 in the F4
for a total of 101. The coat color of 73 specimens was exactly
Mapping Iljin's findings to modern day canine coat
It appears that what Iljin referred to as the 'zonar-grey' coat color is the
expression of the same allele that Burns and Fraser (1966) refer to as the 'ag'
agouti (wolf grey) allele in the Agouti (A) series. The black and tans that
appeared in Iljin's F1s were either expressions of the 'as' (saddle marking)
alleles or the 'at' (bicolor) alleles in the same Agouti series. The
leading (most dominant) member in the Agouti series is the A (dominant
black) allele and the third most dominant member allele is the 'ag' agouti
(wolf grey) allele. However, interestingly enough it appears that the
solid black color that the zonar-grey wolf male carried as a recessive and
the matching allele in the black female sheepdog are indeed evidence of
*another* black allele (recessive) 'a' in the same series. The 'ag' agouti
(wolf grey) allele is dominant to the 'a' black (recessive) allele. A
good reference to get a handle on these coat color series is, "Genetics of
the Dog"(1989) by Malcolm Willis which on p. 65 explains the Agouti (A)
Recessive Black allele found in wolves, German
Shepherds, and Belgian Shepherds:
If black were dominant then all black dogs would have at least one black
parent. Willis (1976) showed that only 55 in 115 black German Shepherds
had at least one black parent. In Iljin's experiment, 2 black F1 hybrids
(apparently a/a) produced only black offspring, while 2 F1 zonar-grey
hybrids (apparently ag/a) produced both zonar-grey (ag/a or ag/ag) and black
(a/a) offspring which is indeed evidence of the recessive black
allele 'a' in the wolf.
Carver's (1984) data show clear evidence of a recessive black allele in the
A series using data from 564 black German Shepherds, which in addition to
the Belgian Shepherds are 2 of the very few dog breeds known to carry the
recessive black allele. This may be considered circumstantial evidence of
the influx of wolf heritage into the early dog lines of the Belgian Shepherd
and the German Shepherd.
Doberman markings and liver-colored wolfdogs:
Iljin found the tan markings in the F1 offspring (probably at/at) to be
identical to that of the Doberman (at/at) and the Gordon Setter (at/at)
which is the expression of the 'at' (bicolor) allele in the same Agouti
series. These markings were 'covered by' the recessive black 'a' allele
which makes it dominant to the 'at' (bicolor) allele.
Iljin describes 'brown-fulvous' colored F2s with light whitish-blue irises
that were the progeny of 2 black (F1) wolfdogs. The manifestation of this
color was independent of other characters indicating an independent
allelomorphic site/series. It is not clear to me whether this is the
expression of what is in the present day referred to as the liver/chocolate
allele 'b' in the B series (Little 1914), the Dilution allele 'd' in the D
series (Little and Jones 1919), or the chinchilla allele 'ch' in the C
(Albino) series (Burns and Fraser 1966). The definition of fulvous is
"yellowish-brown, tawny, dull yellow, or yellow"; however, the black and
white pictures of this cross looked quite dark - nearly as dark as the black
coat color and the nose color could not be determined from the
This 'brown-fulvous' color would be confirmed to be an expression of the B
series if Iljin would have noted that the nose color of his 'brown-fulvous'
offspring was not black but brown or liver; however, there was no mention
of nose color at all. He did mention that the coat color was similar to
Newfoundlands, Pointers, and Dashhunds, all of which can occur in the
liver/chocolate color phase indicative of the B series allelle 'b'.
Iljin mentioned that the eye color of all these 'brown-fulvous' wolfdogs was
similar to the eye color of the 'marbled' Great Dane, which I think is what
we call the Harlequin Great Dane today. However, no Great Dane today
(Harlequin or otherwise) is *supposed* to carry the 'b' allele, so the light
eyes are likely the effect of the Dilution or D series. The 'd' dilution
allele is recessive to the 'D' (regular intensity) allele and often lightens
the eyes as well as the coat which could ALSO explain the light eye and
('brown-fulvous') coat color of these F2 offspring.
The last possibility is that the base coat color of the offspring in
question was lightened by the 'ch' chinchilla allele (in the C series) which
does not allow the full expression of coat and eye color to take place.
Canines whose coat is lightened by the 'ch' allele which is recessive to the
C allele (full expression of coat color) can still have black noses. There
is also a postulated allele in this series 'cb' cornaz coat/blue-eye allele
(Pearson and Usher 1929) which can give rise to blue eyes and is recessive
to 'ch' chinchilla allele.
'Brown-fulvous' coat color, whitish-blue eyes,
spots, and wavy coats are likely dog contributions:
It should be noted that regardless of the cause of the lighter
('brown-fulvous') coat and eye color the source of the recessive allele(s)
does NOT have to be the wolf since 2 F1s were bred together to produce these
lighter coat and eye colors. Each F1 could have inherited the recessive
'brown-fulvous' source allele from their dog mother and when bred
together this recessive allele was expressed (ie. doubled up) in their
In much the same way as the 'brown-fulvous' coat color was produced,
white-spotted offspring and longer wavy-coated F2 offspring appeared. When
2 black F1s were bred together, some F2 offspring with white spots occurred
although there were no occurrences of white spots in the F1 generation.
This points to the expression of the 'si' (Irish Spotting) allele in the
Spotting (S) series (Warren 1927 and Little 1957) which was
probably inherited from the dog mother. This allele is recessive to the 'S'
(Self) allele. Most wolves are likely S/S although there are sometimes
small white chest and feet marks in wolves which could be caused by either
the 'si' allele or modifiers at other gene loci that act on the S loci.
There is also heavier spotting in the S series through the 'sp' (Piebald)
and the 'sw' (Extreme White Piebald) alleles which are virtually never seen
in the wolf.
When 2 straight regular-coated F1s were bred together, some F2 offspring
with longer wavy-coats occurred. This is consistent with the modern
description of long (wavy) coats as recessive to straight regular coats.
Again there were no occurrences of long (wavy) coats in the F1s so the long
(wavy) coated allele likely came from the dog mother alone as wavy coats are
virtually never seen in wolves.
'Blue' wild wolves:
Ashen 'blue' wolves were mentioned as being caught (but not bred by Iljin)
in the northern region of Tobolsk in the USSR. The were described as being
a dilute zonar black or a zonar black with reduced black intensity. It is
my guess that this is actually the expression of the 'd' dilution allele
in the D series discussed above which may occasionally occur in certain
geographic regions or among certain subspecies of wolf. Since it is a
recessive allele, it would be nearly impossible to eliminate from the wild
population of wolves unless a single dose of it conferred some maladaptation
upon its carriers as well as upon those that expressed it.
The segregation of coat color in wolfdogs proceeds
in the same way as in dogs:
Characters recessive in dogs are recessive in wolfdogs. Characters
dominant in dogs are dominant in wolfdogs. The genes of dogs introduced
into crosses behave in just the same way as when they meet their normal
allelomorph. Coat color in wolves and dogs result from identical genes in
the same linkage groups.
End Part I (History, Literature Search, Coat Color Results)
See Part II (External Characteristics Results)