DP Sponenberg

MC Weise

Genetics Selection Evolution

Crossref

NoteDominant black in horsesDP Sponenberg MC WeiseDepartment of Biosciences and Pathobiology,Virginia-Maryland Regional College of Veterinary Medicine,Virginia Tech, Blacksburg, VA 240611-2714 Wocjik Ln, Sobieski, WI 54171, USA(Received 20 January 1997; accepted 7 March 1997)Summary - The existence of dominant black in horses is supported by a black stallionproducing 12 black or near black and no other color of foals from bay mares, and 16 blackor nearly black and no other color of foals from chestnut mares. This allele is suggestedas being dominant black, ED, at the Extension locus. This allele does not always causecompletely eumelanic phenotypes, since some offspring (which were heterozygous) werenear black rather than completely black. The dam of this stallion was of a near blackor brown, rather than black, phenotype. The sire of this stallion was black. Foals weresometimes born a color close to that of bay foals, but these ultimately turned completelyor nearly black at maturity.color inheritance / Extension locus / dominant black / horseRésumé - Noir dominant chez les chevaux. L’existence d’une coloration noire dominantechez les chevaux est mise en évidence par un étalon noir produisant 12 poulains de couleurnoire ou presque à partir de juments bai et 16 poulains de couleur noire ou presque noireà partir de juments alezan. Cet allèle est suggéré être le noir dominant, ED, au locusExtension. Cet allèle ne cause pas toujours des phénotypes eumélaniques puisque certainsdescendants (qui étaient hétérozygotes) étaient presque noirs plutôt que complètementnoirs. La mère de cet étalon était presque noire ou brune, plutôt que noire. Le père decet étalon était noir. Les poulains naissaient quelquefois avec une couleur proche de celledes poulains bai mais devenaient complètement ou presque noirs à la maturité.hérédité de la couleur / locus Extension / noir dominant / chevauxINTRODUCTIONThe inheritance of color in horses has been studied for much of this century. Earlystudies were largely based on studbook data, and the accuracy of subtle distinctionsin color nomenclature were sometimes lacking (Gremmel 1939, Castle 1940a, b,* Correspondence and reprints: Veterinary College, Virginia Tech, Blackburg, VA 24061USA1951a, b, 1954). This is especially so when horses are registered as foals, since foalcoats and adult coats can be very distinct from one another. Later studies weremostly based on segregation data from selected families of horses, and provideda sound basis for the testing of hypotheses concerning the inheritance of horsecolor. These studies also tended to use the principles of homology in assigning lociand alleles for the various genetic phenomena controlling color (Odriozola, 1951;Adalsteinsson, 1974, 1976).Black horses are usually considered to be uniformly covered with black hairs, andin many breeds the presence of even a few reddish or tan hairs (usually in flanksor muzzle) results in the designation of such horses as brown, or even bay, ratherthan black (Sponenberg, 1996). Horses with reddish brown bodies and black lowerlegs, manes and tails, are consistently designated as bay in most breed registries.While such red horses are consistently referred to as bay, the term bay can alsoinclude darker colors, some of which approach black. Horses with a mixture ofblack and reddish brown body hairs and black lower legs, manes and tails arevariably considered brown or bay, depending on the breed or the observer. Thebrown classification can therefore include horses that are nearly black, as well asthose that are nearly bay. An additional type of brown is seal brown, which has blacklower legs, mane and tail, and a black body with conspicuous lighter hairs on themuzzle, over or around the eye sockets, and in the flanks and axillary regions. Thegenetic differences between these various sorts of brown have never been elucidated,largely because the nomenclature in the English language is ambiguous enoughthat it is difficult to accurately designate horses into one color group or the other(Sponenberg, 1996).Few studies have been accomplished that finely split the various browns accu-rately. Abeles (1979) presents an hypothesis for the interrelationships of horses withblack lower legs, manes and tails, and bodies of reddish brown, mixed reddish brownand black, or uniform black. Although no segregation data are presented, her expe-rience is consistent with the reddish brown and black mixtures (which she refers toas brown) dominating the clear reddish browns (her bay), which in turn dominatethe uniformly black bodies (her black). This is perhaps explained by the interactionof the Agouti and Extension loci. These loci have long been postulated as impor-tant in controlling common horse colors, although the hypotheses concerning theirinteractions have varied over the years (Odriozola, 1951; Jones, 1982; Sponenberg,1996).The existence of a uniform black in horses that is recessive to bay has long beenacknowledged, and accounts for the majority of black horses. This black is assignedto the Agouti locus in keeping with principles of homology (Wilson, 1910; Wright,1917; Adalsteinsson, 1976; Lauvergne et al, 1991).The existence of a dominant black in horses has also long been speculated(Gremmel, 1939; Castle, 1940a, b, 1951a, b, 1954). The evidence for a dominantmechanism producing black has been the production of bay or brown foals fromthe mating of two black parents. Such instances have usually been from studbookdata, which are suspect owing to the possibility of inaccuracies in the registrationsof animals, or in mistakes of parentage. Another problem with studbook datafor the colors bay, brown and black is one of definition. Various breeders mayhave different mental images of these three and consequently studbook records areprobably inaccurate at this level of detail. Dreux (1966) presents evidence for theexistence of a dominant gene contributing to black or dark bay phenotypes in theFrench Trotter breed, and mentions that this is rare in horses of European breeding,having been introduced through an imported American Trotter stallion.Currently, documentation best fits with the view that the Agouti locus comprisestwo alleles, with bay dominant to black (Sponenberg, 1996). A third allele, black andtan (!4! is fairly poorly documented as a cause of seal brown, and is intermediatein dominance to the other two alleles. The Extension locus likewise has twowell-documented alleles, wild type dominant to chestnut (Sponenberg, 1996). Therecessive chestnut allele, when homozygous, results in a uniformly reddish brownphenotype (including lower legs, mane and tail), and is epistatic to the Agoutilocus. Past literature has postulated intricate relationships with these two loci,with no clear epistasis of one over the other, and with these intricate interactionsexplaining the differences between bay and brown, and between shades of chestnut(Jones, 1982). The Extension locus may well be the site of an allele that providesfor the intermixture of black hairs into the red body coat of some bay horses, whichtranslates into the difference between bay and brown. This is consistent with Abeles(1979), even though segregation data are lacking in that report.This report discusses data that support the existence of an additional Extensionallele, for dominant black, in the horse.MATERIALS AND METHODSProduction data from a black Arabian stallion (Serr Ebony Star) were evaluatedfor foal color production with various colors of dams. The production data fromthe black sire (Shilosmidnight) and the dark brown dam (PRF Gali Gaeraff) of thisstallion were similarly evaluated.RESULTSResults are summarized in table I. The black stallion was mated to mares ofvarious colors. Following mating to bay mares he produced seven black and five’near black’ foals (’near black’ is used to describe horses that are black save for afew tan or red hairs in muzzle or flank, or those horses that are very dark but forwhich it is possible to discern a subtle difference in the color of body and the lowerlegs, mane and tail). Following mating to black mares he produced nine black foals.Following mating to chestnut mares he produced ten black and six ’near black’foals.The sire of this stallion was black and was mated to black mares to producetwo black foals. Matings to bay mares resulted in three black and three bay foals.Matings to chestnut mares resulted in one black and two bay foals.The dam of this stallion was dark brown, and was mated to a black stallion toproduce one black foal and three ’near black’ foals. A mating to a chestnut stallionproduced a ’near black’ foal.DISCUSSIONThe problem of nomenclature concerning dark horses is a concern for this study.In the Arabian breed many breeders are very strict in their use of the term black,and reserve it for horses with black hairs only, excepting white marks due to spottingpatterns. Arabian breeders do not use the term brown, and so horses that are nearlybut not quite black are usually referred to as black bays, or even simply as bay.These are, however, very distinct from the usual bright reddish brown bay thatis typical in the breed. Black bays in the Arabian breed are easy to confuse withblack horses, especially without close and detailed examination, and are difficult toconfuse with bright reddish brown bay horses, which are common in this breed. Thegrouping of blacks and near blacks, or black bays, as one group is therefore logical,since this is how these colors appear to most observers. These three are very closeto one another visually, if not in terms of nomenclature.The results of the mating of the horses of this study is consistent with thesegregation of a dominant allele for black. The mating of the stallion to bay maresfor 12 foals can be tested for goodness of fit for the hypothesis that this stallion is arecessive black. If that were the case, and if all the bay mares were heterozygous forthis usually rare allele, then X2 = 12, 1 df, P < 0.005. This deviates significantlyfrom the expectation, even postulating the most favorable condition of all the baymares being heterozygous for a rare allele. These results are therefore inconsistentwith the segregation of a recessive allele for black, but are consistent with thestallion being homozygous for a dominant allele causing black color.The sire of this stallion was black, while the dam was dark brown. Both neededto have had the allele causing dominant black if the stallion is indeed homozygous,as is supported by the segregation data. The mare always produced black or nearblack foals when mated to a black stallion. She also produced a near black foalwhen mated to a chestnut stallion, even though she is an obligate heterozygote fordominant black by virtue of having had a bay sire. The sire of the stallion producedequal numbers of bay and black foals when mated to bay mares, which is furtherevidence that a dominant gene is segregating in this family of horses. The sire of thestallion is heterozygous for this gene, even though both of his parents were black.A few of the foals in this study were born a color resembling that of foals thatultimately prove to be bay. These foals darkened with age, ultimately becomingblack or nearly so. Some of the blackest of the horses in this study were born withthis light bay foal coat, only to darken later. It is fairly typical for black horses(assumed to be of the recessive Agouti genotype) to be born an ashy grey color, todarken later. Whether or not the differences in foal coat are consistent with thesetwo different genetic mechanisms that cause black horses is undetermined.The allele causing the black phenotype in these horses is most likely the dominantblack (ED) at the Extension locus. This allele is documented in numerous otherspecies as a cause of uniformly eumelanic phenotypes (Searle, 1968). It appearsto be arguably incomplete in its action in some instances, causing dark brown ornear black instead of truly black individuals. In foxes, for example, the ED alleleis frequently incompletely dominant resulting in three color classes from this alleleand the wild type allele (Adalsteinsson et al, 1987). The dominant black allele maybe the genetic mechanism behind some of the very dark phenotypes in horses thatare not truly uniformly black. The differences in genotype between the black andnearly black horses are uncertain, since in this data set it is conservative to assumethat most, if not all, of these had the Agouti locus bay allele, and its presence orabsence can therefore not be responsible for the subtle differences in the black andnear black horses. Likewise, most of these foals had to have been heterozygous forthe dominant black allele, and this genotype includes both black and near blackindividuals. The dominant black allele appears to be rare in most breeds, since inmost breeds black does segregate as a recessive.REFERENCESAbeles HM-S (1979) A coat of many colors. Equus 17, 30-38Adalsteinsson S (1974) Inheritance of the palomino color in Icelandic horses. J. Hered. 65,15-20Adalsteinsson S (1976) Colour inheritance in Icelandic Ponies. In: First Int. Synzp. onGenet. and Horse Breed., Royal Dublin Soc. Dublin, 17-18 September 1975, 42-49Adalsteinsson S, Hersteinsson P, Gunnarsson E (1987) Fox colors in relation to colors inmice and sheep. J. Hered. 78, 235-237Castle W (1940a) Mammalian Genetics. Harvard University Press, Cambridge, Mas-sachusettsCastle W (1940b) The genetics of coat color in horses. J. Hered. 31, 127-28Castle W (1951a) Dominant and recessive black in mammals. J. Hered. 42, 48-50Castle W (1951b) Genetics of the color variety of horses. J. Hered. 42, 297Castle W (1954) Coat colour inheritance in horses and in other animals. Genetics 39,35-44Dreux P (1966) Contribution a 1’etude du gene E chez le cheval domestique. Ann Genet,9, 168-170Gremmel F (1939) Coat colors in horses. J. Hered. 30, 437-45Jones WE (1982) Genetics and Horse Breeding. Lee and Febiger Publishers, PhiladelphiaLauvergne JJ, Silvestrelli M, Langlois B, Renieri D, Poirel D, Antaldi GGV (1991) A newscheme for describing horse coat colour. Livest. Prod. Sci. 27, 219-229Odriozola M (1951) A los colores del caballo. Publicaciones del Sindicato Nacional deGanaderia, MadridSearle A (1968) Comparative Genetics of Coat Colour in Mammals. Logos, LondonSponenberg DP (1996) Equine Color Genetics. Iowa State University Press, Ames, IowaWilson J (1910) The inheritance of coat colour in horses. Proc. Royal Dublin Soc. 12,341-348Wright S (1917) Color inheritance in mammals VII: The horse. J. Hered. 8, 560-64.

English

Dominant black in horses

Note
Dominant black in horses
DP Sponenberg MC Weise
Department of Biosciences and Pathobiology,
Virginia-Maryland Regional College of Veterinary Medicine,
Virginia Tech, Blacksburg, VA 240611-
2714 Wocjik Ln, Sobieski, WI 54171, USA
(Received 20 January 1997; accepted 7 March 1997)
Summary - The existence of dominant black in horses is supported by a black stallion
producing 12 black or near black and no other color of foals from bay mares, and 16 black
or nearly black and no other color of foals from chestnut mares. This allele is suggested
as being dominant black, ED, at the Extension locus. This allele does not always cause
completely eumelanic phenotypes, since some offspring (which were heterozygous) were
near black rather than completely black. The dam of this stallion was of a near black
or brown, rather than black, phenotype. The sire of this stallion was black. Foals were
sometimes born a color close to that of bay foals, but these ultimately turned completely
or nearly black at maturity.
color inheritance / Extension locus / dominant black / horse
Résumé - Noir dominant chez les chevaux. L’existence d’une coloration noire dominante
chez les chevaux est mise en évidence par un étalon noir produisant 12 poulains de couleur
noire ou presque à partir de juments bai et 16 poulains de couleur noire ou presque noire
à partir de juments alezan. Cet allèle est suggéré être le noir dominant, ED, au locus
Extension. Cet allèle ne cause pas toujours des phénotypes eumélaniques puisque certains
descendants (qui étaient hétérozygotes) étaient presque noirs plutôt que complètement
noirs. La mère de cet étalon était presque noire ou brune, plutôt que noire. Le père de
cet étalon était noir. Les poulains naissaient quelquefois avec une couleur proche de celle
des poulains bai mais devenaient complètement ou presque noirs à la maturité.
hérédité de la couleur / locus Extension / noir dominant / chevaux
INTRODUCTION
The inheritance of color in horses has been studied for much of this century. Early
studies were largely based on studbook data, and the accuracy of subtle distinctions
in color nomenclature were sometimes lacking (Gremmel 1939, Castle 1940a, b,
* 
Correspondence and reprints: Veterinary College, Virginia Tech, Blackburg, VA 24061
USA
1951a, b, 1954). This is especially so when horses are registered as foals, since foal
coats and adult coats can be very distinct from one another. Later studies were
mostly based on segregation data from selected families of horses, and provided
a sound basis for the testing of hypotheses concerning the inheritance of horse
color. These studies also tended to use the principles of homology in assigning loci
and alleles for the various genetic phenomena controlling color (Odriozola, 1951;
Adalsteinsson, 1974, 1976).
Black horses are usually considered to be uniformly covered with black hairs, and
in many breeds the presence of even a few reddish or tan hairs (usually in flanks
or muzzle) results in the designation of such horses as brown, or even bay, rather
than black (Sponenberg, 1996). Horses with reddish brown bodies and black lower
legs, manes and tails, are consistently designated as bay in most breed registries.
While such red horses are consistently referred to as bay, the term bay can also
include darker colors, some of which approach black. Horses with a mixture of
black and reddish brown body hairs and black lower legs, manes and tails are
variably considered brown or bay, depending on the breed or the observer. The
brown classification can therefore include horses that are nearly black, as well as
those that are nearly bay. An additional type of brown is seal brown, which has black
lower legs, mane and tail, and a black body with conspicuous lighter hairs on the
muzzle, over or around the eye sockets, and in the flanks and axillary regions. The
genetic differences between these various sorts of brown have never been elucidated,
largely because the nomenclature in the English language is ambiguous enough
that it is difficult to accurately designate horses into one color group or the other
(Sponenberg, 1996).
Few studies have been accomplished that finely split the various browns accu-
rately. Abeles (1979) presents an hypothesis for the interrelationships of horses with
black lower legs, manes and tails, and bodies of reddish brown, mixed reddish brown
and black, or uniform black. Although no segregation data are presented, her expe-
rience is consistent with the reddish brown and black mixtures (which she refers to
as brown) dominating the clear reddish browns (her bay), which in turn dominate
the uniformly black bodies (her black). This is perhaps explained by the interaction
of the Agouti and Extension loci. These loci have long been postulated as impor-
tant in controlling common horse colors, although the hypotheses concerning their
interactions have varied over the years (Odriozola, 1951; Jones, 1982; Sponenberg,
1996).
The existence of a uniform black in horses that is recessive to bay has long been
acknowledged, and accounts for the majority of black horses. This black is assigned
to the Agouti locus in keeping with principles of homology (Wilson, 1910; Wright,
1917; Adalsteinsson, 1976; Lauvergne et al, 1991).
The existence of a dominant black in horses has also long been speculated
(Gremmel, 1939; Castle, 1940a, b, 1951a, b, 1954). The evidence for a dominant
mechanism producing black has been the production of bay or brown foals from
the mating of two black parents. Such instances have usually been from studbook
data, which are suspect owing to the possibility of inaccuracies in the registrations
of animals, or in mistakes of parentage. Another problem with studbook data
for the colors bay, brown and black is one of definition. Various breeders may
have different mental images of these three and consequently studbook records are
probably inaccurate at this level of detail. Dreux (1966) presents evidence for the
existence of a dominant gene contributing to black or dark bay phenotypes in the
French Trotter breed, and mentions that this is rare in horses of European breeding,
having been introduced through an imported American Trotter stallion.
Currently, documentation best fits with the view that the Agouti locus comprises
two alleles, with bay dominant to black (Sponenberg, 1996). A third allele, black and
tan (!4! is fairly poorly documented as a cause of seal brown, and is intermediate
in dominance to the other two alleles. The Extension locus likewise has two
well-documented alleles, wild type dominant to chestnut (Sponenberg, 1996). The
recessive chestnut allele, when homozygous, results in a uniformly reddish brown
phenotype (including lower legs, mane and tail), and is epistatic to the Agouti
locus. Past literature has postulated intricate relationships with these two loci,
with no clear epistasis of one over the other, and with these intricate interactions
explaining the differences between bay and brown, and between shades of chestnut
(Jones, 1982). The Extension locus may well be the site of an allele that provides
for the intermixture of black hairs into the red body coat of some bay horses, which
translates into the difference between bay and brown. This is consistent with Abeles
(1979), even though segregation data are lacking in that report.
This report discusses data that support the existence of an additional Extension
allele, for dominant black, in the horse.
MATERIALS AND METHODS
Production data from a black Arabian stallion (Serr Ebony Star) were evaluated
for foal color production with various colors of dams. The production data from
the black sire (Shilosmidnight) and the dark brown dam (PRF Gali Gaeraff) of this
stallion were similarly evaluated.
RESULTS
Results are summarized in table I. The black stallion was mated to mares of
various colors. Following mating to bay mares he produced seven black and five
’near black’ foals (’near black’ is used to describe horses that are black save for a
few tan or red hairs in muzzle or flank, or those horses that are very dark but for
which it is possible to discern a subtle difference in the color of body and the lower
legs, mane and tail). Following mating to black mares he produced nine black foals.
Following mating to chestnut mares he produced ten black and six ’near black’
foals.
The sire of this stallion was black and was mated to black mares to produce
two black foals. Matings to bay mares resulted in three black and three bay foals.
Matings to chestnut mares resulted in one black and two bay foals.
The dam of this stallion was dark brown, and was mated to a black stallion to
produce one black foal and three ’near black’ foals. A mating to a chestnut stallion
produced a ’near black’ foal.
DISCUSSION
The problem of nomenclature concerning dark horses is a concern for this study.
In the Arabian breed many breeders are very strict in their use of the term black,
and reserve it for horses with black hairs only, excepting white marks due to spotting
patterns. Arabian breeders do not use the term brown, and so horses that are nearly
but not quite black are usually referred to as black bays, or even simply as bay.
These are, however, very distinct from the usual bright reddish brown bay that
is typical in the breed. Black bays in the Arabian breed are easy to confuse with
black horses, especially without close and detailed examination, and are difficult to
confuse with bright reddish brown bay horses, which are common in this breed. The
grouping of blacks and near blacks, or black bays, as one group is therefore logical,
since this is how these colors appear to most observers. These three are very close
to one another visually, if not in terms of nomenclature.
The results of the mating of the horses of this study is consistent with the
segregation of a dominant allele for black. The mating of the stallion to bay mares
for 12 foals can be tested for goodness of fit for the hypothesis that this stallion is a
recessive black. If that were the case, and if all the bay mares were heterozygous for
this usually rare allele, then X2 = 12, 1 df, P < 0.005. This deviates significantly
from the expectation, even postulating the most favorable condition of all the bay
mares being heterozygous for a rare allele. These results are therefore inconsistent
with the segregation of a recessive allele for black, but are consistent with the
stallion being homozygous for a dominant allele causing black color.
The sire of this stallion was black, while the dam was dark brown. Both needed
to have had the allele causing dominant black if the stallion is indeed homozygous,
as is supported by the segregation data. The mare always produced black or near
black foals when mated to a black stallion. She also produced a near black foal
when mated to a chestnut stallion, even though she is an obligate heterozygote for
dominant black by virtue of having had a bay sire. The sire of the stallion produced
equal numbers of bay and black foals when mated to bay mares, which is further
evidence that a dominant gene is segregating in this family of horses. The sire of the
stallion is heterozygous for this gene, even though both of his parents were black.
A few of the foals in this study were born a color resembling that of foals that
ultimately prove to be bay. These foals darkened with age, ultimately becoming
black or nearly so. Some of the blackest of the horses in this study were born with
this light bay foal coat, only to darken later. It is fairly typical for black horses
(assumed to be of the recessive Agouti genotype) to be born an ashy grey color, to
darken later. Whether or not the differences in foal coat are consistent with these
two different genetic mechanisms that cause black horses is undetermined.
The allele causing the black phenotype in these horses is most likely the dominant
black (ED) at the Extension locus. This allele is documented in numerous other
species as a cause of uniformly eumelanic phenotypes (Searle, 1968). It appears
to be arguably incomplete in its action in some instances, causing dark brown or
near black instead of truly black individuals. In foxes, for example, the ED allele
is frequently incompletely dominant resulting in three color classes from this allele
and the wild type allele (Adalsteinsson et al, 1987). The dominant black allele may
be the genetic mechanism behind some of the very dark phenotypes in horses that
are not truly uniformly black. The differences in genotype between the black and
nearly black horses are uncertain, since in this data set it is conservative to assume
that most, if not all, of these had the Agouti locus bay allele, and its presence or
absence can therefore not be responsible for the subtle differences in the black and
near black horses. Likewise, most of these foals had to have been heterozygous for
the dominant black allele, and this genotype includes both black and near black
individuals. The dominant black allele appears to be rare in most breeds, since in
most breeds black does segregate as a recessive.
REFERENCES
Abeles HM-S (1979) A coat of many colors. Equus 17, 30-38
Adalsteinsson S (1974) Inheritance of the palomino color in Icelandic horses. J. Hered. 65,
15-20
Adalsteinsson S (1976) Colour inheritance in Icelandic Ponies. In: First Int. Synzp. on
Genet. and Horse Breed., Royal Dublin Soc. Dublin, 17-18 September 1975, 42-49
Adalsteinsson S, Hersteinsson P, Gunnarsson E (1987) Fox colors in relation to colors in
mice and sheep. J. Hered. 78, 235-237
Castle W (1940a) Mammalian Genetics. Harvard University Press, Cambridge, Mas-
sachusetts
Castle W (1940b) The genetics of coat color in horses. J. Hered. 31, 127-28
Castle W (1951a) Dominant and recessive black in mammals. J. Hered. 42, 48-50
Castle W (1951b) Genetics of the color variety of horses. J. Hered. 42, 297
Castle W (1954) Coat colour inheritance in horses and in other animals. Genetics 39,
35-44
Dreux P (1966) Contribution a 1’etude du gene E chez le cheval domestique. Ann Genet,
9, 168-170
Gremmel F (1939) Coat colors in horses. J. Hered. 30, 437-45
Jones WE (1982) Genetics and Horse Breeding. Lee and Febiger Publishers, Philadelphia
Lauvergne JJ, Silvestrelli M, Langlois B, Renieri D, Poirel D, Antaldi GGV (1991) A new
scheme for describing horse coat colour. Livest. Prod. Sci. 27, 219-229
Odriozola M (1951) A los colores del caballo. Publicaciones del Sindicato Nacional de
Ganaderia, Madrid
Searle A (1968) Comparative Genetics of Coat Colour in Mammals. Logos, London
Sponenberg DP (1996) Equine Color Genetics. Iowa State University Press, Ames, Iowa
Wilson J (1910) The inheritance of coat colour in horses. Proc. Royal Dublin Soc. 12,
341-348
Wright S (1917) Color inheritance in mammals VII: The horse. J. Hered. 8, 560-64.


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Dominant black in horses

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