CAVY GENETICS

Peter and Cell Herman

Caviaria Rusticana
Las Cruces, NM
Please note that this article was written for the American Cavy Breeders Association Guide Book
and is copyrighted. It may be printed/copied for personal use. It may not be reprinted in any
publication whether distributed free or for profit without the express written consent of the authors and
the ACBA.

The word genetics causes panic in many cavy breeders. This need not be the case. A few terms
and concepts need to be understood first, then all genetics problems become variations or
derivatives or the original principles.
The first concept is that every trait is controlled by a pair of factors called genes. One gene
comes from. the father and the other from the mother. The genes arecarried on microscopic structures
called chromosomes. There are many genes on a chromosome, and many chromosomes in each
cell of an organism. Every cell in a cavy (except sperms and eggs)
contains 64 chromosomes, 32 from the mother and 32 from the father. The genes are arranged on the
chromosome in a specific order so thatevery trait has its own particular spot on a chromosome. This
spot is called a locus. At each locus there are two or more choices called alleles. For example, at the
coat length locus, there are two alleles possible, one for long and one for snort. When the two alleles
at a given locus are the same, the animal is homozygous. When they are different the animal is
heterozygous. In breeders terms, homozygous equals pure breeding and heterozygous
equals hybrid for the trait in question.
As many of you are aware, when a Peruvian is crossed to an Aby, all the babies will have
short coats. This occurs because of dominance. Very frequently, one allele will mask or dominate
the expression of another allele at the same locus. The masking allele is called dominant and the
masked allele, recessive. Thus, in our example, the babies had a gene for long from the Peruvian
parent, and a gene for short from the Abyssinian parent. The gene for short is dominant over
the genefor long, making all the babies appear short coated. You can see from this ttat an animal
can look like a purebred and be carrying genes that make it otherwise.
In general, a capital letter is used to represent a dominant trait and a small letter to represent
the recessive. We will use the letter L to represent this locus. L = short, and I = long. If we diagram the
cross made in our example, it would look like this:
Peruvian (ll) x Abysslnian (LL)
I
V
Abyssinians (Ll)

If we were to take two of the so-called "Abyssinian" babies from this cross and breed them
together, we would get both Abyssinians and Peruvians in an approximate ratio of three
Abyssinians to one Peruvian.
 In genetics, the way an animal looks is called its phenotype. The genetic make-up of an animal
is its genotype. The ratio of phenotypes is 3 short : I long, while the genotypic ratio is 1 LL:2 Ll:1 ll.
At this point, let us warn you that we use the term "Peruvian" and "Aby" only loosely in describing
the phenotypes of offspring of such a mating. There are many modifying factors other than the gene
L which distinguish a good show Aby from a good show Peruvian.
When looking at Peruvians and Abyssinians can see that both breeds have rough coats,
whereas Americans and Silkies both have smooth coats. This rough versus smooth character
is controlled at the rpugh, locus. The two alleles are the dominant rough (R) and the recessive
smooth (r). If we cross an American (rr) with an Abyssinian (RR) we will get all rough coated babies.
If we then cross these babies, we will have offspring produced in the ratio of three Abyssinians to one
American, or three rough to one smooth.
Thus you can see that an animal which shows a dominant trait can either be pure for that trait
(homozygous) or carry the recessive as well as the dominant (heterozygous). In order to express
the recessive trait, an animal must be homozygous for it. This should help explain why occasionally
a breeder will breed two Abyssinians together and come out with an American, or two Peruvians
together and come out with a Silkie. In these cases, both the mother and father must carry the
smooth gene (r). We diagram these crosses

Finally, after much introduction, we get to the genetic makeup or each of the breeds. As you
can see from our examples in the introduction, each breed has its own particular combination of
alleles at the rough and long loci. Since some contain dominant characteristics, more than one
combination of genes (genotype) can result in the appearance (phenotype) that we associate with
each breed.
First, let us consider the American, with its smooth, short coat. As we have seen,
smooth Is recessive, so all Americans are rr. They can be either Li or LL at the long locus. Thus, a
purebred American is rrLL, whereas a Silkie-carrying American is rrLl.
Since toe Peruvian has a long coat which is a recessive, all Peruvians are ll. A pure Peruvian
is RRIl, while a Silkie-carrying Peruvian is Rrll.
A purebred Abyssinian is RR for rough and LL for short. Other combinations which look like
Abyssinians are: RRLI (Peruvian-carrying), RrLL (American-carrying), and RrLl (it is possible to get
the four breeds by crossing two of these together). The quality of rosetting in Abys is controlled, in
part, by a locus known as rough modifier. This gene, M, suppresses the formation of rosettes. Show
Abyssinians are RRmm. RrMM animals can look completely smooth as the result of the rosette
suppressing activity of MM.
Silkies are genetically true-breeding. Since they are both long and smooth coated, they must
be homozygous recessive for both characters, making their genotyperrll. There are some rare
cases where Silkie-looking animals contain R genes, having the genotype RrMMlI.
The newer breeds, Cresteds, Teddies, and Satins are all genetically related to Americans in
that they are rrLL. Each carries an additional, characteristic allele at other loci. Cresteds carry the
gene St for star. This gene is a dominate character which produces a rosette (star) in the center of
the forehead. Star also produces a white spot on the crest about 50% of the time in the absence of
other spotting factors. Star alone does not give you a show Crested as any Crested breeder will
woefully tell you. Teddies carry the recessive alleles tt. This recessive trait causes the hair shaft to
kink. Satins carry the recessive alleles snsn. The sn allele, when homozygous, causes a thinner
hair shaft with a cleared shell, as in the Satin rabbit or mouse. Since all this gets pretty confusing,
we'll make a table of just those loci which affect breed.

LOCUS

BREED Rough Long Star Teddie Satin

American rr LL stst TT SnSn

Silkie rr ll stst TT SnSn

Crested rr LL StSt TT SnSn

Teddie rr LL stst tt SnSn

Satin rr LL stst TT snsn
Abyssinian RR LL stst TT SnSn
Peruvian RR ll stst TT SnSn

Variety, or coat color, is controlled by the same sets of genes in all breeds. Remember that
the way the animals actually look will not necessarily be the same among breeds, even if they carry
the same major genetic information for color. For example, in Americans you are looking primarly at
top coat. In Please note that these genotypes only apply to pure breeding animals. As we saw
before animals with a breed phenotype are not necessarily pure breeding. In Abyssinians, you see
top coat, but the undercolor is exposed by the rosetting. In Peruvians and Silkies almost all you see
is undercolor since the hairs are continually growing.
All cavy colors are made from two basic pigments, black and red. The Black series colors are
black, chocolate, lilac, and beige. The red series colors are red, orange, and cream. All the varieties
are comprised of these colors or white, the absence of color. We'll start by giving a list of the major
coat color genes with a brief description of what they do. After that we will go through the varieties
and venture a guess as to possible genotypes that yield desirable show phenotypes.

LOCUS Gene (allele) Symbol Function

Name
AGOUTI Agouti A A causes ticked hairs to be produced in the presence of b
base derived from the black series of colors and a tip deri
belly color is the same as the tip color, appearing untippe
Ticked Belly Agouti Ar Ar produces the solid agouties. They have ticked hairs all o
back color are the same. Ar is recessive to A
Non-agouti a Animals with black series pigment that are non-agouti are
is aa, Aa or AA because black pigment must be present t

EXTENSION Extension E Allows The production of black pigment to extend throug

animals of black series colors or agoutis.
Partial Extension ep Allows black pigment to extend partially through the coa
series and red series pigments

Non- Extension e Restricts black pigment to the eye, leaving a red series c
and eP.
BLACK Black B Causes normal black pigment granules resulting in black
.
Chocolate b Modified black pigment granules are produced by bb an
chocolate. Gives eye a ruby cast.

PINK-EYE Non-Pink-Eye P Normal pigment is produced.

Pink-Eyed Dilute p The presence of pp causes an 80% reduction in the amo
produced, with little effect on red. The reduction of black
lilacs, and chocolates to beiges. The eye color is pink be
pigment allows the blood in the eye to show through. Re
REQ and RE Creams respectively.

ROAN Roan Rn Show Roans are Rnrn. Homozygous RnRn animals are
frequently deformities.

Non-Roan rn Non-roaned animals are rnrn

WHITE SPOTTING White Spotting s Animals with ss usually have over 50% white. Animals wi
white
Non Spotted S Animals that are SS are not usually spotted. On occasio
a white blaze.

These genes, and those of another locus, the C locus, interact as the major factors that
produce the color varieties that are in the show standard. We should emphasize that there are
many unknown, or uninvestigated modifiers which can cause an animal to be close to the show
standard while another animal of the same basic genotype without the modifiers might be pet stock.

The C locus is quite complex because there are five possible alleles. To make matters worse,
there is no simple dominance. There are a number of different heterozygotes of lower C alleles that
have the same phenotype. Below is a table that shows basically what effect the different C alleles
have on black and red pigment production.

ALLELE SYMBOL RED BLACK EYE COLOR

Full Color C ++++ +++++ Dark

Dark Dilute ck ++ ++++ Dark

Light Dilute cd +++ +++ Dark

Ruby-Eyed Dilute cr - Dark with ruby

cast
Albino/Himalayan ca - ++ Pink
(on points only,
in presence of E
or eP)
 The alleles of the C series interact in sometimes strange ways. Below is a table of the relative
amount of red and black pigment (percent relative to full color) produced by the various
combinations of C alleles in the absence of other modifiers. (extracted from Searle, 1964)

GENOTYPES
PIGMENT caca crca crcr cdca cdcr cdcd ckca ckcr ckcd ckck C_
Red 0 0 0 30 30 40 30 30 40 35 100
Black 20 40 80 35 70 60 80 90 90 95 100

The genotypes we have listed are our best guess of the desired show genotypes based on
extensive reading, a dozen years of breeding experience, and lengthy conversations with other
knowledgeable breeders. they are by no means the only possible genotypes to achieve the desired
show phenotypes. As we have said before, unknown or unstudied modifiers can have tremendous
influence on an animals appearance.
For convenience we have grouped the varieties genetically rather than by the Standard of
Perfection (SOP). The Solid varieties of the SOP have been moved to Agouti or marked, depending
on their genetic affinities.

SELF COLORS
The Standard lists nine self colors. All the other varieties are made up of some combination
of these. In the chart below we have listed only the important loci for each variety. The
notation cx indicates an unspecified lower C allele.
Variety Genotype Comments
Black EE BB SS CC PP C produces the darkest color, unknown modifiers
strengthen the under color
Chocolate EE bb SS CC PP bo converts black to chocolate
Lilac EE BB SS cxcx pp pp dilutes black in eye and coat
Beige EE bb SS cxcx pp pp dilutes chocolate in eye and coat
Blue (note no There is no agreed genotype for blue. It
longer in the SOP probably involves heterozygosityat the C locus
and modifiers
as of 2000
Red ee bb SS CC PP bb tends to produce proper ear and foot pad
color, C produces most intense top color,
unknown modifiers influence undercolor
Red- ee bb SS cxcx pp lower C allele desirable for orange color
Eyed Orange
Cream (Dark- ee bb SS cxcx PP The presence of pp converts Dark-eyed to Pink-
eyed) ee bb SS cxcx PP eyed Cream.
Cream (Pink- The best color is produced by one of these
eyed) genotypes cdca, ckca. cdcr. Creams with the
genotype cdcd, ckck & cdck are usually buff
colored or "hot creams".
White ee caca ee produces clear points

AGOUTI VARIETIES
All agouti varieties must carry E to express the tipped agouti hairs throughout the coat.
Animals with the genotype epep or epe will have spots of agouti color. There are two basic agouti
colors, Golden and Silver. There are Solid Goldens and Solid Silvers as well. Below are the
genotypes
Variety Genotype Comments
Golden Agouti AA EE BB SS CC the basic agouti color and the genotype of
wild cavies
Silver Aoouti AA EE BB SS crcr cr eliminates the red color leaving a white tip.
It also dilutes the black base color to silver
Solid Golden ArAr EE BB SS CC Ar causes the tipped belly hair
Solid Silver ArAr EE BB SS crcr cr eliminates the red color leaving a white tip.
It also dilutes the black base color to silver.
. Ar causes the tipped belly hair.
Dilute Agouti These animals have any of the diluting
factors (pp,cx,bb) which act on black or red
pigment. Note: the combination of tip and
base color must appear in the SOP to
be showable

MARKED VARIETIES
There are several genetically related sets of varieties in this group. An Examination of the
chart below will point them out.
MARKED VARITIES
Variety Genotyp~ Comments
Himalayan caca EE BB PP EE. EE and SS are required to insure a
complete set of points. BB and PP improve
point intensity
Tortoise/Brindle ep_ BB PP SS Unknown modifiers determine the
distribution of patches. Animals that
areepe have more red on the average
than epep animals.
Tortoise Shell ep_ BB PP ss or Ss The presence of s increases the clarity of
and White patching. Animals with ssusually have more
white than Ss animals.
Broken Color Any color and sS or ss Animals with ss usually have more white
(with White) than Ss animals.

Dutch Any Color with Ss Dutch pattern is not fixed by a known locus.
There are unknown modifiers that increase
the tendency toward getting the patches in
the right place
Broken Color ep with A_ or bb,or pp, (s) These are basically modified Tortoise Shells
(2 colors no that carry genes that modify the black or
White, or 3Colors) red. White is added by s
Roan Rnrn with any color Amount of roaning and pattern controlled by
modifiers. Heads are usually not roaned
Dalmation Rnrn with any color Amount and clarity of spotting controlled by
modifiers.

Cavy Genetics - Basics of the Breeds

Finding good information on breed and color genetics for cavies is MUCH more difficult than
rabbits. The following is a compilation of what I've managed to pull together. Additional
information/sources GREATLY appreciated! Written for the reader who already knows rabbit color
genetics (or has at least read my rabbit color genetics 101).
The ARBA/ACBA Standard of Perfection recognizes cavies in 13 breeds. Differences among the
breeds are not as pronounced as the differences among rabbit breeds -- in many cases a single
gene mutation is responsible for the distinction between the breeds. Basic genetics suggest that
crossing breeds would thus result in perfectly showable animals, but doing so is not generally
recommended as hybrids often results in animals with intermediate characteristics or mismarkings
(such as too few rosettes).

Coat Length - L
Short coat (as in Americans, white crested, and teddies) is a dominant trait - designated L. Long
coat (as in Silkes, Peruvians, Coronet and Texel) is recessive, designated l. Hybrids - Ll - usually
have short coats, but may be slightly longer than the purebred LL.
Rosettes - R
The dominant R allele causes rosettes (as in Abyssinians and Peruvians). Breeds without rosettes
(American, white crested, teddy, Silkie, texel, and coronet) are rr. Modifiers genes (separate,
unknown genes) are needed to get the correct number and placement of rosettes. Abyssinians
should have 10 rosettes. The long-haired 'sweeps' of Peruvians are also a modified rosette - only 6
are typical.
Star - St
The dominant Star allele (St) creates a single 'rosette' in the middle of the forehead (as in white
crested and coronet). Breeds without the crest or coronet have the recessive genotype
(stst). Additional modifier genes are responsible for the color and shape of the star. Hybrids (Stst)
will have a coronet, but it is often poorly shaped due to the loss of the modifiers.
Teddy - T
The recessive teddy allele (t) causes a crimping in the hairs, resulting in a plush coat that stands on
end. All other breeds have the dominant T- genotype.
Rex - Rx
The recessive rex allele (rx) causes a fur type similar to that of a rex or minirex rabbit. Longhaired
cavies with the rxrx genotype (essentially silkies with rxrx rather than RxRx) are texels, which have
long ringlets for fur. Shorthaired cavies with the rxrx genotype are called rexes - these are accepted
in Britain and other countries, but not (yet) by the ACBA. Similarly, coronets with rxrx are called
merinos and Peruvians with rxrx are called alpacas - none are (yet) accepted by ACBA.
Satin - Sn
The recessive satin allele (sn) imparts a glowing satin sheen to the coat - over and above the
normal luster of the healthy coat. Satins are accepted in American satin, Abyssinian satin, Silkie
Satin, Peruvian satin, and Teddy satin.
Breed Genotypes
Taking American as the 'base type':
American - L-rrststT-Sn-Rx-
Abyssinian - L-R-ststT-Sn-Rx- (American plus R [rosettes] and modifiers)
White crested - L-rrSt-T-Sn-Rx- (American plus St [star, crest or coronet] and modifiers)
Teddy - L-rrststttSn-Rx- (American plus tt [teddy])

Silkie - llrrststT-Sn-Rx- (long-haired American)

Peruvian - llR-ststT-Sn-Rx- (long haired Abyssinian with reduced rosettes)
Coronet - llrrSt-T-Sn-Rx- (long-haired white-crested [except other crest colors are allowed])
Texel - llrrststT-Sn-rxrx (Silkie with a ringlet coat, a cobby body and a broad face) [NOT a long-
haired teddy, different gene responsible]
American satin - L-rrststT-snsnRx- (American plus the satin gene)
Abyssinian satin - L-R-ststT-snsnRx- (Abyssinian plus the satin gene)
Teddy satin - L-rrststttsnsnRx- (Teddy plus the satin gene)
Silkie satin - llrrststT-snsnRx- (Silkie plus the satin gene)
Peruvian satin - llR-ststT-snsnRx- (Peruvian plus the satin gene)

Other genetic combinations exist, but are not accepted by ARBA. Some are accepted elsewhere in
the world. A few of the more common ones...

English crested -- unlike our White-crested, English crested are not required to have the modifier
which turns the crest white. So the crest appears the same color as the body.

Rex - L-rrststT-Sn-rxrx (similar to a teddy, but slightly rougher texture)

Merino - llrrSt-T-Sn-rxrx (a coronet with ringlets - or a texel with a crest)
Alpaca - llR-ststT-Sn-rxrx (a curly Peruvian - or a texel with rosettes)
Somali - L-R-ststT-Sn-rxrx (a rex with rosettes)

Crested satins - L-rrSt-T-snsnRx-

Coronet satin - llrrSt-T-snsnRx-
Texel satin - llrrsrstT-snsnrxrx

Sheba Mini-Yak - llR-ststT-Sn-Rx- (a variation on Peruvian retaining more rosettes due to a

recessive modifier resulting in a more 'tousled' appearance) crosses between abyssinians and
longhaired cavies often result in an appearance closer to the sheba than to the Peruvian.
Ridgeback - L-R-ststT-Sn-Rx- (a variation on Abyssinian in which modifier genes restrict the
rosettes to produce a single ridge along the spine)

Lunkarya - llrrststT-Sn-Rx- +Cu- a new gene discovered in Sweden (Cu) is the basis of the
lunkarya, which looks like a texel with a looser curl. This is a separate gene from Rex or teddy,
though with a similar 'rexoid' look.

Swiss (aka Swiss teddy) - L-rrststT-Sn-Rx- TWO separate novel genes. Recessive Sw (swiss) and
a dominant semi-longhair gene combine to create a 'teddy' with an intermediate hair length with a
'puffball' effect -- often the hair on the rear is longer than on the front.

Skinny - a separate recessive gene which causes cavies to be born (and remain) hairless. Skinny
pigs often grow some sparse hair on the nose and feet. This gene originated in teddies, so many
skinnies are hiding teddy genotypes.

Baldwin - a separate recessive gene which causes cavies to lose their hair (usually at a few weeks
old). Baldwins originated in white-crested cavies and often hide white-crested genotypes. Skinny
and Baldwin are separate genes -- crossing skinnies and baldwins results in cavies that have hair
(but are carriers for both types of hairlessness).

Lakeland - a guinea pig which is born with teddy fur, loses it's hair at ~3 weeks old, and then
regrows its hair. This is a carrier for the skinny gene. Not all skinny carriers are Lakelands, but all
Lakelands are skinny carriers.

Werewolf - a guinea pig which is born hairless, but grows a significant amount of hair on its head,
possibly shoulders and feet. This is a genetic skinny pig, but has modifiers that allow some hair
regrowth.

Why don't we have rexes, merinos, alpacas, coronet

satins or white crested satins?
ARBA/ACBA decided that a single gene difference (e.g., Rx and Sn) which are easily bred to the
base type (as in breeding an American to an American satin) should not be sufficient to constitute a
separate breed. Those breeds which had been accepted prior to this ruling (texel and the accepted
satins) were grandfathered in - but no new satin or rex breeds will be accepted unless the rule
changes.
Cavy Color Genetics
Way more genes, alleles and co-dominant traits, and gene interactions than with rabbits...hang on.
Black/Chocolate (B/b)
Simplest first. For those used to rabbit color genetics, let's do the B gene first. It comes in two
forms (alleles). B for black and b for chocolate. Cavies that are BB or Bb are black, bb are
chocolate (chocolate also adds a ruby glow to the eyes). Just like with rabbits, we can do lots of
things with other genes to modify these two base colors.
Pattern Gene (A series)
Similar to the pattern series in rabbits, but we add an extra allele and change the rules just a bit.

A is the most dominant gene and results in an agouti animal very similar to the agouti pattern in
rabbits -- rings of chestnut and black (with B-) or tan and chocolate (with bb) and a solid 'belly band'
with just the chestnut/tan color.

Ar is recessive to A but dominant over the following alleles. Ar- animals look a lot like agoutis - until
you flip them over. The ring pattern in these animals extends through the belly region, so the hair
on the entire body looks the same. These animals are called solids.

At is the tan pattern allele. It gives you an animal that looks a LOT like a tan pattern rabbit - solid
colored back (e.g., black or chocolate), tan belly, eyes circles and tan spots by the ears. Tan is
dominant over self (the last allele in the series - below) but CO-dominant with A and Ar. Thus an
animal with genotype AAt looks like an agouti except that you still get the tan eye circles and spots
by the ears (a DQ'ed type). ArAt also looks like an agouti except you get the eye circles and pea
spots (also a DQ). AtAt and Ata both give the tan pattern. Tans are currently accepted ONLY in
American and teddy cavies.

a is the final allele in the series. Just like in rabbits, it is the self gene, resulting in a solid colored
cavy.

So far:
AAB- and AArB- and AaB- = agouti
AAbb and AArbb and Aabb = chocolate agouti (accepted as one of several dilute agoutis)
ArArB- and AraB- = solids
ArArbb and Arabb = chocolate solid (accepted as one of several dilute solids)
AAtB- and AAtbb and ArAtB- and ArAtbb = unaccepted mismarked agouti and solids
AtAtB- and AtaB- = Black Tan (accepted in Americans and Teddies only)
AtAtbb and Atabb = Chocolate and Tan (accepted in Americans and Teddies only)
aaB- = self black
aabb = self chocolate

Prior to the addition of tan, the three A-alleles in cavies were treated as a simple dominance
series. The A-series is actually entirely co-dominant. If the two alleles of a heterozygous pair (two
different alleles in the same animal) give different instructions -- one to produce eumelanin (base
color) and the other to produce phaeomelanin (the tan or white markings) - phaeomelanin always
wins. The 'recessive' a allele codes only for eumelanin and will be overwritten by any other allele
that calls for phaeomelanin. Since the solid allele (Ar) has phaeomelanin in fewer locations than
agouti (A) and no phaeomelanin in any locations that A has eumelanin, the Ar allele acts as though
it were recessive to A. But in the case of the tan allele (at), the tan pattern has phaeomelanin in
places where the A and Ar have eumelanin (most notably the peaspots) as well as eumelanin in
places where A and Ar have phaeomelanin (most of the coat except tipping).

CAUTION - with the rise of popularity in tan pattern cavies, breeders of agoutis and especially
solids should be on the lookout for mismarked animals with peaspots. This is especially true for
breeders of agoutis -- it is now increasingly possible to have a cavy sold as an agouti that is a
genetic solid carrying tan -- Arat cavies appear agouti (except that it has peaspots which are small
enough to be easily overlooked) even though they do NOT have any agouti genes!

Pink-eyed Dilute (Pp)

Unlike with rabbits, pink eyes aren't restricted to white animals. The recessive p gene lightens the
black portions of the coat color (but not all the way to white) and causes pink eyes. It doesn't
lighten red portions of the coat.

Lets start with the selfs (aa) - Black becomes lilac (unlike rabbits, where lilac is the dilute of
chocolate, in cavies lilac is the pink-eyed dilute for black) and chocolate becomes beige (with pink
eyes).

So...
Black = aaB-P-
Chocolate = aabbP-
Lilac = aaB-pp
beige = aabbpp

Let's tackle tan combinations next..

AtAtB-P- and AtaB-P- = Black tan

AtAtbbP- and AtabbP- = Chocolate tan
AtAtB-pp and AtaB-pp = Lilac tan
AtAtbbpp and Atabbpp = beige tan

[tans accepted only in Americans and teddies]

Solids with dark eyes

A-B-P- = agouti
A-bbP- = dilute agouti (dark eyes) (chocolate agoutis)
Ar-B-P- = solid
Ar-bbP- = dilute solid (dark eyes) (chocolate agoutis)

Solid Pink-eyed dilutes

A-B-pp = dilute agouti (pink eyes) (lilac agoutis)
A-bbpp = dilute agouti (pink eyes) (Beige agoutis)
Ar-B-pp = dilute solid (pink eyes) (lilac agoutis)
Ar-bbpp = dilute solid (pink eyes) (Beige agoutis)

Note - Dilute agoutis and solids are shown as a group with only a subset of color combinations
acceptable.

Please note - In cavies the eye color caused by the pink-eyed dilute gene (pp) or the himalayan
gene (below ch) is referred to as PINK-eyed - even though it looks pretty much identical to what we
would call 'RED-eyed white' in rabbits. This is most likely because in cavies the cr gene causes
RED eyes of a deep red shade (closer to what we call 'ruby cast' in rabbits). In rabbits, red eyes
are found only in whites and himalayans and are due to the action of either the himalayan (ch) or
albino (c) gene -- both on the C-series. In cavies, pink-eyes are due to EITHER the p gene
(described here) or the himalayan gene (ch - described below) - the true albino gene c does not
exist in cavies.
The blue tan color recently accepted in American cavies is attributed to a genotype pgpg - which is
an intermediate allele in the P series.
P>pg>p

The pg allele lightens black to a slate-blue color intermediate between black and lilac. It lightens
chocolate to a caramel color intermediate between chocolate and beige. The pg allele lightens eye
color to a slate-grey or deep ruby (NOT blue). In the US, ACBA accepts the pgpg genotype ONLY
in blue tan and marten americans and teddies. NO caramels are accepted.

The US is the only place in the cavy world to refer to this color as blue. Elsewhere it is called
slate. The rest of the cavy world reserved the term 'blue' in the hopes that someday a D-mutation
(giving blue-grey eyes) would arise. Their patience has paid off as a new mutation found in
tortoiseshells in Sweden appears to be true blue (dd) - but lots of work still needs to be done to
confirm that.
Non-extension - E series
Similar to rabbits in that this is how to get red color, but works a bit differently!

E is the dominant allele - EE results in the normal colors above.

ee is the recessive. As with rabbits, it extends the middle band and removes the base
(black/chocolate) color. Does not remove dark eye color.

As in rabbits, A-ee gives red. So does Ar-ee. Unlike rabbits, so does At-ee (no fox, just red). Also
unlike rabbits, so does aaee (no tort, just red).

In cavies, aaee is the preferred genotype for red-based colors. Reds with A, Ar or at often have
some dark pigment (smut) on their ears.

Under the ACBA standard, the non-extension 'reds' are broken into subgroups based on the
'dilution' (combination of B, C and P genes): red, red-eyed orange, gold, and cream (see below)

We've also got an intermediate allele in play - Ep or partial extension.

aaB-P-EpEp gives a color that is called tortoiseshell in cavies (but is what we would call harlequin in
rabbits) in which the animal has patches of black and orange (requires some additional modifiers to
make the patches look right) OR (depending on modifiers) can give brindle (interspersed black and
red hairs).

With chocolate, dilutes (pink-eyed above or C series below), agouti, or solid, EpEp is called broken
(broken in cavies can be any two or more colors, not just a color and white). Adding EpEp to tan
(At-) usually leaves small tan marks, which are not accepted.

Note that so far, I've just given the 'pure' E series genotypes - EE, EpEp and ee. That's because
the dominance in this series isn't so simple.

EEp animals often (but not always) have stray red marks (like Vienna carriers and Vienna marks in
rabbits).
Ee can also (less commonly and smaller when present) also result in stray red marks.

Epe pretty much looks like EpEp -- since the Ep already causes red patches, its hard to tell whether
or not the e is adding a few extra - but sources argue that an Epe cavy will be 'more red' than the
EpEp.

Recap - Like rabbits, the E series gene is responsible for red color. But it works much more like the
rabbit Vienna (BEW) gene - completely covering other colors and patterns and, when 'carried,'
occasionally showing though as stray marks.

The Dilution Gene - C series

Not at all like either the dilution gene (D series) nor the C series in rabbits. But it has an effect
somewhat similar.

Point #1 - the C series alleles all have a similar effect - lightening pigment color. They can operate
on the fur color, the eye color or both.
Point #2 - C, the 'normal color' allele is dominant and can hide any of the others.
Point #3 - the rest of the whole series is co-dominant - mixing alleles gives you intermediate
degrees of dilution.

The alleles
C is the normal color (as dictated by ABE and P genes)
ck is dark dilute. It lightens the coat a bit, but not the eyes (unless combined with pp).
cd is the light dilute (aka buff dilute). It lightens the coat significantly more but leaves the eyes dark
(unless combined with pp, which lightens the eyes)
cr is the ruby eyed dilute (aka Dark-eyed white dilute). It lightens the coat to white or near white
and shifts the eyes to a ruby red (dark eyes with a ruby cast, not pink).
ch is the himi gene, responsible for both himalayans and pink-eyed whites.

I'm going to start with the 'most recessive' lightest color first - ch.

Black + chch = aaB-P-E-chch = himalayan - as with the rabbit of that name, the result is a white
cavy with black points (nose, ears, feet).
Chocolate + chch = aabbP-E-chch = chocolate himalayan (can be shown, but the black preferred)
lilac/beige + chch = aa--ppE-chch = eye color fades to pink - point color often fades to white.
Red + chch = white (pink eyes acceptable in white). [I believe this includes all the reds that were
agouti, solid and tan as well as the selfs]
Agouti/solid +chch = Not showable (himalayan with very poor points)

I find precious few mentions of the ruby-eyed gene (cr). It's definitely responsible for the dark-eyed
whites (aabbP-eecrcr and aaB-P-eecrcr). It is also responsible for the silver agoutis and silver
solids. Added to a tan pattern, it results in the marten. It may also be responsible for the ruby cast
mentioned as acceptable in the following varieties: chocolate, cream, red, some dilute agoutis and
solids, dalmatian.

crch genotypes result in a slight lightening of the coat except over the points and a ruby cast to the
eyes. In it's purest form, this color is referred to as sable. Sables are not accepted in the US (here
they are just considered to be 'uneven colored selfs' and often disqualified for the ruby cast eyes) -
but are accepted elsewhere in the world.

The ck and cd alleles produce color dilutions that don't affect eye color and are definitely co-
dominant (interacting with one another).
ck/cd dilutions change black to silver and lemon and chocolate to cinnamon and cream (not
accepted as selfs, but possible as dilute agoutis - below)
ck/cd dilutions lighten the shade of lilac and beige selfs - these lighter shades are typically
preferred.
They interact with the non-extension series to produce lighter shades that are recognizably (and
accepted as) different...
black & lilac based reds lighten to golds (--B---eecxcx)
chocolate-based reds (--bbP-eecxcx) lighten to cream.
beige-based reds (aabbppeecxcx) lighten to red-eyed orange (ckcd) or pink-eyed cream (cdch)
A quick note on albino - The true albino gene (a recessive usually symboled c) which all by itself
(without the assistance of any other gene) completely shuts down the production of pigments (both
eumelanin and phaeomelanin) is unknown in cavies. Although the red-eyed white of rabbits
LOOKS identical to the pink-eyed white of cavies, REW rabbits are true genetic albinos, while PEW
cavies are not.

A note on the dilute agoutis and solids

ACBA accepts 10 color combinations in the category of dilute agouti (and dilute solid) these are
achieved by combining the chocolate gene (b), pink-eyed dilute gene (p), the buff dilute gene (cd),
and the ruby-eyed dilute gene (cr). All must be E- (ee turns completely red or white!).

Black series (BBP-)

with C- is not diluted at all - golden agouti = A-B-P-C-E- (shown as its own group)
lemon agouti (black with cream tips and dark eyes) - A-B-P-cd-E- (shown as part of the dilute agouti
group)
Note the cr dilution here is called silver agouti - A-B-P-cr-E- (shown as its own group rather than
part of the dilutes)

Chocolate series (bbP-)

chocolate agouti (chocolate with red tips and dark eyes with a ruby cast) - A-bbP-C-
cream agouti (chocolate with cream tips and dark eyes with a ruby cast) - A-bbP-cd-
cinnamon agouti (chocolate with white tips and dark eyes with a ruby cast) - A-bbP-cr-

Lilac Series (B-pp)

gold/lilac argente (lilac with orange tips and pink eyes) - A-B-ppC-
lemon/lilac argente (lilac with cream tips and pink eyes) - A-B-ppcd-
white lilac argente (lilac with white tips and pink eyes) - A-B-ppcr-

Beige series (bbpp)

gold/beige argente (beige with orange tips and pink eyes) - A-bbppC-
lemon/beige argente (beige with cream tips and pink eyes) - A-bbppcd-
white/beige argente (beige with white tips and pink eyes) - A-bbppcr-

For solids, replace the word agouti/argente with solid and the A- with Ar-. Keep in mind a dilute
solid (Ar-) is NOT a dilute self (aa)! Solids have the same color hairs over their whole body, but
each hair is made of two colors - base and tip.

White Spotted - S/s

The white spotted gene is a co-dominant gene used in the Dutch, broken and tortoiseshell & white
varieties. Its a pretty straightforward co-dominant system...SS is the 'normal' genotype with no
white spots. Ss produces an animal with less than 50% white -- anywhere from a small spot up to
half white. ss produces an animal that is more than 50% white. Rarely, this may give an all white
animal, but most often you get a white cavy with spots of color. Exactly how much white (within the
ranges above) as well as the pattern of the white patches are determined by modifier genes.

Broken is the most basic color pattern to rely on the white-spotted gene (note for cavies, color
combinations NOT including white - and so not using this gene - are also called and shown as
broken). Showable broken cavies in white + color (and the color can be any self, red, agouti or
solid - but not tan) must have a white patch at least as large as a 50 cent piece. The preferred
(i.e., needed to win) patterns are those close to 50-50 (half white, half colored) with large
patches. The truly ideal approaches the pattern we call harlequin in rabbits (only in this case
colored and white). Winning broken cavies can thus be either Ss or ss but are unlikely to be
produced at random (requires very careful selection from broken lines to get those ideal patches).

Tortoiseshell and white cavies (and other tri-colored cavies which are shown as brokens) also have
the white-spotted gene in play along with the Ep gene (harlequin - see above). As tri-colored cavies
should ideally have just 1/3 of their color white, they should be all Ss. Again modifier genes are
critical to get the preferred (winning) patterns.

Dutch marked cavies look a lot like Dutch rabbits. The Ss genotype gives the Dutch pattern, but
again, modifiers are needed to get the marks in the right places, so you are unlikely to produce a
Dutch-marked cavy out of brokens that aren't out of Dutch lines. Black Dutch are EE and red Dutch
are ee.

Roan - Rn
Roan is a co-dominant gene. The 'normal' cavy colors all have the recessive genotype rnrn. Rnrn
produces a roan kit in which the colored and white hairs are mixed together producing a speckled
pattern. RnRn produces white pups with ruby-blue eyes, called 'lethal whites'. Unfortunately, RnRn
kits earn the name 'lethal' -- often blind, with twisted teeth and other deformities. In most cases,
these deformities are severe enough to limit quality of life and lethals should be put down. For this
reason, roans should always be bred with a solid colored cavy (in the color you are trying to include
in the roan) and never with another roan. This is the same type of breeding pattern we use in
broken rabbits (breed broken to solid) when we try to avoid producing the unshowable charlies. But
here it is much more serious as those deformed pups have to be put down.

Dalmatian is a variation on roan - addition of some modifier genes cause the colored hairs to cluster
into spots rather than mix evenly through the coat. Because the roan gene is involved, you should
never breed dalmatian to dalmatian either.
Cavy Color Genetics 102
A quick reminder on terms...
Phenotype is the color you see. Genotype is a code for the genes you see AND the set you don't
(which might show up in later pups).

A gene is a segment of DNA coding for a particular trait. Genes come in pairs - one from the
mother and one from the father. A locus is a location on the DNA - for example the "A" locus is
where the A, Ar, At or a genes reside. An allele is a different form of a gene - for example the
melanin gene comes in two forms - black (coded B) and chocolate (coded b). Even though each
individual can only have two alleles for a locus, more than two alleles can exist in the population -
for example C, ck, cd, cr, and ca are all alleles for the C locus. Gene pairs can be homozygous in
which both genes in the pair are the same allele (e.g., aa) or heterozygous in which two different
alleles are included in the pair (e.g., Aa).

In a heterozygous gene pair, usually one gene shows and the other is hidden; the gene which
shows is said to be dominant and the one which doesn't is said to be recessive. Sometimes the
pair is codominant such that a heterozygous pair (e.g., Rnrn = roan) looks different than either
homozygous pair (RnRn is lethal white, rnrn is solid colored).

A purebred animal is one that (when crossed with another purebred of the same type) will produce
offspring that look like the parent. The term is most commonly used in reference to the breed - a
pair of purebred Abyssinians will produce only Abyssinian pups - they don't carry the genes for long
hair or no rosettes. The term can also be applied to most colors - a pair of purebred blacks will only
produce black pups - they don't carry the genes for chocolate, red or other colors. A handful of
colors - such as roan - can never be purebred for color.

The ACBA standard divides cavies into 5 groups - self, agouti, solid, marked and tan/marten. For
each breed, different colors of a group may be showable (or not) and they may be shown as a
single group, subdivided into separate varieties or subgrouped somewhere in between. For
example, Abyssinians are shown in 6 classifications: self, agouti, brindle, roan, any other solid, and
marked - while Americans are shown in 19 classifications: black, cream, red, white, any other self
(but only beige, chocolate, lilac, red-eyed orange, and gold), brindle, roan, dilute solid, golden solid,
silver solid, dilute agouti, golden agouti, silver agouti, dalmatian, dutch, himalayan, tortoiseshell &
white, any other marked (including broken and tortoiseshell), and tan/marten pattern.

Patterns called for in the breed standard - such as the white crest of the white crested are controlled
by separate genes and not considered in the discussion below.
Self Group
Self cavies have only one color in their fur - no markings, no rings. All nails must match and
generally the entire skin must also be a single color (e.g., whites with dark pigment in the skin of
their ears are disqualified). Cavies can be shown in 9 self colors: beige, black, chocolate, cream,
lilac, red-eyed orange, red, gold and white. Not all phenotypic selfs are genetic selfs! I divide the
selfs into two subgroups for discussion of genetics. Black, chocolate, lilac and beige are true
genetic selfs which always have aa as part of their color genotype. Crosses made within the true
self group (e.g., black to beige) will always produce selfs and can be said to be 'purebred for
self'. Red, red-eyed orange, gold, cream and white are non-extension selfs which alway have ee as
part of their genotype and which may or may not be genetic selfs - the non-extension genotype (ee)
can hide the effect of any of the pattern alleles causing a self appearance (red can hide agouti, solid
or tan). This group can be said to be 'purebred for non-extension'. Crosses made within the non-
extension self group will produce only non-extension selfs. However, crosses made between the
two 'self' subgroups may produce animals which are NOT self (e.g., crossing a black and a red can
give agouti!).

Black - Blacks are true genetic selfs (aa). They have at least one copy of the black pigment allele
(B) but may hide the chocolate allele (Bb). They have at least one copy of the dark-eyed allele (P)
but may carry the pink-eyed allele (Pp). They have at least one copy of the full intensity allele (C)
but can hide any of the other "C" alleles (Cck, Ccd, Ccr, Cca). They have at least one copy of the
full extension allele (E) but may carry non-extension (aka red as Ee). They cannot have the white
spotting allele (they are ss) or the roan gene (they are rnrn). The base genotype for black is aaB-P-
C-E-ssrnrn - if you have a black you know this much of its genotype for certain. A purebred black is
aaBBPPCCEEssrnrn. As a relatively dominant color, it is very difficult to establish that a particular
black cavy is purebred - but easy to disprove. If the parents of the black cavy were any color other
than black, the black cavy is NOT purebred for black. If any color other than black appears in the
last 3 generations, it is unlikely that the black cavy is purebred for black. If the cavy ever has a pup
that is any color other than black (when crossed to black, chocolate, lilac, or beige), it is NOT
purebred for black. A red, orange, gold or cream pup indicates the black parent (both parents) is
carrying non-extension (Ee). Bi-color pups (ejej or eje) indicate the black parent is carrying the
partial extension allele (Eej) OR the non-extension allele (Ee). Pink-eyed pups (of any color)
indicate the black parent (both parents) is carrying the pink-eyed allele (Pp). Chocolate, beige, gold
or cream pups indicate that the black (both parents) is carrying chocolate (Bb). Himalayan or pink-
eyed white pups indicate that the parent is carrying both pink-eyed and himalayan alleles
(PpCca). Dark-eyed white pups (crcr or crca) indicate the black parent is carrying the silver allele
(Ccr) or the himalayan allele (Cca).

Chocolate - Chocolates are true genetic selfs (aa). They have two copies of the recessive
chocolate pigment allele (bb) and can never hide black. They have at least one copy of the dark-
eyed allele (P) but may carry the pink-eyed allele (Pp). They have at least one copy of the full
intensity allele (C) but can hide any of the other "C" alleles (Cck, Ccd, Ccr, Cca). They have at least
one copy of the full extension allele (E) but may carry non-extension (aka red as Ee). They cannot
have the white spotting allele (they are ss) or the roan gene (they are rnrn). The base genotype for
chocolate is aabbP-C-E-ssrnrn - if you have a chocolate you know this much of its genotype for
certain. A purebred chocolate is aabbPPCCEEssrnrn. As a relatively dominant color, it is very
difficult to establish that a particular chocolate cavy is purebred - but easy to disprove. If any color
other than chocolate appears in the last 3 generations of the pedigree, it is unlikely that the
chocolate cavy is purebred for chocolate. If the cavy ever has a pup that is any color other than
chocolate (when crossed to chocolate or beige), it is NOT purebred for chocolate. A orange, gold
or cream pup indicates the chocolate parent (both parents) is carrying non-extension (Ee). Bi-color
pups (ejej or eje) indicate the chocolate parent is carrying the partial extension allele (Eej) OR the
non-extension allele (Ee). Pink-eyed pups (of any color) indicate the chocolate parent (both
parents) is carrying the pink-eyed allele (Pp). Himalayan or pink-eyed white pups indicate that the
parent is carrying both pink-eyed and himalayan alleles (PpCca). Dark-eyed white pups (crcr or
crca) indicate the chocolate parent is carrying the silver allele (Ccr) or the himalayan allele (Cca).

Lilac - Lilacs are true genetic selfs (aa). [for those who came to rabbits first, in cavies lilac is the
pink-eyed dilute of black, NOT the dilute of chocolate as in rabbits!] Lilacs have at least one copy of
the black pigment allele (B) but may hide the chocolate/beige allele (Bb). They have two copies of
the pink-eyed allele (pp) and never carry the dark-eyed allele. They have at least one copy of the
full extension allele (E) but may carry non-extension (aka red as Ee). They cannot have the white
spotting allele (they are ss) or the roan gene (they are rnrn). The base genotype for lilac is aaB-pp--
E-ssrnrn - if you have a lilac you know this much of its genotype for certain. The intensity of the
color in a lilac coat can be modified by the c alleles. C- animals will be a dark, rich color. ck- cavies
will be slightly lighter. cd- animals will give the preferred pale lilac shade. cr- lilac cavies are often
too light to show. A top show-quality purebred lilac is aaBBppcdcdEEssrnrn. Except for the "C"
locus, lilacs are usually purebred (though chocolate and partial/non-extension alleles can be
carried). If the parents of the lilac cavy were any color other than lilac, the lilac cavy is NOT
purebred for lilac. If chocolate, beige, orange, cream, gold, or white appear in the last three
generations of the pedigree, it is unlikely that the lilac cavy is purebred for lilac. A red, orange, gold
or cream pup indicates the lilac parent (both parents) is carrying non-extension (Ee). Bi-color pups
(ejej or eje) indicate the lilac parent is carrying the partial extension allele (Eej) OR the non-
extension allele (Ee). Chocolate, beige, gold or cream pups (all bb) indicate that the lilac (both
parents) is carrying chocolate (Bb). Himalayan or pink-eyed white pups indicate that the parent is
carrying the himalayan allele (ca) or the silver allele (cr).

Beige - Beige cavies are true genetic selfs (aa). They have two copies of the recessive chocolate
pigment allele (bb) and can never hide black. They also have two copies of the pink-eyed allele
(pp) and can never hide the dark-eyed allele. They have at least one copy of the full extension
allele (E) but may carry non-extension (aka red as Ee). They cannot have the white spotting allele
(they are ss) or the roan gene (they are rnrn). The base genotype for beige is aabbpp--E-ssrnrn - if
you have a beige you know this much of its genotype for certain. The intensity of the color in a
beige coat can be modified by the c alleles. C- animals will be a darker shade. ck- cavies will be
slightly lighter. cd- animals will give the preferred pale beige shade. cr- beige cavies are often too
light to show. A top, show-quality purebred beige is aabbppcdcdEEssrnrn. Except for the "C"
locus, beige cavies are usually purebred for beige - though they may hide partial and non-extension
alleles. If any color other than beige appears in the last 3 generations, especially non-extensions,
brokens or whites, it is unlikely that the beige cavy is purebred for beige. A gold or cream pup
indicates the beige parent (both parents) is carrying non-extension (Ee). Bi-color pups (ejej or eje)
indicate the chocolate parent is carrying the partial extension allele (Eej) OR the non-extension
allele (Ee). Himalayan or pink-eyed white pups indicate that the parent is carrying a the himalayan
allele (ca).

Red - Red is a non-extension self (ee) which may be a genetic self or may be masking any of the
other A series alleles (despite the self appearance, genetically red can be agouti, solid, tan or
self). Reds have two nonextension alleles (ee) and at least one dark-eyed allele (P-). They cannot
have the white spotting allele (they are ss) or the roan gene (they are rnrn). Because the non-
extension combined with crcr, crca, or caca turns the animal white, reds cannot have these
genotypes even though they may carry either the cr or ca allele (but not both). The base genotype
for red is ----P---eessrnrn - if you have a red you know this much of its genotype for certain. A
homozygous chocolate genotype (bb) usually dilutes the red color to a brighter orange. This color
is allowed to be shown as red (or possibly gold), but is not preferred. C- produces the darkest red
color. ck- (with sufficient modifiers) also produces a red color sufficiently dark to be shown. cd-
dilutes the red color too signficantly to be shown as red. Agouti, solid and tan based reds (A- and
Ar- and at-) may have darker black shading on the ears, which is not preferred. A purebred red
must have the genotype ----PP--eessrnrn - with the "C" series genotype CC, Cck, or ckck. This
genotype crossed with itself will produce only reds. The top show-quality purebred reds will have
genotype aaBBPPCCeessrnrn. They will also have modifier genes in play to darken the red color
to the preferred rich 'Irish setter' color.

Red-eyed orange - Red-eyed orange is a non-extension self (ee) which may be a genetic self or
may be masking any of the other A series alleles (despite the self appearance, genetically orange
can be agouti, solid, tan or self). Red-eyed oranges have two nonextension alleles (ee) and two
pink-eyed alleles (pp). They cannot have the white spotting allele (they are ss) or the roan gene
(they are rnrn). Because the non-extension combined with crcr, crca, or caca turns the animal
white, red-eyed oranges cannot have these genotypes even though they may carry either the cr or
ca allele (but not both). Likewise, genotypes cdcd, cdcr, and cdca will lighten the color to gold or
cream so red-eyed oranges cannot be these genotypes though they can carry any one of these
alleles (cd, cr or ca, but not two from this group). The base genotype for red-eyed orange is ----pp--
eessrnrn - if you have a red-eyed orange you know this much of its genotype for certain. You also
know that one of the "C" series alleles must be C or ck. A homozygous chocolate genotype (bb)
dilutes the color to a brighter orange - which is preferred over the darker black base (though black-
based oranges are allowed). Likewise, the ck- genotypes are also brighter than the C- genotypes
and are preferred. Agouti, solid and tan based red-eyed oranges (A- and Ar- and at-) may have
darker black shading on the ears, which is not preferred. A purebred red-eyed orange must have
the genotype ----pp--eessrnrn - with the "C" series genotype CC, Cck, or ckck. This genotype
crossed with itself will produce only reds. The top show-quality purebred red-eyed oranges will
have genotype aabbppckckeessrnrn. They will also have modifier genes in play to brighten the
orange color.

Gold - Gold is a non-extension self (ee) which may be a genetic self or may be masking any of the
other A series alleles (despite the self appearance, genetically gold can be agouti, solid, tan or
self). Gold in the ACBA standard combines two colors shown in Britain (and elsewhere) as buff
(with dark eyes) and saffron (with pink eyes). They cannot have white spots (they are ss) or
roaning (they are rnrn). The cdcd genotype is needed to remove the 'orange shading' as called for
in the standard. Thus a gold has the base genotype ------cdcdeessrnrn - if you have a gold you
know this much of its genotype for certain. That said, pink-eyed reds, dark eyed oranges, and non-
extension cavies with darker C alleles but lacking modifiers, or creams with the wrong modifiers
darkening the color can easily be mistaken for (and even shown as) golds. All golds with the proper
base genotype will produce only golds (when bred to each other) and can be considered purebred.
The homozygous chocolate genotype dilutes the color - which is preferred. Agouti, solid and tan
based golds (A- and Ar- and at-) may have darker black shading on the ears, which is not
preferred. The preferred purebred gold genotype is aabb--cdcdeessrnrn. Gold is accepted in both
dark-eyed and pink-eyed variants, so genotypes PP, Pp and pp can all be considered purebred
golds.

Cream - Cream is a non-extension self (ee) which may be a genetic self or may be masking any of
the other A series alleles (despite the self appearance, genetically cream can be agouti, solid, tan
or self). Creams must have a homozygous chocolate genotype (bb) to lighten the color
sufficiently. Creams must also have the genotype cdcr or cdca to lighten the color. Because cdcd
is too dark (gold), and crcr too light (white), all creams are heterozygous. This means that creams
are NEVER purebred - breeding cream to cream will always throw the occasional gold or
white. Although sometimes difficult to see, creams cannot have white spots (they are ss) nor
roaning (they are rnrn). The base cream genotypes are --bb--cdcreessrnrn and --bb--
cdcaeessrnrn. Creams are accepted with either dark or pink eyes - they can have genotype PP, Pp
or pp. Agouti, solid and tan based creams often have darker ears, making the self genotype (aa)
preferred.

White - White is a non-extension self (ee) which may be a genetic self or may be masking any of
the other A series alleles (despite the self appearance, genetically white can be agouti, solid, tan or
self). Either the cr or the ca allele can lighten non-extension cavies to the point that they appear
white. The based genotypes for white are ------cr-ee and ------cacaee. Because white patterns are
invisible on a white coat, white cavies can hide the whitespotting gene (they can be SS, Ss or ss)
and the roan gene (they can be Rnrn or rnrn - note RnRn is lethal). With the exception of roan
carriers (which might produce the 'deformed lethal whites') whites can be considered purebreds
which will only produce whites (when bred to whites). Whites are accepted in pink-eyed or dark-
eyed and can be PP, Pp or pp. Black-based whites (B-) and whites hiding agouti, solid or tan
patterns are more likely to have dark-tipping on the ears (a disqualification) - especially with cr. and
dark eyes. Thus the preferred genotypes for whites are aabbppcacaeessrnrn (pink eyes) and
aabbPPcrcreessrnrn (for dark eyes).

Solid Group
Solid cavies have fur of one pattern on their body with no other markings or patterns. Stomachs are
colored the same as backs without patches, eye spots, pea spots, etc. Most solids have a speckled
appearance - either due to relatively even mixing of two colors of hair (as in brindles and roans) or
individual hairs which include multiple colors (as in golden, silver and dilute solids). Three totally
separate genes can result in a solid appearance. Ar is an A series allele resulting in the 'true' solids
in which each hair is comprised of rings of color. Ep is an E series allele which, with the correct
modifiers, results in brindle which is an even intermixing of the base color (black, chocolate, lilac,
beige) with non-extension pigmented (red, orange, gold, cream, white) hairs. Animals which have
both genes at play, genotype Ar-Ep-, will have the base solid pattern with intermixed non-extension
pigmented hairs. Roans result from the Rnrn genotype which speckles a colored coat with
white. Although dalmatians are also caused by the roan genotype, their spotted pattern is not even
and they are shown as marked rather than solids. Roans can never be purebred as the required
genotype is heterozygous. Breeding roan to roan is not advised as it produces 25% 'lethal
whites'.

Golden solid - The golden solid is a true solid color caused by the Ar allele. Each hair of a golden
solid consists black hairs with red/gold tips and the eyes are dark. Unlike the golden agouti (below)
this includes the entire belly. The base genotype for the golden solid is Ar-B-P-C-E-ssrnrn. The
solid allele can hide the self allele (Ara). It cannot hide the tan pattern gene (Arat results in red/gold
eye circles, belly and pea spots) or the agouti allele (AAr is agouti). Golden solids can hide
chocolate (b) and any of the pink-eyed (p allele) or c series (ck, cd, cr, ca) dilutions. Golden solids
can also hide the alleles for brindle/broken (Eep) and non-extension (Ee). A purebred golden solid
is genotype ArArBBPPCCEEssrnrn. As a relatively dominant color, it is difficult to prove that a
particular cavy is a purebred for golden solid - but easy to disprove. When bred to anything other
than an agouti, a purebred golden solid will produce only golden solids.

Silver solid - The silver solid is a true solid color caused by the Ar allele. Each hair of a silver solid
consists of black hairs with silver/white tips and the eyes are dark with a ruby cast. Unlike the silver
agouti (below) this includes the entire belly. The base genotype for the silver solid is Ar-B-P-cr-E-
ssrnrn. The solid allele can the self allele (Ara). It cannot hide the tan pattern gene (Arat results in
silver/white eye circles, belly and pea spots) or the agouti allele (AAr is agouti). Silver solids can
hide chocolate (b) and the pink-eyed dilutes (p allele). Silver solids can also hide the
white/himalayan allele (crca). Silver solids can also hide the alleles for brindle/broken (Eep) and
non-extension (Ee). A purebred silver solid is genotype ArArBBPPcrcrEEssrnrn. As a relatively
dominant color, it is difficult to prove that a particular cavy is a purebred for silver solid - but easy to
disprove. Purebred silver solids can produce only silver solids, agoutis (only when bred to agoutis),
and golden solids (when bred to anything with higher C alleles - C, ck, cd).

Dilute solids - Dilute solid is actually a group of several colors in which the coat (and possibly
eyes) are of a lighter (dilute) shade than the golden or silver. All dilute solids are true solids caused
by the Ar allele. The solid allele can hide the self allele (Ara). It cannot hide the tan pattern gene
(Arat results in silver/white eye circles, belly and pea spots) or the agouti allele (AAr is
agouti). Dilute solids come in 10 ACBA recognized colors as follows:
Golden/lilac solid - lilac hairs with orange tips and pink eyes - base genotype Ar-B-ppC-E-ssrnrn
Chocolate solid - chocolate hairs with red tips and dark or ruby eyes - base genotype Ar-bbP-C-E-
ssrnrn
Gold/beige solid - beige hairs with orange tips and pink eyes - base genotype Ar-bbppC-E-ssrnrn
Lemon solid - black hairs with cream tips and dark eyes - base genotype Ar-B-P-cd-E-ssrnrn
Lemon/lilac solid - lilac hairs with cream tips and pink eyes - base genotype Ar-B-ppcd-E-ssrnrn
Cream solid - chocolate hairs with cream tips and dark or ruby eyes - base genotype Ar-bbP-cd-E-
ssrnrn
Lemon/beige solid - beige hairs with cream tips and pink eyes - base genotype Ar-bbppcd-E-ssrnrn
White/lilac solid - lilac hairs with silver/white tips and pink eyes - base genotype Ar-B-ppcr-E-ssrnrn
Cinnamon solid - chocolate hairs with silver/white tips and dark or ruby eyes - base genotype Ar-
bbP-cr-E-ssrnrn
White/beige solid - beige hairs with silver/white tips and pink eyes. - base genotype Ar-bbppcr-E-
ssrnrn

Eye color of the dilute solid tells you the genotype at the P locus. Pink-eyed dilute solid cavies are
always pp. Dark-eyed and ruby-eyed dilute solids are genotype PP or Pp. The ruby cast to some
dark-eyed cavies can be caused by any of several dilution genes - genotype Pp, genotypes Bb or
bb, or any visible or hidden cr.

Tip color of the dilute solid tells you the genotype at the C locus. Dilute solids with red/orange/gold
tips to their fur are C- (rarely ck-). Dilute solids with cream tips are cd-. Dilute solids with silver tips
are cr-.

Base color of the dilute solid tells you the genotype at the B locus. Chocolate and beige base
colors are genotype bb. Black and lilac base coats are genotype B-.

Because the dilution effects are caused by separate genes, the dilute solid group cannot be said to
breed true -- breeding dilute solids to one another without regard for the particular color will
frequently result in gold or silver solids (or shades different from either parent). Each of the
individual colors within the dilute agouti group, however, can be purebred.

Brindle - Brindle is a solid pattern evenly mixing two colors of hair - red and black. Brindling of
other colors is possible (e.g., chocolate and orange) but these are not showable in the ACBA
standard. Brindle is not a 'true solid' and does not have the Ar allele. Brindle results from the
substitution of the EpEp or Epe genotype in the self black genotype along with modifier genes to
achieve the desired even mixing of the two colors. Because the modifiers are easily lost when
crossing to any other color or pattern (where they cannot be seen and so cannot be selected for),
most show quality brindles are purebred aaBBPPCCEpEpssrnrn + modifiers.

Roan - Roan is a solid pattern evenly mixing any self (black, lilac, chocolate, beige, red, orange,
gold or cream) or solid color (golden, silver or dilute solid) with white. The ideal roan has white
evenly mixed over the entire coat giving a solid appearance -- but most roans are significantly
darker (with less white) on their heads and this is allowed in the ACBA standard. Roaning is not a
result of the Ar allele, it is a result of the roan gene. All roans are genotype Rnrn. Thus roans can
never be purebred. Roans should never be bred to each other because it can result in lethal white
pups (with genotype RnRn) which are deformed. Breeding roans to whites or very light dilute solids
(cr-) is discouraged because the roaning is too easily hidden. Instead, roans with good pattern
should be bred to selfs or solids of the desired (preferrably matching) background color.

Agouti group
Agouti cavies have fur with a ring pattern (ticking) over their entire body EXCEPT the belly, which
should be the same color (tan to white) as the tip color. Agoutis come in golden, silver and dilute -
the same colors as solids (except NOT in brindle or roan) Agouti is the result of a dominant A
allele. Purebred agoutis are AA, but agoutis can carry the Ar, at or a alleles.
Golden agouti - Each hair of a golden agouti consists black hairs with red/gold tips - except the
belly hairs, which are red/gold - and the eyes are dark. This color pattern is considered the original
pattern of wild cavies - with all other colors being mutations of this original pattern. The base
genotype for the golden agouti is A-B-P-C-E-ssrnrn. The agouti allele can hide the self allele (Aa)
and the solid allele (AAr). It cannot hide the tan pattern gene (Aat results in red/gold eye circles
and pea spots). Golden agoutis can hide chocolate (b) and any of the pink-eyed (p allele) or c
series (ck, cd, cr, ca) dilutions. Golden agoutis can also hide the alleles for brindle/broken (Eep)
and non-extension (Ee). A purebred golden agouti is genotype AABBPPCCEEssrnrn. As the most
dominant color, it is difficult to prove that a particular cavy is a purebred for golden agouti - but easy
to disprove. A purebred golden agouti will produce only golden agoutis. Interestingly, British
breeders state that the Aa genotype golden agoutis have better show color than the purebred.

Silver agouti - Each hair of a silver agouti consists of black hairs with silver/white tips - except the
belly which is silver-white - and the eyes are dark with a ruby cast. The base genotype for the silver
agouti is A-B-P-cr-E-ssrnrn. The agouti allele can hide the self allele (Aa) and the solid allele
(AAr). It cannot hide the tan pattern gene (Aat results in silver/white eye circles and pea
spots). Silver agoutis can hide chocolate (b) and the pink-eyed dilutes (p allele). Silver agoutis can
also hide the white/himalayan allele (crca). Silver agoutis can also hide the alleles for
brindle/broken (Eep) and non-extension (Ee). A purebred silver agouti is genotype
AABBPPcrcrEEssrnrn. As a relatively dominant color, it is difficult to prove that a particular cavy is
a purebred for silver agouti - but easy to disprove. Purebred silver agoutis can produce only silver
agoutis, and golden agoutis (when bred to anything with higher C alleles - C, ck, cd).

Dilute agoutis - Dilute agouti is actually a group of several colors in which the coat (and possibly
eyes) are of a lighter (dilute) shade than the golden or silver. The agouti allele can hide the self
allele (Aa) and the solid allele (AAr). It cannot hide the tan pattern gene (Aat results in eye circles
and pea spots). Dilute solids come in 10 ACBA recognized colors (the same as dilute solids) as
follows:

Golden/lilac agouti - lilac hairs with orange tips/belly and pink eyes - base genotype A-B-ppC-E-
ssrnrn
Chocolate agouti - chocolate hairs with red tips/belly and dark or ruby eyes - base genotype A-bbP-
C-E-ssrnrn
Gold/beige agouti - beige hairs with orange tips/belly and pink eyes - base genotype A-bbppC-E-
ssrnrn
Lemon agouti - black hairs with cream tips/belly and dark eyes - base genotype A-B-P-cd-E-ssrnrn
Lemon/lilac agouti - lilac hairs with cream tips/belly and pink eyes - base genotype A-B-ppcd-E-
ssrnrn
Cream agouti - chocolate hairs with cream tips/belly and dark or ruby eyes - base genotype A-bbP-
cd-E-ssrnrn
Lemon/beige agouti - beige hairs with cream tips/belly and pink eyes - base genotype A-bbppcd-E-
ssrnrn
White/lilac agouti - lilac hairs with silver/white tips/belly and pink eyes - base genotype A-B-ppcr-E-
ssrnrn
Cinnamon agouti - chocolate hairs with silver/white tips/belly and dark or ruby eyes - base genotype
A-bbP-cr-E-ssrnrn
White/beige agouti - beige hairs with silver/white tips/belly and pink eyes. - base genotype A-
bbppcr-E-ssrnrn

Eye color of the dilute agouti tells you the genotype at the P locus. Pink-eyed dilute agouti cavies
are always pp. Dark-eyed and ruby-eyed dilute agoutis are genotype PP or Pp. The ruby cast to
some dark-eyed cavies can be caused by any of several dilution genes - genotype Pp, genotypes
Bb or bb, or any visible or hidden cr.

Tip/belly color of the dilute agouti tells you the genotype at the C locus. Dilute agoutis with
red/orange/gold tips to their fur are C- (rarely ck-). Dilute agoutis with cream tips/belly are cd-
. Dilute agoutis with silver tips/belly are cr-.

Base color of the dilute agouti tells you the genotype at the B locus. Chocolate and beige base
colors are genotype bb. Black and lilac base coats are genotype B-.

Because the dilution effects are caused by separate genes, the dilute agouti group cannot be said
to breed true -- breeding dilute solids to one another without regard for the particular color will
frequently result in gold or silver solids (or shades different from either parent). Each of the
individual colors within the dilute agouti group, however, can be purebred.
Tan/marten group
The tan/marten group is the result of the fourth 'A series' allele - at - which is intermediate in the
dominance series. The tan allele (at) is dominant over self - ata is a tan pattern. At is co-dominant
with the agouti (A) and solid (Ar) alleles - Aat agoutis will have eyes circles and pea spots. Arat
'solids' will have eyes circles and peaspots AND tan/white bellies - causing them to be easily
mistaken for agoutis.

The tan/marten group is divided into 10 colors comprising two groups:

Tans have a self appearing coat (black, 'blue', lilac, chocolate or beige) with red/tan eye circles, pea
spots (above/behind the ears), and bellies. Base genotype at-C-.

Black tan - at-B-P-C-E-ssrnrn

Lilac tan - at-B-ppC-E-ssrnrn
Chocolate tan - at-bbP-C-E-ssrnrn
Beige tan - at-bbppC-E-ssrnrn

Blue tan - Blue in the ACBA standard is the color called 'slate' elsewhere in the world - not a true
blue dilution (D gene) seen in rabbits and other mammals. The blue/slate color is caused by an
intermediate allele in the P series -- pg is recessive to dark eyes (P) and dominant over pink eyes
(pp) - and results in cavies with a ruby-blue cast to their eyes - similar to but usually slightly darker
than the shade seen in the eyes of lilac rabbits. The ACBA standard currently accepts blue only in
the tan/marten group - though slates are certainly possible in other patterns and are shown as self
slate, slate broken, etc elsewhere in the world. Blue tan is base genotype at-B-pg-C-E-ssrnrn.

Martens have a self appearing coat (black, 'blue', lilac, chocolate or beige) with silver/white eye
circles, pea spots (above/behind the ears), and bellies. Base genotype at-cr-.
Black marten - at-B-P-cr-E-ssrnrn
Lilac marten - at-B-ppcr-E-ssrnrn
Chocolate marten - at-bbP-cr-E-ssrnrn
Beige marten - at-bbppcr-E-ssrnrn
Blue marten - at-B-pg-cr-E-ssrnrn

No other C series dilutions (e.g., ck, cd) are currently allowed in the ACBA standard.

Marked
A catch-all group for cavies that do not appear a single even color but rather have a defined pattern
determined by genes other than the A series alleles.

Marked includes the following subgroups - himalayan, dalmatian, dutch, tortoiseshell, tortoiseshell
and white, and broken.

Himalayan - Himalayan cavies exhibit a pattern similar to the pattern seen in himalayan or pointed
rabbits in which the ears, nose, and feet are a dark color and the rest of the coat white - eyes are
always pink. With the darkest points preferred, most himalayans are black-based, though dark
chocolates are allowed. The ca allele is responsible for the himalayan pattern - like the ch gene in
rabbits, this gene is temperature-sensitive so animals which grow their coats in colder temperatures
will have darker points (and possibly more dark color or 'smut' extending into the white portions of
the coat) while coats grown in warmer temperatures will have lighter points. The ca allele is the
most recessive of the C series alleles - himalayans are always caca. Himalayans must also be full
extension (E-) - cavies which have the non-extension genotype (ee) will lose the point color on the
ears to be pink-eyed whites. The caca genotype also strips the eye color -- although they always
have pink eyes, himalayans can hide the darker-eyed genotypes (they can be PP, Pp, Ppg, pgpg,
php or pp). Good himalayans always have a self genotype on the A series (aa) - as agouti, solid or
tan genotypes fade the point color. Introduction of roan genes into himalayans is to be avoided as
they may be invisible roans (cacaRnrn) that could later have lethal white (RnRn pups). White
spotting (SS or Ss) is generally invisible on the himalayan genotype, but may cause mismarkings
(e.g., a white ear or foot) and so is to be avoided as well. Thus the preferred base genotype for
himalayan is aaB---cacaE-ssrnrn.

Dalmatian - Dalmatian cavies are a variation on roan in which modifier genes cause the colored
hairs to cluster together in spots rather than be evenly intermixed. Like roans, dalmatians are never
purebred because the RnRn genotype gives lethal whites. Instead, the base Dalmatian genotype is
Rnrn (+modifiers) and they are produced by crossing a dalmatian with a self of the desired
color. SS and Ss genotypes may be invisible, but may result in large patches without spots, so are
usually avoided. ACBA accepted dalmatian colors include:
Black aaB-P-C-E-ssRnrn +modifiers
Chocolate aabbP-C-E-ssRnrn + modifiers
Lilac aaB-ppC-E-ssRnrn
Beige aabbppC-E-ssRnrn
Red --B-P-C-eessRnrn
Orange ----ppC-eessRnrn (or ck-)
Gold ------cdcdeessRnrn

Although technically allowed, cream dalmatians do not show spotting well enough to do well on the
show table and are usually avoided. Dalmatian spotting is invisible on white selfs, so mixing
dalmatian genes with whites is avoided.

Dutch - Dutch are the result of a white spotting gene (S) with modifier genes that place the 'spots' in
the correct pattern - white blaze and collar. Dutch are accepted in any self, true solid (not roan or
brindle) or agouti base color. Dutch can be SS or Ss, though only the SS will breed true and the Ss
much more frequently throw mis-marked pups.

Tortoiseshell - Unlike 'tort' rabbits, tortoiseshell cavies are not shaded, rather they exhibit a pattern
similar to harlequin rabbits with a pattern of red and black patches. As with harlequins, the
responsible gene is a partial extension - ep allele on the E series which is recessive to full extension
but dominant over non-extension. Also as with harlequins, modifier genes are needed to get the
preferred alternating patch pattern. Tortoiseshell cavies are all selfs -- agouti and solid patterns
would show in the black patches (and are shown as broken rather than tortoiseshell). Similarly,
tortoiseshell cavies are also shown only in black - chocolate, lilac, beige and other dilutions are
shown as broken. Tortoiseshells are genotype aaB-P-C-ep-ssrnrn + modifiers.

Tortoiseshell and White - The equivalent of 'tri-color' in rabbits. Tortoiseshell and white cavies
have patches of black, red and white. The white is added to the base harlequin genotype by the
addition of the S allele. Unlike tri- rabbits, the ideal pattern is the alternating patches - with all three
colors present on each side of the body (at least 2 patches of each color). Achieving the ideal
pattern is difficult and relies on modifier genes for both the ep (as in tortoiseshell) and S ( but
different from the dutch modifiers). Tortoiseshell and whites are all self blacks without dilution
(other color patterns with three colors are shown as broken) - aaB-P-C-ep-S-rnrn + modifiers.
Broken - Unlike rabbits, where broken means a single color pattern plus white, in cavies broken
can be any two or three colors. Unlike the spotted and blanket pattern brokens preferred for rabbits
with a range of 10-50% color, broken cavies are preferred in large patches (greater than a 50-cent
piece) of color as close as possible to 50-50 (or 33-33-33 for three-color brokens). Brokens can
include any combination of self, true solid or agouti colors - evidence of tan, brindle, dalmatian,
dutch, himalayan or roan patterns are disqualifications. (a poorly marked dutch cannot be shown
as broken, a dalmatian with one large patch cannot be shown as broken, etc). Likewise, the
black/orange and black/orange/white coloration of tortoiseshell and tortoiseshell & white cannot be
shown as brokens. Brokens come in 3 basic genetic types:

Agouti, solid and/or dilute tortoiseshells - achieved by the ej- genotype with any color or pattern
other than self black. These will have 2 colors - one from the non-extension series (orange, gold,
cream or white) and one true self, true solid, or agouti (lilac, chocolate, beige, golden solid/agouti,
silver solid/agouti or dilute solid/agouti). Agouti based and solid-based can only be told apart if the
solid patch happens to extend into the belly (which will only happen with the solid).

White patched - achieved by the S- genotype with any other color or pattern. These will usually
have two colors, one of which is white and the other can be any self, true solid, or agouti color
(black, lilac, chocolate, beige, red, orange, gold, cream, golden agouti/solid, silver agouti/solid or
dilute agouti/solid). 'Agouti color' here can be achieve with either the A or Ar allele. They may
appear to have three colors if the agouti patches (A) appear on the belly (where they will be tan
rather than tipped).

Tri-colors - achieved by the combination of the ej- and S- genotypes. These have three colors - one
from the non-extension series (red, orange, gold, cream or white), one true self, true solid or agouti
(black, lilac, chocolate, beige, golden agouti/solid, silver agouti/solid, or dilute agouti/solid) and
white (from the white-spotted gene). (note that it is possible for both the non-extension series and
the white spotted to give white, so possible to have a two-color 'tri-').
Genetics

Genetics are often given a bad wrap - with most people thinking that they're too
complicated to understand. I think it's a fair statement to make that the majority of guinea
pig owners (and even breeders) don't have a good understanding of the genetics behind
the breeds they either own or are trying to breed.

If you're just a happy 'piggy-slave' to non-breeding pets then you really DON'T need to
know what their genetic make up is. But if you are planning to breed or have a registered
 stud, I think this is a really important bit of information to comprehend - even if you just
grasp the basics.

I'm not going to assume a basic level of understanding here, but rather explain things from
the beginning in common terms with as many images as I can to make this as simple as
possible. You can also get some more detailed genetic/inheritance background information
on this page (it relates to plants, but the basics of dominance and recessive are there) OR
if you do already have a good basic understanding, you might benefit more from a guinea
pig specific e-book written by another brisbane breeder that you can purchase and
download here.

GUINEA PIG GENETICS

Guinea pigs make babies through sexual reproduction. This means that you need one
male and one female to produce offspring. (please visit my Breeding Info page for more
information on cavy pregnancy/birth/newborn care if you're looking for more of the 'nuts
and bolts' type info ).

When the guinea pig's body is making the reproductive cells (eggs for girls, sperm for
boys), the usual 64 chromosomes (DNA make-up found in every cell in their body) are split
in half to make the sex cells (which will contain 32 chromosomes). This means that when
the guinea pig's sperm meets the egg, the two 'half' cells join and create the perfect
amount of full, new DNA for the new baby .This also means that only half of a parent's
genetic make up is passed onto their offspring.

In this extremely simplified example above you can see that the mum's DNA consists of
two different genes 'D' and 'd', whereas the dad is 'pure' - both of his genes are the same
('D' and 'D').

When the eggs are made by the mother guinea pig, half of them will be genetically capitol
'D', while the other half will be lowercase 'd'. When the sperm is formed by the father, they
can be nothing other than capital 'D' as this is all he can offer genetically. This is why true,
pure-breed animals will produce identical babies when paired together, and why cross-
breed animals can produce a whole host of different babies depending on what genes they
are carrying.

This is when we get into the dominant genes and the recessive genes. Dominant genes
mean just that - they dominate their corresponding recessive gene. This usually means
that there only needs to be one copy of the gene at the cellular level to produce the
physical appearance in the animal's coat (with some exceptions we can get to later). With
recessive genes you need two copies to get that particular appearance in the coat. The
example I've drawn below is for a rosetted (Abyssinian) coated guinea pig crossed with a
smooth coat guinea pig. Rosettes are dominant in guinea pigs - so I've used a capitol 'R' to
represent a rosette gene and a lowercase 'r' to represent the lack of rosette, or smooth
coat.

In the first generation we have a 'pure' rosetted guinea pig (RR) crossed with a 'pure'
smooth coated guinea pig (rr). The genes are split in half to form the sperm and eggs and
 are paired up to produce 4 possible pairings - all end up being genetically half rosetted,
half smooth (Rr). BUT - because the rosette gene is dominant, these babies will have
rosettes in their coats, and will look a lot like a pure Abyssinian. (The rosette placement
might not be as nice as on the parent guinea pig though due to the presence of the
recessive gene/complicating factors from other genes).

If we were to then take two babies from the second generation and breed them together,
we would get different results again. This time, because both of the guinea pigs in this
pairing are genetically half/half (Rr), there are 4 possible pairing combinations that could
occur: RR, Rr, Rr, and rr. The 'RR' baby would likely look just like it's siblings who are 'Rr' -
so there would be no way to tell what it's genes were until it came time for it to breed. The
baby with the 'rr' genes would look smooth coat like it's grandparent. This is how different
breeds can 'pop' up in crossbreed litters. Only recessive characteristics pop up
unexpectedly. Dominant genes are seen in the animals coat/colouring.

Dominant genes in guinea pigs include (but are not limited to):
* Rosettes
* Short hair
* Dark eyes
* Straight hair

Recessive genes in guinea pigs include (but are not limited to):
* Smooth coat
* Long hair
* Red eyes
* Curls

Partial Dominance

Long and short coat genes can act a little less clear cut than the simple dominant and
recessive roles. Generally the short coat is dominant and the long coat is recessive. If you
have an animal with hair that grows to the ground, that animal is guaranteed to have two
copies of the recessive long-coat gene (which I'm referring to with a lowercase 'l' in the
diagram below). It becomes less clear with the short-coat gene.

A pure short coat animal with two copies of the dominant gene - LL - (here we're using a
capital 'L' to represent this dominant gene) can often look exactly like a cross-bred half
long-coat/half short-coat animal (Ll). BUT - the short-coat gene is not always completely
dominant, and a lot of half/half animals will have telltale longer strands of hair on the lower
half of their bodies while having short hair everywhere else.
 In the pairing shown above we've taken a pure sheltie (long-haired guinea pig) and
crossed it with a pure short-coated guinea pig. The resulting offspring will all be genetically
half/half. They would likely either look just like their short-coated parent, or have a few
wispy longer stands on their rumps.

If we then crossed two of these second generation guinea pigs, we would be left with 4
possible genetic combinations - LL (pure short coated), Ll and Ll (both half/half again) and
ll (pure long-coated) - this animal would end up looking like it's grandparent. Again
showing how recessive genes can pop up generations later.

Obviously the possible pairings are just statistical odds - it just means that if there were 4
babies born from the litter, chances are that 1 would be LL, 2 would be Ll and 1 would be
ll. You may well get 4 babies who are all genetically LL, or all of any of the other combos.
You could have a single litter who is ll... or 2 babies that are both Ll like their parents... you
really have no idea which eggs and which sperm are going to meet.

We work out the statistical odds by using a probability grid (see below:). This will obviously
look different for different pairings. For example if both parents were pure for long coats (ll
x ll), you'll have a 100% chance of babies also being ll as the parents have nothing else
but the long-hair gene to pass on.
 Impossible Outcomes

There are many urban myths out there of offspring born to parents who could not have
possibly produced their specific coat-type. For example, you cannot breed an abyssinian
(RR or Rr) from two smooth coated guinea pigs (rr) - even if the smooth coat came from
aby parents as they do not have the rosette gene to pass on.

Likewise, you cannot breed a short-coated guinea pig (LL or Ll) from two pure long-haired
parents (ll). Nor can you produce a straight-coated animal from two curly parents (unless
both parents had different curl-genes, not very common in Australia, but has happened in
the past when imported pigs were paired with Australian pigs - this is something that would
need a whole other page devoted to it). We will discuss basic curls and straight-coats
below.
 The Illusive Curl

I've always had a soft spot for the curly-pigs. Specifically the long-coated curlies like
Texels, Merinos and Alpacas, but Rex pigs are gorgeous too! In AUSTRALIAN guinea
pigs, the curly gene is recessive and the straight hair is dominant. This is how curly texel
babies can be born to straight-coated sheltie parents. (You cannot get a sheltie from a
pairing between two texels as texels have nothing but the curly gene to pass on to their
offspring).

In the diagram below I have shown how it is possible to get texels and shelties from
different pairings. For the purposes of this information I am referring to the dominant
straight coat gene as capital 'S' and the recessive curly gene as lowercase 's'. In the first
generation we have a sheltie who carries the texel gene (but would otherwise just look
sheltie in appearance) paired to a texel. If you don't know the ancestry of your sheltie, a
pairing like this would show you if he/she was carrying the texel gene. If the resulting
babies were all shelties, you could assume that there were no recessive curly genes to
pass on and your sheltie was just sheltie (obviously with the laws of probability you may
not see a curly bub even when the parent is carrying the curl - so you would never be able
to be 100% certain.)
 In the diagram above we have hypothetically paired two of the sheltie-looking offspring
from the second generation who both carry the texel gene. The results of a pairing like that
would statistically produce 25% 'pure' sheltie (SS), 50% Sheltie (carrying texel) and 25%
texel. Due to the complete dominance of the straight coat over the curl, you would not
know which babies were genetically 'SS' versus 'Ss'. If any babies came out with curls
though, you would be guaranteed that their genetics would be 'ss' for curl.

The above diagram could also be substituted for rex guinea pigs and smooth-coat guinea
pigs (substitute the shelties for the smooth-coats and the texels for the rex).

(Two sheltie, texel carrier babies. Texel mum, sheltie dad)

Obviously the complexities of guinea pig genetics are a lot more intricate than I have
explained here. You don't have only one genetic factor to take into account with each
pairing, but countless subtle additional ones (body shape, type, coat density, colouring,
rosette positioning etc). However, if you can grasp the basic fundamentals of the coat-type
and the dominance and recessive nature of the genes it may help you in your future
pairing decisions.

Cavy Coat Color Genetics

Please excuse our mess while we work on these sections!!!

Okay, for all the non-genetics folk out there, a little terminology
101. Genetics won't be quite as painful once you know some of the
jargon. Think of it like starting new in the fancy. I'm fairly sure there were
a lot of breed specific and species specific words you had to
learn. Genetics is no different. So with that in mind, there are just a few
words you need to know the definitions for. Once you've got those, the
rest will seem easy. These are going to be my paraphrasing for you for
ease of communicating.

Genetics is the study of inheritance. That's about as basic as I can get

with that one. Of course there's more to it, but that's generally it (as far as
animal breeders are concerned lol).

a Trait is anything we select for based on an animal's appearance. Such

as coat length, coat color, eye color, crown, shoulder width, etc.

Phenotype is how an animal looks to us on the outside. We show

according to this.

Genotype is the animal's specific lineup of genes on the inside.

a Gene is the mode of inheritance. Merriam-Webster dictionary has this

really long definition of a gene, which includes several much longer words
that I refuse to utter here. Most of you probaby know what a gene is
without having to put it into words. Examples would be coat length, pink-
eye color, black coat color. They are quite specific.

In simple inheritance, one gene controls one trait. Most of the things we
select for in cavies, are simply inherited.

In complex inheritance, one trait is controlled by many genes. Most of

these we are not consciously aware of, as they are difficult to trace. Most
of the type traits are complex inherited. Progress is slow as a
consequence of the many genes involved.

a Locus is the place where a gene is known (or believed) to exist. (plural
is loci)

an Allele is each "type" or alternative form of a gene that can occur at a

single locus. So if the gene is the pink-eye gene, there would be 2
alleles, P for non-pink, and p for pink. There are some genes that have
more than 2 possible alleles, such as the color locus, where you can have
C, full-color; ck, dark dilute; cd, light dilute; cr, ruby-eye dilute; and ca,
himalayan. Only 2 alleles can be present at any single locus. So even
though there are 5 possible alleles for the color locus, only 2 will be found
in any one animal.

Dominant is one allele that asserts it's effects over others. The Black (B)
allele is dominant to the brown (b) allele. Agouti (A) is dominant to the
solid (Ar) allele, both those are dominant to the tan (at) allele, and all
three are dominant to self (a).

Recessive is one or more alleles whose effects may be hidden by other

alleles. Pink-eye (p) allele is recessive to the non-pink-eye (P) allele.

(these 2 are actually a little more complex than this, but I'm trying not to
confuse)

Homozygous (also homozygosity and homozygote) is where both alleles

at a locus are the same. A self is homozygous aa, a Teddy is
homozygous wv wv.

Heterozygous (also heterozygosity and heterozygote) is where the two

alleles at any given locus are different. A roan is heterozygous Rsrs.
Complete Dominance is where both the homozygous dominant and the
heterozygote exhibit the same phenotype. At the agouti locus, AA and Aa
will both be agoutis, while only aa will be self. We say that agouti is
completely dominant over self.

Incomplete Dominance is where the heterozygote acts as an

intermediate between the two homozygotes. At the roan locus, roan is
the intermediate between non-roan and lethal. We say then that the roan
gene is incompletely dominant.

I think that pretty much covers the basics of the terminology. Next I'm
going to mention a few specific genes of interest.

The first locus to look at is the agouti locus. There are 3 possible alleles
that can exist here: A is agouti, Ar is solid, and a is self. Since only 2
alleles can co-exist at the locus, you can have AA, AAr, ArAr, Aa, Ara,
and aa.

Because agouti is dominant to both solid and self, then AA, AAr, and Aa
will all appear as agoutis. We say then that agouti masks the other
alleles, they are hidden from out view and could only appear in breeding
to a solid or self (or to another agouti that is also masking those alleles)

Solids would appear only as ArAr or Ara. Solids are dominant to self and
mask and hide that allele. A solid would only be shown to carry self if
either one parent was a known self, or they produce one themselves.

Self is only aa. If you have a self, you know they cannot produce
anything but self when bred to other selfs. To produce agouti or solid
using a self, you would have to breed to an animal with agouti or solid at
that locus.

The next locus to look at is the gene that controls black color. Here the
black allele (B) causes normal black pigment, while the recessive brown
(b) causes modified black pigment resulting in brown or "chocolate"
pigment. (Claus explains though that you need more than this one set of
recessives to get good chocolates, that's why I'm using brown and not
chocolate!).
So this gene is a simply inherited gene, with only two alleles and is
completely dominant in that both BB and Bb will appear black. Only bb
will exhibit a brown animal.

So coupling the first locus with this second. Any of the agouti
combinations (AA, AAr, Aa) with either BB or Bb will give you the black
based agoutis, golden, silver, & lemon. Either of the 2 solid allele
combinations (ArAr or Ara) coupled with black (BB or Bb) will give you the
black based solids (golden, silver, lemon). And self coupled with the
black combinations will give you black. Substitute brown (bb) at the black
locus, and you'll get chocolate based agoutis and solids, and chocolate
self. This is completely ignoring the pink-eye dilution locus. One step at
a time. lol

Black based agoutis: Chocolate based agoutis:

AA BB AA bb
AAr BB AAr bb
Aa BB Aa bb
AA Bb
AAr Bb
Aa Bb

Black based solids: Chocolate based solids:

ArAr BB ArAr bb
ArAr Bb Ara bb
Ara BB
Ara Bb

Black Self Chocolate self:

aa BB aa bb
aa Bb

Bear in mind that you would not be able to simply look at an animal (with
no pedigree or progeny information) and tell whether it is AA, AAr, Aa or
whether it is BB or Bb. So when you are faced with a black-based agouti
animal, you would simply say that it is A- B- where the dashes account for
the fact that it could carry any of the other recessive or dominant
allele(s). The only ones you could be absolutely certain of just by looking
are the homozygous recessives (self and
brown).
I was going to skip over the color (C) locus for now, but I think I'll just hit it
head on, and anyone with more information than I have can jump in and
comment, clarify, or correct.

Okay, so the color locus is one of the most complicated of cavy coat color
genes. It's not completely understood how everything works with this
one. There are five alleles associated with this locus. The various
combinations of the five alleles are what give rise (in large part) to the
multitude of varieties (and shades!) we see in domestic cavies. There is
no simple dominance associated with the lower alleles, and several of the
recessive heterozygote combinations will exhibit the same
phenotype. Thus making it that much more difficult to differentiate which
of these alleles each individual might carry.

The five alleles are full color (C), dark dilue (ck), light dilute (cd), ruby-
eyed dilute (cr), and himalayan (ca). The one dominant allele, full color
(C), will mask the four remaining recessive alleles. So in the presence of
C, all else is carried. Therefore CC, Cck, Ccd, Ccr, and Cca will all exhibit
full color. An example would be red vs orange, the best red will be CC,
while a good orange will have some combination of heterozygote
recessive alleles.

For the recessives, let's talk about them in reverse order. So ca is

himalayan (this is strictly for ease of discussing this particular allele, in
reality there is another gene involved in the determination of an animal to
be himalayan or white), you could think of this allele as a primary factor of
an animal being either himalayan or white. Since cavies have no actual
albinism gene, this allele is largely the cause of the lack of
body pigmentation in himis and whites. Incidentally, this is also why a
himi-smut will result from the crossing of a white to a dark animal. The ca
allele also assists in the creation of good creams. Hence the reason
many good cream breeders will tell new breeders to cross whites to
creams. Okay, so caca will give rise to himalayan and white, and coupled
with one of the other recessive color alleles (cd or ck), tends to "lighten"
the pigmentation. ca removes all red pigmentation and most of the
black. Pink eyes are associated with this allele.

The next recessive color allele to talk about is the ruby-eyed dilute allele
(cr). This one seems to have a somewhat similar effect as the ca allele,
but rather than removing a large majority of the color, resulting in a "clear"
animal, it removes only the red pigmentation (leaving black), resulting in a
"silvering" effect. The homozygote of this allele can be found in silvers
(agouti and solid), and DEW. A cr allele coupled with a ca allele tends to
have similar effects as if you had two ca alleles. Incidentally, ruby cast
eyes are associated with this allele. This explains why silvers (not
goldens!) should have a ruby cast eye.

The remaining 2 alleles (cd & ck) should be talked about together as they
each have very similar effects on the phenoptype of the animal. The
difference between the two would be that light dilute (cd) would show
about an even amount of red and black pigmentation, while dark dilute
(ck) shows just slightly more black than red pigmentation. However, that
being said, an ordinary animal breeder would be very unlikely to be able
to differentiate these two by merely looking at a cavy's phenotype. The
two together and the homoygotes of each (coupled with a few other
genes) largely result in buff colored animals (dark cream). These alleles
also assist in the production of good chocolates, and good
REO's. Matching one of these alleles with a himalayan allele (ca) gives
rise to the nicely colored light creams.

So a summary of the color gene:

CC found in good reds & blacks
Cck, Ccd, Ccr found in good chocolates
cdcd, ckck, & cdck result in buffs (does not act alone, requires another
gene)
cdca & ckca result in good creams
caca results in himalayans and whites
crcr results in DEW or silvers

Other unknown modifiers may be involved with these, but that's about as
basic as I can get with the color locus and still be understood (and
understand myself)

Genetics doesn't have to be complicated. I'm trying to make it as simple

as I can and still make it understandable. If I don't quite manage it,
question! lol I question everything, and believe me, some of this stuff is
stumping me. lol I'm going to be referring to other papers and another
group to try and get some answers. I find that Harry Claus is causing
more holes than filling them with his book. lol (I expected that though)
...one other thing, don't get too caught up with all this stuff I'm writing
down. I'm trying to break it down first so can understand what each gene
does individually. But remember, there is no such thing as a cavy with
only one or two of these genes, so most of these interact to give us the
various colors. So first I'm breaking it down, then I will put it together
(might make more sense then), and then I'll go through some examples.

btw, I'm probably a day late on asking this....but if anyone would prefer I
not continue this thread, I will stop. I know some people probably would
just prefer to skip all this stuff. Just say so, if you aren't comfortable
saying so in the thread, send me a pm.

Reply to : Falls-Acre

Please continue! It is really helping me understand cavy

genetics more. Keep it coming!

I am sure you have these links, but if not, here are a few
places on the internet to find more papers written on cavy
genetics.
Cavy Genetics by Peter and Cell Herman (this is also found in the Claus book)
Basic Color Genetics by Pip Squeak Caviary
Sigi's Sauhaufen Cavy Genetics
Eva's Cavy Genetics Page
Cavy Color Breeding Advice
Cavy Genetics: An Exploration by Nick Warren
Color Genetics of Cavies
Colour Genetics of the Cavy by Catherine Whiteway (can also be found in the "file" section
of the Cavy Colors group)

Some great links Leslie, thanks.

Alright, moving on, the next gene to consider would be the Pink-Eye (P)
dilution gene. This one is another simply inherited gene, completely
dominant. So both PP and Pp would have dark (non-pink eyes). This
gene affects the eye color of the animal separate from the effects of the
color locus (so you can have a white or albino that is PP or Pp at this
locus, and they would still have pink eyes! This is partly why it Is possible
to produce dark eyed animals from whites and himis)
The allele P causes normal coat color pigmentation to be produced, it
completely masks the effects of the recessive allele, so an animal
carrying one pink-eye allele will still have dark eyes and normal coat
pigmentation.

The presence of the homozygous recessive at that gene causes a large

reduction in the amount of black pigment that is made, with no effect on
red pigment. So black gets diluted to lilac and chocolate gets diluted to
beige. Red becomes REO, and DE cream becomes RE cream, but
because there is no effect on the red pigmentation of the animal, the coat
color of these two varieties would remain virtually unchanged. This is
why it is possible to have a REO with a coat as dark as a red, and a RE
cream, with a coat as pale as butter. The only change caused by the P
gene in those animals is to change the eye color.

So now, putting the the agouti, black, and pink-eye genes together, we
can have:

black based agouti = A- B- P-

black based solid = Ar- B- P-
black self = aa B- P-

chocolate based agouti = A- bb P-

chocolate based solid = Ar- bb P-
brown self = aa bb P-

lilac based agouti = A- B- pp

lilac based solid = Ar- B- pp
lilac self = aa B- pp

beige based agouti = A- bb pp

beige based solid = Ar- bb pp
beige self = aa bb pp

For right now I'm going to be skipping both the extension locus and the
white spotting locus and will come back to those a little later. You'll see
why when I do get to them. lol
For now, I want to discuss the two genes that are responsible for
roaning. Yes I said two, because there are two genes that are accepted
as being able to cause white and color intermixing. However, in
heterozygote form only the actual roan spotting (Rs) gene will give
showable roans. The other gene is the silvering gene (showable roans
with this one are homozygous recessive). In this post I'll go in depth with
the roan spotting gene, and cover silvering in the next post.

Most breeders that have ever owned roans for show and breeding know
the basics of the roan spotting gene. But there's more to this one than
meets the eye. The roan spotting gene is incompletely dominant,
meaning that there are three phenotypes associated with the various
allele combinations. The heterozygote is intermediate between the two
homozygotes. There are only two alleles, the non-roan allele (rs) and the
roan allele (Rs), which means that rsrs will be our ordinary non-roaned
pigs, Rsrs will be roans, and RsRs will be the lethal micropthalmics. (I
know, old hat to many of us)

This gene will have the same effect on all colored pigs, regardless of what
is found at the other loci. So we can have agouti roans (little or hard to
see roaning on the belly due to the belly band), solid roans, and self
roans. Even whites can be "roaned" but we cannot see it becuase white
intermixed with white will still be white.

So what's so different and interesting about this gene? Well, it is unclear

which modifiers cause the differences between roans and dalmatians. It
has been shown that the same gene (Rs) causes both, but the modifiers
which result in spots rather than intermixed hairs have not been
identified.

BTW, when we cross in a broken to a roan, we cause spotting in the

roans, essentially a broken roan, but I'll get more into that a bit later.

Hey, you guys are going to love this! There are three, not 2 different
ways for a cavy to be roan in color. I was going to talk about silvering
here, and I will, but first I want to talk about the other way a cavy can be
roan, although this one would be a "false" roan, since the only way to
perpetuate it would be to breed two "roans" together! See,
genetics can be fun! lol
The "false" roan could also be called a silver brindle (per Catherine
Whiteway). If you look back to the post on the color locus, one of the
alleles (cr) removes all red pigmentation, leaving nearly all black
pigmentation, giving the animal a "silvered" appearance. This is seen in
silvers, where it removes the red tips of the golden and makes them
white. A brindle is red and black intermixed. So if a brindle has crcr at
the color locus, it would remove all red pigmentation, leaving a black
animal with white intermixing hairs. I love genetics! lol (incidentally, a
brindle pig with crcr and white patching are our magpies)

BTW, this will only work with red/black brindles and the derivatives
(chocolate/red, lilac/reo, & beige/reo) If you had a cream/black, well
remember that cream is a lower c heterozygote, that would be replaced
by crcr and you'd still have a false black roan (silver
brindle). Chocolate/red would yield a false chocolate roan, lilac/reo would
be a false lilac roan, and beige/reo would be a false beige roan. You
would know that strawberry roan, reo roan, and cream roan could NOT be
crcr (remember that the effects of cr is to remove all red
pigmentation!). In those cases, you'd have whites.

Okay, now for the silvering gene (Si). This one is incompletely dominant
(just like Rs), where the heterozygote will yield some intermediate
between the two homozygotes, but rather than the intermediate giving the
best roaning, the homozygote silvered (sisi) pig usually has the best
roaning. Once upon a time, it would seem that most roan cavies were
silvered, roan spotting was apparently a mutation that grew in popularity
(first reported by Catherine Whiteway in 1973). Si is the normal, non-
silvered allele of this gene, the si allele causes a dark face, and white
hairs intermixed over the body. It's important to note that silvering cannot
yield a dalmatian. Only the roan spotting gene can cause spots (and
dapples). So if you've got a roan on your hands that is spotty, it's
probably a good bet that it's Rsrs, while a complete lack of spots may (but
is not guaranteed) be sisi. Of course, the two may be present together as
well, and we would not know what these were by phenotype alone.

We already know that RsRs produces micropthalmics. sisi all by itself will
not produce lethals. However, sisi coupled with another gene called
Diminished (Dm) can result in sickly whites that die young. Dm is the
normal form of the gene, dm in homozygous form and coupled with sisi is
what yields the sickly whites. If you are fortunate to have a line of perfect
roan cavies, that happen to be sisi DmDm, breed them together and you'll
never get a lethal! (yeah right, like there are really any of these left out
there! lol)

In other words, when dealing with the roan spotting gene, it's best to
cross roan to non-roan. When dealing with the silvering gene, it's best to
cross roan to roan (to maintain homozygosity of the recessive and
maintain the best roans), however, crossing these without knowing what
they carry at the dimished locus may also yield sickly whites. lol Aren't
roans a blast?! I never even realized how much fun they could be until I
started looking into this stuff. lol

There are two other genes that give the "appearance" of being roans, but
aren't. They are the grizzling gene (gr) whose homozygous recessive can
cause white hairs to begin to appear on cavies after about 4 months of
age (not born with it). And also the whitish gene (w), which causes white
hairs to appear in black and brown cavies, but which may come and go
with hair growth. I think what we all know as "grizzling" of chocolates is
actually the result of this whitish locus, since it usually appears around
weaning and is gone by senior age (generally).

Now for genetics terminology 102. :)

Wild Type is the (believed) original genetic code, no mutations. It

generally consists of all dominant alleles. In cavies, the believed Wild
Type is the golden agouti. So the dominant allele may also be referred to
as the Wild Type or WT.

Pleiotropy is the

Epistasis is the

Inbreeding is the act of breeding two related individuals, regardless of

what that relation might be. Mother x son, father x daughter, aunt x
nephew, full-sibs, half-sibs, cousins, etc.

Linebreeding is generally believed to be the act of breeding within a

family line. Father x daughter and mother x son. Also grand-mother x
grandson and grand-father x grand-daughter.
Crossbreeding is the act of breeding two different breeds. Such as
Peruvian to Silkie, American to Teddy, or (according to ARBA) Satin
Abyssinian to Abyssinian.

Outcrossing is the act of crossing two genetic lines within the same
breed. Lines are created through linebreeding (inbreeding), and
outcrossing is when you introduce non-related individuals into the line.