Untitled

FUNDAMENTALS
OF COLOR GENETICS IN CANARIES

Reproduction and Control
by Octavio Perez-Beato
PITTSBURGH, PENNSYLVANIA 15222

The contents of this work including, but not limited to, the accuracy of events, people, and places depicted;
opinions expressed; permission to use previously published materials included; and any advice given or
actions advocated are solely the responsibility of the author, who assumes all liability for said work and
indemnifies the publisher against any claims stemming from publication of the work.
All Rights Reserved

Copyright © 2008 by Octavio Perez-Beato
No part of this book may be reproduced or transmitted
in any form or by any means, electronic or mechanical,
including photocopying, recording, or by any information
storage and retrieval system without permission in
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ACKNOWLEGMENTS
I want to express m y love and gratitude to m y wife and m y daughter for all the rem arkable effort
designing the pictures and illustrations; without their help this book would not be possible. My
son, whose encouragem ent and support is invaluable. To the loving m em ory of m y father; he
taught m e so m any things about birds.
Special thanks to all those canary fanciers that I have m et during tim es of m y life, and from
whom I have learned so m uch. I am especially indebted to Antonio Sanchez Berm udez, friend
and colleague. He passed away long ago; a great lost for all those who learned so m uch from his
wise advises. Alfredo Rovere generously provided valuable inform ation in the 70 ’s, when spe-
cialized canary m agazines were not affordable for m e. Dr. Lidia Berm udez has been a trem en-
dous support in the com pletion of this book. Silvio del Valle and Maria Elena Rodriguez were
the pioneers in the first attem pts to get canary pictures for m y book. Karelia Llanes was of a
great help in com puter assistance.
My form er teachers in all courses of Genetics and Anim al Genetics at the University of Havana,
Faculty of Biology, im planted in m e the unbreakable decision to go deeply in knowledge, no
m atter how hard it m ight seem to be at first.
iii
CONTENTS
Acknowledgem ents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii

Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii
Chapter 1: Basic Concepts on Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Chapter 2: Mating System s . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Chapter 3: Pigm ents and Feathers in the Canary. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Chapter 4: The Dom inant White and the Recessive White Canary . . . . . . . . . . . . . . . . . . . . . 17
Chapter 5: The Yellow and the Lem on-Yellow Canary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Chapter 6: The Red Factor Canary. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Chapter 7: The Green and the Brown Canary. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Chapter 8: The Dilution Factor: the Agate and the Isabelle Canary . . . . . . . . . . . . . . . . . . . . 38
Chapter 9: The Pastel Canary: the super dilution factor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Chapter 10 : The Opal Canary: the extra dilution factor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Chapter 11: The Ivory Factor: dilute lipochrom es . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Chapter 12: The Pied Canary Genetics. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Chapter 13: New Mutations. The Incredible Eighties . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Chapter 14: Genetic Possibilities of a True Black Canary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Chapter 15: Obtaining Varieties through Sim ple Pairing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Chapter 16: Concom itant Factors in Color Canaries: other m utations . . . . . . . . . . . . . . . . . . 76
Chapter 17: The Nom enclature of Color Canaries . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Chapter 18: Quantitative Canary Breeding . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
v
INTRODUCTION
Canary breeding has been historically one of the oldest hobbies in the world, taking into account
that the wild canary bird was introduced in Europe, particularly in Spain, as far as the 15 th or
16 th century. At that tim e, that m elodious little bird created a very special predilection am ong
European aristocrats. The canary rapidly spread all over Europe, and consequently variations
soon appear in its genom e. It seem s that at the end of the 17th century two im portant m utations
took place, dom inant white and yellow. Surprising avalanches of m utations have occurred ever
since. Nowadays, we can enjoy a lot of different colors, hues, and shades probably inconceivable
for those forerunner canary breeders of the 17th century.
Today, in our highly technological world canary breeding has not been left behind. As a rule, that
sophisticated beehive of color variations due to mutations, special breeding combinations, and a
very accurately selection can only be achieved properly, with technical and basic scientific knowl-
edge. Is each and every canary breeder in such position and possibility? I’m afraid not.
Unfortunately, many fanciers obtain good birds after struggling through several breeding seasons;
trial and error has been the only tool available in the absence of better knowledge. If good
results could be obtain in three breeding seasons applying technical and basic genetic knowledge,
it shall probably take six or more to obtain the same results, if we ignore such methods.
I have been reading books about canary breeding, as well as m any articles available on the inter-
net. What I have found as m y own conclusion is that, no possible flawless application could be
achieved if the reader is not prepared in basic genetics, anim al breeding techniques, and a prop-
er control of what he/ she is doing during the breeding season, no m atter what the com plexity
of the color variety is, fanciers need to know how things are going; they are drifting, otherwise.
Of course, I know for certain that there is a kind of breeder that rather spend a good am ount of
m oney buying excellent birds, avoiding basic genetic knowledge, breeding techniques, and a
proper technical control on each breeding season. Here, a logical question arises: Is it worth-
while? Furtherm ore: Is it the m erit of the buyer obtaining good and excellent birds next season,
or is it the m erit of the seller who bred the parental strain? The answer is sim ple: the seller is.
In m y own opinion, it is better for any fancier to apply as m uch knowledge as possible, since this
is the core of canary breeding. Better birds based on better knowledge. If you have only regu-
lar birds, and you apply technical knowledge properly, it is possible to obtain outstanding birds
vii
in two or three breeding seasons. You’ll be proud to say: “those are m y very own birds, I m ade
them m yself.” You will be satisfied without spending a lot of m oney, and those birds being bet-
ter than their parents, are exclusively obtained by your personal effort as a canary breeder; a
technically true canary breeder.
This is exactly the purpose of this little book, to give canary fanciers that basic knowledge (and
a little bit m ore) to accom plish canary breeding in a m odern, technical, and m ethodological
way. There are not special secrets in obtaining excellent birds. Applying knowledge properly is
the answer. Canary breeding is only a particular specialty in anim al breeding. High perform -
ance results are based on this. Hence, specific procedures m ight be taken into account.
Som e sim ple suggestions are m andatory to those readers that take this book having in m ind to
thoroughly learn and apply its content. First of all, it is a m ust to understand the content of the
first chapter, sine qua non, it is im possible to go further. Besides, it is im portant that every sin-
gle chapter be analyzed carefully, in order to m aster its content in full. I do not advise reading
a chapter if the previous one has not been thoroughly understood. These rem arks cannot rep-
resent a discouragem ent at all; on the contrary, sm art precautions that guarantee the proper
developm ent of your knowledge.
The author
viii
CHAPTER 1
Basic Concepts on Genetics
The first approach in Genetics is the cell, as it is the biological unit that encloses and guards the
nucleus, in which genetic m aterial is included. Only basic structures of the cell will be consid-
ered, not beyond the concerns of basic Genetics.
Every single tissue, in any anim al or plant, is built by m icroscopic structural and functional
units called cells. They show different sizes and shapes, depending on the function accom -
plished. As an exam ple we can m ention the kidney cells which are com pletely different from the
brain cells, as they perform a very different function. In general term s we m ay say that a cell is
basically form ed by a nucleus, nucleus m em brane, cytoplasm , and cell m em brane, as it is shown
on Fig. 1.
Fig. 1 Basic parts of an anim al cell.
1
Octavio Perez-Beato
The nucleus is the true reproduction center of the cell. Inside it chromosomes are found, and
depending on the cell stage, they appear as very thin filam ent structures, like m icroscopic
threads in a com plex hank. The size, num ber and shape of chrom osom es are different in every
anim al species. The num ber of chrom osom es in the cell nucleus is constant in every species. In
the chrom osom es genes are located, which are the functional units bearers of the genetic traits
in all individuals. Between 1910 and 1928 well conducted investigations docum ented the lin-
earity of genes in the chrom osom es; it is, genes are placed in chrom osom es as beads in a neck-
lace: one after the other.
Cell Division
The production of som atic cells (cells of the body; from soma: body) is accom plished by a
process of cell division called mitosis (from the Greek mitos: thread). Each single cell is divid-
ed in two identical cells, and at the end of two divisions there are four cells as it is shown in Fig.
2. In the process of m itosis different phases are considered: prophase, m etaphase, anaphase,
and telophase; each one corresponding to im portant stages in cell division. Next, those steps
will be roughly explained.
Fig. 2 Two cell divisions in a regular m itosis process.
Prophase: is the starting point of cell division. Each single chrom osom e is longitudinally divid-
ed in two perfect halves, nam ed chromatids; in fact they are strands resulting from chrom oso-
m al duplication. These strands are linked together in just one point called centrom ere, located
at one end of the chrom osom e, in the m iddle or in any other position between the center and
the extrem e end of the chrom osom e, as shown in Fig. 3a. Toward the end of the prophase the
nucleus m em brane disappears, and chrom osom e are grouped in the equatorial plate of the cell.
The so called m itotic spindle is form ed. This is the beginning of the next stage, the m etaphase,
as shown in Fig. 3b.
2
Fundamentals of Color Genetics in Canaries
Fig. 3 Mitosis process in an anim al cell.
Metaphase: the m itotic spindle is com pleted, and chrom osom es are placed just in the m iddle of
that structure coincident with the equatorial plate of the cell. Now, each chrom osom e is seen as
a pair of chrom atids, joined by the centrom ere. At the end of the m etaphase the centrom ere is
gone, but chrom atids are still one next to the other.
Anaphase: now the chrom atids set apart and m igrate to each pole of the m itotic spindle. In fact,
they are true and individual chrom osom es, as shown in Fig. 3c. From now on they are genuine
chrom osom es. We m ay realize that a fundam ental part of cell division is the own division and
duplication of chrom osom es, them selves.
Telophase: the m itotic spindle disappears, and chrom osom es becom e m uch longer, it is not
possible to tell them apart. The cell starts a narrowing process in the center of its whole body,
as shown in Fig. 3d. , ending up with cell com plete division, and creating two daughter cells
which bear the sam e am ount of chrom osom es as the m other cell, due to the m igration of chro-
m atids-in fact, true chrom osom es at this tim e- to opposite poles of the m itotic spindle being
included in equal am ounts in both daughter cells. See Fig. 3e.
All of the above, as was m entioned before, is regarding som atic cell division. Som ething differ-
ent occurs when division involves the creation of gam etes – egg-cells and sperm atozoids- which
is known as m eiosis or reduction division. Germ line cells only bear half of the am ount of chro-
m osom es com pare with som atic cells. If the gam etes of a certain anim al species bear n chro-
m osom es, then its som atic cells bear 2n chrom osom es, known in Biology as the diploid num -
ber, while the gam etes own a haploid num ber n. The reason for haploidy and diploidy phe-
nom ena shall be understood at the end of the following.
3
Octavio Perez-Beato
Meiosis
During the normal reproductive animal cycle, testis and ovaries play an outstanding role form-
ing reproductive-oriented cells. First, they create a special kind of cells, which bear in their nuclei
a diploid number, the same amount of chromosomes as in any regular somatic cell. When these
special cells are transformed in a very particular way to create spermatozoids and egg-cells – male
and female gametes- then, meiosis starts. It ends when spermatozoids and egg-cells are com-
pleted, depending if the process is performed in a male or in a female. The so called special cells
bear the same amount of chromosomes as in any somatic cell; the biological process starts as in
the mitosis that was explained before; but there is a big difference. During the mitosis, in the step
known as metaphase all chromosomes are located one next to the other, on an individual basis,
but in the meiotic metaphase chromosomes are located in two different groups, and in each par-
ticular group only one chromosome from each pair, is found. Remember that chromosomes exist
in pairs. Fig. 4 shows the difference between metaphase in mitosis, and the one in meiosis. Once
they are located in such positions, the first division is accomplished which in fact is oriented to
totally separate both groups of chrom osom es, which in turn will create two cells, being each one
loaded with only one single group of chrom osom es. Nevertheless, we m ay observe that these
chrom osom es are longitudinally differentiated in two chrom atids. Fig. 5 shows the first and sec-
ond m eiotic division. In the last one each chrom atid is separated from the other, and in turn
becom es an individual true chrom osom e, in each of the four cells originated at the end of the
second division. From here on these cells are in fact egg-cells or sperm atozoids, accordingly.
Fig. 4 In (a) chrom osom es are one next to the other. In (b) there is one single chrom osom e of
each pair in each group; typical m eiotic m etaphase.
All this biological process has been explained in a very sim ple and schem atic way on behave of
sim plicity and easy understanding, but the biological reality is so m uch m ore com plex.
As you m ay see in Fig. 5, not four egg-cells are produced at the end of the second m eiotic divi-
sion, only one egg, instead. The other three cells undergo a degenerative process and stop
developm ent. On the other hand, it is not the case regarding the four sperm atozoids since they
really com plete full developm ent, from each cell that reaches m eiotic m etaphase. It is also
4
im portant to take into account that in m eiosis two cell divisions are accom plished, but only
one single chrom osom e division is perform ed.
Now, one single question arises: What is the objective of the meiosis process to allow spermato-
zoid and egg-cells to carry only half of the chromosomes that normally bears any somatic cell? A
very simple answer exists if we consider what the objective of spermatozoids and eggs is. The
objective is nothing but fecundation, from which a new individual shall be born. This new biolog-
ical being will bear in each and single somatic cell the same amount of chromosomes as its par-
ents, and as the rest of the individuals of its same species. This could be accomplished exclusively
if the spermatozoid contributes with half of the chromosomes, and the egg-cell with the other half.
Fig. 5 Meiotic process to create m ale and fem ale gam etes.
The cell form ed with these contributions is called zygote, and of course will bear the sam e
am ount of chrom osom es as is custom ary in any individual of its sam e species. Such a process is
clearly shown in Fig. 6.
Fig. 6 The approach of a sperm atozoid to an egg-cell is depicted, together with the inclusion of
the sperm atozoid inside the egg m ass.
5
Octavio Perez-Beato
The contribution in term s of chrom osom es from both gam etes readily creates the zygote. Soon,
the zygote will start m itotic division, and a new living thing is form ed here on. This new indi-
vidual has inherited traits from its parents, loaded in the chrom osom es located in each and
every one of its cells, which are nothing but replications of the original chrom osom es from the
sperm atozoid and egg-cell, from which the zygote was originated.
Terminology and Definition of Some Basic Genetic Concepts
As it was dem onstrated in previous paragraphs, chrom osom es are paired in every som atic cell,
and in an unpaired fashion in the gam etes. If we recognize the existence of pair of chromo-
somes, this m eans that each pair is carrying a whole series of genes, that in turn are also locat-
ed by pairs on specific locations along the chrom osom e, except in the chrom osom es called XY.
From this, it is quite sim ple to assum e that there are specific locations along each single chro-
m osom e where genes are placed. The specific location for a gene on any chrom osom e is known
as locus; plural loci. Each single gene located in the sam e chrom osom e pair, is called allele.
In m odern term s and sim ple way, the gene m ust be defined as a factor that determ ines the
inheritance of a specific trait in a biological organism .
When both alleles of a gene that determine a certain trait are identical in both chromosomes
– homologous chromosomes- it is said that the individual bearing this combination is homozygous
for that specific trait. On the other hand, if the alleles of a gene are different, then it is said that the
individual in heterozygous for that trait. In Fig. 7 all the aforementioned concepts are depicted, in
a very schematic way. If we refer to this figure in a hypothetic sense, we may express the follow-
ing: the gene that determines eye color is located at locus H comprising two alleles, A and a. Then
again, if a gene comprises two alleles three different combinations are possible: AA, Aa, and aa,
being the first and third combinations the homozygote stage. The Aa combination represents the
only possible combination for the expression of heterozygosis. See Fig. 7a, b and c.
Fig. 7 Graphic representation of a single locus (H) for a theoretical gene with two alleles.
6
Dominance, Expressivity, and Penetrance of a Trait. Phenotype and Genotype.
It is said that an allele is dom inant regarding the other allele in the gene, when the heterozygote
– for exam ple, Bb- only shows the trait determ ined by the allele B.
Then we m ay say that b is recessive regarding B. Consequently, all individuals bearing BB or Bb

are no different at a glance. Nevertheless, BB is hom ozygous regarding trait B, and Bb is het-
erozygous, since b is the other allele in this pair. Taking this into account we sim ply reach the
concept of phenotype, which is precisely what we observe externally in any living thing. This
m ean that based on phenotype only, BB and Bb are identical, since both show the sam e trait B
(dom inant trait), but from a genetic point of view they are different, since Bb bears the b allele,
and BB does not. That’s why we m ight say that their genotypes – genetic types-are different. Not
always dom inance is shown in this sim ple form , since there are cases in which the heterozygous
develops and shows characteristics in between. A little bit of B, and a little bit of b. Then we m ay
say that it is an intermediate dominance. Som e other tim es the dom inance is incomplete, since
the recessive allele gets som e expression on the phenotype. Furtherm ore, we have to recall two
im portant phenom ena regarding any specific trait. It is possible that a dom inant allele will not
be expressed the sam e all over the population. Som e few differences are readily seen from one
bird to the other. These ranges of signs are called variable expressivity. Such is the case regard-
ing the frost character in canaries. We m ay see birds with a m ild frost up to heavily frost. This
range of appearance is telling about a variable expressivity.
The other phenom ena that I want to m ention is the reduced penetrance which refers to the pro-
portion of birds expressing a particular trait. For exam ple, let us suppose we have 50 canaries
bearing a certain genotype; only 40 show the corresponding phenotype. Consequently, the pen-
etrance of the trait is 40 / 50 = 0 .80 ; this m eans penetrance = 80 %.
We ought to consider that expressivity and penetrance are both affected by environm ental fac-
tors such as: tem perature, light, nutrition level, etc. and by intrinsic factors as age, sex, health
condition, etc.
Sex Linked Inheritance and Sex Determination
When a certain gene is located in the sexual chrom osom e X, we say that the inheritance of the
trait is sex linked. That is why a m ale canary could be heterozygous or hom ozygous for any sex
linked trait, since the genetic form ula for a m ale canary is XX and then the m ale m ay carry the
sam e allele in both chrom osom es, or different ones in each X chrom osom e for any particular
trait. On the other hand, the genetic form ula for the fem ale canary is XY. On the Y chrom osom e
there is not any allele of our concern, and we usually say that Y chrom osom e is em pty. Then
again, for a fem ale it is only possible to bear sex linked traits (genes) on the only one X chro-
m osom e that she possesses. It is clearly seen that a fem ale canary never ever could be het-
erozygous for any sex linked trait.
The opposite situation is for the traits located in non sex chrom osom e and then we say the spe-
cific trait autosomic.
7
Octavio Perez-Beato
Proportion of Genotypes Expected in a Particular Mating
It is of the m ost im portance for any fancier to precisely predict the genotypes of descendants,
considering the genotypes of the selected parents. Obviously, fanciers in general are only inter-
ested on one, two or three traits in the breeding accom plished. Hence, all efforts are oriented
on the calculation to accurately predict the descendants’ genotypes. Let us assum e that a m at-
ing is accom plished between an AA genotype m ale canary and an Aa genotype fem ale canary.
The purpose of the fancier is to ascertain which genotypes will be found in the offspring, and
what the proportion of such genotypes shall be. This is accom plished by m aking a sim ple chart
widely used in genetics, known as the Punnett squares.
Fig. 8 Exam ple of a Punnett squares. Depicted are four groups of descendants – one in each
cell- for a theoretical locus with two alleles.
Fig. 8 shows such chart where colum ns are depicting the fem ale canary gam etes – one allele per
gam ete-and then rows represent the m ale canary gam etes. Each cell represents the allele input,
one from the m ale and one from the fem ale. Now we have two cells with the genotype AA, and
two cells with the genotype Aa. If we obtained two cells with the genotype AA, out of four, then
2/ 4 = o.5o, which m eans that 50 % of the offspring will bear genotype AA. This is a sim ple hypo-
thetic exam ple where a single gene with only two alleles has been taken into account. If we need
to consider two genes with two alleles each, this could be this way: m ale canary’s genotype Aa
BB; fem ale canary’s genotype Aa Bb. Then again, the Punnet squares is shown in Fig. 9.
Fig. 9 Punnett squares showing eight offspring groups

considering two loci with two alleles each.
As in the form er exam ple, and as in any other case, each single cell represents a particular geno-
type which could be obtained in the expected offspring.
8
Based on this, if we are interested in obtaining the m ale canary genotype in the offspring for
both involved traits, the inform ation from the Punnet squares will provide us with the expect-
ed proportion of that specific genotype in the expected offspring. As we got two cells with geno-
type AaBB out of eight cells, the expected proportion of that genotype in the offspring will be:
2/ 8 = 0 .25 which indicate that 25% of the offspring shall bear the sam e genotype as the m ale
canary regarding the considered alleles. In the upcom ing chapters the reader will learn the alle-
les com prising the genes involved in the color inheritance in the canary.
Using the Punnet squares the fancier can m ake calculations to know the proportion, of every
single genotype, to be expected in the offspring.
At this point, I realize it is quite convenient to m ake clear som e few aspects, for those readers
that are not fam iliar with genetics; it is, if the total expected genotypes in the offspring is 4, 5 or
m ore, and the num ber of chicks born is sm all, chances are that som e genotypes will not appear
in the expected offspring, since all these proportions are based on probabilities.
9
CHAPTER 2
Mating Systems
There are two basic m ating system s: inbreeding system , in which related birds are bred, and out
breeding system in which not related birds are used in breeding. Besides, the relation m ight be
a close one, or a collateral one, depending on the purpose of the fancier, and the breeding pos-
sibilities based on the available stock. Below, the schem e for such m ating is offered as a guide
for the fancier to accom m odate the objectives of breeding procedures. First, it is im portant to
establish som e sim ple definitions to better understand what is printed in the schem e below.
Line-breeding: mating in the

direct line of descent. In line-
breeding, what is usually
obtained is a discrete homozy-
gosis for selected traits.
Lineage: group of individu-

als belonging to a specific
breed (in our context: color
anaries) which are hom ozy-
gous for certain traits.
Breed: a specific group of

individuals which bear a rel-
ative hom ozygosis for specif-
ic traits, but that hom ozygo-
sis condition is m ore dis-
crete than in the lineage.
Som etim es, it is difficult to

establish a definition between
lin eal or lin eage con dition ,
since lineal relation is a genet-
10
ic trend toward lineage. Inbreeding tends to hom ozygosis. Then, if we are interested in creating
a lineage we ought to use inbreeding procedures.
Since inbreeding creates hom ozygosis, it is expected that recessive traits not frequently seen in
the population will suddenly appear, just because now they are in a hom ozygous condition,
openly expressed in the phenotype. The great advantage is that all those unseen traits now go
up on the surface. But careful, they could be desirable or undesirable ones. Now culling is
m andatory. We cannot be tolerant, not with the slightest fault. The m ost m eticulous selection
ought to be applied. Do not hesitate about culling.
If you proceed this way, you can be sure good traits will be incorporated in your birds.
Nevertheless, inbreeding has a well known disadvantage, characterized by what is known as
depression due to inbreeding. This problem is evident by the presence of a low fertility level,
viability in hatchlings is low, som e cases of sterility m ay suddenly appear, and other distur-
bances related to reproductive traits. Fortunately, experim ents have dem onstrated that, norm al
reproductive levels are recovered as soon as we start m ating inbred lineages am ong them selves.
On the other hand, out-breeding leads to heterozygosis which in turn produces a phenom enon
known as heterosis; it is the counterpart of depression due to inbreeding. We m ay say that het-
erosis is present when m ating individuals of different lineages, breeds, etc. the F1 offspring
obtained is better than the parental average, or superior to the best of the parents.
Any experienced canary breeder has observed, for sure, the outcom e of a bird sharply superior
com pare with its parents. This is heterosis.
Breeding not related individuals has the disadvantage of possible introduction of traits not con-
venient to our goals. Furtherm ore, it is highly dangerous from a genetic point of view to intro-
duce new birds, if the fancier is already working to establish a lineage for certain specific traits.
Next, som e m ating schem es are shown for the breeder to have an illustrated reference, in order
to set up breeding program s according to what is available and possible.
Fig. 10 shows m ating that start with m ale (1) and fem ale (2), producing an offspring (3), which
bears 50 % from the father (1), and 50 % from the m other (2). All m ales from group (3) will be
selected according to what the breeder wants; the best of those m ales will be back-crossed to
fem ale (2), then obtaining offspring group (4), which possesses 75% from her m other (2); the
best m ale from group (4) will be back-crossed again to fem ale (2) obtaining the offspring group
(6), which possesses 87.5% from fem ale (2).
Then again, the best m ale from group (6) will be back-crossed to fem ale (2) to create the off-
spring group (8), which now possesses 93.75% of the phenotype from fem ale (2). This group (8)
is alm ost a copy of fem ale (2), very near to a clone. Using this m ating procedure we replicated
the phenotype of fem ale (2), in approxim ately 94% in a whole group of descendants.
What is said before regarding fem ale (2), is true for m ale (1) represented in the left hand side of
the schem e and fully developed in a parallel fashion. In this way the breeder will have descen-
dants in both lines of the breeding lineage: m aternal and paternal, which is m andatory for the
continuity of the breeding system and genetic im provem ent, as well.
11
Octavio Perez-Beato
Finally, the best birds from group (8) will m ate the best from group (9), to obtain group (10 ) off-
spring, which is sim ilar to the first offspring (group 3) from m ale (1), and fem ale (2). Besides, if
group (10 ) m ates to group (8) and (9), then we will obtain groups (11) and (12), which turn to
be quite sim ilar to groups (4) and (5) from previous years, since group (4) and (5) represent
24/ 32 of fem ale (2) and m ale (1) genotype, respectively, and then groups (12) and (11) represent
23/ 32 from fem ale (2), and 9/ 32 from m ale(1), respectively.
We m ay continue m ating groups in analogous form , taking into account that birds from group
(8) are very sim ilar to fem ale (2), and birds from group (9) are very sim ilar to m ale (1). Keep on
breeding in a sim ilar way to obtain groups sim ilar to (6) and (7), and (8) and (9), accordingly.
I have explained in full details Fig. 10 , since it is the basis to understand Figs. 11 and 12; both
are also based in linebreeding procedures; interpretation rests on sim ilar considerations.
It is of the m ost im portance to start with birds carefully selected, and if possible of known pedi-
gree, no m atter what the linebreeding schem e is applied, rem em ber that inbreeding itself is a
tool to surface faults, and then again culling m ight go to great length if the m ating starting-line
is not attem pted with good quality birds.
Fig.10 Inbreeding schem e to obtain

nine offspring groups
Fig. 11 Inbreeding system where

m ales A and B are siblings,
and fem ales C and D are not related.
12
Fig.12 Both original fem ales are

not related. On the last m ating a
heterosis effect m ust be evident
in the offspring. Arrow heads are
used as in the previous figure.
13
CHAPTER 3
Pigments and Feathers in the Canary
The plum age color in the canary is biologically conditioned by m eans of two pigm ents: the
melanins and the lipochromes. The com bination of these two substances in the feathers of the
canary, give birth to all com binations that we adm ire today in our beloved feathery pet. Besides,
m elanins and lipochrom es are genetically affected by other factors, which will be the topic of
upcom ing chapters. These substances are fully studied by the science of Biochem istry, which is
in charge of the research and study of all the chem ical processes in living things.
The Melanins
The canary bird possesses two types of m elanins, though som e authors state that are m ore than
two. For the sim plicity of explanations, and better understanding of basic concepts on this
topic, I will consider only two m elanin types throughout the book. These m elanins are: eume-
lanin or black m elanin, readily seen in Green, Bronze, and Blue intensive canaries. The
pheomelanin (brown m elanin) is a pigm ent very well seen in Yellow Brown and White Brown
canaries, am ong others. These m elanins are biochem ical elaborated by the canary itself, based
on the genetic inform ation of the bird. It is not enough that a canary could elaborate its own
m elanins, it is m andatory that very special and specific substances have to be present in this
biochem ical process. These substances are known as oxidative enzym es – oxidases-, which act
on the m elanin, and then developing the appearance of each type of pigm ent. If this enzym e
would not be present to carry on the oxidative action, it will be im possible to see eum elanin and
pheom elanin in the plum age of our canary.
The Lipochromes
It is necessary to explain the etymology of the word lipochrome. Lipo means oily, and chrome
means color, it is a colored substance prone to be dissolved in oil, or oil chemical-affinity substance.
14
These pigm ents are present in the canary diet, then m etabolized and fixed in its skin; as feath-
ers are epiderm al products any canary with the proper genetic inform ation shall deposit
lipochrom es on its feathers. Lipochrom es range from yellow to red, and canaries get them by
m eans of the m etabolic process of carotenes and xanthophylls, which are substances fully
em bedded in green vegetables. Carotenes are abundant in carrots, and in a lot of green leave
vegetables. Xanthophylls, yellow color substances, are also present in green leaves and green
parts of m any plants. The darker the green color the better as a canary food.
The genetic aspect regarding the way the canary transform s carotenes and xanthophylls into
pigm ents to color its plum age, will be considered in detail in the chapter about White Dom inant
and Recessive White. As a previous exam ple we m ay say that the Red Siskin has the necessary
genetic inform ation to transform carotenes into red pigm ent, red lipochrom e, which is deposit-
ed in the m ale’s feathers and on a discrete fashion, in the fem ale’s, too. On the other hand, the
yellow canary is able to transform xanthophylls into yellow lipochrom e, and then deposit it in
its feathers. I strongly agree with other authors that have stated four fundam ental steps to set
up the proper color in the canary; these steps are:
a) Vegetables substances – carotenes and xanthophylls- m ust be assim ilated by the bird.
b) Those substances m ust be transform ed into true pigm ents, due to the m etabolic action
of the canary organism itself.
c) The pigm ents already elaborated m ust be concentrated in the feathers. In fact, these
pigm ents are concentrated in the skin of the bird; as I said before feathers are of epi-
derm al origin; these pigm ents will be deposited in the prim ordial feather structures in
the skin. Finally, they are seen in the full-grown feathers, if the canary has the proper
genetic inform ation to do so.
d) As the last step, other substances acting according to the genetic inform ation present
in the genotype, will determ ine in what way and how m uch of those pigm ents will accu-
m ulate in the feather structures. The genetic aspect of these four steps will be explained
in the upcom ing chapters.
The Frost Feather and the Intensive Feather in the Canary
There are two feather structures in the dom estic canary; the frost feather and the intensive
feather. The frost feather presents the barb extrem es with no lipochrom ic pigm entation at all,
which m ake the plum age to appear like frosted. At the sam e tim e this kind of feather is soft, like
a down. On the other hand, the feather known as intensive reveal lipochrom e pigm ent up to
edge of the barbs, being shorter and m ore rigid than the frost feather.
The frost and intensive feathers are controlled by two alleles; the intensive is dom inant, the
frost if recessive. This m eans that three different genotypes are possible regarding this kind of
feather structure. Let I be the dom inant allele for the intensive feather, and i the recessive allele
that yield the frost feather. Then there are three possible genotypes: II, Ii, and ii. Genotype ii
canaries are frost, and II and Ii are intensive. Now, is m andatory to fully explain what to expect
15
Octavio Perez-Beato
in genotype II birds. This hom ozygous dom inant is a bird with long fingers and nails, extrem e-
ly nervous, and cases have been reported of birds suffering from seizures. These characteristics
are seen only if the bird reaches full grown stage, since it is frequently observed that they die
during the em bryonic process; sudden death in the nest. If it survives, definitely will be a degen-
erated biotype. Besides all previous characteristics, its plum age is short and poor, absence of
feathers around the eye circles, and its general appearance is ragged. Based on all of the above
is certainly stated that allele I in hom ozygous condition is sub-lethal. There is som ething else,
allele I shows an incom plete dom inance, and variable expressivity since birds with genotype Ii
ranges from a true and harm onic intensive appearance up to birds with a very m ild frost. These
m ight be excellent breeders but not recom m ended as show quality birds.
On the other hand, if two frost birds are m ated, the offspring will be com posed of excessively
feathered anim als, rather lazy, and not really good in rearing their chicks. On the average, there
will be no problem s regarding sub-lethal effects, but they do not m atch with any standard.
Fig. 13 Mating frost and intensive will yield an offspring com prising frosts and intensives
in the sam e proportion.
According to what was explained in the previous paragraphs regarding frost and intensive
feathers, m ating has to be carried between frost and intensive, and results will be in accordance
with what is shown on Fig. 13. Theoretically, 50 % of the offspring will be intensive, and the
other 50 % will be frost. The balance is perfect.
16
CHAPTER 4
The Dominant White and the Recessive White Canary
The dom inant white is the m ost com m on white canary, is the one that is seen in any aviary, fully
represented in any bird show, as a rule. Its color is really white, but a lipochrom ic tint (yellow,
orange or red) is apparent in the prim ary wing feathers, and som etim es extended to the coverts.
It is believed that this canary appeared at the end of the XVII century. It was a m utation. These
birds are white color just because they are unable to deposit lipochrom es in its feathers, which
is determ ined by a gene F that, in heterozygous condition (Ff) produces a dom inant white phe-
notype; if it appears in hom ozygous condition FF, the bird bearing such com bination will die
during the em bryonic stage. That is why it is said that, the gene that determ ines the dom inant
white is lethal in hom ozygous condition. As a result, any dom inant white canary is heterozygous
for that specific trait. This is a dom inant autosom ic gene. Nevertheless, there are other loci
where im portant genes involved in different m etabolic steps regarding lipochrom es, are locat-
ed. I will explain in detail the theory that supports this genetic event.
Locus K
The dom inant allele K allows the norm al assim ilation of carotenoids. The recessive allele k does
not allow the norm al assim ilation of these substances. All birds with genotypes KK or Kk will
assim ilate the carotenoids on a norm al basis. Unfortunately, the genotype Kk does not get
enough carotenoids to be lately transform ed into yellow pigm ent. This aspect will be discussed
in the next chapter.
Locus G
The dom inant allele G allows the conversion of carotenoids into yellow pigm ents.
On the other hand, the recessive allele g does not allow such conversion. All canaries with geno-
type Gg will readily transform the carotenoids into yellow pigm ent, though they are bearing the
recessive allele g. This allele g is just a convenience, and the theoretical hom ozygous gg is called
Recessive German white, never obtained as a biological reality, and only accepted nowadays as
17
Octavio Perez-Beato
a plausible theory. In fact, the Recessive Germ an white is a potential m utation. It has never
shown up, unfortunately.
Locus F
The recessive allele f is responsible to elaborate an enzyme that transfers the lipochromes into the
feather structures, no matter what if they are red or yellow substances. The dominant allele F inhibits
the deposition of the lipochromes in the feathers. This is exactly the allele that carries the dominant
white canary, which genotype is Ff responsible for not allowing the deposition of lipochromes in the
feathers, except in the primary wing feathers and upper coverts, as mention somewhere before.
The Recessive White Canary (English-White)
This beautiful bird is the product of a mutation that appeared in New Zealand by the year 190 8. It
showed up from the mating of a couple of yellow canaries. The well known canary fancier A. K. Gill
from England, stated in his book New Coloured Canaries that also in 190 8 the same mutation
appeared in London. In a periodical magazine specialized in canary breeding issued in 1967 was
reported again the outcome of the mutation ten years before, from a couple of yellow canaries.
Many authors take for granted that after World War I recessive white canaries alm ost disap-
peared. Taking into account all of the above, it is obvious that the m utation responsible for the
recessive white canary has occurred several tim es in the past. This m eans that it seem s to be a
recurrent mutation.
This mutation inhibit the normal assimilation of carotenoids, rupturing the metabolic chain for
the appearance of lipochromes in the plumage, giving way to the purest white color in the canary.
The plum age of the recessive white does not show any trace of lipochrom e, as is the case in the
dom inant white canary. Besides, the skin of the recessive white canary is bluish color.
These birds as being unable to assim ilate the carotenoids, cannot either elaborate the vitam in
A, which is a norm al biochem ical process in the rest of the canaries, from the carotenoids in
their food. This is the reason why it is necessary to supply vitam in A in the diets of the recessive
white canaries.
Genetic Characteristics
Based on all that has been discussed regarding the dom inant white and the recessive white
canaries, now we are able to discuss the genetic form ulas that characterized these birds.
The dom inant white canary does not deposit lipochrom e pigm ents in its feather structures,
except on the areas already discussed in previous paragraphs. But certainly does assim ilate the
carotenoids, and also transform s them into lipochrom e pigm ents. Then, according to the func-
tion of the previously explained loci in this chapter, now we have that a dom inant white canary
18
has the genetic form ula: KKGGFf; and a recessive white canary is form ulated as kkGGff. As I
have previously expressed, the allele F is lethal under hom ozygous condition; enough reason to
explain that m ating two dom inant white birds the obtained offspring will show a 25% of em bry-
onic m ortality. See the chart below:
Fig. 14 The offspring group (1) is hom ozygous for allele F, which is detrim ental to the
em bryonic developm ent. The offspring group (3) are yellow canaries, which genetic form ulas
will be used and explained in the next chapter.
Using the Punnett reticulum it is possible to check that 25% of the offspring will be hom ozygous
regarding allele F. Next chart shows the results:
Fig. 15 The Punnett reticulum shows the genotype proportion as a result

of m ating a Dom inant White m ale and fem ale.
Cell (1) represents hom ozygous FF, which will not com plete their em bryonic developm ent. At
the sam e tim e this group represents 25% of the offspring. Besides, cells (2) and (3) represent
50 % of the offspring which are dom inant white canaries. Finally, cell (4) represents yellow
color offspring.
It is very im portan t for the reader to un derstan d every sin gle detail explain ed in this partic-
ular chapter, sin ce it is the basis to really un derstan d the gen etic prin ciples regardin g the
in heritan ce of classic white can aries. Besides, if the reader takes hold of these fun dam en tal
con cepts the followin g chapters will be easy to un derstan d, m ain ly when readin g will reach
all the theory an d statem en ts on the Red Can ary, which is the m ost attractive approach to
lipochrom e gen etics.
19
CHAPTER 5
The Yellow and the Lemon-Yellow Canary
The yellow canary appeared as a m utation between 1680 and 1713, as a descendant of canaries
from the Canary Islands, introduced in Europe in the XVI century.
In a previous chapter has been explained the biochem ical action of carotenoids in the canary
plum age. Besides, in chapter 4 the genetic form ulas of the white canaries have been established.
Now again, the yellow canary as a counterpart of the white recessive canary, fully assim ilate the
carotenoids, in particular those pigm ents that will be transform ed into yellow lipochrom e.
In Chapter 4 the genetic function of alleles located at K, G, and F loci have been stated.
Consequently, we should advice the reader to go over that chapter again. In it, the genetic for-
m ula of the yellow canary (KKGGff) was established as the result of m ating two dom inant white
pair. See Fig. 15 as a reference. As the allele K is dom inant over allele k , it is possible to say –
from a genetic point of view- that a yellow canary m ay also bear, theoretically, the genetic for-
m ula KkGGff, which will not affect the norm al assim ilation of carotenoids. This heterozygous
canary regarding locus K will show a poor yellow lipochrom e in its plum age, since the dom i-
nance of allele K is incom plete then producing a partial assim ilation of carotenoids.
This could be a first hand explanation for those poor colored yellow canaries, though they have
been properly fed with carotenoids. Now it is m andatory to consider a special point about locus
K. If we take a closer look to the previous form ula KkGGff though it is obviously characteristic
of a poor yellow color plum age, it is also indicating a white recessive split bird, since it is het-
erozygous Kk. At this point of the analysis I particularly do not think it is the genetic explana-
tion for the poor yellow color in the plum age. Nonetheless, it could be the genetic form ula that
bore the parents of the white recessive birds that appeared in different tim es in the past, which
pointed to the possibility that the white recessive trait is a recurrent mutation, as was stated in
Chapter 4.
20
The lem on yellow canary is widely known am ong canary breeders. This bird does not possess
different lipochrom es than those bears by the com m on yellow canary. The special appearance
of its lem on-yellow color is due to a special feather structure. When sunlight strikes on that kind
of feather, its special structure acts as a filter to allow the reflection of certain wave-lengths only,
which are responsible for the appearance of the well known lem on-yellow color. This feather
structure is genetically controlled. Let us call allele s the one that determ ines such special struc-
ture, and S the allele for the com m on feather structure.
The allele s is known in canary breeding as blue optic factor. Taking into account what is known
for recessive alleles, a lem on-yellow color canary has to carry ss in its genotype to be a visible
lem on-yellow color bird. In other words, that canary has to be hom ozygous ss to show such a
color as its own phenotype.
As the reader surely knows the genetic form ula of the yellow canary from a previous chapter, let
us add the optic factor gene to that basic form ula to obtain the lemon-yellow canary: KKGGffss.
It is im portant to know that the allele S that determ ines the com m on structure of the feather,
acts based on an interm ediate dom inance, hence the heterozygous canaries Ss show an inter-
m ediate phenotype between com m on yellow and lem on-yellow, known as m id-lem ons. It seem s
that this denom ination was form erly acquainted in France by the French term demi-citron
(half-lem on).
Nowadays, in many bird shows rules establish no difference between yellows and lemon-yellows;
they all concourse under yellow, (frost and intensive) though lemon-yellow is a genetic reality.
An interesting visual experience that we m ay observe to understand what the special structure
is, occurred when an Indigo Bunting (Passerina cyanea) takes a bath, usually in a bathing dish
available in m any backyards during the fall and winter m igration. When it gets wet, the irides-
cent blue color vanishes, and the bird plum age turns into a rather dark brown color. The char-
acteristic blue color will not return until the feathers are thoroughly dry. What really happened
is that when the feathers got wet, the special structure to reflect the specific blue wave-length is
m om entarily altered in its surface, due to the water, and then is unable to properly reflect such
a color. What we see at that very m om ent is the true pigm ent underneath the surface structure,
since the blue color in birds, no m atter what the species is, does not exist as such, it is always
the effect of that special structure on the surface, which m ay produce from a very light to an iri-
descent m etallic blue color.
It is an interesting rem ark that the dim orphism apparent between the m ale and fem ale of the
Indigo Bunting is due to this optic blue factor only present in the m ale plum age. Unfortunately,
our canaries do not posses such an efficient feather structure to reflect the true blue color, and
by the m om ent tim e we only can adm ire that m ild blue optic factor in the lem on-yellow and
other colors that will be discussed in upcom ing chapters.
Now, the reader can add another allele to the general genetic form ula of the com m on yellow
canary, since it has been established the norm al allele that denotes absence of the blue optic fac-
tor; the com plete genetic form ula is KKGGffSS for the com m on yellow canary, and for the
lem on-yellow is KKGGffss
21
Octavio Perez-Beato
From the m ating of a lem on-yellow bird to a non-lem on-yellow, the offspring will be as shown
on Fig 16.
Fig. 16 Mid-lem on proportion offspring -10 0 %- from a yellow and lem on yellow m ating.
As readily seen in the previous chart, the whole offspring will bear the optic factor, and then
they will be m id-lem on birds. If the conducted m ating is m id-lem on and lem on yellow, the off-
spring will be : 50 % m id-lem on , and 50 % lem on yellow. See Fig. 17
Fig. 17 Offspring proportion from Ss and ss genotypes.
The reader has to be acquainted that the optic factor for blue color is an autosom ic trait, accord-
ing to what has been shown on the previous paragraphs and charts.
22
CHAPTER 6
The Red Factor Canary
It was in 1926 that the first attem pts to create a red canary were m ade. I m ay say, and I think I
am certain that it was Dr. Duncker the first to be interested about genetics in the canary.
Nevertheless, by m eans of Dr. Duncker him self it was known that the first canary breeder that
carried experim ental m ating with the red siskin (Spinus cucullatus) and the fem ale canary was
nam ed Mr. Adam s, but the m ethodology to get the hybrids was stated by Dr. Duncker.
I also have to m ention B. Matern and F. M. Durham ,that m ade im portant contributions regard-
ing the Red Siskin and the canary hybridization process. A doubtless top contribution was the
book New Coloured Canaries by A. K. Gill offering in its tim e full inform ation regarding all con-
cerns about the red canary.
I think it is necessary to m ake a review, though it m ight be sim ple, about theories regarding the
red canary genetics. According to Dr. Duncker theory the yellow lipochrom e was absent in the
red siskin. Then in the m ating of a red siskin and a yellow fem ale canary, the hybrid will get the
red factor from the red siskin, and the yellow color from the fem ale canary, being this com bi-
nation responsible for the “copper color” shown in F1. Next it follows a back-cross F1 m ale with
a yellow fem ale canary; the offspring cam e to be: copper color, pied-oranges, pied-yellows, light
oranges, and light yellows. The m ales were fertile, but not the fem ales. Strikingly, dissections
m ade by Dr. Duncker on these F1 fem ales, dem onstrated and absent of reproductive organs.
In his book Mr. Gill provides a very interesting inform ation from his own experience about the
F1 cross to fem ale canaries; the figures are as follow:
Total m ates 52
Total eggs laid 215
Infertile eggs 171
Dead in the shell 4
Total chicks hatched 40
Total chicks reared 30 (22 m ales, 8 fem ales)
23
Octavio Perez-Beato
From this inform ation it is possible to conclude that fertility was really low, hatching level is
also low, as well as surviving chicks, under the standards of canary breeding. Nonetheless,
though all these figures were really low we have to take into account that, it was an attem pt to
hybridize two different species, in which the reproductive results are usually very low.
On the other hand, we m ay observe that the num ber of eggs per nest is on the average expect-
ed for a regular brood in the canary. This reproductive trait seem s not to be affected in this
cross, which is certainly logical since it is the fem ale canary and not a hybrid fem ale.
Let us keep on discussing about the crossing schedule run by Dr. Duncker, to know exactly what
was the conclusion he arrived to. The crossing red siskin x yellow fem ale canary yielded the so
called “copper color”, which in turn when backcrossed to yellow canary nothing better than light
oranges were obtained in the offspring. At this point Dr. Duncker stated that the m ain problem
to obtain the red canary was precisely the yellow color contribution from the yellow fem ale
canary. Then, he accom plished the red siskin x dom inant white fem ale canary cross, expecting
that the white dom inant gene shall be an obstacle for the yellow to show up, but not for the red
factor which would be fully expressed in the F1 plum age. Unfortunately, it was not that way
either. The F1 offspring were copper and ash grey color, in both sexes. So, it was confirm ed that
introducing the dom inant white did not do any better in obtaining a good red, and not a copper
color. From this last experim ent it was obvious that the dom inant white gene inhibit the expres-
sion of the red color as it does with the yellow color.
Dr. Duncker’s opinion suddenly changed course; then he stated that it was m andatory to
accom plish the cross red siskin x recessive white fem ale canaries, since these fem ales has no
gene for yellow color. According to his new theory it was expected a 10 0 % red offspring. This
theory was fully stated in 1929. In 1933 Mr. Gill accom plished this cross, then obtaining all
m ales copper color after m olting, and all fem ales were grey. Not even one single red F1 was
obtained; besides, all copper color m ales were exactly the sam e as those obtained during the
crosses red siskin x yellow or dom inant white fem ale canary. These F1 copper color from red
siskin x recessive white, when in turn backcrossed to recessive white fem ale canaries, yielded an
offspring (R1) com prising oranges, yellows, and recessive whites.
Then again , all theories were facin g a true practical reality that turn ed them in to a doubtful
groun d. If the recessive white an d the red siskin were yellow free gen otypes an d accordin g to
Dr. Dun cker the copper color was a by-product of red-yellow com bin ation …where the yellow
factor cam e from to yield copper color in F1 offsprin g, from red siskin x white recessive
fem ale can ary?
In fact, Dr. Dunker’s suppositions as the red siskin and the recessive white were yellow free
birds, and that the yellow factors m ixed with red factor yielded a copper color F1, both were
wrong. This has been evidenced by the practical experience, which has fully enriched the knowl-
edge that we have today about the red factor canary.
Today we know that the red siskin possesses yellow lipochrome. Besides, this lipochrome is not an
obstacle to fully express the red color. It is well known to fanciers that many red plumage birds in
captivity turn to yellow, including the red siskin. This is the result of environmental factors, con-
clusively nutrition quality. From a genetic basis the parents only provide the offspring with genes
responsible for the proper assimilation of certain pigments, the transformation of those plant pig-
24
ments into animal pigments, and finally the onset of these in the feather structures. If a bird is not
properly supplied with a diet rich in beta-carotene to be transformed into red lipochrome, but is
supplied with xanthophylls, then yellow lipochrome will be deposited in its feathers.
Now it is quite sim ple to understand that the red plum age phenotype is not only an open expres-
sion of its genotype, but also of the environm ent, too. This is com pletely true. In genetics it is
clearly stated that:
Phenotype = Genotype + Environm ent
Today, we enjoy beautiful red color canaries due to a diet where carotenes are present in con-
centrated quantities, easily absorbed during the m etabolic process for those birds genetically
able to transform them into red lipochrom e. Nevertheless, up to this m om ent the red color
present in the red siskin has not been really and com pletely accom plished in the red canary,
according to the criteria of several specialists.
Records of Fertility Level in Hybridizing Red Siskin and Female Canaries.
At the beginning, it was believed that the F1 (red siskin x canary), and the R1 (F1 x canary) only
yield fertile m ales in a sm all proportion in the offspring, with no possibilities for fem ale fertili-
ty, at all. Nowadays, in the light of all gathered knowledge over the years around the world, it is
well known today that F1 m ales are fertile in a high proportion, and that fem ales are m ainly
infertile. Besides, in the R1 cross (F1 x canary) fertility level is a little bit m ore reduced, m uch
m ore in fem ales than in m ales, since m ales could be fertile to a larger extent. When R1 m ales
are backcrossed to fem ale canaries, then a rem arkable difference is observed in the offspring
fertility levels, since m ales and fem ale are fertile, as well; nevertheless, sporadic cases of infer-
tility m ay show up. This offspring level is known as R2. When these birds are backcrossed to
canaries to obtain the fourth generation R3, all previous fertility problem s will be over, since the
whole offspring in norm ally fertile. The chart below shows what has been explained:
25
Octavio Perez-Beato
Today, canary breeders all over the world use red fem ale canaries to hybridize with the red
siskin in all crossbred levels, according to the previous diagram . Nonetheless, a deeper red color
has not been accom plished, at least from a genetic point of view. Nowadays we enjoy beautiful
red canaries based on special diets where beta-carotenes are present in a very concentrated
form ula, together with a very easy assim ilation process, which outcom e is an excellent red color
com pare with canaries fifty years ago. Many specialists hold the opinion that the red canary has
rendered its full potential, and nothing else on a genetic basis, could be expected.
Next paragraphs will be devoted to explain the m ost popular theories on the red factor canary,
according to its theoretical genetic form ula. A portion of this form ula has been previously
explained in the chapter dealing with white canaries and in the chapter about yellow canaries,
all these genes are also com m on in the red factor canary.
The red factor canary genetically needs to bear the K gene to norm ally assim ilate the carotenoid
substances, plus the G gene in order to transform these substances into yellow pigm ents. If the
red factor canary would bear the recessive g gene in heterozygosis condition, a proportion of the
brood would be hom ozygote for that allele, and this is only possible in theory, since the reces-
sive german white canary has never been obtained; it is only and exclusively a theoretical pos-
sibility, as has been explained som ewhere before in this book. Besides, the red factor canary
possesses the f gene in hom ozygosis (ff), since the heterozygote condition (Ff) pertains to a
dom inant white canary, in which case its red color shall be fairly hidden. Based on all of the
above, we m ay arrive to the conclusion that the genetic form ula for the red factor canary could
be KKGGff. Unfortunately, with these genes we only can obtain a yellow canary. Obviously, we
need to add som ething else to obtain a red factor canary.
In m odern genetics we assum e that the m ost plausible issue is the existence of a locus R with
two alleles, R and r. The R allele is responsible to elaborate a specific enzym e that transform s
the carotenoids into red pigment inherited from the red siskin. On the other hand, the r
allele genetically m eans, the im possibility to transform carotenoid into red pigm ent. Then
again, the genetic form ula for a red factor canary could be theoretically: KKGGffRR. Besides,
crossbreeding experim ents point to a variable expressivity of the trait, since from parents with
excellent red factor expression on the plum age, part of the brood are not so good.
Now, as a by-product, we can better com plete the genetic form ula of the yellow canary, based
on what was previously discussed: KKGGffrr since the yellow canary is unable to transform
carotenoids into red pigm ent, as does the red factor canary.
I would like to m ention other sim ple experim ent as well. In the cross-breeding of a red factor
canary x yellow canary the brood obtained runs from light orange to properly red factor
expression, as long as you supply carotenoids in the diet. These results point to the aforem en-
tioned condition of variable expressivity in the red factor gene.
If mating two red factor birds bearing the genetic formula KKGGffRR, only a brood of red factor
descendants will be obtained with the same genetic formula. Besides, based on all of the above, the
previous formula is also applicable to the red siskin, but the red factor canary has never achieved
the extremely pure red color of its red siskin ancestor. Then, if that genetic formula is convenient,
to determine and explain, what must be expected to some extent in breeding the red factor canary,
it is just that, a formula to solve a practical need, but for me it is incomplete from a biological point
26
of view, which is more apparent when we have to admit that it is also applicable to the red siskin;
from a critical and logical thinking, the red factor canary, in fact, is not a genetic copy of the red
siskin, regarding the gene pool for the expression of the red color plumage. Furthermore, nowa-
days we obtained wonderful red factor canaries based on special addition to canary foods of high-
ly concentrated beta-carotene, which are well known to all canary breeders, but that concentrated
additive is not available for the red siskin in the wild, though it exhibits an extremely scarlet color
on its plumage. Red siskin includes in its regular diet a generous amount of food containing beta-
carotene from the wild. Experiments have been conducted to establish the mating preferences of
red siskin females, and conclusively results indicate that deep red males are preferred for mating.
I stick to the criteria of other authors that Spinus cucullatus extrem ely deep scarlet color is
determ ined by several gene pairs, and not by only one. We m ay discuss about the chrom osom al
stabilization that F1 hybrids go through, as well as the subsequent descendant lines R1, R2, etc.,
but that topic is not in the objective fram e of this book. I do really believe this topic together
with other biochem ical aspects, m ust be studied for those who are in research work about the
gene com plex of the red siskin.
My opinion is that hybridization process with Spinus cucullatus has rendered all what was
expected, and based on today evidence we cannot expect a deeper red factor canary to em erge,
unless a very specific m utation occurs, which is a lot im probable.
The Red Mosaic Canary
This bird em erged after the hybridization between the canary and the red siskin. The latter
shows an apparent dim orphism between both sexes; the fem ale only shows the red color on the
lesser coverts, prim ary coverts, breast and rum p. The dim orphism in the m osaic fem ale canary
is a by-product from the aforem entioned hybridization with the red siskin, though the red
lipochrom e is not exactly located as in the red siskin fem ale, since the genom e of the canary
species probably induced variations to this pattern, to som e extent.
Several experim ents were conducted by Matern in Germ any, Gill in England, and by Kerrison
and Bennett in the United States, regarding the m osaic trait in the fem ale canary. They con-
cluded that by using these m osaic fem ales (form erly: dimorphic), it was possible to obtain a
deeper red color in the canary. It was Gill who, unquestionably, stated that the m osaic trait in
the canary was an autosomic gene, and not a sex link one.
The adult red mosaic female canary shows a chalk-white color in its plumage, except in the areas
where the red lipochrome is fully shown. These areas are as follows: upper eye-ring feathers, less-
er and primary coverts, and in some birds the greater coverts and primaries are also tinted, rump
and breast. In contrast, fledging shows a common lipochrome color, except in the aforemen-
tioned areas, which usually are a little bit more colored, though it is not always the case.
After the first m olt, the new growth shows the typically chalk-white plum age with all red m ark-
ings well visible, and with an outstanding deep red color. Nonetheless, it is im portant to say that
the well known m osaic phenotype in not exclusively shown by the fem ale canary. Any canary
breeder knows that m osaic m ale canaries are a fact, and that they are really beautiful birds.
27
Octavio Perez-Beato
Before their first m olt, the m ale canaries that bear this m osaic trait are not different from any
com m on lipochrom e bird, as discussed in a previous paragraph concerning the m osaic fem ale.
When they com plete their first m olt and the new growth show up, an increased red color is
apparent alm ost all over the plum age, turning into a pure white the rest of them , at the sam e
tim e that the critical areas, as in the fem ales, are of a deeper red color. After the m olt process
has been com pleted, the m ale m osaic canary seem s very m uch like any other red canary, but a
closer look will dem onstrate that certain areas are uncom m only white, not seen in any frost red
m ale canary. Those areas are: upper part of the neck, around the vent, and the head. These are
technical visual m arks that differentiate the m osaic m ale canaries, from the com m on red frost
canary.
Genetic Aspects of the Mosaic Trait
The m osaic phenotype in the canary is the direct consequence of the com bined action of a gene
and horm one levels. In the m ajority of bird species where the fem ale plum age differs from the
m ale, we m ust take for granted that an interaction of sex horm ones and genetic factors, are
involved. From this concept, the m osaic m ale canary cannot show the sam e m osaic phenotype
as the fem ale canary, since the sex horm ones are com pletely different; consequently, the inter-
action should differ from one another.
Experim ents have shown that extirpation of the ovary from a m osaic fem ale canary, lead to a
non- m osaic phenotype after the next m olt, since no ovary can segregate the sex horm one nec-
essary to interact with the gene responsible for the m osaic trait. Then, such fem ale will becom e
a com m on red factor bird.
When aging, the m osaic fem ale m ay becom e a com m on red factor canary, since the ovary will
not produce the sex horm one; or will, but in an insufficient level.
E. H. Kerrison stated that the m osaic trait is controlled by an autosomic dominant gene, that
he called H. This gene is activated by the sex horm one, to finally produce the m osaic phenotype.
As the m osaic inheritance is due to the action of an autosom ic dom inant gene, the birds with
genotype HH, and Hh, will show the sam e phenotype. In fact, that gene has a variable expres-
sion, since the technical red m arkings in the m osaic birds differed from one canary to the other.
Canary breeders are always concerned in selecting the best m arked birds for the shows, as well
as for breeding purposes. Regarding the hom ozygous recessive birds hh, being h the allele
responsible for the absence of dim orphism , no interaction is possible between this allele and the
sex horm one, since the recessive allele h, has no response to the action of the sex horm one. The
hom ozygous birds hh, will be the com m on, non-m osaic canaries.
At this point it is convenient to say that Dr. Bennett conclusively stated that it was necessary to
m ate red frost birds and red m osaics, and never attem pt to m ate m osaics to intensive, since the
intensive factor – allele I- is an inhibitor for the m osaic expression in the plum age. Rigorously,
the dom inant I allele (intensive factor) probably produce an epistatic action on the dom inant H
allele, and this is the reason why the m osaic factor is never expressed on intensive birds, no
m atter if the canary is hom ozygous – HH- or heterozygous – Hh- for the m osaic trait.
28
Once in a while, som e regular variations in m osaic m ales prevent the identification of such birds
as m osaics. If in doubt if a m ale canary is really a m osaic bearer, just let it m ate to a non-m osa-
ic fem ale. If you obtain any m osaic bird in the brood, then it is a clear dem onstration that the
suspect is in fact a m osaic bearer canary.
The m osaic trait is com m on in m elanin birds, in all types as: bronze, brown, agate, pastel, opal,
etc. These birds show, together with m elanins, the lipochrom e color rem arkably deep in the
typical areas m entioned before for m osaic fem ales. I would like to express that it is not only
the red lipochrom e which is expressed in the plum age of a m osaic bird; yellow lipochrom e
could be also present, instead. Unfortunately, canary breeders in general, are prone to take red
factor breeding, and not yellow lipochrom e in their m osaic birds. It is not uncom m on to hear
a surprised rooky canary breeder when he/ she gets in touch with yellow m osaic canaries. In
m y opinion these birds are as good as the red ones, it depends on how we project canary breed-
ing purposes. We m ust rem em ber that, prim arily, a good show is determ ined by the diversity
of birds registered.
Those canary breeders devoted to the raising of yellow m osaic birds, ought to be very careful in
m ating their canaries. A rule of thum b is to select the excellent, am ong the best, and try not to
go out of the technical procedures recom m ended in breeding m osaic canaries.
Next, some guidelines and mating examples are shown in order to obtain yellow mosaic canaries.
If any fancier wants to obtain yellow m osaics, and by the m om ent tim e only possesses a red
m osaic heterozygous fem ale – as the worst-it is possible to m ate this fem ale to a yellow frost
non-m osaic m ale, then:
Yellow frost m ale x red m osaic fem ale

KK GG rr ff ii hh KK GG RR ff ii Hh
The F1 offspring is:
KK GG Rr ff ii hh (a) and KK GG Rr ff ii Hh (b)
We are only interested in the F1 offspring (b), since they are m osaic for they bear the genotype
Hh which is typical of a heterozygous m osaic canary. These F1(b) descendants have to be back-
crossed to a yellow frost m ale. We have selected a m osaic fem ale from F1(b) group since they
are easily recognized. Besides, all F1(b) m ales have to be back-crossed to yellow frost fem ales,
and if any m osaic bird from this brood is obtained, it is a proof that the F1 m ale is for certain a
m osaic canary.
The selected fem ale from the F1(b) group is back-crossed to a frost yellow m ale as stated in the
previous paragraph, and then:
Yellow frost m ale x F1(b) m osaic fem ale

KK GG rr ff ii hh KK GG Rr ff ii Hh
29
Octavio Perez-Beato
The R1 offspring will com prise the following:
KK GG Rr ff ii hh
KK GG Rr ff ii Hh
KK GG rr ff ii hh
KK GG rr ff ii Hh
From group (d) we obtain m ale and fem ales yellow m osaic heterozygous canaries, and then
again, phenotypically they are true m osaic canaries. Consequently, we can introduce the m osa-
ic factor in m elanin canaries; in short, we will be handling the m osaic factor in both, lipochrom e
and m elanin lines at the sam e tim e.
I think it is convenient to dem onstrate that, if the original m osaic fem ale that yielded the F1 off-
spring, was hom ozygous for the m osaic factor, instead, the whole F1 brood would be heterozy-
gous for the m osaic trait, and then phenotypically m osaic birds:
Yellow frost m ale x Red m osaic fem ale
KK GG rr ff ii hh KK GG RR ff ii HH
The F1 offspring is:
KK GG Rr ff ii Hh (m osaic m ales and fem ales)
Some Important Remarks on the Red Siskin
It was Swainson in 1820 that reported and described for the first tim e the red siskin, then sci-
entific nam e was Carduelis cucullatus, today accepted and also known as Spinus cucullatus.
The red siskin inhabits certain territories in the northern part of Venezuela, Trinidad, north-
eastern part of Colom bia, and probably Monos and Gasparee islands to som e extent.
Nevertheless, it is extrem ely rare in its native locations, and has been included in the list of
endangered species. Besides, today ornithologists know that this bird has been reported from
Puerto Rico, in a very specific area, and probably com prising a really sm all wild population.
Facts point to the event that in the 1930 ’s the establishm ent of the species occurred, due to a
heavy im portation of the bird at that tim e, augm enting the possibility of escaped birds, and
released ones.
According to history, the Spaniards appeared in Venezuela by 1530 , and being traditionally
good trappers and bird traders, soon sailed the red siskin, together with other birds, to Spain
and Canary Islands.
Inconceivably, the red siskin remained unknown for the rest of Europe during the next three cen-
turies, being responsible the Spaniards for keeping a strong grip on this and other finches as well.
Swainson’s description of the red siskin was based on a bird in possession of a bird keeper.
Around 1870 , some red siskins were imported to Europe due to the cage bird breeding booming.
It is a fact, that several canary breeders successfully accom plished the red siskin x canary cross,
at the beginning of the 20 th century. Precisely, it was the extrem e encouragem ent for Dr.
30
Duncker to start in a short period of tim e his experim ents team ing with Reich, ending up with
a “red canary” in few years.
In the 1940 ’s red siskin populations were dangerously decim ated, and this trend continued
through the 1980 ’s. After m any efforts, today a Red Siskin Recovery Project is approaching the
goals of conservation. The Am erican Federation of Aviculture, Inc. is involved in this project.
Further inform ation could be obtained at:
http://www.afabirds.org/afa_rsp.shtml
31
CHAPTER 7
The Green and the Brown Canary
The green canary is the variety that closely resem ble the appearance of the wild canary; that lit-
tle bird from the Fringillidae fam ily, introduced in Europe about 50 0 hundred years ago, which
popularity gained the favor of ornithologists and am ateurs, as well.
The green canary, according to m any specialists, is typically a rustic bird, hence is the variety
that less problem s m ay introduce in breeding and raising procedures. From this green canary,
m any m utations have been developed, being the origin of whites, yellows, agate, etc.
In this very chapter, for the first tim e, we face a m elanic canary: the green canary. This bird pos-
sesses the two m elanins present in the species, it is, pheom elanin and eum elanin; the ground
color is yellow lipochrom e; these three pigm ents m ake the com bination for the green color, as
it is in the wild canary, and inherited by our dom estic bird.
Though it is a biological fact, the presence of pheom elanin m ust not be visible in a good green
canary, which back streaks, wing feathers, and tail feathers, m ust be black (eum elanin), with no
trace of brown color (pheom elanin).
It is mandatory to say that melanins in the canary are sex link genetic factors, and they follow the
inheritance pattern of such genes. Then, if we call B the gene that is responsible for the brown
melanin, and N the gene that is responsible for the black melanin, we may approach basically an
important part of the melanic inheritance in the canary. To be completely manifested these
melanins on the plumage, it is necessary the action of an oxidative enzyme, that I will call O.
Fig 18 graphically shows the genetic form ulas and the locations for genes in sex chrom osom es,
from a green m ale and fem ale. Observe that the Y chrom osom e is empty, not bearing any gene
of our concern.
32
Fig. 18 Sex chrom osom es carrying the genes for m elanin and the oxidative enzym e,
typical of green m ale and fem ale canaries.
The Brown Canary
This type of canary appeared before 170 8 according to som e specialists, since in that year
Hervieux de Chanteloup included the brown canary in his treatise on canaries titled: Noveau
trait des serins de canarie, printed in France. In fact, the nam e for this type of canary has not
been brown, until recently; since its first description this color has been known as cinnamon.
It was not until 190 8, that two researchers nam ed Durham and Marryat discovered that the
brown color (then known as cinnamon) was the response to a sex link allele due to a m utation.
Besides, this m utation produced pink eyes in nestlings, and a darker pink in adult birds. The
article describing their findings was titled: Note on the Inheritance of Sex in Canaries. Many
years after this publication, A. K. Gill, already m entioned in previous chapters, published an
article on the color inheritance of the cinnam on tint: Cinnamon Inheritance in Canaries. In this
article the m echanism of inheritance of the cinnam on color (brown color), was explained in full
details.
This brown color on the plum age appeared as a genetic consequence of a m utation, which
inhibits the build-up of black m elanin (eum elanin) in the feathers, leading to the solely expres-
sion of the pheom elanin, hence the brown color. At the sam e tim e, the com plete suppression of
eum elanin, allows the reddish or pinky eye color in the nestlings, which is readily seen in their
unfeathered heads.
Nowadays, instead of using the term cinnamon, which is exactly how the bird looks like, and
besides being the way how this type of canary was originally described with a very accurate def-
inition, a m odern tendency led to call them “brown,” as the accepted adjective to nam e such
m utation; probably based on the strong influence of Europe, on canary breeding. I say this
because in Italy they used the term bruno, from the Italian language, of course. In French the
sim ilar term is brun. Undoubtly, brown is the closest phonem e to these Italian and French
appellatives. Let us say that it is a m atter of snobbish term inology, m ore than a true approach
to the canary color phenotype. It is not the intention of this book to change term inology, but we
need to know why we call a bird the way we do.
33
Octavio Perez-Beato
To approach the genetic form ula of the brown canary, let us call n the allele that denotes the
absence of eum elanin; then we m ay face the structure of the genetic form ula corresponding to
a brown canary, as shown in Fig. 19. The interpretation is exactly like the one on Fig. 18, but
som ehow sim plified to have it easier.
Fig. 19 Sex chrom osom es from a m ale and a fem ale brown canaries,
with alleles for the expression of pheom elanin and the oxidative enzym e.
The reader should com pare Fig. 18 and 19 to fully understand how the brown canary cam e to
be. Observe that the allele N is dom inant to n, which is absolutely necessary to understand the
following steps.
If a brown m ale m ates a brown fem ale, the whole offspring will be brown, as shown in Fig. 20 .
Fig. 20 Brown m ale and brown fem ale m ating,

and offspring obtained which is 10 0 % brown.
Likewise, the reader m ay verify that m ating a green m ale to a green fem ale, the offspring will be
10 0 % green. J ust follow the sam e procedure as above to test this assertion.
At this point, we are ready to continue setting up couples to m ate these m elanic canaries. Next
experim ent will be m ating a green m ale to a brown fem ale, as shown in Fig. 21. A closer look
will dem onstrate that all m ales in the offspring are carrying the brown factor; on the other hand,
34
all fem ales are hom ozygous green, as they only posses one X chrom osom e cannot carry any
other allele, since the Y chrom osom e is em pty as long as we are concerned.
Fig. 21 Results from a green m ale and brown fem ale m ating.
All m ales are green carrying the brown factor. All fem ales are green.
What has been stated before is a genetic principle, which is an unconvertible rule regarding
m elanin traits in canaries, and in any other sex link trait in birds: females will always be pure
for any sex link trait, since they can carry only one single allele of this type. Now, it is con-
venient to analyze the opposite m ating, it is, brown m ale and green fem ale. See Fig. 22.
Fig. 22 Results from a brown m ale and a green fem ale m ating. All m ales are green carrying
the brown factor. All fem ales are brown.
35
Octavio Perez-Beato
In this m ating we have obtained all green m ales carrying the brown factor, as we obtained
before in the previous m ating (Fig. 21), but in the present m ating all fem ales are brown, not
green as in the previous one.
We can see that fem ales always receive the chrom osom e X from their father; then again, they
will be pure for the trait included in their father’s X chrom osom e.
We will continue with another m ating: green m ale carrying brown factor and brown fem ale.
Results are shown in Fig. 23.
H ere we have four differen t offsprin g types: green m ales carryin g brown as their father,
brown m ales, green fem ales, an d brown fem ales. As the father possesses two types of chro-
m osom e X, on e carryin g the brown factor, an d the other carryin g the green factor, the fem ale
offsprin g are com posed of brown an d green fem ales, depen din g on what chrom osom e they
will received, (1) or (2).
Fig. 23 Results from a green m ale carrying brown factor and a brown fem ale.
Males are green carrying brown factor and brown.
Fem ales are green and brown.
The reader m ight recall that, the com bined effect of m elanin and lipochrom es was m entioned
in the chapter devoted to such pigm ents. Precisely, now is the opportunity to talk about the
results of lipochrom es, as a ground color for green and brown canaries. Again, these ground
colors could be yellow, white, and red. Besides, keep in m ind that regarding the yellow
lipochrom e, if carried together with the allele s, the phenotypic effect is lemon yellow, though
in som e countries the shows put together all yellow canaries, without distinction. But as I said
before, the allele s is a genetic reality, not a m ere speculation. It does exist, and then creates
the lemon yellow.
36
It is alm ost m andatory to m ention specifically, the varieties that could be obtained according to
the type of lipochrom e as a ground color. Such varieties for the green and the brown canary are
listed below:
Lipochrome Green Brown
Yellow Green (com m on) Yellow Brown

Lem on Yellow (allele s) Green (lim e green) Lem on Y. Brown
White (absent lipochrom e) Grey White Brown
White (plus allele s) Blue White Brown
Red Bronze Red Brown
In fact, the so called Blue is nothing but the Green with white ground color plus the allele s;
the canary will look as a poor grey color, otherwise. By no m eans can we call it “blue”. If we
accept this as a fact… why not to accept the lem on yellow as a ground color? Then again, we
are not dealing with a subjective criteria, the optic factor expressed by m eans of the allele s, is
a genetic reality.
Not accepting the presence of the lem on yellow canary is detrim ental to the list of varieties
that really exist, and the allele s contributes to the so called blue canary as it does to the lemon
yellow.
37
CHAPTER 8
The Dilution Factor: the Agate and the Isabelle Canary
In the previous chapter it was explained the need for an oxidative enzym e for the com plete and
full expression of m elanins in the green and brown phenotypes, which are both oxidized
m elanic form s.
In this chapter, the diluted melanic forms will be the topic. These form s are yielded by an inher-
ited factor that produces an incom plete oxidative process on the m elanins, turning them into
diluted form s.
If in the previous chapter the oxidative factor was called O, in the present chapter the dilute fac-
tor, or incom plete oxidation will be called o. In this sim ple way, we are inside the genetics of the
dilute canaries: the Agate and the Isabelle.
The Agate Canary
This type of canary, in som e of the varieties, is not exactly a beautiful bird; nevertheless, in other
varieties the com bination of colors is really beautiful. This dilute factor appeared for the first
tim e in the Agate, and it was a turning point for canary breeding at the end of the 19 th century.
Not long before 190 0 , a Germ an canary fancier by the nam e of Helder raised a fem ale canary of
an unknown color at that tim e, from a pair of green birds. That fem ale was of a grey tint
plum age, and the lines on the back were reduced, light tinted, and m ore pale than those of a reg-
ular green canary. The first nam e for this color was ash-grey.
Som e late considerations led the criterion toward the fact that, this potential new color was
nothing but the one that Hervieux described as No. 12 in his treatise, published in 170 8, m en-
tioned som ewhere before in this book. Hervieux called this color “Com m on Agate Serin”. From
that on, this variety was called Agate, and not Ash-Grey.
38
Posterior conducted research, dem onstrated that the Agate was in fact a Dilute Green, and that
it was a sex link factor. This factor is expressed as the im possibility for the oxidative enzym e to
act at full strength, hence the m elanin show itself dilute, since it is not com pletely oxidized. It
is sim ple to say that the dilute plum age is the one in which the m elanic pigm ents are pale, due
to the action of the dilute gene o.
As a result, the genetic form ula for the Agate is:
Agate male Agate female

XBNo XBNo
XBNo Y
Now com pare these form ulas to the green canary’s from the previous chapter. It is readily seen
that the green canary differs from the agate just because the green carries the oxidative factor
O, and the agate the non-oxidative factor o.
The Isabelle Canary
This type of canary is nothing but a Dilute Brown. It is, if a brown canary carries the non-oxida-
tive factor o, instead of the oxidative factor O, then we have an Isabelle instead of a Brown one.
The Isabelle is not the product of a m utation, it showed-up from the Agate, as a result from cer-
tain breeding, transferring the non-oxidative factor to the Brown genotype, which turned to be
the Isabelle.
I th in k it is in terestin g to explain to th e reader, h ow is th e m ech an ism by wh ich , th e gen e

th at determ in es th e dilute m elan in , gen e o, is tran sferred from th e Agate to th e Brown to
create th e Isabelle. Th is m ech an ism is un derstood th rough th e gen etic ph en om en a kn own
as crossing-over.
Let us suppose that a m ale with green phenotype is in fact a Brown carrying Agate, according to
the form ula X Bno X BnO. If you take a closer look, this genetic form ula will yield a phenotype
according to a green canary. Then, in a very specific m om ent during the m eiosis, the hom olo-
gous chrom osom es overlap, each one cut-along in two chrom atids. The hom ologous chrom atids
exchange hom ologous segm ents, producing a real exchange of genes. Let see the application of
all this in our particular case:
39
Octavio Perez-Beato
Fig. 24 Steps in the process of a crossing-over, ending-up with two new chromosomes that will
give the offspring new characteristics.
Two chrom osom es of Brown canary carrying Agate genes, at a precise m om ent during m eiosis,
exchange genetic m aterial, creating two new chrom osom es. In (a) it is observed a pair of hom ol-
ogous chrom osom es of a Green phenotype canary carrying Agate genes. In (b), the crossing-
over is depicted, showing the exchange of the lower ends of the chrom osom es involved, where
alleles O and o, are located. In step (c), the chrom osom es are getting apart after exchanging
their segm ents. In step (d) the chrom osom es are set apart already, having each one a new seg-
m ent in their lower portions. Chrom osom e 1 is carrying Isabelle genes, since the allele B is pres-
ent, which determ ines brown m elanin (pheom elanin), plus the allele n which indicates absence
of black m elanin, finally the allele o, dilute factor. Consequently, this chrom osom e will be
responsible for a dilute brown canary, which is nothing but an Isabelle. Chrom osom e 2 is typi-
cal for a green canary. Each one of these two chrom osom es will go to a different gam ete, lead-
ing to an offspring of Isabelle and Green fem ales, besides Brown and Agate fem ales, as well.
These last two phenotypes originate from those original chrom osom es not having a crossing-
over process. Now, it is very clear how an Isabelle fem ale originates from Green m ale carrying
Agate genes, due to the crossing- over process. Furtherm ore, it is convenient to recall that, a
canary showing a Green phenotype, could be genetically Brown, carrying Agate genes. This kind
of canaries is highly valuable, since according to what was previously dem onstrated, they pro-
duce four different kinds of gam etes, which generate the four classical m elanic canaries: Green,
Brown, Agate and Isabelle. In France, they usually call “pass-partout”(crow bar) this kind of
canary, since it opens the possibility to obtain four different types of m elanic birds.
The Agate Phenotype
Fig. 25 depicts schem atically, the head of an Agate canary, stressing the designs that best char-
acterize this type of bird. The crown shows a striated design that resem bles the one on the back,
but m iniaturized. The eyebrow is m elanin-free, so exhibiting the lipochrom e ground color at its
best. The so called whiskers are well visible in a genuine Agate. This design is the direct result
of m elanin concentration in the feathers of this area, producing a very strong contrast with the
rest of the feathers that show a dilute m elanin. Besides, legs and bill, are pink in the Agate, no
trace of m elanin is accepted.
40
Fig 25 Agate head. Melanin and lipochrom e areas are shown.
The classic varieties for the Agate are: White Agate, Yellow Agate and Red Agate. Of course, the
list m ay continue adding White Agate Ivory, Yellow Agate Ivory, Red Agate Ivory, Red Agate
Mosaic, Red Agate Opal Mosaic, and so forth. The m ost outstanding White Agates are those car-
rying the optic factor, which provide a delicate silvery appearance, only detectable with peering
eyes. For the Isabelle, the sam e varieties are also true; and the sam e consideration for the optic
factor, as well.
It is well known since long ago, that the best m ating is accom plished between Agate and
Isabelle, which produce a better quality offspring, than m ating the sam e type. The m isguided
breeding, regarding dilute canaries, m ay result in an im poverished type in the offspring. It is
highly detrim ental in bird shows and judgm ent of the birds, which hardly qualify in the corre-
sponding category; extrem e cases lead to disqualify such birds, resulting in a frustrating feeling
for the breeder, besides an em barrass situation am ong his/ her peers.
41
CHAPTER 9
The Pastel Canary: the super dilution factor
In 1960 , a m utation appeared affecting m elanins, and m ainly the pheom elanins, since the
eum elanin did not suffered any substantial change. More than fifty years had passed without a
m utation affecting the dilute condition in the canary plum age. This new m utation affected the
Isabelle, diluting the tint of the already diluted brown color in these canaries, but brown
m elanin was apparent in wing feathers, tail, back, and head. On the back, the lipochrom ic
ground color was rem arkable, different from the com m on Isabelle.
In the Brown canary, this new m utation becam e particularly different, in such a way that pro-
vided the nam e to this new m utation in the canary. The ground lipochrom e color is delicately
blended with the pheom elanin, just like as it is in a pastel painting, therefore, the nam e pastel.
The lines or streaks on the back and flanks are alm ost im perceptible, and the canary shows that
sam e blending of lipochrom e and m elanin, all over the plum age.
If we can talk about the Brown Pastel and the Isabelle Pastel, som ething som ehow different
occurs regarding the Green Pastel, Blue Pastel, Bronze Pastel, and Agate Pastel, since the Pastel
Factor does not dilute the black m elanin (eum elanin), only the brown m elanin (pheom elanin)
is affected. It has been a controversial issue, regarding the existence of true pastel tones in the
aforem entioned eum elanic types, ever since. Besides, it is not a sim ple task to determ ine if a
Bronze, a Green or a Blue canary are carrying the Pastel Factor or not. Then again, this issue
has been based on the fact that, a true Pastel phenotype cannot be assigned to Bronze, Green,
Blue and Agate, though at international shows these have been registered and still shall be.
In the sam e way that the Agate m utation occurred, the Pastel factor is a recessive and sex link
gene, too. At this point, we m ay consider the genetic form ula for the Pastel factor, taking into
account that it is a m utation able to partially inhibit the production of brown m elanin, through
a different biochem ical reaction step that the one responsible for the appearance of the Agate
canary. Therefore, the non-pastel canaries are considered the norm al m orph, represented by an
allele P, and then, the Pastel trait m ight be defined by the allele p, as it is a recessive one. Based
on all of the above, the genetic form ula for the Brown Pastel is:
42
Brown Pastel male Brown Pastel female

XBn Op XBn Op
XBn Op Y
It is apparent that adding the genes for lipochrom e colors to the previous genetic form ulas, the
classic Pastels will be obtained. What follow are the genetic form ulas for those classic Pastel
canaries, m ost com m only obtained in regular breeding procedures.
Yellow Brown Pastel intensive Yellow Brown Pastel frost

X BnOp KKGGffIi X BnOp KKGGffii
White Brown Pastel intensive White Brown Pastel frost

X BnOp KKGGFfssIi X BnOp KKGGFfssii
X BnOp X BnOp
Yellow Isabelle Pastel intensive Yellow Isabelle Pastel frost

X Bnop KKGGffIi X Bnop KKGGffii
X Bnop X Bnop
One im portant aspect to get the best is the optic factor on the white ground color.
White Isabelle Pastel intensive White Isabelle Pastel frost

X Bnop X Bnop
X Bnop KKGGFfssIi X Bnop KKGGFfssii
Note that in all white varieties the alleles ss have been included, since they determ ine the optic
factor on the white color (either dom inant or recessive). In fact, if the optic factor is not present
on the white ground color, it turns to a dull and lack of appealing condition. It is som ething that
som e fanciers do not know, being very concerned why their birds do not get good m arks in bird
shows
As in any other sex link factor, the pastel factor carried by a m ale, will be inherited by its daugh-
ters. A straight inference leads to the conclusion that, a heterozygous m ale for the pastel factor,
will transfer it to half of its daughters, and they will be Pastel. It is now apparent that if a m ale
carrying the Pastel factor m ates any m elanic fem ale canary, we will obtain Pastel fem ales, by
m eans of which we m ay set up a com plete line of this type.
Now the reader is alm ost ready to conduct the necessary breeding to get Pastel canaries, com -
prising different varieties.
43
CHAPTER 10
The Opal Canary: the extra dilution factor
Chapter 8 was devoted to a type of canary which recalls the nam e of a gem : the Agate. The pres-
ent chapter will deal with another gem nam e: the Opal. According to certain records, this m uta-
tion appeared in 1949 in Germ any, probably not well understood at that tim e. Through the
fifties, it took a long journey, and is exactly during the sixties that becam e well known and pop-
ular, all over the world. It appears that, with the first Opal canary fanciers thought it was m ere-
ly, and nothing but a m ore dilute Agate. Then, the exact recognition of a new m utation was not
a sim ple decision.
Between the Agate and the Opal, we m ay establish a com parison. The dilute factor that m ade
possible the Agate type, affected both, the pheom elanin and the eum elanin. The sam e is true for
the Opal, in which an extrem e dilution is accom plished to affect both kinds of m elanins, m uch
m ore than occurred in the Agate phenotype. The Opal feather is darker on the inferior vane por-
tion; it stands as one rem arkable characteristic of the Opal phenotype. Besides, it is stiff, som e-
how fracturable, and not flexible, m ainly in the Green, Blue, and Agate. It is precisely a draw-
back in the Opal type, poor plum age quality, on the average.
Why that nam e Opal? In fact, the back of this canary shows certain iridescences when red
lipochrom e is present, and in m elanic types, if of good quality, bluish shades could be seen.
These unusual changes of color and shades, are typical of the gem we call Opal.
The Way the Opal Looks
The gene responsible for the Opal phenotype produces a m ore acute dilution than the observed
in a com m on Agate, m ainly on the back of the bird, on brown and black m elanins. Nevertheless,
the Opal canary shows bill and legs fairly dark, since such gene is not able to affect such parts
of the body in the Green Opal, Blue Opal, and Bronze Opal.
44
The Opal gene swipe off the m elanin from the feather edges, without touching the bill, legs and
under-plum age area, in m elanic types. Besides, the Opal factor com pletely dilute the brown
m elanin (pheom elanin) so it disappears from the plum age, then the under-plum age appears
bluish, by the solely presence of eum elanin, extrem ely diluted. As the pheom elanin is not pres-
ent, the ground lipochrom e shows up, shiny and lustrous.
It is so a strong dilution on the m elanins that, the Opal canary with black and brown m elanins,
looks like a lipochrom e canary, but with the iridescences and bluish tones of the Opal factor,
itself. Only very faint streaks of m elanin on the back and on both flanks could be seen.
On a Brown canary, it is so strong the effect of extra-dilution, that the bird looks like a very
dilute Isabelle, though certain basic designs rem ain, m aking possible to establish the difference
between the Brown Opal and the Isabelle Opal. On the back of the Isabelle, the Opal factor
action is so strong that, to the eyes of an inexperienced observer, the bird looks like a com m on
lipochrom e canary, except the duvet, which is black pigm ented and partially dilute.
The Opal factor acting on the Agate, swipes off the pheom elanin, com pletely; nonetheless, the
typical Agate design rem ains, including: whiskers, back and flank streaks, and the crown
design, but all these are reduced to a m inim um expression. They are shortened on half of their
width, and sm aller in length.
Genetic Features on the Opal Canary
The dilution factor that created the Opal phenotype is recessive and non-sex linked; it is auto-
som ic recessive. Knowing this, inheritance characteristics can be established; then we are in
good shape to m ake a forecast from the breeding schem e we plan to m ake. Let p be the allele
that produces the Opal condition, and P the non-Opal phenotype.
A Green m ale carrying the Opal factor has the genetic form ula:
X BNO Pp
X BNO
A Green fem ale genetic form ula, also carrying the Opal factor, is:
X BNO Pp
Y
Now, let us num ber the gam etes to establish the possible offspring:
1 X BNO Pp m ated to: 3 X BNO Pp
2 X BNO 4Y
1 + 3 PP = non-Opal Green m ale 1 + 4 PP = non-Opal green fem ale

Pp = Green m ale carrying Opal factor Pp = Green fem ale carrying Opal factor
pp = Green Opal m ale pp = Green Opal fem ale
2 + 3 = 1 + 3 (already explained) 2 + 4 = 1 + 4 (already explained)
45
Octavio Perez-Beato
Now, a sim ple conclusion is: from this m ating we m ay obtain m ales and fem ales, as follow: non-
Opal, carrier of the Opal factor, and pure Opal canaries. It is im portant to stress that, as the
Opal factor is not a sex-link gene, it segregates independently of the sex chrom osom es, then it
is possible to obtain the previously discussed offspring.
46
CHAPTER 11
The Ivory Factor: dilute lipochromes
During the fifties, the date is not very exact though som e records point to the year 1950 , for the
first tim e in the history of canary breeding a m utation showed-up that affected the lipochrom es,
not the m elanins. This new m utation appeared from a couple of yellow Hartzer Roller, from
which all the fem ale offspring showed the new color, as a delicate “cream ”, which enriched a lit-
tle bit m ore the already up going genetic background of our feathered little friend, and so open-
ing new possibilities to create color com binations not im aginable, just a short tim e before.
Ivory, as a color, is well known from the teeth of m am m als, m ainly from the elephant tusks,
which have been used in works of art for centuries. There are good reasons to nam e the new
color m utation as Ivory. To better approach this new color in the canary plum age, it is neces-
sary to recall that in the yellow canary, the wing and tail feathers, are not com pletely colored
with yellow lipochrom e, and then som e white areas are com m only seen. On the contrary, in the
Ivory canary the cream y color is extended to wing and tail feathers, com pletely.
In fact, the Ivory factor acts on the lipochrom es as a dilute gene, besides extending the pigm ent
to wing and tail feathers.
The fact that the first two Ivory canaries were fem ales, pointed to the certainty that it was a sex
link factor, as it was confirm ed lately, besides being a recessive gene.
In this m utation, there is som ething new added to all of the above. It was alm ost a rule that the
form er m utations affecting the m elanins in the canary also inhibited the trait to som e extent.
That is why som e authors assessed that the m utation which created the Brown canary, is such
just because inhibited the eum elanin expression in the canary plum age. In addition, the appear-
ance of the Agate is possible just because that particular m utation is able to inhibit the total oxi-
dation process on the m elanins. Furtherm ore, the Isabelle reached on the stage, sim ply because
of the convergence of two form er inhibitory processes: the one that created the Brown canary,
and the incom plete oxidation process that originated the Agate variety.
47
Octavio Perez-Beato
Now, let us exam ine the Ivory factor at the light of the inhibitory records that, characterized the
m utations on m elanins before the appearance of the Ivory m utation. First, we m ay say yes, this
m utation diluted the lipochrom e pigm ents not allowing deep colors, but at the sam e tim e the
wing and tail feathers are now fully lipochrom ed, not seen before in the canary breeding histo-
ry. These feathers are so well pigm ented in the Ivory, as any other feather all over the body of
the bird. It is evident that now we have two faces of the sam e phenom ena: dilution of the
lipochrom e (inhibitory effect), and extension of the color to wing and tail feathers (anti-
inhibitory effect). This apparent opposite effects produced by the sam e m utation, is recorded
for the first tim e in the canary breeding history.
Advanced canary breeders went beyond the Yellow Ivory. The Red Factor lovers wanted to
experim ent on the red canary. Eventually, the efforts ended with the appearance of the Red
Ivory (Rose), a new outstanding color not seen before in the canary plum age. It is a very deli-
cate pink color, evenly extended on all feathers.
In English language it was not a problem to nam e these two new colors, but in Spanish speak-
ing countries, canary breeders deliberated about a fair nom enclature for both new colors. It was
then agreed that, the Ivory plus yellow lipochrom e becam e Marfil-Crem a (Ivory-Cream ) accord-
ing to the exact look of the plum age as a whole. For the red factor Ivory, it was nam ed Marfil-
Rosa (Ivory-Rose) in rem em brance of the delicate color of the rose flower. Nowadays, Spanish
nom enclature m atches with English equivalent, for a better international understanding of the
always growing color listing, in our m odern canaries.
Genetic Features of the Ivory Canary
Now, it is necessary to focus on the genetic form ulas for the Ivory phenotype, since it will be
used shortly in breeding procedures to obtain a better knowledge regarding the Ivory canary.
Let a be the allele responsible for the Ivory trait:
Yellow Ivory Frost: X BNOa KKGGffrrii

X BNOa
Rose (Red Ivory) Frost: X BNOa KKGGffRRii

X BNOa
It is evident now, how com plex the genetic form ulas m ight be, describing colors in the canary.
Nevertheless, they are necessary by all m eans to a dedicated canary breeder, who wishes to
com pletely m aster in all details, the color genetics on this little bird.
The reader, at this point, m ust be in good shape to fully interpret the m ating procedures that
follow. They will be useful to have a full view of the genetic features of the Ivory factor.
48
Green carrying Ivory factor male x Green Ivory female
1 X BNOa 3 X BNOa
2 X BNOA 4Y
Offspring:
Green Ivory m ale Green Ivory fem ale

1 + 3 = X BNOa 1 + 4 = X BNOa
X BNOa Y
Green carrying Ivory Factor m ale Green fem ale

2 + 3 = X BNOA 2 + 4 = X BNOA
X BNOa Y
Green carrying Ivory factor male x Brown Ivory female

1 X BNOa 3 X BnOa
2 X BNOA 4Y
Offspring:
Green Ivory carrying Brown factor m ale Green Ivory fem ale
1 + 3 = X BNOa 1 + 4 = X BNOa
X BnOa Y
Green carrying Brown and Ivory factors Green fem ale

2 + 3 = X BNOA 2 + 4 = X BNOA
X BnOa Y
The reader surely realized that allele A represents the non-Ivory factor, which is the dom inant
original allele. The last m ating, depicted in the previous diagram , is the key to understand the
com binations Ivory-Melanin phenotypes, which m any canary fanciers believe are som ehow
com plex to obtain. In fact, it is not. It is just a m atter of applying correctly the knowledge
obtained about, dom inant and recessive genes, autosom ic and sex link, together with the cor-
responding dom ain of lipochrom es.
49
CHAPTER 12
The Pied Canary Genetics
In this chapter, not only the genetics of pied individuals will be considered, also the eye color
will be included, since eye color and pied condition are tightly related, for both are produced by
m elanin pigm ents. Ultim ately, the eye color in the canary can provide an extrem ely im portant
inform ation regarding m elanin present in the genotype, and not shown in the phenotype. What
is needed is to closely look at the eyes of our birds, with the help of light reflection. In this way,
chances are that we m ay succeed in detecting that im portant inform ation, needed in the genet-
ics of the pied canary.
Locus P bears two alleles: P and p. The dom inant allele P allows the developm ent of m elanin in
the plum age, while the recessive allele p inhibits such developm ent. The m elanic canary has the
genetic form ula PP, a pied canary has the genetic form ula Pp. Finally, a lipochrom e canary has
the form ula pp.
Nonetheless, the spot and shape of m elanin patches on the plum age depend upon other genes.
Gene P only allows the m elanin to show up on the plum age, nothing else. Now, a question aris-
es: Is there a com plete lack of m elanin in lipochrom e canaries, just because their sex chrom o-
som es do not carry genes for the synthesis of those pigm ents? The answer is, NO. The
lipochrom e canaries do carry m elanic traits; their sex chrom osom es have the inform ation for
such inheritance. The issue is that a lipochrom e canary cannot develop m elanic color in its
plum age, since its genotype is pp, which precisely inhibits the developm ent of m elanin. A com -
m on lipochrom e canary m ay be carrying Green, Brown, Agate or Isabelle. How do we know? It
is not always sim ple. As an exam ple, a Green, Bronze or Blue canary will show black eyes, since
eum elanin and pheom elanin are deposited in the eyes, as they m ight be on the plum age. An
Agate, has dark eyes, but not exactly black. In fact, it is not an easy task to establish a difference
between a Green and an Agate, based on the eye color.
On the other hand, a Brown canary has reddish eyes, seen on light reflection; Isabelle eyes are
rather pink, also seen on light reflection. When I say “rather pink” I do not m ean albino or luti-
no, it is not the case. The Isabelle eye color is pink, just in the pupil; albino, lutino and rubino
canary’s eyes, are as red as in an albino m ouse’s eyes.
50
The reddish eyes in the Brown canary and the pink eyes in the Isabelle, is just because these two
types of canaries do not carry eum elanin, and then that pigm ent cannot be present in their eyes,
either. Based on all of the above, it com es that a lipochrom e canary with “pink” eyes (on light
reflection) is carrying the Isabelle factor; it is not seen on the plum age because the canary does
not carry the gene P, which allows m elanin to be seen. In analogous way, a lipochrom e canary
with black eyes is carrying the Green factor, and so forth.
Paradoxically, it is very clear how the eye colors denote the melanic genotype, of a lipochrome
canary. Based on this, and on previous chapters, it is possible to study the results of som e
im portant couplings. If a breeder has a Yellow m ale with reddish eyes, and a Green fem ale, of
course, then the following m ating is possible:
Yellow male with reddish eyes m ated to: Green female

X BnO KKGGffpp XBNO KKGGffPP
X BnO Y
the offspring is as follow:
Yellow-Green pied m ale Yellow-Brown pied fem ale

X BnO KKGGffPp X BnO KKGGffPp
X BNO Y
What is im portant here is the outcom e of the Yellow-Brown pied fem ales, not seen in the par-
ents. The breeder in this exam ple was keenly interested in obtaining birds to start a line of
Brown canaries. The m ale parent with reddish eyes was unequivocally a Brown carrier, but lack-
ing the allele P to express the m elanin in its plum age.
It was then necessary a specific fem ale to provide the offspring with such allele. The chosen
Green fem ale provided the P allele, ending up with an offspring of Yellow-Brown pied fem ales.
From these fem ales, now it is possible to develop a whole line of Brown canaries. To accom plish
this is sim ple, just selecting the best of the Yellow-Brown pied fem ales and m ake a back-cross
using its father (inbreeding):
Yellow male with reddish eyes m ated to: Yellow-Brown pied female
X BnO KKGGffpp X BnO KKGGffPp
X BnO Y
Offspring:
Yellow-Brown pied m ale (A) Yellow m ale (B)

X BnO KKGGffPp X BnO KKGGffpp
X BnO X BnO
Yellow-Brown pied m ale (C) Yellow-Brown pied fem ale (D)

X BnO KKGGffPp X BnO KKGGffPp
X BnO Y
51
Octavio Perez-Beato
Yellow fem ale (E) Yellow-Brown pied fem ale (F)

X BnO KKGGffpp X BnO KKGGffPp
Y Y
From this entire offspring is now possible to choose from three groups of m ales, and three of
fem ales. Our choices increased, and then our possibilities to focus on the creation of a Brown
genetic line from the initial m ating. It is a good choice to select m ales from group (C), and
fem ales from group (F) since both groups show hom ozygote birds for the Brown factor. From
these m ales and fem ales, it is possible to successfully develop a whole lineage of Brown
canaries. At this point it is advisable to use inbreeding m ethods. By all m eans,
there is not a better choice to create lineages as the one discussed in this chapter. The reader
has been provided with inbreeding m ethods in a previous chapter.
Now, is the tim e to go directly to the practice of inbreeding m ethods, together with a strict selec-
tion of the best, to ensure a good quality lineage to begin with.
As the reader m ay see, the m ost im portant action is to start with the chosen m elanin hidden in
a lipochrom e phenotype, focused on the point that the offspring could get the allele P, in order
to express the m elanin on the plum age, this allele has been obtained from the non-lipochrom e
parent, and introduce in the offspring by m eans of the right m ating system .
In this very chapter only the exam ple of a Yellow carrying the Brown factor, has been discussed,
but it is easily projected to lipochrom e canaries carrying Green, Isabelle, etc. depending on the
decision of the canary fancier. The ultim ate goal is to obtain birds with phenotype PP, to fully
express the m elanin on the plum age.
The Variegated Symmetrical Canaries
According to well docum ented records, during the XIX century sym m etrical variegated canaries
were highly valued, probably due to the fact that it has never been easy to obtain such varieties,
not to m ention the hard task in keeping a whole lineage of such canaries.
It seem s that, com m on pied canaries were kept as good pets. Since breeds as the Hartzer Roller
were m ostly pied, and the song rem arkable sweet. Then, when a m ore technical approach was
placed on the stage of color canary breeding em ergence, the variegated ones were set aside as
less attractive. Nevertheless, the lovers of variegation fought back to obtain a different look in
pied-bird grounds. Probably, that was the surfacing of the sym m etrical variegated canary; an
outstanding bird difficult to cultivate and breed successfully, then becom ing a highly valued
canary ever since. In this way, variegation lovers were at please, with a new m odality of varie-
gated canaries that lead to a very accurate and patient pairing of selected birds. In the very
scarce literature dealing with that type of canaries, it is com m on to find descriptions such as:
“…brown tail and yellow body, dark crown in a light plum age”, etc. Sym m etrical variegation is
also a well established feature in Type Canaries, as in the lost London Fancy. It is clear that in
m any countries, in one way or the other, the fanciers looking for sym m etrical variegated birds,
really abounded.
52
Dr. Duncker also contributed in the genetics of sym m etrical variegation in canaries. He con-
ducted som e studies to determ ine the inheritance of such fashion. I m ay say that it is not an easy
task to get a hold on this type of birds, and the knowledge regarding the inheritance of such
traits is m eager. Based on Dr. Duncker studies the m ain nom enclature details on these canaries
are known from Germ any, and so the term s to nam e them . Exam ples to m ention are: Sattel,
which m eans saddle, and Mucken which m eans m osquito. These Germ an words are applied to
specific designs on sym m etrical variegated canaries. It is called point or mark where the
m elanin shows up in a regular distribution to create a sym m etrical design. In this way, a bird
with m elanin around both eyes, and on every tail feather, is said to be a three-point bird. Figures
26 and 27 depict som e classical sym m etrical variegated canaries; by no m eans are they a com -
prehensive picture-list, other com binations are also possible. They represent only schem atic
approaches, to give an idea of what a sym m etrical m elanin distribution m ight be.
Fig. 26 Upper: Mucken (forehead and throat)

two-points bird. Lower: Also two-points bird,
crown-forehead and tail. Nowadays, color possi-
bilities to create sym m etrical patterns in canaries
are really huge, the lim it is your im agination and,
of course, the technical possibilities to carry on
m ating procedures to achieve what the genetic
projects in our m inds are.
Fig. 27 Upper: Typical Sattel. According to

England definition, the sym m etrical m elanin sec-
tion is fram ed on the m id dorsal portion. Lower:
Sym m etrical variegated of four points, m elanin
patches on both eyes, and on both wings
53
Octavio Perez-Beato
As a general rule, it is recom m ended to use inbreeding procedures in case we detect an individ-
ual with acceptable characteristics of sym m etrical variegation. Only using inbreeding tech-
niques, it is possible to achieve certain results in term s of the offspring showing the desired
sym m etrical patterns. Then again, this is not an easy task to get along with.
54
CHAPTER 13
New Mutations. The Incredible Eighties
From 1981 to 1985 three new colors appeared on the stage of canary breeding. In just have a
decade, the em ergence of three new varieties broke all the records known before. Besides, the
new colors are based on the sam e inheritance pattern: autosomic recessive. They are really new,
so m uch a new that they are not seen in m any bird shows around the world, yet. Perhaps,
Europe is the top of the line leader in m odern colors. Nothing new, Europe has ever been. Is it
just because the wild canary was first introduced in the old continent, and their am ounts have
been large enough to develop a generous genetic variability? I have no answer, but chances are
that the am ount counts, since probability is a m atter of large num bers.
The Eumo
In 1981 the Eum o was born, reported from Holland. The new canary eye color is red, though it
gets a little darker with age in Greens, Blues, and Bronzes. Pheom elanin is alm ost gone. There
is som ething that seem s to be characteristic: the darker the streaks on head, back and flanks,
the darker the red eye color. Melanin runs along next to both sides of the quill, exclusively; the
rest of the vane is washed-off.
In general term s, the Eum o m utation dim inish the am ount of m elanin; both, in length and
width regarding the streaks. The overall design is som ehow streaky, and the non-m elanic por-
tion am ong the streaks is rem arkable sparkling, and shows a neat lipochrom e color.
The Brown, Isabelle, and Agate Eum o keep their eyes really red, and brightly. The eum elanin is
shown as a very peculiar grey color, including the duvet, but not black. Beak, legs, and nails are
pink.
Eum o is an autosom ic recessive m utation. There are som e few problem s with Eum o phenotype;
if it has too oxidized m elanin, the eyes of a non-well trained observer, m ay judge the Eum o as
a very dilute Topaz.
55
Octavio Perez-Beato
The Onyx
Another autosom ic recessive m utation. It showed-up in Spain between 1983 and 1984. This
“m utation” is in fact an allele of the Opal gene; now the Opal gene has two alleles. Onyx is not
a standardized genotype, yet. It is co-dom inant to the Opal allele, each one located in hom olo-
gous chrom osom es, on the sam e locus. Then, the Opal is a m ulti-allelic gene.
The history of the Onyx canary is by no m eans the sam e as for the rest of alleles affecting the
m elanin. All began m ore than twenty years ago, when a South Am erican bird of the genus
Spinus hybridized with a Green Opal fem ale canary in captivity. In fact, it is not very clear if it
was a Green or a Bronze Opal fem ale canary; a Blue Opal could be another possibility.
According to what is known today, som e hybrid m ales were fertile. The back-cross to fem ale
canaries, during a certain period of tim e, cam e up with the first Onyx canary. As the reader m ay
see, it is not a m utation at all, but the direct result of hybridization with a species of the sam e
genus (Spinus) of the red siskin. Records do not specify what the species was. J ust for the read-
er to have a rough idea about what num bers are involved in the genus Spinus, let us say that at
present twenty-one species are recognized with 36 sub-species as well. You m ay im agine that it
is really difficult to guess what the species involved was.
Years later, the Onyx breeding enthusiasm took place in the city of Valencia, Spain. More indi-
viduals were obtained, and obviously, an im provem ent in the stabilization of the Onyx pheno-
type, was achieved.
In fact, the Onyx is the opposite of a dilution. The Onyx plum age is very dark, som ehow a blend
of black and brown m elanins, with som e ash-grey color in the lighter areas. When young, they
look as classic m elanin canaries. The appearance of m elanin is som ehow delayed, but finally
spreads all over the plum age, in a non-evenly distribution. The tail and wing feathers do not
show an even allocation of m elanin, on the contrary, it looks like a dash-line design, on a trans-
verse fashion. There is an evident concentration of eum elanin on the head, from neck up, but
this eum elanin does not look exactly pure black. The rest of the plum age shows a disperse
m elanin color, m ore than in any other phenotype known today. Onyx is not a black canary, as
its nam e m ay suggest; lipochrom e is seen am ong the streaks, though som ehow darker due to
the spread of m elanin. The pheom elanin is really scarce; the eyes are black as well as the duvet.
Beak, legs, and nails are dark to black.
From a genetic point of view, it seem s reasonable that as the hybridization went on, a crossing-
over took place; exchange of genetic m aterial between both species – the canary and the South
Am erican Spinus-was consum m ated. Then, a new allele from the Spinus species was added to
the Opal locus. Now, the Opal and the Onyx are co-dom inant alleles.
Som e exam ples will help to fully understand how this allelic relation works. Let L, be the nor-
m al dom inant, non-opal allele; then l is the recessive allele responsible for the Opal phenotype,
and l’ the recessive allele that yields the Onyx phenotype. According to all of the above, there
are three alleles in the Opal locus: the norm al dom inant alleleL, the recessive l responsible for
the Opal, and finally the recessive allele l’ responsible for the Onyx. Do not forget they are all
autosom ic, and l and l’ are co-dom inant.
56
Now the m ating of an Opal m ale and an Opal-Onyx fem ale will bring into being the following
offspring:
As seen in the Punnett squares, 50 % of the offspring will be Opal, and the other 50 % will be
Opal-Onyx, it is, an interm ediate phenotype between both, since the allele l and the allele l’ are
co-dom inant.
Now, the m ating of a m ale carrying the Onyx factor, with a norm al fem ale, will be:
Then again, the Punnett squares show that 50 % of the offspring will be norm al (not carrying
Onyx factor). The other 50 % will be phenotypically norm al carrying the Onyx factor. Now, from
the 50 % carrying the Onyx factor, select the best fem ales to be back-crossed to the original
m ale, then:
Twenty-five percent of the offspring will be hom ozygous Onyx (l’l’). Now, it is very clear you
would obtain Onyx phenotype canaries, in a term of two breeding seasons though you would
only have a m ale carrying the Onyx factor, if you follow the previous procedures.
The Topaz
According to reliable records, in 1985 a new m utation cam e into sight, again in Europe. This
tim e the genetic factor was called Topaz, in accordance to another precious gem . The m ain fea-
57
Octavio Perez-Beato
ture of the Topaz is an apparent m odification of the eum elanin, which is concentrated border-
ing the quill, and leaving a wide pearl color contour all around tail, wing and covert feathers.
The pheom elanin is also reduced, but beak, legs and nails exhibit a light brown color, depend-
ing on the variety. At present, only Green, Blue, Bronze, and Agate are standardized as Topaz.
The design in the first three types shows a brown color on the head, back, and flank streaks,
which are long and sym m etrical, neat and well delineated. Eyes are reddish when very young,
and darker when adult. The gene responsible for Topaz m utation is autosom ic recessive, as it
was the case in the Eum o and Onyx phenotypes, previously discussed.
The Topaz was the last new color in the eighties, but surprisingly it was not the last in the row.
Next, it will be discussed briefly, the newest color of all.
The Kobalt
Once again, another autosom ic recessive m utation on the m elanins cam e into view. It was
around the year 1994, in Germ any. In a tight period of thirteen years, four new expressions of
color were in the hands of the fanciers. As in the Onyx, the Kobalt shows an extension and dark-
ening process of m elanins. It is apparent that, those areas where lipochrom es are seen through
a light cover of m elanins (throat, chest and abdom en), as is the case of the classic Greens, Blues,
and Bronzes, in the Kobalt it is not a light cover of m elanin, it is a m ore dense dark patina that
com pletely changes the appearance of the bird, besides a well streaked throat, chest, and
abdom en are apparent. The beak, legs, and nails are very dark. In general, the streaks are cut,
som ehow fragm ented in the areas m entioned before, and on the flanks and head. In well select-
ed birds these short streaks look like an irregular dashed-design.
This m utation has been officially accepted during 20 0 6, and in m y opinion it m ay be im proved
in the years to com e. Now, at the sight of four new colors under the autosom ic recessive condi-
tion, all affecting the m elanin expression and design, a very sim ple question arises: Are these
four last m utations autosomic recessive just because, the possibilities of another sex link m uta-
tion affecting m elanins is extrem ely restricted nowadays, and nothing m ight be expected in that
direction? The answer is not a sim ple one. If we just carefully observe what the events have been
so far, it is possible to conclude, with a certain degree of uncertainty, that perhaps gene provi-
sion on the sex chrom osom es is not enough to enter a new m utational process affecting
m elanins, as happened before with Agate and Opal.
The Kobalt is in fact, a dark canary, but it is not Black. On the next chapter, a discussion on the
possibility of a true black canary is developed, up to the point that is possible at this tim e.
58
CHAPTER 14
Genetic Possibilities of a True Black Canary
At present, we already know about the so called “Onyx” color variation in the canary. Oddly
enough, the gem known by this name, is completely black, no doubt about it. In Mexico, is com-
mon to find rings and other jewelry with onyx on it. But our Onyx canary is not that black. In the
present chapter, the possibility of a true black canary will be discussed to the best of my
knowledge. Some historical facts have been recorded since decades ago, but the surfacing possi-
bilities in those occasions in the past, have not been successfully achieved in the stabilization of
a black canary, despite of the occurrences the phenotype appeared, recurrently, in former times.
Unfortunately, the story of the red canary has not been projected in obtaining the black canary.
Not enough breeding leaders devoted to pursue the goal of the black canary, has ever appeared on
the stage of genetic experimental efforts, with a real commitment, and decided to go to great
lengths, no matter how difficult the task might be. Recently, some attempts in hybridizing
Carduelis atrata (Negrito de Bolivia or Black Siskin), also classified as Spinus atratus with the
female canary, have been performed to some extent. Besides, on the same scene is also another
bird, the Yellow Bellied Siskin (Spinus xanthogaster). Crossing between the Black siskin and the
Yellow Bellied Siskin have created hybrids as well. Those hybrids are all black, except on the
throat, chest and belly which are completely white. These hybrids have been back-crossed to an
Onyx female canary. What seems certain is that fertility is not so good in those attempts, besides
the problem with white color on throat, chest and belly, which seem to be stubborn to disappear
from the phenotype. Nevertheless, keep on working is necessary. Hybridization is not an easy task,
many random genetic factors act on those crossings. The more attempts, the higher the possibili-
ty in obtaining the black canary. Sharing experiences among all fanciers involved in this goal, is a
must. We probably need emulous of Hans Duncker and Karl Reich teaming to accomplish on the
project of the Black Canary, as they did many years ago in the project of the Red Canary.
Of course, for many a simple and genuine question may arise: is it really possible a true black
canary? Up to present, colors on the plumage of the canary have been a mutational surprise, except
the Red Canary, which was, no doubt about it, the first genetically engineered animal, conceived,
performed, and brought into reality thank to the extraordinary effort of dedicated, well informed,
and scientifically stubborn ongoing fanciers, during several decades of extolling efforts.
59
Octavio Perez-Beato
I will try to answer the question from the preceding paragraph. First, I may say that two possible
ways may be taken into account, but we need to know first certain historical facts, and then draw
your own conclusion. Second, technically we assume the genetic possibility of a black canary,
based on the knowledge of melanin inheritance learned in previous chapters, not beyond.
What has happened in the history of color canaries in the past is exactly the basic to understand
what the future m ay bring, regarding new colors. The sam e is true for a real black canary. It
m ay show up from hybridization, or from a m utation. What is the m ost plausible one? At this
point, we need to know what the history is about black canaries in the past. In m any instances,
the facts in the past m ay be the answer to the present. Let us roll the historical facts, and then,
you m ay draw your very own conclusions. Perhaps, at least a dim light on the possible achieve-
m ent of a black canary m ay com e true.
According to certain records it appears that, at the tim e the fever of the Red Canary was at a
peak (1930 ), others focused on the possibility of a Black Canary. And then again, the Red and
the Black, one hundred years before (1830 ), it was the title of the outstanding novel written by
Stendhal; what a coincidence!
That was the way that between the twenties and the thirties, canary breeders from different
countries were im m ersed in hybridizing the canary and the Black Siskin. This species belongs
to the sam e fam ily of the canary (Fringillidae). Negrito de Bolivia or Black Siskin is of a deep
black color, with yellow belly and lower tail coverts, with two wing bars of the sam e color. Its
behavior as a cage bird, is not easy to handle, since it shows som etim es itself as a very nervous
bird, very sensible to aviary or cage conditions and m anagem ent. Finally, m ost of them do not
survive in captivity, according to certain fanciers.
From all of the hybridizing attem pts, nothing was clearly obtained. Nevertheless, the general
opinion is that the Black Siskin x fem ale canary’s hybrid, is fertile, but in a very low proportion.
Around 1930 , a canary breeder from Argentina succeeded in obtaining several fertile hybrids
from the aforem entioned crossing. It is said, not confirm ed, that in the first bird show in that
country, that canary breeder showed the first true black canary, together with several Black
Siskins x fem ale canary’s hybrids. After that bird show, no other records have been available.
In 1954, at the Turin Bird Show, Italy, a black canary was registered. Unfortunately, very few
facts were known about the bird only that it sang like a com m on Roller. The owner declared that
it was the product of hybridization.
In 1955, at the National Exhibition of Cage Birds and Aquaria, in England, the judges carefully
watched an individual registered as a black canary. They finally declared the bird as a true black
canary. When the owner of that outstanding bird was asked about the way he obtained such a
color, he openly declared that it was not his intention to obtain such a color, he was trying to
obtain green canaries from a couple of Borders, the m ale being dark green pied, and the fem ale
white with black eyes. That canary was com pletely black, except the tail and a sm all patch on
the neck, both being white color; besides, som e few brown wing feathers.
Also a m utation gave birth to a black canary; the bird was registered at the Chelm sford Bird
Show. The owner A. J . Spooner declared that he m ade nothing to obtain such a bird, which
cam e out of a brood from a green fem ale. As a fledging it was not so dark, but after the first m olt
it turned to be black.
60
Other attem pts regarding hybridization were perform ed using the Cuban Bullfinch (Melopyrrha
nigra nigra) a sm all black finch from Cuba, with a sm all white bar on the wings. I personally
watched a Cuban Bullfinch and fem ale canary,s hybrid. It was very dark, larger than a bullfinch
but sm aller than a canary, having a beak resem bling that of the Bullfinch. Around 1957, hybrids
were obtained from such m ating. One of those hybrids was registered at a Canary Show in
Havana. The bird had black head and chest, som e few black patches on the wings, and the rest
it was chocolate brown. It was larger than F1-Red siskin and canary, the tail was tip-squared,
and the beak wide at the base.
According to the available records, birds from the Fringillidae and Emberizidae fam ilies that
have been used in hybridization experim ents with canaries are: The J acarina Finch or Blue-
back grassquit (Volatinia jacarina); Negrito de Bolivia (Carduelis atrata), rem em ber, also clas-
sified as Spinus atratus; Alario finch (Serinus alario) belongs to the sam e genera as the canary;
Cuban Bullfinch (Melopyrrha nigra nigra).
As far as I can see, Spinus atratus and Melopyrrha nigra nigra are good candidates for
hybridization experim ents with fem ale canaries. Both could be the first genetic step up to obtain
hybrids carrying true genes to approach the so long expected black canary. The Yellow Bellied
Siskin (Spinus xanthogaster) is another good candidate.
Nowadays, there are many specialized fanciers around the world on Siskin species breeding; I do
not mean hybridization, just breeding to keep siskins in captivity. To mention some facts, the
leading countries in Siskin breeding are: Italy, Belgium, Germany, Spain, Netherlands, U.K., New
Zealand, Australia, and Brazil. Counting on this, it is expected that the availability of many Siskin
species may impulse the experiments to obtain the Black canary. I may say that most of the Siskin
species are easy to breed in captivity as is the Yellow Bellied Siskin mentioned before, one of the
good candidates to be involved in obtaining the Black canary. Those well informed breeders, and
well trained in Genetics, are the first runners in the hard racing to obtain the expected goal.
Be advised, this is not an easy task to successfully perform . It shall take tim e, probably a long
tim e, and a lot of patient work with no dism ay. Of course, this is not for a rookie. Only experi-
enced canary breeders m ay face such a challenge.
A Theoretical Approach to the Black Canary
I will exclusively use just what has been discussed in previous chapters. I will not go beyond that
fram e in genetic grounds, since the objective of this topic is precisely, to fit out the reader with
all previous knowledge to share.
About hybridizing fem ale canaries with other species, enough said. It is tim e to consider what a
m utation m ight be for the real outcom e of a black canary, based on the gene system s already
discussed in previous chapters.
The first approach or theory is to consider that a Black Canary m ust be the opposite of a
Brown Canary, just because the Brown Canary is unable to synthesized eum elanin, and a Black
Canary m ight be unable to synthesized pheom elanin. Then the first m utation step could be the
change of gene B to a gene b, which m eans no pheom elanin is genetically structured, never
61
Octavio Perez-Beato
recorded before in canary breeding history. It is a fairly good genetic approach, and then the
expected gene structure in sex chrom osom es shall be:
Male: X bNO Fem ale: X bNO

X bNO Y
It is clearly seen that eum elanin is com pletely oxidized with no presence of pheom elanin. But
this is not enough. It is necessary for the eum elanin to be extended all over the plum age, to real-
ly create a black feathered canary. We all know that a good Green canary shows wing and tail
feathers com pletely black; unfortunately, this spread pigm ent does not cover the rest of the
bird, just tail and wing feathers.
Now we need another m agic touch to com pletely fulfill the dream of a Black Canary. As I said
in the previous paragraphs, the black color (eum elanin) needs to be spread all over the plum age.
Let us assum e that the allele responsible to deposit eum elanin in canary feathers, as we know it
today, is the autosom ic dom inant allele D. Then, the m utant allele to cover the whole plum age
with eum elanin is d, being an autosom ic recessive allele. According to all of the above, two
m utations are necessary to produce a Black canary, which is not very feasible, since a single
m utation is a very sporadic event in genetic history; it is too m uch to consider two m utations in
a row to produce the black phenotype.
The second approach or theory is: if we refer to the Black Canary registered in England
(1955), we m ay recall that brown feathers were visible in that individual; this m ay indicate that
it is not com pletely necessary to inhibit the Brown allele to produce a Black canary, but a larg-
er am ount of eum elanin, instead. If it is correct, then only one single m utation is necessary,
allele d, and nothing else.
As I said before, let D be the regular gene responsible to deposit eum elanin in the regular canary
plum age, then d is the m utant recessive allele which is able to deposit enough eum elanin in all
feathers, to create a Black Canary. Then again, a possible genetic form ula for the black canary
m ight be:
X BNO
dd
X BNO
As the allele d is recessive, this m eans that a Black canary m ust be hom ozygous for that allele.
Besides this allele is not sex link, on the contrary, it is autosom ic.
According to historical facts already discussed som ewhere in this book, the em ergence of black
canaries seem s to be associated with a recurrent m utation, since it has been repeated over a cer-
tain period of tim e, as recurrent m utations do. Nonetheless, the outcom e of a Black Canary
needs the concom itant occurrence of m ale and fem ale carrying the recessive autosom ic allele d,
if this theory is correct. The expected outcom e of such m ating shall be 25% black canaries,
either m ales or fem ales, or a com bination of both.
Tim e and effort will solve the problem of the “waited so long bird”; tim e and effort go together
in m ost of m ankind goals, and the Black Canary cannot be different.
62
CHAPTER 15
Obtaining Varieties through Simple Pairing
Taking into account that new color canaries are som ehow difficult to obtain, since they are not
widely available, it is the purpose of this chapter to present to the reader som e sim ple pairing
procedures and schem es, that lead step by step in obtaining the desire varieties having only one
individual of the kind.
It is necessary for the reader to be in com plete knowledge of the preceding chapters, in order to
apply it and reach the goal in obtaining the varieties he/ she is looking for.
To fully develop the exam ples in the following pairings, genetic sym bols will be created for the
new varieties to m ake possible a true understanding of different genetic com binations, accord-
ing to what each variety represents. Besides, only basic procedures will be shown as a guideline,
which can be extended to other pairings as necessary.
Obtaining an Eumo Lineage
Let allele E, be the autosom ic norm al non-Eum o factor, and allele e the autosom ic recessive
allele responsible for the Eum o factor. Now, let us suppose that the fancier only possesses one
Eum o m ale or fem ale canary, as the Eum o is an autosom ic factor, it does not m atter if we start
with a m ale or a fem ale canary.
If the fancier has an Agate Eum o (m ale or fem ale) and the goal is to obtain an Agate Eum o lin-
eage, then sim ply by pairing this Agate Eum o with a com m on Agate (or Isabelle, which is a bet-
ter choice), the starting point of the lineage is achieved.
In the following, sim plified genetic form ulas will be used; for this
purpose, it will not be considered the ground lipochrom e.
63
Octavio Perez-Beato
Agate male carrying

the Eumo factor X Agate female
1 X BNo Ee 3 X BNo EE
2 X BNo 4Y
Offspring:
1 + 3 = X BNo EE or 1 + 3 = X BNo Ee
X BNo X BNo
Agate non-Eum o m ale Agate carrying Eum o m ale
1 + 4 = X BNo EE or 1 + 4 = X BNo Ee
Y Y
Agate non-Eum o fem ale Agate carrying Eum o fem ale
2 + 3 and 2 + 4 yield the sam e results as the previous ones.
For the next breeding season, the original Agate m ale carrying the Eum o factor will be paired
with the best fem ale(s) from the 1 + 4 or 2 + 4, groups. Then, what shall be obtained from such
backcross is as follow:
Original Agate male x Any female from groups

carrying the Eumo factor 1 + 4 or 2 + 4
1 X BNoEe 3 X BNo Ee
2 X BNo 4Y
Offspring:
1 + 3 = X BNo EE or 1 + 3 = X BNo Ee or 1 + 3 = X BNo ee

X BNo X BNo X BNo
Agate non-Eum o m ale Agate carrying Eum o m ale Agate Eum o m ale
The gam ete com bination 2 + 3 is identical to 1 + 3.
2 + 4 = X BNo EE or 2 + 4 = X BNo Ee or 2 + 4 = X BNo ee

Y Y Y
Agate non-Eum o fem ale Agate carrying Eum o fem ale Agate Eum o fem ale
It is evident that on the second breeding season, Agate Eum o m ales and fem ales are both
obtained. From now on, the fancier m ay establish a whole lineage of Agate Eum o canaries,
always using a line breeding procedure to ensure the purity of descendants and of course, selec-
tion ought to be applied, rigorously.
This exam ple is valid for any autosomic recessive inheritance as is the case of the Onyx
(already explained) and the Topaz birds. Next, an exam ple using an autosomic dominant
allele will be discussed. A good choice could be the m osaic canary, which is an autosom ic dom -
inant gene.
64
It is relevant to say that when you are looking at a m osaic canary, you cannot tell if it is hom ozy-
gous or heterozygous for this particular trait, as it is a dom inant allele. Then, let be allele h the
non-m osaic norm al allele, and allele H the one responsible for the m osaic trait.
Now, suppose you only have a m ale or fem ale m osaic canary, as this is not a sex link gene, no
m atter what the sex of your selected bird is. Let us suppose you have a m osaic fem ale, then:
Example 1:
The offspring is com posed exclusively of heterozygous birds (Hh), which is logically expected
from this kind of m ating. Then again, nobody can tell they are heterozygous, since their pheno-
types are typically m osaic.
Example 2:
The offspring is com posed 50 % heterozygous m osaic birds, which look like com m on m osaic
canaries, and 50 % norm al non-m osaic birds. These norm al non-m osaic birds in the offspring
show them selves as an irrefutable proof that the parental fem ale is a heterozygous bird, for the
m osaic factor.
Now, the last exam ple is devoted to a sex-link recessive factor to com plete a genetic trilogy,
which encom passes the three m ain categories of genes in the color genetics of canaries. Our
m odel factor in this exam ple is the Brown canary.
If you want to obtain a lineage of Brown canaries and only possess a m ale Green canary carry-
ing Brown factor, you m ay pair this canary to a Green fem ale, which are readily available alm ost
anywhere; the procedure is as follow:
Green male carrying Brown factor m ated to: Green female

1 X BNO 3 X BNO
2 X BnO 4Y
65
Octavio Perez-Beato
Offspring:
1 + 3 = X BNO 1 + 4 = X BNO
X BNO Y
Green m ale Green fem ale
2 + 3 = X BnO 2 + 4 = X BnO
X BNO Y
Green m ale carrying Brown factor Brown fem ale
For the next breeding season, a Brown fem ale is already available to be back-crossed to the orig-
inal Green m ale carrying Brown factor.
The results are as follow:
Original Green male carrying Brown factor m ated to: Brown female
1 X BNO 3 X BnO
2 X BnO 4Y
Offspring:
1 + 3 = X BNO 1 + 4 = X BNO
X BnO Y
Green carrying Brown factor Green fem ale
2 + 3 = X BnO 2 + 4 = X BnO
X BnO Y
Group 2 + 3 is com posed of Brown m ales, and group 2 + 4 of Brown fem ales. In a couple of
breeding seasons, Brown m ales and fem ales have been available. From now on, it is possible to
create a com plete lineage of Brown canaries.
Then again, autosom ic recessive, autosom ic dom inant, and sex link recessive alleles represent
the core of color genetics in canary breeding. The previous exam ples constitute a practical les-
son in obtaining varieties, when not all the parental stock is readily available.
About the Shape in Color Canaries
Though a good color could be obtain following right breeding techniques, one im portant
aspect cannot be overlooked: the quality of a nice profile and a harm onious shape, as an unde-
niable com plem ent to the attributes of color, both flow together in pursue of the excellence. It
is, to the perfection of color you ought to add the perfection of shape; nothing valuable is
obtained, otherwise.
66
When one talks about shape, the m eaning is straightforward: harm ony that m ust exist in all
parts of the body and plum age contour, in any good color canary. What follows is just a guide-
line to be considered, when breeding color canaries. These aspects m ake a big difference
between an ordinary bird, and a high quality one.
The first aspect to be considered is the size of the color canary. Som e fanciers think “the larger
the better”. It is not exactly the truth. Color canaries are qualified between 13.5 cm and 15 cm .
Beyond this size, m ight not be considered to m eet the standards of the color canary category.
The beak should be short, cone-shaped, well im planted and its m iddle line runs along theoret-
ically tangent to the lower edge of the eyes. Head m ight be rounded, the eyes being the gentle
center of it. The neck m ust follow a cylindrical contour, harm onious, linking together the head
and the body, following a soft curve line. The chest should be rather short, sm oothly rounded to
continue in a soft curve to conform the abdom en, which ends up in a well packed tail, to an M
ending-tip and lateral straight lines. Besides, the tail should be neither short nor long, just visu-
ally proportional to the length of the body. Dorsum m ust continue in a straight line with the tail.
Wings well tighten to the body, with upper edges slightly touching each other and running par-
allel on the back, without overlapping at the tip.
The longitudinal body axis m ust be 45 degrees regarding the horizontal line. Legs well settled,
slightly positioned beyond a right angle. No deform ities in fingers or nails. The num ber of tail
feathers m ust be 12, and wing feathers 18, as a m inim um .
67
CHAPTER 16
Concomitant Factors in the Color Canary: other mutations
The Ino Factor
The m ain characteristic of the Inos is red eyes, due to an autosom ic recessive allele. Individuals
with pheom elanin as the only m elanic pigm ent are called Phaeo-inos; different varieties are
known. Besides, for those non-m elanic canaries, if theplum age is white they are nam ed Albinos,
if the lipochrom e is yellow they are nam ed Lutino, and if the lipochrom e is red, Rubino. It is not
worthless to say that Inos are highly diluted birds; hence eye pigm ents are gone. There is a par-
ticular characteristic in Lutinos; it is a certain light greenish shade, usually all over the plum age.
The Satine Factor
If there were a really quite polemic canary, it was the Satine. At least, in the last fifty years in canary
breeding scenario no other mutation has been so controversial, and strong arguments have so risen
about. In addition, the nominative Satine arose to a great debate in its time. That is why lots of arti-
cles written by specialists and true authorities, have been devoted to Satine, all over the world.
This peculiar m utation em erged in Argentina, at the end of the 60 ’s. Based on the m utation
hom e country nam e and the fact of red eyes in these birds, the first appellative was Argent-ino.
What was really new in these canaries? Very sim ple, they have red eyes, with identical pheno-
type as the Europeans’ Inos, but Argent-inos, nowadays Satine, are sex link recessive, besides
showing a lustrous patina all over the plum age, which has been ground enough to nam e them
Satine, a word from French language (Satine: satin-like). Exactly, it was France the country
where they becam e Satine; appellative that they hold today.
The Satine is a canary that shows certain problems. One of them is the fact that the Satine factor com-
pletely dilute the eumelanin, almost does not touch the pheomelanin. As a consequence, it is some-
times really difficult to discern if we are looking at an Agate Satine or a Bronze Satine, taking into
account the phenotype, exclusively. Then for the rest, both birds may look as a common lipochrome
canary. Besides, looking at an Isabelle Satine and a Brown Satine, the decision is neither simple.
68
The Greywing
This variety, as long as I know, cam e to light from the Bronze Pastel. Nevertheless, Greywing
m elanins are diluted, except those on the wing feathers, which are tinted with an ostensible grey
color. In som e individuals tail feathers are grey tinted, as well.
The aforem entioned m utations appear just like desirable additions to the m utation catalogue
already in existence. It is evident that a m eticulous breeding, and a very accurate selection m ay
create valuable birds from a genetic point of view. Of course, the m ore factors in a single bird,
the m ore valuable that bird is. Nonetheless, the fancier has to be very careful and plan in
advance what the genetic objective is, in a short and a long term basis. It is not m erely to have
canaries with a lot of factors com posing their genotypes, it is com pletely necessary the harm o-
nious arrangem ents of those factors, to obtain birds that are phenotypically appealing.
Besides, the m ore com plex color m utations present, the m ore excellence m ight be foreseen
regarding the shape of the bird, otherwise the m utation-loaded canary showing a bad confor-
m ation, m ay seem som ehow like a patchy deform ed individual, since the color com binations
enhance any defect, on the body shape and plum age.
Som ewhere in this book, som e guidelines were written to offer a basic idea of what a propor-
tional and harm onious body shape m ight be, in our canary. Read those lines again, it is worth-
while to keep in m ind som e sim ple concepts about color and shape.
Another aspect I want to stress is the extrem ely strict selection when dealing with several m uta-
tions at a tim e, to produce really valuable birds. The slightest defect m ight becom e a real prob-
lem in breeding seasons to com e. Do not hesitate in culling, that is the rule.
69
CHAPTER 17
The Nomenclature of Color Canaries
Along the whole book, all aspects have been treated from a genetic point of view, naming in some
cases the specific color according to the accepted nomenclature. Now, in the present chapter the
classification of color canaries will be dealt with, stressing the principles in which are based all
the terms used in the so called nomenclature. It is of extreme importance, since the newcomer
breeders might themselves be lost in a jargon that seems quite difficult, if not well explained.
What is explained hereof is just a guideline for those who, for the first time, are entering the “has-
sle” of so many new terms used in color canaries nomenclature. To be a canary breeder means to
handle all aspects of this activity, and naming the birds in the right way is really important, since
it is the language to establish an unambiguous communication among canary breeders. It is of
supreme importance when taking part in a bird show, to deeply understand how and why the
birds are prized. Without knowing the nomenclature such an approach is almost impossible.
Nom enclature itself is not a hard issue; on the contrary, it clarifies every single detail and sub-
tle features, nowadays so com m on in color canary breeding. In every science or in any branch
of technology, it is necessary to establish a precise nom enclature that m akes com m unication
run sm oothly, to express concepts and technical aspects understandable worldwide. Canary
breeding is not an exception.
Internationally accepted, nom enclature of color canaries is a m ust, not only involving dom estic
bird shows, but all around the world these shows are feasible just because an international way
of nam ing is respected and accepted, based on technical phenotype characteristics, as well as on
organization ethics.
The Basis of Color Canaries Nomenclature
Three elem ents are the support of the m ainfram e of nom enclature, for this particular case of
color canaries. These elem ents are: Variety, Type, and Category.
70
This sim ple division allows locating and including all possible color com binations existing at
present, being flexible enough to include future m utations to com e. Besides, it is quite sim ple
for novices to understand and apply.
Variety
This partition includes all lipochrom ic m anifestation on the plum age, such as: Red, Yellow,
Dom inant White, Recessive White, Rose, and Yellow Ivory. It is precisely the Variety which
grade the quality of lipochrom e distributed on the plum age, or its absence, com plem enting the
Type from a nom inal basis. In lipochrom e canaries, nom enclature starts on Variety.
Type
Under this division all m elanin-related nouns are included, such as: Green, Brown, Bronze,
Agate, Isabelle, Isabelle Pastel, Green Opal, Agate Opal, and Brown Pastel, just to m ention few.
As it is apparent, Type could be single or double. Single Types are assigned to classic m elanin:
Green, Bronze, Brown, Agate,and Isabelle. Double Type are: Isabelle Pastel, Green Opal, Agate
Opal, etc. It is obvious that the Type cannot include any lipochrom e, since lipochrom es are
assigned to Variety. As a consequence, we cannot talk about Type in strictly lipochrom e
canaries. It does not exist in those birds.
Category
The category is the third and last com ponent in color canaries’ nom enclature structure. At pres-
ent, only three categories are recognized to describe the feather quality in the color canary; they
are: frost, intensive, and m osaic.
Perhaps, for som e canary breeders it m ay sound weird to include the m osaic condition into the
category, as a feather structure. The only possible explanation that we have at hand is that the
m osaic condition is the opposite of the intensive feather; you m ust rem em ber what was already
explained in a previous chapter about the opposite relation between intensive and m osaic. It is,
never m ate m osaics to intensives.
Knowing what the three components of the color canary nomenclature are, you are in a very good
shape to approach the classification of classic colors, as a means of exercising in a practical way
what has been explained up to this point. What follow are simple examples of classification.
Variety Type Category Variety Type Category

Red Brown m osaic - Bronze mosaic
Yellow - intensive Yellow Brown Opal intensive
White Isabelle frost White Isabelle Pastel m osaic
Red Agate intensive Yellow Brown Pastel intensive
- Green intensive White Agate Opal frost
Red - frost Rose Bronze Opal intensive
- Green Opal frost
It is apparent that som e individuals are lacking the Variety and others are lacking the Type, but
all bear the Category. The previous exam ples can be taken as a pattern to classify other colors
71
Octavio Perez-Beato
of our canary. If you take a closer look at the previous exam ples, you will find that under the
Type com ponent those nam es with two words description, the first nam e is always the oldest
m utation and the second, the newest one.
Next, som e Varieties and Types will be described for the reader, m ainly for the new com ers to
this activity, to get acquaintance of the peculiarities of the m ost com m on color canaries. What
follow is not a com prehensive list, it is just a guide, a grosso modo approach to learn som e few
details about the m ost com m only seen color canaries.
Melanin Types without Dilution
Blue Dominant Intensive

To talk about a “Blue” Type, the optic factor m ust be present; the canary will look dull grey, oth-
erwise. No pheom elanin traces on the plum age, but well defined and recognizable eum elanin on
the streaks of the back and flanks, and wing and tail feathers, as well. Exposed to natural light
an undeniable bluish tint is apparent on the chest, neck and head, as well as on the rum p. Beak,
legs and nails are fairly dark, to denote the presence of eum elanin.
Bronze
These canaries m ust reveal red lipochrom e evenly distributed all over the plum age. Black
streaks on the back and flanks m ust be evident; wing and tail feathers black, with very dark legs,
beak and nails. Any trace of pheom elanin in unacceptable.
Dilute Melanin
White Agate Intensive

In these canaries the presence of the optic factor adds a very delicate silvery shade, which
enhances the beauty of the bird. Streaks on the back and flanks show a lead shade, and are nice-
ly delineated on the whitish ground typical of this variety. The m inute design on the crown and
whisker, are readily seen, all of which is superim posed on a very light grey color. The coverts
show the Agate typical design, which is basically form ed as an almond-like contour on each of
these feathers. This drawing is precisely known as the Agate Almond Design. Beak, legs, and
nails are pink.
In those intensive high quality birds, the set of plum age, flanks and back resem ble a tiger-like
design.
Yellow Agate Intensive

In these birds, the yellow lipochrom e is the ground color for the dilute m elanin. All the streaks
on back, flanks and head m ust be neat and clean cut. Any trace of pheom elanin is a true dem er-
it, since only dilute eum elanin m ust be seen, which shows itself as a lead color design. In this
particular variety, the tiger-like appearance is exceedingly shown.
72
Yellow Isabelle Frost

The pheom elanin is diluted in these birds, and only a very light shade of brown is apparent on
the plum age. Dilute designs on back and flanks are openly seen. Wing and tail feathers show a
very pale brown tint, it is such that when individually observed, a sem i-transparent vane is all
you can see. The yellow ground is utterly seen, m ainly on the chest, belly, and rum p.
Super Dilute Melanin
White Brown Pastel

The m ost appealing birds in this variety are those bearing the optic factor in hom ozygous con-
dition, which confer a true silvery shade to the carrying bird.
Streaks on the flanks and back are still visible in this variety. The overall effect of the Pastel fac-
tor with a ground white color turns the plum age toward a lead-look appearance.
White Isabelle Pastel

The Pastel factor creates a m ore attractive visual effect, acting on the dilute brown of the
Isabelle. A soft Pastel tinge blended with the optic factor turns out to be a hue of silvery appear-
ance, m uch genuine than the lead-resem blance appearance of the White Brown Pastel.
Extra-Dilute Melanin
Green Opal
The Green canary, with the genetic addition of the Opal factor, still keeps the beak, legs and
nails well pigm ented. The sam e is true for the Bronze and Blue varieties. Pheom elanin is alm ost
gone, and eum elanin is severely reduced. Iridescent reflections are seen on the plum age, and
the general appearance is that of a lipochrom e individual.
Bronze Opal
On these birds, the typical iridescent reflection of the Opal canary is highly enriched, due to the
red ground color. Regarding m elanin dilution, the effect and appearance are the sam e as
described for the Green Opal.
The description of the aforem entioned birds is just an approach, for the reader to have an idea
of the phenotype details that have to be considered, when breeding these com m on varieties. It
is not recom m ended to start breeding any variety without knowing exactly what details and fea-
tures should be considered in those birds. Objectives have to be very clear from the start; unde-
sirable characteristic m ay show up, otherwise.
In m any countries, the nom enclature does not consider the presence of the optic factor. This
m eans, for them no m atter if the white or yellow ground is com bined with the optic factor, or
not. In fact, the optic factor has been very controversial in the classification of color canaries;
73
Octavio Perez-Beato
nonetheless, as it was expressed in a previous chapter, the optic factor is a genetic reality inde-
pendent of any classificatory criteria, or any checklist convenience.
Nowadays, the am ount of m utations already seen in our feathery friend, m ake possible an
incredible num ber of color com binations in Type and Variety that show them selves as really
com plex. At the sam e tim e, all these com binations confer an outstanding genetic value to those
birds bearing them . It is possible to bring into being birds that com bine in their genotypes four,
perhaps five of all available m utations known today, together with factors inherited from the
Red Siskin, such as the red color, and the m osaic factor. J ust to m ention a few of these possible
genotypes, as exam ples of real possibilities, let us take a look at this: Rose Isabelle Opal Mosaic,
Rose Brown Pastel Mosaic, Rose Bronze Opal Frost, Yellow Agate Ivory Frost, and Yellow Ivory
Satine Intensive.
It is evident that the aforem entioned canaries are beautiful m asterpieces of genetics, obtained
through a very careful and technical breeding, not com m only seen in bird shows; they are rather
scarce. The m ore genetic factors put together in just one individual, the m ore accurate breeding
and selection procedures have to be accom plished, sim ply because the com bination of m ultiple
factors m ight create a conflict in the phenotype expression for each individual factor, and for a
whole harm onious phenotype, as well. Today, canary breeding is not just an attractive hobby; it
turned out to be a genuine branch of zoo technology, with its own m ethods and principles, due
to the particular aspects of canary genetics, being com parable to any other dom estic anim al
breeding, at a world wide level. I would like to encourage all fanciers involved in color canary
breeding to m ake a try in obtaining those types and varieties seldom seen around, just because
those birds are beautiful. It is not a secret that m any fanciers do not attem pt breeding the com -
plex genotypes, due to lack of enough knowledge, m ainly on genetics ground. I really hope that
every topic included in all previous chapters, could be a real encouragem ent to every color
canary breeder, up in the way to better birds at each breeding season.
74
CHAPTER 18
Quantitative Canary Breeding
As it was stated in the previous chapter, canary breeding should be considered at present, a true
specialty in the dom estic anim als breeding activity. Nonetheless, though outstanding advance-
m ents have been achieved in practical m anagem ent procedures, and reproductive aspects of
canary breeding, just few…probably very few fanciers all over the world, have been involved in
a num erical evaluation of their breeding results. Honestly, this is the im pression I have after
checking published texts, and lots of references and articles in the internet. Most of that infor-
m ation, not to say all, is based on individual practical experiences, which m ay differ from one
breeder to another. Then a question arises: What are the quantitative results of all these prac-
tical advises? To answer this question is dedicated all of the following.
The Quantitative Idea
Quantify is nothing but put results as num erical values; otherwise, only qualitatively shown and
of course, incom plete. The m ost sim ple and outstanding advantages of quantifying are:
Concentrate in just few figures the work results of several years
Interpretation sim plicity
Precise inform ation without am biguities, at a single glance.
To take sm art decisions according to quantified results obtained.
Nevertheless, som e breeders m ay state that, if quantifying is taken as a procedure, Mathem atics
is involved in som e way. The answer is, yes. But only very basic Math concepts are going to be
used, and the results are worthwhile based on so sim ple Math procedures. I do agree that the
m ethod of quantifying, no m atter what the application area m ight be, is not for everybody. A
basic Math knowledge is necessary. For those that are willing to use those basic m athem atical
skills, this chapter has been written.
75
Octavio Perez-Beato
Quantitative Information
The so called quantitative inform ation is usually envisioned in two basic form s:
1.- Relative figures (usually percents)
2.- Absolute figures
Relative figures are a very popular m anner to offer results, being com prehensible for everybody,
since in m odern tim es it is a com m on issue that TV program s offer a lot of inform ation and
results of advertised products, based on percentages.
Let us not forget that a percent is telling us what proportion has been considered, from a base total.
On the other hand, absolute figures offer concrete information, such as: chicks hatched, number
of mating pairs, etc. or a combination of such figures, which are also a target for this chapter.
Premises for Quantitative Results
In order to quantify results, basic inform ation is needed. This inform ation is the raw m aterial
of the analysis, and ought to be collected in the m ost reliable way. It is supposed that the breed-
er keeps all this inform ation in his/ her breeding log. Such inform ation is as follow:
Total fem ales used in breeding
Total nests
Total eggs laid by each fem ale, with dates.
Total chicks hatched
Total chicks full-fledged
Laying range
All of the above could be registered in sim ple cards designed for such a purpose, or could be
com puterized if an ad hoc program is available. No m atter what the registration procedures
m ight be, what is really im portant is to register the necessary inform ation.
It is unacceptable an aviary without registering the necessary inform ation. Besides, this is part
of the delightful hobby of canary breeding; no records, no control. I personally have known
breeders that leg-band every single bird, no doubt about it, but they are unable to keep proper
records other than leg-bands.
Now, it is necessary to state certain definitions regarding the breeding log m entioned before, so
no am biguities m ay arise. At the sam e tim e, it is necessary to provide the reader with a stan-
dardized m ethod, in order for everyone to apply procedures in the sam e way. No drifting inter-
pretations could be allowed for a proper understanding of m ethods, and then results.
Definitions
Total females used in breeding: it is the num ber of fem ale canaries that took part in the breed-
ing season, constructing their nests, laying eggs (from 3 to 5 eggs per nest; as an exception 6
76
eggs), and then, incubation. If from those incubated eggs hatching was obtained or not, that is
another issue. The point is that the fem ale canary took part in the breeding season.
Total nests: it is the total am ount of nests where eggs were laid, and then hatched.
Total eggs laid by each female: it is the total num ber of eggs laid by each single
fem ale canary during that breeding season. It is m andatory that egg-laying infor-
m ation includes at least, the date of the first and last egg for each nesting.
Total chicks hatched: it is considered “chick hatched” each single chick that gets
off the egg-shell, com pletely.
Total chicks full-fledged: it m ust be considered as a prim ary concept that, every
single fledging set apart from its parent to strive for itself is a full-fledged chick.
Laying range: it is the tim e interval (days) from the first and last egg laid in that
breeding season. No m atter what hen laid the first or the last.
Once all the elem ents that will be used as raw m aterial in the analysis of the results have been
thoroughly stated, it is tim e to go directly to the basic form ulas for quantitative purposes.
CHICKS PER NEST (Cn):
Cn = total chicks hatched / total nests
This form ula tells about the average chicks per nests in each breeding season. It is an indicative
m easure of hatching potential for each breeding pair per nest. Theoretically, the value of this
form ula (Cn) m ust be from 3 to 5; being 3 the m inim um acceptable value, and 5 the optim um .
Any value near or larger than 4, is considered good. Then again, 5 is optim um .
CHICKS PER HEN (Ch);
Ch = total chicks hatched / total fem ales used in breeding
Ch is an average fertility m easure for each hen that took part in the breeding season. What fol-
lows is a sim ple table as a reference for the breeder to com pare his/ her results with the theo-
retical expected values. The table com prises from 1 to 4 broods; this last figure is not recom -
m ended, since it is detrim ental to the health condition of breeding pairs. Results are divided in
three categories: poor, good, and excellent.
77
Octavio Perez-Beato
Ch Values Considering from One to Four Broods in a Single Breeding Season
Interpreting the Values from the Table
The table offers typical values as a reference. It is likely that the Ch value obtained by any fanci-
er is not exactly referred on the table. Let us put an exam ple to illustrate the practical issue: a
hypothetical canary breeder used four hens in the breeding season, and each run three
broods for a total of 47 chicks, then Ch = 47/ 4; then Ch = 11.75, but this value is not in the
table. Then, for the colum n indicating 3 broods, the Table value for good results is 12. The fig-
ure the canary breeder obtained is very close to 12, since his/ her result is 11.75; it is interpreted
as good results for 3 broods per hen.
ASSESSED POTENTIAL (Pa):
There are three possible form ulas for the Assessed Potential, depending on the goal stated by
the canary breeder: it is, 3 chicks per nest, 4 chicks per nest, or 5 chicks per nest. Next, the three
form ulas are fully explained.
For three chicks per nest the Assessed Potential is calculated by:
Pa = [ 33.3 x Cn] – 10 0
For four chicks per nest the Assessed Potential is calculated by:
Pa = [25 x Cn] – 10 0
For five chicks per nest the Assessed Potential is calculated by:
Pa = [20 x Cn] – 10 0
Now, the interpretation and application of this form ula will be discussed in detail. First of all,
this form ula result is a percentage value. Then, the Assessed Potential is expressed as a percent,
and accurately m easures the goal fulfillm ent of the breeder in obtaining 3, 4, or 5 chicks per
nest. That is why the value of Cn is a crucial elem ent in the form ula to determ ine the value of
Pa. There are three constant values 33.3, 25 and 20 for each alternative in the form ula.
78
Whichever the constant is, its value is m ultiply tim es the Cn value, and then 10 0 is subtracted
from the result.
When any canary breeder decides that his/ her goal will be three chicks per nest, in fact is fixing
the reference constant that is im posed to his/ her aviary for that particular breeding season,
regarding the calculation of Pa. The sam e is true for 4 or 5 chicks per nest. That is why the exis-
tence of the three alternatives in the form ula.
The interpretation for values of Pa is quite sim ple. If the value obtained after calculating Pa
equal zero (Pa = 0 ), this m eans that the objective has been achieved 10 0 %. If the result is larg-
er than zero, then indicates that the objective has been over-achieved in that sam e value. On the
other hand, if the value of Pa is sm aller than zero (negative sign -), this indicates that the objec-
tive has not been achieved, and the negative value is the m easure below the 10 0 % fulfillm ent.
It is m y advice to use the alternate form ula: Pa = [25 x Cn] – 10 0 , since the goal of 4 chicks per
nest is feasible and rational, as an expected result, fram ed into acceptable biological lim its.
Anyway, each breeder is com pletely free to decide what alternate form ula is better according to
his/ her projected goals. That is the reason why the three alternatives are offered.
For those interested to go deeply into the developm ent and origin of this form ula, what follows
m ight be interesting:
Pa = 10 0 [Cn / 4 (– 1) ] = [ 10 0 Cn / 4] – 10 0 = 25 Cn – 10 0
Being Cn chicks per nest.
I developed the basis of the form ula considering 4 chicks per nest (constant = 4), which could
be substituted by 3 or 5, accordingly.
SURVIVORSHIP (S):
S = [total chicks full-fledge / total chicks hatched] x 10 0
The output value from this form ula is a percent. The inform ation obtained is understood
straightforward: how m any full-fledged chicks were obtained from the total chicks hatched. In
other words, it is the percent that represent full-fledged chicks of the total hatched. This sim ple
form ula directly points to certain possible problem s in the aviary. A low S value m ay indicate
som e problem s in every day routine handling, such as:
inadequate feeding procedures

health problem s in the breeding stock
inefficient hygiene
parasitism
genetic problem s (possible m ishandled inbreeding)
Next, rational value intervals for S are offered, intended for the breeder to have an idea of the
way results m ight be interpreted.
79
Octavio Perez-Beato
It is convenient to say that one of the m ajor problem s that could negatively affect Survivorship
(S), are m isguided inbreeding procedures. Inbreeding itself is not a wrong m ethod; on the con-
trary, breeds of any im aginable anim al species have been obtained by rational and well direct-
ed inbreeding procedures. What is unsafe is to apply it m istakenly. Chick m ortality during the
first days after hatching could be the result of im properly m anaged inbreeding procedures.
FERTILITY (F):
F = [chicks hatched / total eggs laid] x 10 0
F value is also a percent, and tells us what proportion of all the eggs laid in the breeding season,
were bearing chicks able to hatch. The value of F gives inform ation about the breeding stock,
m ales and fem ales as well. If a relatively low num ber of chicks hatch, com pare with the am ount
of eggs laid, this m eans that som e kind of fertility problem is arising in our breeding stock. But
not only fertility is involved, am ong other problem s that decrease the hatching level, the fol-
lowing could be m entioned:
fem ale canaries not properly incubating

sudden tem perature changes
certain toxicity levels in the food supply
m ale canaries disturbing and bothering the fem ales, not allowing them to be
properly devoted to incubation
After discussing all of the above, it is apparent that the F value is just and indicator connected
to the incubation and hatching issues, that set an alert flag about possible problem s that really
exist in the aviary.
80
DEVELOPMENT POTENTIAL (Pd):
Pd = [ S x F ] / 10 0
The form ula that expresses the Developm ent Potential (Pd) has two m ain elem ents already
explained: Survivorship (S) and Fertility (F). The product of both factors divided by 10 0 turns
out the value of Pd. This value in term s of a percent, express the relation between chicks full-
fledged and total eggs laid in the breeding season. Consequently, the Developm ent Potential
express the perform ance of the laying potential, that in turn depends on how m any full-fledge
chicks are obtained from the total eggs laid.
The Developm ent Potential (Pd), globally expresses how efficient is the m anagem ent in the
whole aviary, as well as it includes possible problem s in chicks developm ent, from hatching to
full-fledged tim e.
Next, a reference table is offered with Pd values.
As it was stated before, the Developm ent Potential is a global indicator, offering a general view
of how efficiency shows up in the aviary, as a whole. A clear evidence of this statem ent is that
Pd includes the Fertility (F) and the Survivorship (S), as structural elem ents of the form ula.
REPRODUCTIVE FREQUENCY (Rf):
Rf = laying range/ total chicks hatched
This form ula tells us how frequently chicks are hatching in the aviary. This figure point to the
progress and production level of full-fledged chicks. An exam ple will be explained to better
understand the inform ation that could be obtained using this form ula.
81
Octavio Perez-Beato
Let us suppose that there are three canary breeders A, B and C. The results they obtained dur-
ing the breeding season are as follow:
The interpretation for the Rf values as shown in the table above, are as follow:
For canary breeder A, the value of 5.2 m eans that on the average, every 5 days a
chick will hatch, taking into account that for his/ her aviary the breeding season
com prised 78 days, m easured from the first to the last egg laid.
For canary breeder B, every third day a chick hatched, for a laying range of 95
days. Then, for canary breeder C, on the average, every second day a chick
hatched. It is obvious that the best Reproductive Frequency perform ance was
achieved by canary breeder C.
It is not an uncom m on issue that, for highly developed aviaries the hatching frequency would
be just a few hours, on the average. This m eans that the values for Rf are in the order of deci-
m al figures, just like Rf = 0 .83, and less.
USING THE INDICATORS IN TIME SERIES
Indicators offer inform ation for a particular breeding season, as was seen before, and the cum u-
lative record of all those indicators from one year to the next, will enable us to exactly know if
our aviary is progressing, or not. Next, a com parative cum ulative table is offered for the breed-
er to post the inform ation collected every year, and then a chronological log could be kept,
which is the basis to establish a com parison of his/ her results in a tim e series basis. When at
least five years have been recorded uninterruptedly, then the fancier would be ready to carry on
another kind of analysis, which will be explained in full details after a brief explanation about
the com parative table tim e series-oriented.
82
Comparative Table for Annual Indicators
The year where our records start is called base year. This base year will be the tim e reference
to com pare all years to com e, from which a very im portant inform ation will be obtained about
the progress or retreat, for every single indicator on the table. This inform ation is obtained
through the so called index numbers, which are nothing but quotients obtained in a successive
fashion using as the unique divisor the base year, m entioned before. Then, the GENERAL FOR-
MULA to obtain index num bers is as follow:
Index = [target year / base year] x 10 0
As an exam ple, the index num bers for Developm ent Potential (Pd), will be expressed:
Ipd = [target year / base year] x 10 0
Based on this exam ple, any convenient notation could be used for index num bers applied to any
specific indicator. All index num bers are expressed as percent values. What follows is a good
exam ple using Ipd (Index for Developm ent Potential) in a hypothetical tim e series:
Years Pd
20 0 3 94.3
20 0 4 94.8
20 0 5 96.4
20 0 6 98.3
20 0 7 92.5
The year 20 0 3 is considered the base year, according to what was discussed in a previous para-
graph. Then, applying the form ula for Ipd, the results are as follow:
Ipd = [94.8 / 94.3] x 10 0 = 10 0 .5 for the year 20 0 4. In a sim ilar way Ipd values are calculated
for the rest of the years, always using 20 0 3 as the base year, since it was the first recorded year
for the analysis. Next, are shown the Ipd values according to the applied form ula for each year
on the previous table:
Year Ipd Year Ipd

20 0 4 10 0 .5 20 0 6 10 4.2
20 0 5 10 2.2 20 0 7 98.1
83
Octavio Perez-Beato
It is evident that years 20 0 4, 20 0 5 and 20 0 6 were better than the base year (20 0 3), since the
Ipd values are larger than 10 0 %. On the other hand, the year 20 0 7 shows a decline for Ipd value.
Som ething happened that year in the aviary, since the trend was increasing in the previous
years, this m eant progress. In cases like this, the breeder m ust carefully analyze what really hap-
pened, in order to take sm art actions to benefit from .
All of what com prises this chapter, put num erical tools of trem endous practical value in the
breeder’s hands, as guide lines in the m anagem ent of the breeding activity, year by year and on
a tim e series basis, as well. The success of this special task depends on the enthusiasm and
endeavor of the breeder, regarding the num erical aspects in m anaging his/ her canaries. Those
who feel them selves prone to enjoy the sim ple m ath aspects in applying these controls,
undoubtly will adjoin another appealing aspect to canary breeding wonderful hobby; besides,
the helping addition of a better control on the breeding doings, and technical tools that are well
effective. From now on, not only will you enjoy CANARY BREEDING, you will expand your
technical knowledge doing QUANTITATIVE CANARY BREEDING.
84
BIBLIOGRAPHY
Babra, A. : El Canario Pastel. El Canario Mosaico. Rev. El Canario. No77: 10 -11. Argentina,
1966.
_ _ _ _ : La Tecnica Actual de Coloracion. Rev. El Canario. No. 89: 12- 13. Argentina. 1971.
_ _ _ _ : Disparidad de Pareceres en el Canario Satine. Rev. El Canario.

No. 94: 6-8. Argentina, 1974.
Birkhead, T.: A Brand New Bird. How two am ateur scientists created the first genetically engi-
neered anim al. First Edition. Basic Books. USA, 20 0 3.
Chiny, J .: La Form acion del Color en los Canarios. Translated. Ornithophilie. No. 4: 5-7. France,
1959.
Duncker, H.: Einige Beobachtungen uber die Vererbung des weissen

Farbe bei Kanarienvogeln. Z. ind. Abst. Vererb. No. 32: 363-376. Germ any, 1924.
Durram , F. M. and Marryat, D. C. C.: Note on the Inheritance of Sex in Canaries. Rep. Evol.
Com m . Roy. Soc. No. 4: 57-60 . England, 190 8.
Eyckens, J .: Canarini di Colore. Edizioni Zootechniche, Italia, 1963.
Gill, A. K. : Cinnam on Inheritance in Canaries. First Edition. Ed. Dorset House, London. w/ y.
_ _ _ _ _ : La Cria del Canario Rojo. Translated. Rev. El Canario. No.83: 3-4. Argentina, 1968.
Gill, F. B.: Ornithology. Third Edition. W. H. Freem an. USA, 20 0 6.
Gottlieb, F. J .: Genetica del Desarrollo. Edit. Alam bra S. A. Madrid, 1968.
Hutt, F. B. Genetica Avicola. Salvat Editores, S. A. Barcelona, 1958
85
Hutt, F. B., and Rasm usen, B. A.: Anim al Genetics. J ohn Wiley & Sons. USA, 1982.
Lam b, Bernard C. : Applied Genetics of Hum ans, Anim als, Plants and Fungi. Im perial College
Press. England, 20 0 7.
Materi, C. M. M.: El Canario Opalo. Rev. El Canario. No. 83: 6-7. Argentina, 1968.
Mc Carthy, E. M.: Handbook of Avian Hybrids of the World. Oxford University Press. USA,
20 0 6.
Perazzo, B.: Canarios Blancos (dom inantes y recesivos). Pigm entos y Form ulas Geneticas. Rev.
El Canario. No. 78: 24-25. Argentina, 1967.
Skertchly, A.: Hibridacion. Rev. El Canario. No. 82: 27. Argentina, 1968.
Speicher, K.: Farbiger Rassenatlas Kanarienvogel. Germ any, 1971.
Vaccari, G.: Canarini di Colore. Edizioni ENCIA. Italia, 1971.
86

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FUNDAMENTALS

OF COLOR GENETICS IN CANARIES

PITTSBURGH, PENNSYLVANIA 15222

All Rights Reserved

Printed in the United States of America

For information or to order additional books, please write:

Or visit our website and online catalogue at www.rosedogbookstore.com

Acknowledgem ents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii

Basic Concepts on Genetics

Fig. 1 Basic parts of an anim al cell.

Fig. 2 Two cell divisions in a regular m itosis process.

Fig. 3 Mitosis process in an anim al cell.

Terminology and Definition of Some Basic Genetic Concepts

Dominance, Expressivity, and Penetrance of a Trait. Phenotype and Genotype.

Then we m ay say that b is recessive regarding B. Consequently, all individuals bearing BB or Bb

Sex Linked Inheritance and Sex Determination

Proportion of Genotypes Expected in a Particular Mating

Fig. 9 Punnett squares showing eight offspring groups

Line-breeding: mating in the

Lineage: group of individu-

Breed: a specific group of

Som etim es, it is difficult to

Fig.10 Inbreeding schem e to obtain

Fig. 11 Inbreeding system where

Fig.12 Both original fem ales are

Pigments and Feathers in the Canary

The Frost Feather and the Intensive Feather in the Canary

The Dominant White and the Recessive White Canary

The Recessive White Canary (English-White)

Fig. 15 The Punnett reticulum shows the genotype proportion as a result

The Yellow and the Lemon-Yellow Canary

Fig. 17 Offspring proportion from Ss and ss genotypes.

The Red Factor Canary

Phenotype = Genotype + Environm ent

Records of Fertility Level in Hybridizing Red Siskin and Female Canaries.

The Red Mosaic Canary

Genetic Aspects of the Mosaic Trait

Yellow frost m ale x red m osaic fem ale

The F1 offspring is:

KK GG Rr ff ii hh (a) and KK GG Rr ff ii Hh (b)

Yellow frost m ale x F1(b) m osaic fem ale

The R1 offspring will com prise the following:

The F1 offspring is:

KK GG Rr ff ii Hh (m osaic m ales and fem ales)

Some Important Remarks on the Red Siskin

The Green and the Brown Canary

The Brown Canary

Fig. 20 Brown m ale and brown fem ale m ating,

Lipochrome Green Brown

Yellow Green (com m on) Yellow Brown

The Dilution Factor: the Agate and the Isabelle Canary

The Agate Canary

As a result, the genetic form ula for the Agate is:

Agate male Agate female

The Isabelle Canary

I th in k it is in terestin g to explain to th e reader, h ow is th e m ech an ism by wh ich , th e gen e

The Agate Phenotype

Fig 25 Agate head. Melanin and lipochrom e areas are shown.

The Pastel Canary: the super dilution factor

Brown Pastel male Brown Pastel female

Yellow Brown Pastel intensive Yellow Brown Pastel frost

White Brown Pastel intensive White Brown Pastel frost

Yellow Isabelle Pastel intensive Yellow Isabelle Pastel frost

White Isabelle Pastel intensive White Isabelle Pastel frost